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2015 Bioarchaeology PDF
2015 Bioarchaeology PDF
Now including numerous full color figures, this updated and revised edition of
Larsen’s classic text provides a comprehensive overview of the fundamentals of
bioarchaeology. Reflecting the enormous advances made in the field over the past
20 years, the author examines how this discipline has matured and evolved in
fundamental ways.
Jargon free and richly illustrated, the text is accompanied by copious case
studies and references to underscore the central role that human remains play in
the interpretation of life events and conditions of past and modern cultures, from
the origins and spread of infectious disease to the consequences of decisions made
by humans with regard to the kinds of foods produced, and their nutritional,
health, and behavioral outcomes. With local, regional, and global perspectives,
this up-to-date text provides a solid foundation for all those working in the field.
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Cambridge Studies in Biological and Evolutionary Anthropology
Consulting editors
C. G. Nicholas Mascie-Taylor, University of Cambridge
Robert A. Foley, University of Cambridge
Series editors
Agustín Fuentes, University of Notre Dame
Sir Peter Gluckman, The Liggins Institute, The University of Auckland
Nina G. Jablonski, Pennsylvania State University
Clark Spencer Larsen, The Ohio State University
Michael P. Muehlenbein, Indiana University, Bloomington
Dennis H. O’Rourke, The University of Utah
Karen B. Strier, University of Wisconsin
David P. Watts, Yale University
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67. Bioarchaeological and Forensic Perspectives on Violence: How Violent Death is
Interpreted from Skeletal Remains Debra L. Martin & Cheryl P. Anderson (editors)
978 1 107 04544 6
68. The Foragers of Point Hope: The Biology and Archaeology of Humans on the Edge of
the Alaskan Arctic Charles E. Hilton, Benjamin M. Auerbach, & Libby W. Cowgill
978 1 107 02250 8
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Cambridge Books Online © Cambridge University Press, 2015
Bioarchaeology
Interpreting Behavior from
the Human Skeleton
SECOND EDITION
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Cambridge Books Online © Cambridge University Press, 2015
University Printing House, Cambridge CB2 8BS, United Kingdom
www.cambridge.org
Information on this title: www.cambridge.org/9780521838696
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Cambridge Books Online © Cambridge University Press, 2015
For Chris and Spencer and
In memory of George J. Armelagos
(1936–2014), visionary scientist,
bioarchaeologist, friend, and mentor
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Cambridge Books Online © Cambridge University Press, 2015
CONTENTS
1 Introduction 1
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x Contents
8 Isotopic and elemental signatures of diet, nutrition, and life history 301
8.1 Introduction 301
8.2 Isotopic analysis 302
8.3 Elemental analysis 347
8.4 Methodological issues in bioarchaeological chemistry 355
8.5 Summary and conclusions 356
References 433
Index 593
Color plates are to be found between pp. 320 and 321
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PREFACE TO THE SECOND EDITION
It has been more than 15 years since the publication of the first edition of
Bioarchaeology: Interpreting Behavior from the Human Skeleton. The response
following its publication in 1997 was overwhelmingly positive – in reviews and
comments to me from virtually every corner of the globe. I credit Robert Benfer for
convincing me that a synthesis paper I wrote for Michael Schiffer’s book series,
Advances in Archaeological Method and Theory (Larsen, 1987), should be
expanded into a book-length treatment of the field. He made the case to me that
such a book would serve to define what bioarchaeologists do and give bioarch-
aeology a sense of identity and mission.
Since the publication of the first edition, I have been thrilled to see how the field
has matured and evolved, the increasing scientific rigor, the extraordinary volume
of work published, the high quality of the literature, the appeal that it has had for
new and upcoming generations of bioarchaeologists, the development of new
directions and advances, and the impressive increase in international and multi-
disciplinary collaborative research programs. With regard to new directions, we
have seen expansion in areas relating to links between the social and biological,
what some call “social bioarchaeology” (Agarwal & Glencross, 2011; Gowland &
Knüsel, 2006), and facets of it relating to identity, gender, and social and cultural
forces that leave their impression on the skeletal body (Knudson & Stojanowski,
2008, 2009; Larsen & Walker, 2010; Sofaer, 2006). In addition, there have been at
least two books published with Bioarchaeology as the primary title, one providing
a historical overview with reference to the United States (Buikstra & Beck, 2006)
and the other focusing on practice (Martin et al., 2013).
The advances in methods for the study of ancient skeletal and dental tissues
have expanded our understanding of past population health and lifestyle in ways
unfathomable or just on the horizon when the previous edition of the book was
published. As shown throughout the present volume, applications of the study of
ancient DNA to mobility and residence, disease diagnosis, and biology generally
are breathtaking (Kaestle, 2010). The advances made in genome-wide and sequen-
cing technology have given access to remarkable amounts of data, providing new
insights and perspectives on the human experience in the past. Similarly, imaging
technology has developed at a remarkable pace (Chhem & Brothwell, 2007;
Schultz, 2001). These advances have played a central role in the increasingly
interdisciplinary orientation of bioarchaeology (Armelagos, 2003; Zuckerman &
Armelagos, 2011). Fundamental to the development of bioarchaeology is its
comparative approach and its grounding in the scientific method and its approach
to discovery and problem solving. These strengths provide perspective on present
conditions, such as the human–environment interaction, evolution and adapta-
tion, and success and failure, and understanding of who we are today.
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xii Preface to the Second Edition
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Preface to the Second Edition xiii
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xiv Preface to the Second Edition
Columbus, Ohio
May 1, 2014
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PREFACE TO THE FIRST EDITION
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xvi Preface to the First Edition
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1 Introduction
Many thousands of archaeological human skeletons are currently housed in various insti-
tutional repositories throughout the world. Some of these collections are extensive: The
Natural History Museum in London holds some 10 000 cataloged individuals, the Smithsonian
Institution has at least 30 000 skeletons, and the Bavarian State Collection more than
50 000 sets of human remains (Loring & Prokopec, 1994; McGlynn, personal communication;
Molleson, 2003). These and many other major collections around the world started during the
nineteenth century, mostly for purposes of collecting crania and other remains for investi-
gations of racial classification (Larsen & Williams, 2012; Little & Sussman, 2010) or as
“curiosities” without any manner of context (Taylor, 2014). While these motives for collection
have largely disappeared, these collections in Europe, North America, and elsewhere still
provide today an essential foundation for the study of past human variation and evolution.
The importance of the collections lies in the fund of biological information they offer for
interpreting lifeways of past peoples and for developing an informed understanding of the
history of the human condition generally.
One could argue that earlier generations of anthropologists may not have appreciated the
enormous value of human remains for interpreting the past, especially in view of the high
volume of excavated remains versus the remarkably low frequency of analysis and publica-
tion on those studied by biological anthropologists (Steele & Olive, 1989). Today, however,
human remains have become a key element of archaeological research and have contributed
to a burgeoning understanding of past population dynamics in general and the human
condition in particular. This is especially the case for developing and testing hypotheses and
drawing inferences about the human experience relating to diet, health, and lifestyle at all
levels, from the individual (various in Stodder & Palkovich, 2012; and see Knudson, Pestle
et al., 2012; Tiesler & Cucina, 2006) to large swaths of the globe (Steckel & Rose, 2002;
Verano & Ubelaker, 1992; White, 1999). In addition, the study of ancient human remains is
important in the ongoing discussions in anthropology about social organization, identity,
and the linkages between health and gender and health and status (various authors in
Gowland & Knüsel, 2006; Grauer & Stuart-Macadam, 1998; Knudson & Stojanowski,
2009). The study of these remains has played a fundamental role in investigating key
adaptive shifts in recent human evolution, such as the foraging-to-farming transition and
European exploration and colonization during the post-Columbian era (Bocquet-Appel &
Bar-Yosef, 2008; Cohen, 1989; Cohen & Armelagos, 1984; Cohen & Crane-Kramer, 2007;
Klaus, 2014a; Lambert, 2000a; Larsen, 2006; Larsen & Milner, 1994; Pinhasi & Stock, 2011),
and documentation and interpretation of specific disease histories (Palfi et al., 1999;
Powell & Cook, 2005; Roberts & Buikstra, 2003; Roberts et al., 2002). Especially in the last
decade or so, regional analysis has provided compelling insights into human adaptation in
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2 Introduction
broad perspective from a diverse range of settings globally (Domett, 2001; Fitzpatrick &
Ross, 2011; Hanson & Pietrusewsky, 1997; Hemphill & Larsen, 2010; Hutchinson, 2002,
2004, 2006; Judd, 2012; Lambert, 2000a; Larsen, 2001; Mushrif-Tripathy & Walimbe, 2006;
Ortner & Frohlich, 2008; Oxenham & Tayles, 2006; Pechenkina & Oxenham, 2012; Pietru-
sewsky & Douglas, 2002a,b; Robbins Schug, 2011b; Roberts & Cox, 2003; Ruff, 2010a;
Schepartz et al., 2009; Steckel & Rose, 2002; Stodder, 2008; Tung, 2012a; Weber et al., 2010;
Whittington & Reed, 1997; Williamson & Pfeiffer, 2003; Wright, 2006). These regional
studies have provided new perspectives on variation in human adaptation, including the
foraging-to-farming transition. For example, the experience of this transition in the western
hemisphere has shown a general decline in health, linked in part to maize farming. By
contrast, rice farming in Asia may have promoted better health, especially in relation to oral
health (Domett, 2001; Domett & Tayles, 2007; Douglas & Pietrusewsky, 2007; Oxenham,
2006). It is especially exciting to see the development of insights into past lives and lifestyles
in the regional context. In the mid-1980s, the record of poor health in the later prehistoric
American Southwest was beginning to emerge (Merbs & Miller, 1985). Building on this
record, discussions of conflict, its causes and consequences, and the crucial importance of
human remains in these discussions have developed, creating a forum for anthropologists
and other social scientists to engage in developing new solutions to long-standing problems.
In particular, it is through regional and continental comparisons that we are beginning to
understand the patterns and prevalence of violence and its effects on society, demography,
and health (Schulting & Fibiger, 2012).
Skeletal remains offer an important source of information for the study of human variation.
Archaeological skeletons from specific localities are more homogeneous both genetically and
in terms of the environments from where they came than are dissecting room or anatomical
skeletal series. Skeletons from the latter contexts are from many populations and highly
diverse circumstances. The use of archaeological series becomes especially important when
making conclusions about intra-population variability for a range of topics where sex and age
may be important factors.
Various surveys and manuals of human osteology and application to archaeological
settings are available (Baker et al., 2005; Bass, 2005; DiGangi & Moore, 2013; Roberts,
2009; Scheuer & Black, 2000, 2004; Schwartz, 2006; Ubelaker, 1999; White et al., 2012).
In order to address the incompatibility of different researchers’ methods and results, “stand-
ards” for skeletal data collection have been developed (Buikstra & Ubelaker, 1994). Although
dealing with the interpretive role in the study and documentation of human remains, these
works serve primarily as “how to” guides for bone identification and skeletal analysis and
not as resources for the investigation of broader issues in biological anthropology and sister
disciplines. The present book focuses on the relevance of skeletal remains to the study of the
human condition and human behavior generally; namely, how skeletal and dental remains
derived from archaeological settings reveal life history at both the individual and population
levels. It does not advocate a reliance on only human remains in order to tell the whole story
of the human condition. Rather, human remains represent a part of the broad sweep of data
derived from past settings. Human remains do not simply augment other data sources,
archaeological or historical. Rather, they provide perspectives and understandings of past
societies that pertain to human biology that simply may not be visible in other sources
(Perry, 2007).
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Introduction 3
The goal of this book is to provide a synthesis of bioarchaeology – the study of human
remains from archaeological contexts. Although the term was first applied to the study of
animal remains from archaeological settings (Clark, 1972), the focus of study then surfaced
with reference to the study of human remains in the regional “bioarcheological investigation”
of the lower Illinois River valley, an ambitious and innovative research program directed by
Jane Buikstra (1977a). This set into motion the future course of the field (Buikstra, 2006;
Knüsel, 2010; Larsen & Walker, 2010).
The field emphasizes integrative, interdisciplinary study. By doing so, the wide breadth of
bioarchaeology has engendered cross-fertilization between different disciplines, contributing
to its method and theory in approaching a wide diversity of problems. The field recognizes
the inextricable connection between biology and culture – one simply would not exist without
the other (Goodman, 2013). Just as human remains are a crucial component of study, it is the
context of these remains that provide us with meaning and substance. Context is a broad term
to include all potential sources, archaeological and otherwise, such as burial and social
inference, diet, climate, living conditions, and all else that is inferred or documented that
may inform our understanding of the people the skeletons represent.
The enormous potential of bioarchaeology for understanding the past is growing (Buikstra &
Beck, 2006; Katzenberg & Saunders, 2008; Larsen and Walker, 2010; Martin et al., 2013;
Stojanowski & Duncan, 2014). I believe that this is the case for two key reasons. First, in
contrast to earlier work that emphasized description, often poorly connected to any manner of
context, there is a growing interest in the central role that human remains play in understand-
ing patterns and trends in past societies. Second, the centrality of human remains for
understanding past biological, social, and behavioral dynamics has motivated an emphasis
on integrative research strategies, resulting in excavations of human remains having clear
agendas and questions that guide both their recovery and their study. Today, there is a very
different research profile, one that emphasizes the links between biological variation, health
and well-being, and behavior viewed broadly. This contrasts with the descriptive orientation
of earlier generations of osteologists, but it was the work of these predecessors that provided
many of the tools that served to form the field (Buikstra & Beck [2006], an historical treatment
of bioarchaeology).
This book takes a largely population perspective. However, individual-based case studies
are discussed, especially because collectively they help to build a picture of variability in
earlier societies. The population approach is critical for characterizing patterns of behavior,
lifestyle, disease, and other parameters that form the fabric of the human condition. The
discussion in the following pages also underscores the importance of culture and society in
interpreting population characteristics. Dietary behavior, for example, is highly influenced by
cultural and social norms of behavior. If an individual is taught that a specific food is “good”
to eat, then the consumption of that food item becomes fully appropriate in that cultural
context. Food is also an important indicator of a person’s place in society – access to resources
is influenced not only by where one lives but also by their identity and location within a
society. Hence, cultural and social factors play an essential role in determining diets of
individual members of a social group or society.
As I have made clear, the book is fundamentally driven by context and especially by the
placement of the biological record of skeletons in their archaeological context, recently called
“contextualization,” or the location of human remains within relevant archaeological and
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4 Introduction
historical frameworks (Knudson & Stojanowski 2009; Thompson et al., 2014), and treatment
of the body in funerary settings (Duday, 2009). Contrary to the assertion that those who study
human remains have largely disengaged bones from context, whereby the physical anthro-
pologist “does not necessarily need the archaeologist once the archaeologist has excavated
bone” (Goldstein, 2006:376–377), there has been a growing movement focusing on the links
between archaeology and physical anthropology in the development of research programs
globally. Characterizing bioarchaeology as involving a disengagement of bodies from place
misrepresents the vast majority of research cultivated in this exciting discipline, especially
given the enormous progress of the field today.
Unlike many of the aforementioned guides to osteological analysis, this book is not a book
about methods. Certainly, methodological developments make possible much of the discus-
sion presented in the following chapters (see various authors in Katzenberg & Saunders,
2008). Moreover, technological advances, such as imaging, have provided new insights
unimaginable a decade ago (various in Chhem & Rühli, 2004). However, this book focuses
on how human remains inform our understanding of the past. By doing so, this book is
intended to feature the various insights gained about human behavior and biology rather than
to describe or evaluate specific methods and techniques of skeletal analysis. This approach is
central to the biocultural perspective offered by anthropologists – we must seek to envision
past populations as though they were alive today, and then ask what information drawn from
the study of skeletal tissues would provide understanding of them as functioning, living
human beings and members of populations. Nor is this book a critical review, highlighting the
shortcomings of the field or what bioarchaeologists should be doing, but are not.
Bioarchaeological findings are important in a number of areas of scientific and scholarly
discourse. Within anthropology, the use of human remains in interpreting social behavior
from mortuary contexts is especially fruitful (Artelius & Svanberg, 2005; Beck, 1995; Buik-
stra, 1977a; Chapman et al., 1981; Chesson, 2001; Gowland & Knüsel, 2006; Humphreys &
King, 1981; Knudson & Stojanowski, 2009; O’Shea, 1984; Parker Pearson, 2000). The story
human remains tell is also reaching an audience outside anthropology. There is an increase in
use of bioarchaeological data in history, economics, and nutrition science. Skeletal studies of
nutrition, disease, and related topics, and the importance of human remains in developing a
broader understanding of economic history are opening a new path of research interest
(Steckel & Rose, 2002). The emerging role of skeletal remains in the study of the human
condition was underscored by the eminent historian, John Coatsworth (1996:1), who high-
lights the “masses of evidence” provided from bioarchaeological investigations and the
important role they play in understanding historical developments.
Breakthroughs have been made in the analysis of various body tissues in archaeological
settings, including hair, muscle, skin, and other soft tissues (Arriaza, 1995; Asingh &
Lynnerup, 2007; Aufderheide, 2003; Brothwell, 1987; Chamberlain & Parker Pearson, 2001;
Cockburn et al., 1998; Hansen et al., 1991; Lynnerup et al., 2002; Stead et al., 1986). The
discussions presented in this book focus largely on skeletal and dental tissues. Building on the
study of human remains, the unifying theme in this book is behavioral inference. My
discussion of behavior is not limited to physical activity; rather, it is considered in a wider
perspective, including (in order of appearance in the book) physiological stress, exposure to
pathogenic agents, injury and violence, physical activity, masticatory and extramasticatory
uses of the face and jaws, dietary reconstruction and nutritional inference, population history,
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Introduction 5
and social factors and how they influence health and lifestyle. Simply, almost everything
available in the study of human remains has behavioral meaning.
Bioarchaeology is represented throughout the world. This book draws upon a sample of this
record in illustrating important points and issues. The book deals with all regions of the globe,
but especially in those areas that have been studied in depth by bioarchaeologists. One of the
exciting advances in the field in the last decade or so is the proliferation of bioarchaeological
treatments outside North America, especially Europe, Asia, and the Pacific (Domett, 2001;
Lynnerup, 1998; Murphy, 2002, 2003; Ortner & Frohlich, 2008; Oxenham & Tayles, 2006;
Papathanasiou, 2001; Pechenkina & Oxenham, 2013; Pietrusewsky & Douglas, 2002b; Rife,
2012; Roberts & Cox, 2003; Ruff, 2010a; Schepartz et al., 2009; Whittington & Reed, 1997;
and many others). Reflecting this increasingly international bioarchaeology are the inter-
national, multidisciplinary collaborations around the world, including in Latin America,
Europe, Asia, and Africa.
Various points made in the book are addressed by contrasting and comparing data sets
from skeletal assemblages representing human populations from different levels of socio-
political complexity and differing subsistence regimes. Because of the vagaries of dietary
reconstruction in the archaeological past, anthropologists usually characterize human groups
broadly, using terms such as “foragers” or “farmers.” The reader should recognize that these
terms are often overly simplistic and do not convey the underlying complexity of human
adaptive systems adequately. Nevertheless, these categories help us to understand broad
behavioral and adaptive features of different groups better, and therefore, are the starting
points to facilitate yet more specific and contextual reconstructions and interpretations of
past lifeways. Of far more importance to the focus of this book is that these contrasts and
comparisons add an important dimension to the growing discussion in anthropology oriented
toward the understanding of the causes and consequences of adaptive and behavioral shifts in
the past and present.
A fundamental barrier to understanding health in today’s populations is the very narrow
temporal window in which they have been studied. The prevalence records for osteoarthritis
and oral degenerative conditions, for example, are limited largely to the last several decades
when large-scale epidemiological studies of living populations were first undertaken (Arden &
Nevitt, 2006; Blinkhorn & Davies, 1996; Issa & Sharma, 2006; Jordan et al., 2007; Pilot, 1998;
Sreebny, 1982; Woodward & Walker, 1994), representing approximately only the last 0.1% of
the history and evolution of our species. Biomedical, experimental, molecular, and behavioral
studies of skeletal and oral degenerative conditions have been investigated in great detail in
human populations for this 0.1% window of time. However, these studies are limited in scope,
focusing primarily on remarkably gross categories of human variation. They are deficient
because they underrepresent and mischaracterize human biological variation, reducing the
variation to simple dichotomous (or other limited) comparisons having little to do with real
biological or social variation. Analysis of the recent biomedical and epidemiological literature
characterizes variation according to “race” (e.g., White vs. Black vs. Hispanic), geography
(mostly United States and western Europe), sex (men vs. women), diet (e.g., access to refined
sugar), and socioeconomic status based on income level (upper-, middle-, and lower-income),
in addition to the very narrow temporal window (Allen, 2010; Dominick & Baker, 2004;
Jordan et al., 2007; Pilot, 1998). The (mis)characterization of human variation has important
implications for understanding an increasingly diverse society in the United States, far more
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6 Introduction
diverse than the racial categories so prevalent in biomedical research would seem to imply. By
expanding understanding of diversity, in terms of both broad temporal and geographic
perspectives, bioarchaeology as an anthropological subdiscipline provides a more informed
understanding of health in the present through a consideration of heath in the past.
Anthropologists, economic historians, and other social scientists have long recognized that
humans in the past and present are extraordinarily diverse in their food consumption
practices, social habits, workloads, and other behavioral characteristics that collectively
characterize health and lifestyle. The biomedical and epidemiological literature on degenera-
tive conditions is known from a limited, if not insufficient record. Moreover, this record is
limited to the study of populations that have mitigated some of the conditions that influence
the prevalence of degenerative conditions most strongly, namely industrialized societies
having access to healthcare, adequate nutrition, and labor-saving technology. For example,
prevalence of dental caries in many developed countries has been reduced due to the
introduction of fluoride in drinking water, which is certainly the case in the United States
(McDonagh et al., 2000). In addition, owing to work-saving technology, workload has greatly
declined in developed countries especially. If physical activity is the primary factor in
determining prevalence and pattern of osteoarthritis, for example, then one could predict a
decline in its prevalence, especially in recent societies. That is, as advances in technology
essentially replace what the body used to do in work and activity generally, we should expect
to see long-term trends in terms of a decline in osteoarthritis. This hypothesis has not been
tested using the kind of systematic approach offered by bioarchaeology.
Bioarchaeology offers a unique opportunity to study a much more diverse sampling of
humanity, namely the last 10 000 years of more than seven million years of human evolution,
greatly extending the time framework for characterizing the biocultural context surrounding
some of the most important changes and developments in the evolutionary history of our
species. Arguably, it is this small percentage of our evolution where crucial developments
occurred that set the stage for the rise of modern civilization, namely farming, appearance of
complex societies, urbanization, and industrialization.
Many studies of human remains from archaeological contexts focus on a single population
without actively linking the analysis of these remains to the context (climate, diet, time,
culture, settlement, and economic system) from which they derive. A central aim of bioarch-
aeology is to establish a comprehensive record of skeletal and dental conditions in relation to
prevalence and pattern to develop an understanding of behavior and the costs and conse-
quences of particular lifestyle circumstances and conditions. Bioarchaeology provides an
unmatched record of health and life conditions in the past, thereby extending our understand-
ing of diversity in geography, cultures, and time that is simply not possible with sole reliance
on limited health attributes of living populations.
Human skeletal and dental tissues are remarkable records of lives led in the past, what
Stanley M. Garn referred to as “a rich storehouse of individual historical events” (1976:454).
This book provides a tour of the vast holdings in this storehouse, displaying the knowledge
gained about earlier peoples based on the study of their mortal remains.
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Stress and deprivation during growth
2 and development and adulthood
2.1 Introduction
Physiological disruption resulting from impoverished environmental circumstances – “stress” –
is central to the study of health and well-being and the reconstruction and understanding of
health, adaptation, and behavior in both earlier and contemporary human societies. Stress
involves disruption of homeostasis, or the maintenance of a constancy of conditions that keep
the body’s internal environment stable. Stress is a product of three key factors, including
(1) environmental constraints, (2) cultural systems, and (3) host resistance. Goodman and others
(Goodman, 1991; Goodman & Armelagos, 1989; Goodman & Martin, 2002; Goodman, Martin
et al., 1984, 1988; Klaus, 2012; Martin et al., 1991) modeled the interaction of these factors at
both the individual and population levels (Figure 2.1). This model of health and context
emphasizes the role of environment in providing both the resources necessary for survival and
the stressors that may affect the health and well-being of a population, yet includes the profound
influence of culture in health outcomes. Cultural systems serve as protective buffers from the
environment, such as shelter and clothing as buffers against temperature extremes. Cultural
systems can be highly effective at mitigating behaviors necessary for extraction of important
nutrients and resources from the environment, thus supporting the ability to maintain stability.
It appears impossible for the full spectrum of stressors in an environment to be buffered against;
some inevitably slip through the filters of the cultural system. In these instances, the individual
may exhibit a biological stress response observable at the tissue level (bones and teeth).
Importantly, stress, buffering systems, and tissue-level responses are not linked by a simplistic,
linear relationship. Instead, they can interact with and influence other variables within other
levels of the model. Physiological disruption feeds directly back into environmental constraints
and cultural systems. This model makes clear that health is a key variable in the adaptive process.
Biological stress has significant functional consequences for the individual and for the
society in which they are living. Elevated stress and associated disruption of homeostasis may
lead to a state of functional impairment, resulting in diminished cognitive development and
work capacity. The reduction in work capacity can be detrimental if it impedes the acquisition
of essential resources (e.g., dietary) for the maintenance of the individual, the population, and
the society. If individuals of reproductive age are affected by poor health, then decreased
fertility may be the outcome. Ultimately, the success or failure of a population and its
individual constituents to mitigate stress has far-reaching implications for behavior and the
functioning of society.
Stress and developmental instability in early life, prenatal and postnatal, have important
implications for health outcomes in later life. David Barker’s research on chronic diseases in
middle age reveals that individuals with low birth weight and nutritional deprivation in early
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8 Stress and deprivation during growth and development and adulthood
Host Resistance
Feedback
Loop
Physiological Disturbance
Growth Disruption
Feedback Disease
Loop Death
Poplulation-Level Outcomes
Increased Morbidity: Epigenetic Effects
Decreased Work and Reproductive Capacity
Sociocultural Disruption and Instability
Figure 2.1 Biocultural model for interpreting stress. This model emphasizes the primacy
of environmental constraints and cultural influences on outcomes in health and well-being.
(Adapted from Klaus, 2012; reproduced with permission of author and University Press
of Florida.)
life are more predisposed to earlier death and greater frequency of chronic disease, including
cardiovascular disease, hypertension, and type 2 diabetes, than are individuals with normal
birth weight and sufficient prenatal and early natal nutrition (Barker, 2001, 2012; Barker
et al., 2002; Beltrán-Sánchez et al., 2012; Hales et al., 1991; Harding, 2001; and see Jo-
vanovic, 2000; various in Henry & Ulijaszek, 1996; and others). Similarly, low birth weight
infants show lower bone mass as adults than infants with normal birth weight (Antoniades
et al., 2003; Cooper et al., 1995; Gluckman et al., 2008). Experimental evidence in laboratory
animals suggests that poor prenatal environments are also a risk factor for growth stunting
over the life course in general (Dancause et al., 2012). While the prenatal environment does
not determine health consequences in adulthood, it appears to have a profound role to play in
programing circumstances in later life that promote poor health. A growing record supports
Barker’s developmental origins hypothesis regarding the profound influence of intrauterine
stresses in early life on later health, morbidity, and mortality.
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2.3 Growth and development: skeletal 9
approach stems from the recognition that health is a composite of nutrition, disease, and other
aspects of life history. Simply, simultaneous study of multiple, but independent, markers of
stress provides a more reliable and informed understanding of composite health as docu-
mented in a skeletal series. Contrary to medical models of health, stress and disease (see
Chapter 3) represent a continuum rather than a presence versus absence phenomenon, with
respect to both the population and the individuals that comprise it.
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10 Stress and deprivation during growth and development and adulthood
comparison with other life periods (Bogin, 1999). During adolescence, genetic influence on
growth is more strongly expressed than in childhood (Bogin, 1999; Bogin & Loucky, 1997).
Juveniles have been growing taller over much of the twentieth century in industrialized
countries and in some developing nations. This secular trend in growth is related to a variety
of environmental and cultural changes, including improvement in food availability and
nutrition, sanitation, reduction of infectious disease, and increased access to Western health-
care. As the environment improves, growth increases. On the other hand, declines in growth
velocity are well documented, especially during periods of dietary deprivation in wartime
settings, famines, and economic crises (Eveleth & Tanner, 1990; Himes, 1979; Komlos 1994).
This link between growth status and environment is well documented via analysis of histor-
ical data. Comparisons of heights of British school children from various regions and eco-
nomic circumstances for the period of 1908 to 1950 show that children were generally shorter
in areas experiencing high unemployment (e.g., Glasgow, Scotland) than in other regions with
more robust economies (Harris, 1994). These differences were especially pronounced during
the severe economic depression in the late 1920s when nutritional and general health of
children of unemployed parents declined. Similarly, growth velocity and attainment per age
increased in post-World War II following the amelioration of negative socioeconomic condi-
tions (Cardoso & Gomes, 2009; Malina et al., 2010; Tanner et al., 1982). In post-1945 Poland,
relatively greater increases in growth were documented in higher socioeconomic groups
(Bielicki & Welon, 1982).
Beginning with Francis Johnston’s pioneering work on childhood growth based on the
study of archaeological skeletons (Johnston, 1962, 1969), a range of studies have extended
our understanding of growth rates in past societies. That is, the general pattern of juvenile
growth in archaeological populations is broadly similar to that in living populations (Arme-
lagos et al., 1972; Boldsen, 1995; Edynak, 1976; Hillson et al., 2013; Hoppa, 1992; Huss-
Ashmore, 1981; Johnston, 1962; Lewis, 2002; Mays, 1999; Merchant & Ubelaker, 1977;
Molleson, 1995; Ribot & Roberts, 1996; Ruff et al., 2013; Ryan, 1976; Saunders, 2008; Sciulli &
Oberly, 2002; Storey, 1992a, 1992b; Sundick, 1978; Walimbe & Gambhir, 1994; Walker, 1969;
and see later). The congruence of growth in past and living groups suggests that there has not
been a change in the general pattern of growth in recent human evolution (Saunders, 2008).
That is, patterns and processes of growth in known human populations appear to have been
present for at least the last 10 000 years of our evolutionary history, and likely longer.
Some populations appear shorter for age than others. Analysis of juvenile long bones from
prehistoric North America reveals evidence of growth retardation in agricultural and mixed
subsistence economies versus foragers. In children younger than six years of age in the
prehistoric lower Illinois River valley, matching of femur length to dental age reveals growth
suppression in late prehistoric (Late Woodland period) maize agriculturalists in comparison
with earlier foragers (Middle Woodland period) (Cook, 1979, 1984). Cook (1984) concluded
that the decline in growth was due to a decrease in nutritional status with the shift to a
protein-poor maize diet. Children short for their age during the later prehistoric period tend to
express a higher frequency of stress indicators (e.g., porotic hyperostosis, enamel defects) than
children who are tall for their age, lending further support for nutritional deficiency as a prime
factor contributing to growth retardation.
Lallo (1973; and see Goodman, Lallo et al., 1984) found, in addition, a decrease in the
growth of femur, tibia, and humerus diaphysis lengths and circumferences during the
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2.3 Growth and development: skeletal 11
Mississippian period (AD 1200–1300) in comparison with earlier periods (AD 950–1200) in the
central Illinois River valley. Dietary change during this time involved a shift from mixed
foraging and farming to intensive maize agriculture. Growth during the period between two
and five years of age was especially slow, which Goodman and coworkers (1984) conclude
reflects an increase in physiological stress due to poorer nutrition and the presence of other
stressors during the later prehistoric occupation of the region.
The impact of increased stress loads due to the combined effects of undernutrition,
European-introduced infectious disease (e.g., smallpox, measles), warfare, and increased
social disruption has been investigated in the late prehistoric and contact-era Arikara
Indians of the upper Missouri River valley (Jantz & Owsley, 1984a, 1984b, 1994a; Owsley &
Jantz, 1985). Matching of long bone lengths (femur, tibia, humerus, radius) to dental age in
perinatal (late fetal/early neonatal) and other juvenile skeletons reveals that late postcontact
era (AD 1760–1835) Arikara juveniles are smaller than early postcontact (AD 1600–1733)
juveniles, suggesting declining health status as European influence and encroachment into
the region by other tribes increased. Comparison of postcontact Arikara with healthy upper
middle-class Euroamericans from Denver, Colorado confirms a pattern of slower growth in
early childhood in the Native American group (Hillson et al., 2013; Ruff et al., 2013)
(Figure 2.2). In contrast, growth in young children from Neolithic Çatalhöyük, Turkey is
strongly similar to the Denver growth study sample, indicating relatively better nutrition
and health overall, a pattern consistent with other skeletal indicators (Hillson et al., 2013;
Ruff et al., 2013) (Figure 2.2).
200
180
160
140
Stature (cm)
120
100
80
60
0 2 4 6 8 10 12 14 16 18 20 22 24
Age (years)
Figure 2.2 Fitted curves for estimated juvenile statures for protohistoric Arikara (open
circles and dashed lines), Neolithic Çatalhöyük (solid diamonds and solid line), and modern
Denver population (asterisks and dotted line). (From Ruff et al., 2013; © 2012 Wiley
Periodicals, Inc.)
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12 Stress and deprivation during growth and development and adulthood
The interaction between stress and population mobility has been examined in the compari-
son of Late Archaic-period foragers from the Carlston Annis Bt-5 site, Kentucky (3992–2655
BC) and Late Woodland foragers from the Libben site, Ohio (AD 800–1100) (Mensforth, 1985).
Archaeological evidence indicates that Late Archaic populations were highly mobile and
exclusively dependent on wild plants and animals. In contrast, maize was consumed by the
Libben population, but it was of minor dietary significance. For both groups, nutrition appears
to have been adequate (Mensforth, 1985). Comparisons of tibia lengths reveal a general
similarity between the two groups from birth to six months and from four years to 10 years.
For juveniles aged six months to four years, Libben tibiae are shorter than Bt-5 tibiae. The
growth period between six months and four years – the period differing most between Bt-5
and Libben populations – is highly sensitive to metabolic disruption. During this period, an
infant undergoes weaning, involving the shift from breast milk, a relatively stable, nutritious
food source, to exclusive solid food, often less stable in availability, less digestible, and less
nutritious. Passive immunities derived from consumption of breast milk are lost during
weaning during this period of life (Popkin et al., 1986). These immunities are crucial for early
health and well-being because the child’s immune system is not fully developed until after
five years of age (Newman, 1995). Consistent with this outcome of compromised immunity,
Mensforth (1985) found a high prevalence of nonspecific periosteal infections in the Libben
infants, suggesting that high levels of infectious disease in infancy and young childhood
contributed to growth retardation. Thus, in comparison with the Bt-5 population, the Libben
population experienced the effects of greater sedentism and community size that fostered
poor sanitation, elevated infectious disease, and poor health.
Comparison of archaeological samples with a modern reference population (Denver, Color-
ado; Maresh, 1970) confirms the presence of growth rate suppression in children from
archaeological settings. That is, calculation of the average percentage of adult size attained
in successive age groups for the major long bones reveals a slow-down in most settings
(Humphrey, 2003), consistent with the general finding that children in archaeological con-
texts are shorter than their modern counterparts (Lewis, 2007; Mays, 1999). Lovejoy and
coworkers (1990) argue that widespread infection was the chief cause of growth retardation in
the Libben setting. They suggest that inflammation would result in an increased production of
cortisol, the major natural glucocorticoid, which results in limitation of growth and availabil-
ity of amino acids. Thus, elevation of infection in the Libben population may have had a
strong influence on growth generally (Lovejoy et al., 1990).
Historic-era skeletal series furnish important insights into stress in the recent past. Saunders
and coworkers (1995, 2002; Humphrey, 2003) analyzed growth data available from a large
series of juvenile remains from the St. Thomas’ Anglican Church cemetery in Belleville,
Ontario. The cemetery was used by a predominantly British-descent population during the
period of 1821 to 1874. Comparisons of femur length from juveniles buried in the cemetery
with a tenth-century Anglo-Saxon series from Raunds, England, and modern growth data
from Denver, Colorado (Maresh, 1970), indicate a strong similarity in overall pattern of
growth between the three groups (Figure 2.3). The two cemetery samples are temporally
separate, but share general ethnic origins with the modern US population. Figure 2.3 shows
that the St. Thomas’ series is slightly shorter than the modern series for age. The Raunds series
is considerably shorter than either of the other groups, which is to be expected given the
inferior living standards of tenth-century England. With regard to the St. Thomas’ skeletons,
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2.3 Growth and development: skeletal 13
400
350
Diaphyseal length (mm)
300
250
200
150
100
50
0 1 2 3 4 5 6 7 8 9 10 11 12 13
Age (years)
Figure 2.3 Fitted curves for femoral diaphyseal length for the nineteenth-century
St. Thomas’ Church cemetery (dotted line), tenth-century Raunds Anglo-Saxon skeletons
(dashed line), and twentieth-century Denver, Colorado, living population (solid line).
(From Saunders & Hoppa, 1993; reproduced with permission of John Wiley & Sons, Inc.)
Saunders and coworkers suggest that juveniles died from acute causes and not chronic
conditions (e.g., chronic infections or chronic undernutrition) that would result in a decrease
in skeletal growth. Children younger than two years of age had slightly lower growth rates
than those of modern twentieth-century populations. They regard this as perhaps representing
stresses associated with poor maternal health and prenatal growth.
Similarly, on the north coast of Peru, comparison between late pre-Hispanic- and
postcontact-period femoral growth velocity reveals markedly slower growth among the
indigenous children, especially at ages 5, 10, and 12 years in the colonial Lambayeque Valley
(Klaus & Tam, 2009). This decrease in the rate of growth takes place in a setting of introduced
European disease, increased prevalence of other skeletal stress markers, poor diets, and
sociopolitical marginalization.
Analysis of juvenile cortical bone growth via measurement of cortical thickness provides a
complementary source of information to long bone lengths. In living populations, deficiencies
in cortical bone mass are present in groups experiencing undernutrition (Agarwal, 2008;
Agarwal & Glencross, 2011; Cooper et al., 1995; Frisancho, Garn et al., 1970; Garn, 1970;
Garn et al., 1964; Gluckman et al., 2008; Himes, 1978; Himes et al., 1975). The pioneering
investigation by Garn and coworkers (1964), for example, showed that malnourished Guate-
malan children have reduced cortical bone in comparison with well-nourished reference
groups. Although bone lengths increased during periods of growth recovery, cortical thickness
continued to show deficiencies due to earlier episodes of bone loss. Thus, growth recovery
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14 Stress and deprivation during growth and development and adulthood
may involve an increase in bone length (and attained height), but not bone mass (Antoniades
et al., 2003; Huss-Ashmore, 1981; Huss-Ashmore et al., 1982).
Cortical bone mass is a sensitive indicator of environmental disturbance in archaeological
settings. Comparison of femoral cortical thickness from Middle Woodland (Gibson site) and
from Late Woodland (Ledders site) series from west-central Illinois reveals a reduction in bone
mass in young children (24–36 months), the presumed time of weaning and increased dietary
stress (Cook, 1979). In contrast, Hummert and coworkers (Hummert, 1983; Hummert & Van
Gerven, 1983; Van Gerven et al., 1985) documented increased cortical bone deficiencies in
exclusively older children from the early to late Christian periods in Sudanese Nubia (c. AD
550–1450). Long bone lengths of Nubians are shorter in the early Christian period than in the
late Christian period, which may be due to nutritional deficiencies and bacterial and parasitic
infections (Hummert, 1983; Hummert & Van Gerven, 1983). Increasing political autonomy
during the later Christian period may have served to improve living conditions, resulting in
better growth status and health generally. Cortical bone mass continued to be deficient in the
later period, indicating that stress was present throughout the Christian period, both early and
late. Unlike the long bone lengths that show a recovery during adolescence, there was a
continued decrease in cortical bone mass in older children, suggesting that growth in long
bone length continued at the expense of cortical bone maintenance (Hummert, 1983; and
compare with Garn et al., 1964).
Comparison of individuals interred in high-status brick vaults versus low-status grave pits
in a nineteenth-century setting in Birmingham, England revealed that the former had higher
cortical thickness for age than the latter (Mays, Brickley et al., 2009). These differences likely
reflect greater access to adequate nutrition in higher-status individuals than in lower-status
individuals in this early industrial setting.
Analysis of bone mass in living and past populations underscores an important point about
the plasticity of human skeletal tissue and its relationship with growth and development:
skeletal tissue adapts in a highly dynamic way to physiological requirements of the living
body. These requirements are informed by the environment, including biological, social, and
cultural influences over the life course of the individual, from conception through infancy,
childhood, adolescence, and adulthood (and see Agarwal & Beauchesne, 2011). This life course
perspective is motivated by the understanding that the skeleton reflects the lived experience of
the individual at all stages of life. This contextualized approach provides a powerful tool for
understanding variation in bone mass and the implication for risk of fracture and debilitation
(Glencross & Agarwal, 2011; and see Chapter 4).
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2.3 Growth and development: skeletal 15
terminal height (Bailey et al., 1984; and see Bogin & MacVean, 1983; Frisancho, Newman
et al., 1970; Satyanarayana et al., 1980).
The close ties between environmental stress – especially due to poor nutrition – and adult
height are abundantly documented in research developing out of a growing interest in
anthropometric history (Floud et al., 1990; Heyberger, 2007; Komlos, 1994, 2009; Steckel,
1995, 2005; Steckel & Floud, 1997; Tatarek, 2006; Vercellotti et al., 2011; and many others).
Originally inspired by controversy over the health and well-being of enslaved African-
Americans (Steckel, 1979), current research has broadened greatly to include a range of other
populations in North America, Europe, and Asia (Fogel et al., 1983; Komlos, 1994; Steckel,
1995). Evidence from a wide range of recent historical populations indicates that stature
variability can be explained in large part by environmental factors (Steckel, 1995). This
evidence shows that terminal height is a product of nutritional adequacy and, to a lesser
extent, disease history. Individuals with adequate nutrition tend to reach their genetic growth
potential; those with poor nutrition do not.
Genetic factors are also important in determining terminal height. For example, well-off
Japanese reach only the fifteenth height percentile of well-off British (Tanner et al., 1982), and
genomic studies reveal associations with height (Becker et al., 2011). Climate may be a
mediating factor in determining height, but stature shows little correlation with latitude in
the comparison of a wide range of human populations (Ruff, 1994a). Of much more import-
ance to the issue of climate is body breadth, which plays a crucial role in determination of the
amount of body surface area to body mass in hot and cold climates (Ruff, 1994a).
Like childhood growth, there is a temporal trend of stature increase with economic and
nutritional improvement (Boldsen, 1995; Floud, 1994; Greulich, 1976; Komlos & Kriwy, 2002;
Malina et al., 2005; Őzer et al., 2011; Yagi et al., 1989; and many others) and decline during
times of hardship and deprivation (Bogin & Keep, 1999; Fogel et al., 1983; Kemkes-
Grottenthaler, 2005; Kimura, 1984; Leonard et al., 2002; Price et al., 1987; Steegmann,
1985). These analyses show that although growth and height have a genetic component,
environmental factors play a profound role in these outcomes.
Terminal height data for historical populations are drawn from various archival sources,
including military records (Bielicki & Welon, 1982; Komlos, 1989; Mokyr & Ó Gráda, 1994;
Sandberg & Steckel, 1987; Steegmann, 1985, 1986; Steegmann & Haseley, 1988), military
preparatory schools (Komlos, 1987), prison inmates (Riggs, 1994; Tatarek, 2006), enslaved
African Americans (Steckel, 1979, 1986, 1987), and voter registrations (Wu, 1994). Analysis
of these data sets by economic historians reveals temporal trends in stature that can be
linked with changing economic conditions relating to nutritional adequacy in particular and
health status in general. Terminal stature in Euroamerican populations shows significant
variability in relation to time, geography, and socioeconomic status. Over the last several
centuries, marked improvements in health and nutrition have been documented. Popular
convention holds that adult stature has increased during and after the Colonial period in
North America. Steckel (1994) analyzed stature data for American-born Euroamerican male
soldiers for the period of 1710 to 1950. Contrary to convention, twentieth-century Euro-
american males are not appreciably taller than their predecessors living in the eighteenth
century (Steckel, 1995).
Skeletons from archaeological contexts offer an important complementary data set for
stature analyses based on archival sources. Comparison of stature estimates derived from
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16 Stress and deprivation during growth and development and adulthood
measurements of long bones shows little change from the pre-modern (1675–1879) to the
modern (1950–1975) period in the United States (Angel, 1976; Larsen, Craig et al., 1995)
(Table 2.1). Analysis of military archival records shows a slight rise in stature (1710–1830) in
European-descent, American-born males, followed by a marked decline that continues for
most of the nineteenth century (Costa & Steckel, 1997). This decline appears to coincide with
the movement of the population from a rural to urban setting, and increasingly poorer
sanitation. Simply, the height reductions reflected the health declines of the newly urbanized
people in the United States. Following the passage of sanitation laws in cities in the late
nineteenth century, improved nutrition, and healthier environments overall, there is a steady
rise in stature. The skeletal record for stature in the comparison of medieval and early modern
Europeans reveals a reduction in height, reflecting reduced quality of life, exposure to
pathogens, and decline in nutrition in the shift from rural to urban settings (Kemkes-
Grottenthaler, 2005).
In the New World, the transition to agriculture involved the adoption of maize as a key
component of subsistence. There are several negative aspects of maize that potentially could
lead to growth disruption and reduced height in native populations in the Americas. Although
maize meets energy requirements, it is deficient in the essential amino acids lysine, isoleucine,
and tryptophan (FAO, 1970; Whitney & Rolfes, 2011). Because maize has these amino acid
deficiencies, it provides a poor protein source. Niacin (vitamin B3) in maize is chemically
bound, which reduces the bioavailability of this nutrient to the consumer. In maize-based
diets, iron absorption is very low (Ashworth et al., 1973), methionine and phenylalanine are
minimally represented, and the leucine–isoleucine ratio is inadequate. The nutritive value of
maize is altered by the preparation techniques used to transform it into food. Many native
New World societies enhance the nutritional content of maize via alkali-processing (Katz
et al., 1974; Stahl, 1989). The addition of alkali promotes the availability of niacin during
digestion (FAO, 1953). Some evidence suggests that these treatment protocols actually pro-
mote dystrophic effects (Huss-Ashmore et al., 1982). Additionally, removal of the pericarp
(bran) in the grinding process decreases the nutritive value of maize; important minerals and
some fiber are removed if the pericarp is winnowed from the maize. If the aleurone, the
protein- and niacin-rich layer, and bran are removed simultaneously, important nutrients are
also lost (FAO, 1953; Rylander, 1994). Additionally, thiamine content is affected by the
manner in which the maize is processed.
The study of temporal series of archaeological remains, especially in the comparison of New
World foragers with later farming populations, reveals trends that are consistent with declin-
ing nutritional quality in both maize consumers and populations dependent on other plant
domesticates. Comparisons of prehistoric Georgia coastal foragers (pre-AD 1150) with later
maize farmers (AD 1150–1550) indicate reductions in stature of about 3% for adult females
and 1% for adult males (Larsen, 1982; Larsen, Crosby et al., 2002). Similar reductions in other
New World settings are documented in the American Midwest (Perzigian et al., 1984; Sciulli &
Oberly, 2002; but see Cook, 1984, 2007), Mesoamerica (Haviland, 1967; Márquez Morfín & del
Ángel, 1997; Nickens, 1976; Saul, 1972; Stewart, 1949, 1953a; but see Danforth, 1994;
Márquez Morfín et al., 2002; Wright & White, 1996), and Peru (Pechenkina, Vrandenburg
et al., 2007). Comparisons of agricultural populations with other settings indicate relatively
short statures in Mesoamerica (Storey, 1992a; Danforth, 1999), Ecuador (Ubelaker, 1994), and
Peru (Pechenkina, Vrandenburg et al., 2007), which are linked with chronic malnutrition.
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2.3 Growth and development: skeletal 17
Males
Patuxent Maryland rural 1658–1690 6 170 King & Ubelaker, 1996
Colonial-Civil War Various 1675–1879 21 173 Angel, 1976
Belleview Georgia rural 1738–1756 3 170 Rathbun & Scurry, 1991
Old Quebec Colonial military 1746–1747 30 173 Cybulski, 1988; Saunders, 1991
Walton Connecticut rural 1750–1830 5 176 Bellantoni et al., 1997
Fort William British military 1755–1757 14 177 Steegmann, 1986
Henry
Fort Laurens US military 1779 13 172 Sledzik & Sandberg, 2002
Bradford’s Colonial military 1812–1814 180 174 Saunders, 1991
Company
Snake Hill US military 1814 23 174 Sledzik & Sandberg, 2002
St. Thomas’ Ontario village 1820–1860 127 172 Saunders et al., 2002; Sledzik &
Sandberg, 2002
Prospect Hill Ontario 1824–1879 17 173 Pfeiffer et al., 1989
immigrants
Harvie Ontario rural 1825–1894 5 171 Saunders & Lazenby, 1991
Highland Park New York 1826–1863 94 172 Higgins et al., 2002
poorhouse
Cross Illinois rural 1829–1849 5 175 Larsen, Craig et al.,1995
Uxbridge Massachusetts 1831–1872 9 170 Sledzik & Sandberg, 2002;
poorhouse Wesolowsky, 1989;
Mt. Gilead Georgia rural 1832–1849 5 172 Wood et al., 1986
Voegtly Pennsylvania 1833–1861 32 170 Ubelaker & Jones, 2003
town
Glorieta Pass US military 1862 24 173 Sledzik & Sandberg, 2002
Little Big Horn US military 1876 8 176 Sledzik & Sandberg, 2002
West Point cadets2 US military 1840s–1870s 334 172 Komlos, 1987
Modern US2 General 2003–2006 2331 178 McDowell et al., 2008
population
Females
Patuxent Maryland rural 1658–1690 5 161 King & Ubelaker, 1996
Colonial-Civil War Various 1675–1879 7 160 Angel, 1976
Walton Connecticut rural 1750–1830 8 166 Bellantoni et al., 1997
Harvie Ontario rural 1825–1894 4 161 Saunders & Lazenby, 1991
Highland Park New York 1826–1863 64 161 Higgins et al., 2002
poorhouse
Cross Illinois rural 1829–1849 6 163 Larsen, Craig et al., 1995
Mt. Gilead Georgia rural 1832–1849 3 162 Wood et al., 1986
Voegtly Pennsylvania 1833–1861 14 160 Ubelaker & Jones, 2003
town
Modern US2 General 2003–2006 2477 163 McDowell et al., 2008
population
1
Mean values in cm.
2
Documentary/living statures.
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18 Stress and deprivation during growth and development and adulthood
1.5
Males
1 Females
0.5
Z-scores, mean+/-SE
-0.5
-1
-1.5
-2
Beiliu Jiangzhai Shijia Kangjia Xicun
Yangshao Longshan Dynastic
Figure 2.4 Variations in long bone lengths between Yangshao and Longshan cultures, as
expressed in average Z-scores. (From Pechenkina et al., 2002; reproduced with permission
of authors and John Wiley & Sons, Inc.)
In Mesoamerica, prehistoric heights are greater than in modern Maya (Danforth, 1999;
Márquez Morfín & del Ángel, 1997), which appears to be related to increased reliance on
maize and associated nutritional decline, especially in the last 500 years.
Other archaeological settings show reduction in stature in the shift to agricultural econ-
omies or agricultural intensification. Analysis of height data shows that foragers or incipient
farmers in settings in East Asia were taller and more robust than their farming descendants
(Bulbeck & Lauer, 2006; Kennedy, 1984; Temple, 2008; but see Clark et al., 2014). With regard
to millet farmers in the Wei and Yellow River area of China, earlier less intensive farming
populations are considerably taller than later more intensive farmers (Pechenkina et al., 2002,
2007, 2013) (Figure 2.4). Similarly, comparisons of skeletal series from the Upper Paleolithic
through the Neolithic in western Europe indicate a general reduction in average stature
(Bennike & Alexandersen, 2007; Meiklejohn et al., 1984; Meiklejohn & Babb, 2011; Roberts &
Cox, 2003, 2007; but see Jacobs, 1993). In some settings, reduction in stature coincided with
resource intensification, with either agriculture (Pechenkina et al., 2002, 2013; Van Gerven
et al., 1995) or foraging (Ginter, 2011). However, analysis of the record for Europe reveals
no change in the trajectory of stature reduction with the foraging-to-farming transition
(Meiklejohn & Babb, 2011). Nevertheless, the overall record suggests a link involving diet,
resource acquisition, physiological stress, and terminal height.
Much of the research on body size in children and adults in archaeological settings is
oriented toward tracking the consequences of adaptive transformations, primarily from
foraging to farming; relatively little is known about other dietary transitions. The
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2.3 Growth and development: skeletal 19
consequences of change in dietary focus not involving agriculture are manifested in temporal
comparisons of native populations from the Santa Barbara Channel region of southern
California (Lambert, 1993; Walker, 2006; Walker & Thornton, 2002). In this region, popula-
tions shifted their dietary emphasis from terrestrial resources – especially plant foods – to
marine resources after 500 BC (Glassow, 1996). Over the period of 5500 BC to AD 1250, stature
reduced by about 7 cm (Walker & Thornton, 2002; and see Lambert, 1993, and compare with
Temple, 2008). Height reduction was fostered by decline in health due to the combined effects
of declining nutrition and elevated infectious disease. Protein, mostly derived from fish, was
abundant, but other important nutrients may have been lacking in the diets of later prehistoric
populations. During the latest prehistoric period (post-AD 1250), there was a rebound in
stature, possibly due to improved living conditions.
Similar trends in stature reduction have been documented in the Central Valley of interior
California (Ivanhoe, 1995; Ivanhoe and Chu, 1996). Comparisons of populations spanning the
period of 3000 BC to the mid-nineteenth century reveal statistically significant reductions in
stature for both females and males (2.2% and 3.1%, respectively). Archaeological evidence
indicates an increased reliance on acorns. Thus, stature reductions more likely reflect nutri-
tional stress owing to a focus on carbohydrates and a narrowing of the dietary spectrum in
later prehistory in this setting.
While the foraging-to-farming transition shows a general pattern of stature reduction
(Mummert et al., 2011), stature reductions identified in archaeological contexts are not
universal. A number of regions show either no change, an increase, or a high degree of
regional variability in stature (Danforth, 1994; Douglas & Pietrusesky, 2007; Roberts & Cox,
2007; Temple, 2011). In the lower Illinois River valley, there is no clear trend of stature change
in the comparison of early prehistoric through late prehistoric periods (Cook, 1984). This is
especially significant because it indicates that reduced juvenile height in this setting did not
result in reduced adult stature in later prehistoric agricultural groups. Likewise, temporal
comparisons of stature in a diversity of archaeological populations – from Ontario, northern
Great Plains, Peru, and Chile – show no change in stature with the shift in adaptive strategies
involving agriculture (Allison, 1984; various in Cohen & Crane-Kramer, 2007; Cole, 1994;
Katzenberg, 1992; Steckel & Rose, 2002). From c. 8250 yBP to the colonial period in Ecuador,
there is no evidence of stature decline despite increases in physiological stress (Ubelaker,
1994). All groups in Ecuador are relatively short-statured, thus stress (including poor nutri-
tion) may have been severe throughout the entire sequence (Ubelaker, 1994).
The influence of nutritional deprivation on human growth and terminal height and/or long
bone length is revealed in the study of components of past groups that may have been
differentially buffered against stress. Comparison of elite and non-elite adults from Old World
settings shows evidence of greater height in elites than in non-elites (Angel, 1975, 1984;
Becker, 1993; Buzon, 2006; Vercellotti et al., 2011). In a Maitas-Chiribaya (c. 2000 yBP)
population from northern Chile, shaman males are taller than other, non-elite males, which
may indicate better health and resources in the former (Allison, 1984). High-status adult males
in some Mesoamerican populations appear to be taller than low-status individuals or the
general population (Haviland, 1967; Helmuth & Pendergast, 1986–1987; Storey, 1998; but see
Wilkinson & Norelli, 1981). Likewise, elite males are taller than non-elite adults in several
contexts in prehistoric Southeast, Midwest, and Southwest United States (Buikstra, 1976a;
Cook, 1984; Hatch and Willey, 1974; Malville, 2008; Powell, 1988). These apparent status
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20 Stress and deprivation during growth and development and adulthood
differences in attained height suggest that elites may have had nutritional advantages such as
having greater access to animal protein, resulting in achieving greater height than non-elite
individuals (Malville, 2008). Many of the elite distinctions are in adult males, suggesting that
the burden of stress may be primarily on adult males in ranked societies, at least as it is
exhibited in attained height.
This discussion points to the critical importance of understanding body size in relation to
resource availability and social context. Analysis of elite and non-elite individuals from San
Michele’s church in Trino Vercellese, Italy (c. AD 750–1300) shows considerable biological
variation in stature – high-status adults are taller than low-status adults – that is clearly
linked to relative social position in this Medieval setting (Porro et al., 1999). Additional
analysis of stature, body mass, and body proportions in this series reveals significant differ-
ences in high-status adults (Vercellotti et al., 2011). In particular, the comparison of different
body segments revealed significant differences between low- and high-status males. In
particular, males’ distal lower limb and body mass are significantly different in high- versus
low-status individuals. High-status men had longer distal lower limbs and were taller than
low-status men. However, the taller, high-status men had the shortest relative size of the
lower limb, indicating that height is not necessarily influenced by limb length. Rather, other
factors such as body trunk size may have a more important influence on height, at least for
members of a population having the most positive living conditions. The association between
stature and limb length in males is not shared by females in this setting. This may be due to
greater susceptibility to growth disruption in males than in females. The argument of greater
buffering in females than in males is also supported by the presence of greater prevalence of
enamel hypoplasias in men, especially low-status men, than in women in this setting.
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2.3 Growth and development: skeletal 21
The link between cranial base height and nutritional quality may be more apparent than
real, however. Cranial base cartilages, like epiphyseal cartilages of limb bones, are primary
cartilages. Primary cartilages have intrinsic growth capabilities that are characteristically
resistant to compressive loading. This suggests that a model invoking compression as a causal
factor in determining cranial base form is incorrect. Therefore, the phenomenon of cranial
base flattening, while interesting, is largely unexplained.
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22 Stress and deprivation during growth and development and adulthood
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2.3 Growth and development: skeletal 23
Figure 2.5 Rickets in left juvenile (four to five years old) ulna (left) and radius (right);
Medieval period, Gruczno, Poland. (From Kozłowski, 2012; photograph by Tomasz
Kozłowski.) (A black and white version of this figure will appear in some formats. For the
color version, please refer to the plate section.)
identified. This appears to be associated with a segment of a late prehistoric society that had
experienced relatively elevated levels of physiological stress generally (Palkovich, 2008).
Flattening of femoral and tibial diaphyses has been documented in numerous archaeo-
logical skeletal samples worldwide (see Chapter 6). The primary indices measuring the degree
of flatness of femora and tibiae include the meric index (anterioposterior flattening of
the proximal femoral diaphysis), pilasteric index (mediolateral flattening of the femoral
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24 Stress and deprivation during growth and development and adulthood
diaphysis), and cnemic index (mediolateral flattening of the tibial diaphysis at the nutrient
foramen). Some attribute diaphyseal flattening to nutritional stress (Adams, 1969; Angel,
1984; Buxton, 1938). Buxton (1938) asserted that less bone is required in the construction of
a diaphysis if it is flattened rather than round. He viewed the temporal trend of rounder
diaphyses as representing an increase in amount of bone, inferring a decline in nutritional
deficiency in recent “civilized” populations. Structural analysis of long bone diaphyses
reveals that flattening is related not to the amount of bone present, rather to the manner
in which it is distributed when viewed in cross-section (Ruff, 2008). Mechanical loading, not
nutritional stress, is the primary determinant of flatness of long bone diaphyses (see
Chapter 6). Nutritional deprivation or other physiological stressors certainly have an influ-
ence on amount of bone, but the relationship between nutritional status and diaphyseal
shape is unsubstantiated.
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2.4 Growth and development: dental 25
17 – 25 years have a significantly smaller vertebral neural canal size than older adults (Watts,
2011). This size variation suggests a link between skeletal development and conditions that
predisposed them to earlier death. Simply, individuals stressed during the juvenile years of
growth and development are likely to die earlier than individuals who are either not stressed
or less stressed.
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26 Stress and deprivation during growth and development and adulthood
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2.4 Growth and development: dental 27
Table 2.2 Juvenile and adult permanent tooth size (buccolingual; mm) from Santa Catalina de
Guale, St. Catherines Island, Georgia (Adapted from Simpson et al., 1990; Table 5-1)
Juvenile Permanent
population has been tested by the comparison of permanent tooth size of juveniles and adults
in different settings in the American Southeast, namely in the late prehistoric Averbuch series
from the Middle Tennessee River valley (Guagliardo, 1982a) and colonial Spanish missions
(Simpson et al., 1990; Stojanowski et al., 2007). Both populations were sedentary maize
agriculturalists exhibiting skeletal evidence of high levels of physiological stress and poor
health. In these settings, juveniles have smaller permanent teeth than adults. In the Santa
Catalina series, nine of 14 tooth types examined are smaller in juveniles than in adults
(Table 2.2). The other tooth types show either no difference or slightly smaller size in adults.
All statistically significant differences are for smaller juvenile teeth. In these samples, juvenile
– adult size differences are more common in mandibular teeth than in maxillary teeth,
suggesting that the lower dentition may be more developmentally sensitive to stress than is
the upper dentition.
These studies indicate the failure of teeth to reach their growth potential in circumstances
involving increased stress. This conclusion lends support for Sagne’s (1976) hypothesis that
the Medieval-era Swedish dying young received suboptimal nutrition during the years of
dental development, resulting in smaller teeth (and see Lunt, 1969). These investigations
suggest that individuals with small teeth – those who were most stressed – had a reduced
lifespan, which is consistent with evidence based on vertebral neural canal size, height
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28 Stress and deprivation during growth and development and adulthood
(Kemkes-Grottenthaler, 2005), and other dental indicators (e.g., enamel defects: Goodman &
Armelagos, 1988). As with neural arch canal size and height, it is unlikely that small teeth led
to reduced longevity. Rather, size reduction is symptomatic of environmental stress that
contributed to smaller teeth and early death.
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2.4 Growth and development: dental 29
Kieser and coworkers (1986a, 1986b; and see Kieser, 1990) analyzed fluctuating dental
asymmetry as an indicator of environmental disruption in highly stressed Lengua Indians
presently inhabiting the Chaco region of Paraguay. Application of Euclidean map analysis,
a statistically powerful approach whereby each dentition is considered an independent
variable, produces a measure of asymmetry represented by dividing the sum of Euclidean
distances for tooth antimeres by a product of the mean individual tooth size and the
number of tooth pairs. Comparisons with well-nourished, disease-free Whites reveals much
greater asymmetry in the Lengua population. Younger, more acculturated Lengua with
better diets and greater access to Western healthcare show lower asymmetry values than
those found in more traditional Lengua, experiencing elevated stress. Using the same
methodology, Townsend and Farmer (1998) determined asymmetry scores in a sample of
South Australian children. Most children are healthy, and have correspondingly low
asymmetry scores. A few individuals with low birth weight have relatively high asymmetry
scores.
Similar patterns of fluctuating asymmetry as a biological indicator of developmental
instability have been documented in other hard tissues. Based on the earlier finding that
environmental stress was likely higher in the early Christian period than in the late Christian
period in Nubia (and see earlier), DeLeon (2007) tested the hypothesis that there is a decrease
in fluctuating asymmetry in the craniofacial skeleton. Her analysis of landmark coordinate
data revealed a pronounced decrease in level of fluctuating asymmetry in the later popula-
tion compared to the earlier population (Figure 2.6). Interestingly, the lateral landmarks
show more asymmetry than that seen in the medial landmarks, supporting the notion that
developmental instability is trait specific and not random. Similarly, epiphyseal union in the
earlier, more highly stressed population shows considerably more bilateral asymmetry than
that found in the later population (Albert & Greene, 1999), also consistent with other
bioarchaeological evidence for decline in stress in the temporal sequence in Nubia. The
presence of fluctuating asymmetry in dental and craniofacial tissues would suggest the
likelihood of poor health outcomes in adults. Indeed, analysis of nineteenth- and early
twentieth-century remains from Portugal, having known cause-of-death, reveal that adults
who died from degenerative diseases (e.g., chronic heart disease, diabetes) had higher rates
of fluctuating asymmetry than those with other causes of death (e.g., infectious disease)
(Weisensee, 2013).
Directional asymmetry is another pattern of bilateral variation that has been identified in
analysis of tooth size in human populations. This pattern is characterized by larger teeth on
one side of the dental arch than on the other. Directional asymmetry is infrequently reported
for human populations (Ben-David et al., 1992; Boklage, 1987; Harris, 1992; Lundström,
1948; Mizoguchi, 1986; Sharma et al., 1986; Townsend & Farmer, 1998). Harris (1992)
detected directional asymmetry in a large sample of permanent teeth of Euroamerican
adolescents, with a consistently greater left dominance in one dental arch.
Directional asymmetry is unexplained by current models, but may be an indicator of
developmental instability arising from stress (Harris, 1992). The strong environmental basis
of directional (and fluctuating) asymmetry is inferred by observation of low intraclass
correlations between monozygous and dizygous twins (Townsend, 1992). Additionally,
detection of spurious genetic variance indicates a virtual lack of evidence for a significant
genetic basis.
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30 Stress and deprivation during growth and development and adulthood
LAM
PTP
NAS
AST
GPF
LAM
PTP
NAS
AST
GPF
NAS
PTP
AST
LAM
Figure 2.6 Fluctuating asymmetry observed in Early and Late Period Kulubnarti, Sudan.
More fluctuating asymmetry is observed in the Early Period sample, indicating elevated
physiological stress. (From DeLeon, 2007; reproduced with permission of author and John
Wiley & Sons, Inc.)
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2.5 Skeletal and dental pathological markers of deprivation 31
oxygen. The outcome of such a deficiency and associated compromised access to oxygen is
reduced physical activity in general and work performance in particular (Crouter et al., 2012;
Haas, 2006).
There are three causes of anemia – blood loss, impaired production of red blood cells,
increased red blood cell production, or any combination of these conditions. Some of the
anemias are genetic, such as thalassemia, sickle cell anemia, nonspherocytic hemolytic
anemia (e.g., glucose-6-phosphate dehydrogenase deficiency [favism], pyruvate kinase defi-
ciency), spherocytosis, and rarely, hereditary elliptocytosis. In living and past populations, the
majority of the anemias are acquired, caused by either blood loss or nutritional deficiencies.
There are regions of the world, however, where genetic anemias are highly prevalent, reflect-
ing natural selection in areas where malaria is endemic. Angel (1966a) made the case that
pathological conditions he documented in the Mediterranean basin were largely genetic in
origin. DNA sequence analysis from an eight-year-old from the site of el Akhziv, Israel,
revealed that the child had homozygosity for ß-thalassemia (Filon et al.,1995), thus providing
some support for Angel’s hypothesis. However, it now seems unlikely that the record of
cranial porosities in the region is largely due to genetic causes (and see later).
In the normal individual, the rate of red blood cell production is balanced by the rate of red
blood cell destruction. The balance between production and destruction of red blood cells
requires a number of key micronutrients, including especially the essential amino acids, iron,
and vitamins A, B12, B6, and folic acid (Martini & Ober, 2001; Walker et al., 2009). Iron is an
essential element of hemoglobin, thus serving as a key component of oxygen transport. Iron
deficiency is the most common cause of anemia, owing to blood loss, iron-poor diets, and
gastrointestinal malabsorption of the element.
The absorption of iron is dependent upon its source, from either heme or non-heme foods.
Generally, heme sources of iron are efficiently absorbed, with meat being among the best
(Baynes & Bothwell, 1990). Iron in meat does not require processing in the stomach, and the
amino acids derived from digestion of meat help to enhance iron absorption (Wadsworth,
1992). Iron bioavailability of non-heme sources is highly variable, but plant sources are
generally poorly absorbed. Various substances found in plants inhibit iron absorption, such
as phytates in many nuts (e.g., almonds, walnuts), cereals (e.g., maize, rice, whole wheat
flour), and legumes (Baynes & Bothwell, 1990). Unlike protein in meat, plant proteins (e.g., in
soybeans, nuts, lupines) inhibit iron absorption. Tannates found in tea and coffee also reduce
iron absorption significantly (Hallberg, 1981).
A number of foods enhance iron bioavailability, especially when consumed in combination
with non-heme iron. For example, ascorbic acid promotes iron absorption (Baynes &
Bothwell, 1990; Hallberg, 1981; Wadsworth, 1992). Citric acid from various fruits and lactic
acid from fermented cereal bees are implicated in promoting iron absorption (Baynes &
Bothwell, 1990). Non-heme iron (e.g., in maize) is enhanced considerably by concurrent
consumption with meat and fish (Layrisse et al., 1968; Navas-Carretero et al., 2008).
If either absorption or consumption of iron or the other micronutrients is low, red blood cell
production is reduced, resulting in anemia. In response to anemia, the body first steps up its
production of red blood cells (Stockmann & Fandrey, 2006). If this first response fails in
alleviating the shortfall, the regions of the skeleton responsible for red blood cell production
are activated, resulting in an expansion of the cranial vault marrow (diploë). This elevated red
blood cell production occurs at the expense of the ectocranial surfaces, resulting in resorption
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32 Stress and deprivation during growth and development and adulthood
(a)
Figure 2.7a Porotic hyperostosis on frontal and parietals; Santa Maria de los Yamassee, Amelia
Island, Florida. (Photograph by Mark C. Griffin.) (A black and white version of this figure
will appear in some formats. For the color version, please refer to the plate section.)
of the bone and a characteristic porosity called porotic hyperostosis, especially involving
the cranial vault bones (Figure 2.7).
Marrow expansion, therefore, is clearly caused by anemia as a response to deficiencies in
red blood cells and/or hemoglobin (Moseley, 1974; Ortner, 2003; Schultz, 2001). However,
only some forms of anemia result in the characteristic pathognomic porotic lesions of the
cranial vault. That is, only the anemias that can sustain high and elevated production of red
blood cells likely cause porotic hyperostosis. Walker and coworkers (2009) argue that iron
deficiency anemia per se results in decreased production of red blood cells, anemia certainly,
but not the kind of anemia that could produce cranial porosities. That is, iron-deficiency
anemia is a condition of reduced and inadequate blood production and is insufficient, thus
resulting in the kind of marrow hypertrophy associated with porotic hyperostosis and cribra
orbitalia. Specifically, two forms of anemia lead to elevated red blood cell production and
cranial marrow expansion resulting in porotic hyperostosis, namely megaloblastic and hemo-
lytic anemias. Megaloblastic anemia is commonly caused by chronic nutritional deficiencies
and malabsorption of vitamin B12 or folic acid, or both. In both forms of anemia, there is an
overproduction of red blood cells, resulting in expansion of the marrow (Walker et al., 2009).
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2.5 Skeletal and dental pathological markers of deprivation 33
(b)
Figure 2.7b Histological section from Figure 2.7a. The linear orientation and general
morphology of diploic cavities are consistent with iron deficiency anemia. (Schultz et al.,
2001; reproduced with permission of authors and University Press of Florida.) (A black and
white version of this figure will appear in some formats. For the color version, please refer
to the plate section.)
Similar lesions found in the roof areas of the eye orbits, called cribra orbitalia, are also
frequently observed in archaeological remains (Figure 2.8). There is an extensive body of
bioarchaeological research documenting porotic hyperostosis, largely beginning with Angel’s
(1966a, 1967, 1971a, 1978b, 1984) systematic study of a large series of skeletal remains on a
regional and population basis. Based on his study of some 2200 archaeological crania from
the eastern Mediterranean region, principally Greece, Cyprus, and Turkey, Angel proposed
that porotic hyperostosis resulted from hereditary hemolytic anemias, especially thalassemia
or sickle cell anemia. In this setting where malaria is endemic, individuals who are heterozy-
gous for sickle cell anemia or thalassemia have a selective advantage over normal homozy-
gous individuals lacking the sickle-cell or thalassemia genes. Carriers show lower infection
rates by malarial parasites (genus Plasmodium), thus enjoying greater protection from
malaria.
Other regional studies dealing with large samples of skeletal remains showed that cranial
porosities in past populations are likely due to a variety of nongenetic factors. In Wadi Halfa,
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34 Stress and deprivation during growth and development and adulthood
Figure 2.8 Cribra orbitalia; Santa Catalina de Guale de Santa Maria, Amelia Island, Florida.
(From Larsen, 1994; photograph by Mark C. Griffin; reproduced with permission of John
Wiley & Sons, Inc.) (A black and white version of this figure will appear in some formats.
For the color version, please refer to the plate section.)
Nubia, in the Nile Valley, high prevalence of orbital lesions (21.4%) for the Meroitic (350 BC –
AD 350), X-group (AD 350 – 550), and Medieval Christian (AD 550 – 1400) periods have been
reported (Carlson et al., 1974). Reconstruction of the environmental context based on arch-
aeological, historical, and ethnographic evidence indicates that several factors likely contrib-
uted to anemia. Milled cereal grains (millet, wheat), the focus of diet in this setting, contain
very little iron and are high in phytate. Additionally, as with populations currently living in
the Nile Valley, hookworm disease and schistosomiasis were likely highly endemic. These
factors, combined with chronic diarrhea that is also highly prevalent in the region today, are
consistent with the presence of anemia.
Further to the south in the Nile Valley, a high prevalence of porotic lesions (45%) has been
reported at the Medieval-period Kulubnarti site (AD 550 – 1500) (Mittler & Van Gerven, 1994).
Early and late-period Kulubnarti juveniles have remarkably high prevalences (94% and 82%,
respectively: Van Gerven et al., 1995). Like the Nubian groups down river at Wadi Halfa, the
Kulubnarti population suffered the ill effects of anemia owing to nutritional deficiencies and
other negative influences of sedentism and unhealthy living conditions. Analysis of demo-
graphic profiles of individuals with and without lesions indicates that those with porotic
crania are associated with a shortened life expectancy, with differences greatest in the
subadult years (and see Blom et al., 2005). There is a decline in porotic lesion prevalence
from 51.8% to 39.0% from the early to late Christian periods (AD 550 – 750 to AD 750 –
1500). This apparent improvement in iron status coincides with improvements in health
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2.5 Skeletal and dental pathological markers of deprivation 35
generally that arose following increased political autonomy and improved living conditions
(Mittler & Van Gerven, 1994).
Circumstances involving association between anemia and increasing or elevated stress
have been documented in Medieval and seventeenth-century Tokyo (Hirata, 1990), prehistoric
Iran and Iraq (Rathbun, 1984), third-century BC Carthage (Fornaciari et al., 1981), Neolithic
Levant and Greece (Eshed et al., 2010; Papathanasiou, 2005; Papathanasiou et al., 2000),
third-century AD Moldavia (Miritoiu, 1992), and Romano-British, Medieval, and eighteenth–
nineteenth-century Britain (Grauer, 1993; King et al., 2005; Lewis, 2002; Molleson & Cox,
1993; Stuart-Macadam, 1991; Sullivan, 2005; Wells, 1982). All of these settings have good
contextual evidence for elevated environmental stress, but the particular circumstances for
anemia are regional specific. For example, causative factors for high prevalence of porotic
lesions in the Roman-period Poundbury Camp, and likely in British populations, included
parasitism, infectious disease, and perhaps lead poisoning (Stuart-Macadam, 1991). High
prevalences in eighteenth–nineteenth-century urban London appear to be linked with shifts
in weaning practices, poor living conditions, and low maternal health status (Lewis, 2002;
Molleson & Cox, 1993). Indeed, comparisons of temporal series in England reveal that
industrialization had the highest negative impact on children’s health, more so than events
in any other period (Lewis, 2002). On the other hand, improved environments result in the
decline of prevalence of porotic lesions. For example, decrease in prevalence in modern Japan
reflects decreased crowding, reduction in infectious diseases, and improved hygiene (Hirata,
1990; Temple, 2010).
The most abundant data on porotic lesions is available from the New World, especially
North America. In the American Southwest, cranial porosities are highly prevalent, especially
in late prehistoric and historic-era populations (Akins, 1986; El-Najjar & Robertson, 1976; El-
Najjar et al., 1975, 1976, 1982; Hooton, 1930; Kent, 1986; Martin et al., 2001; Palkovich,
1980, 1987; Stodder, 1994, 2006; Stodder & Martin, 1992; Stodder et al., 2002; Walker, 1985;
Walker et al., 2009; Zaino, 1967, 1968). Among mostly late prehistoric Puebloan samples
studied by El-Najjar and collaborators (El-Najjar et al., 1976) from Canyon de Chelly, Chaco
Canyon, Inscription House, Navajo Reservoir, and Gran Quivira, porotic lesions were docu-
mented in 34.3% of individuals. At Chaco Canyon alone, some 71.8% of individuals display
the characteristic lesions. Similarly, high prevalences have been reported from late prehis-
toric- and contact-period sites, including San Cristobal (90%), Hawikku (84%), Black Mesa
(88%), Mesa Verde (70%), Dolores (82%), Cases Grandes (46%), and La Plata Valley (40%)
(Stodder, 1994; Weaver, 1985). There are some Southwestern samples that have relatively low
prevalences (e.g., 16% for Navajo Reservoir children; see Martin et al., 1991). Martin and
coworkers (1991) note that comparisons of data collected by different researchers is problem-
atical because of the varying methods used in identification and recording of porotic lesions.
For example, some researchers may include slight pitting when analyzing their data sets,
whereas others may not. Unfortunately, this distinction is only rarely noted in bioarchaeo-
logical reports, regardless of geographic or cultural setting.
El-Najjar (1976) links the elevated levels of abnormal cranial porosities in the American
Southwest and other regions of the New World to the effects of over-reliance on maize in
conjunction with food processing techniques that may contribute to iron deficiency. Specif-
ically, he regards the presence of phytate – an iron inhibitor – as well as lime treatment to
decrease the nutritional value of maize.
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36 Stress and deprivation during growth and development and adulthood
Analysis of archaeological samples from other maize agriculturalists in the New World
provides mixed support for El-Najjar’s dietary hypothesis. Relatively high prevalences of
porotic lesions (>15% – 20%) are present in agriculturalists in the American Midwest (Cook,
1984; Garner, 1991; Goodman, Lallo et al., 1984; Lallo et al., 1977; Milner, 1983, 1991;
Milner & Smith, 1990; Perzigian et al., 1984), Southeast (Boyd, 1986; Danforth et al., 2007;
Eisenberg, 1991a, 1991b; Hutchinson, 2002, 2004; Lambert, 2000b; Larsen, 2006; Larsen,
Crosby et al., 2002; Larsen & Sering, 2000; Larsen et al., 2007; Parham & Scott, 1980; Rose
et al., 1984), and Northeast (Magennis, 1986; Pfeiffer & Fairgrieve, 1994), as well as a range of
other settings in Mesoamerica and South America (Blom et al., 2005; Cohen et al., 1994;
Hodges, 1989; Hooton, 1940; Hrdlička, 1914; Saul, 1972; Trinkaus, 1977; Ubelaker, 1984,
1992a, 2002; White et al., 1994; Wright, 2006). For some regions where skeletal remains of
foragers (or less intensive agriculturalists) have been compared with agriculturalists, there are
clear temporal increases in porotic lesion prevalence (Cook, 1984; Lallo et al., 1977; Perzigian
et al., 1984; Rose et al., 1984; but see Hodges, 1989). However, skeletal series from large, late
prehistoric Mississippian centers in the American Southeast (Blakely, 1980; Larsen et al.,
2007; Powell, 1988, 1989), contact-era part-time maize agriculturalists in the Great Plains
(Miller, 1995), a large urban center in Mesoamerica (Storey, 1992a), and the coastal desert of
Peru and Chile (Allison, 1984) all display low prevalences of cranial porosities.
The dietary hypothesis presented by El-Najjar does not account for the relatively high
frequencies of porotic lesions in some foraging populations. A number of Pacific coastal
foraging groups with access to iron-rich marine resources have high prevalences of cranial
porosities. Moderate levels of porotic lesions are present in precontact and contact-era
Northwest coast populations (Cybulski, 1977, 1992, 1994, 2006; Keenleyside, 1998, 2006).
In this setting, European-introduced diseases may have prevented adequate iron metabolism
during the contact period (Cybulski, 1994). Presence of porotic lesions prior to contact
indicates that there may have been other important factors, such as blood loss and parasitism
(Cybulski, 1994, 2006).
Late prehistoric foragers from the islands and mainland of the Santa Barbara Channel
Island region of California have higher prevalences than earlier foragers, increasing from
12.8% in the Early period to 32.1% in the Late Middle period (Lambert, 1994; Lambert &
Walker, 1991; Walker, 1986, 2006). Late period populations living on islands located furthest
from the mainland coast have an extraordinarily high prevalence of porotic lesions (73.1% on
San Miguel Island). Walker and Lambert suggest that water contamination explains the
elevated prevalence of the condition. High prevalence of porotic lesions in island populations
coincides with a period of increasing sedentism and population size and a shift from terrestrial
to marine diets. In the Late period, groups became concentrated around a limited number of
water sources. As a result, diarrhea-causing enteric bacteria may have contaminated these
water sources. Ethnographic evidence indicates that island populations preferred eating raw
(versus cooked) fish (Walker, 1986), thus also increasing their chances of acquiring parasitic
infections.
Prevalences of porotic lesions in prehistoric Australian foragers are consistently high in
tropical/subtropical environments and low in desert environments (Webb, 1995). For
example, in southeastern Australia, prevalences range from 62.5% (<21 years) in the Rufus
Valley to 30.0% (<21 years) in the desert. Half of the juvenile crania from the tropics of
northeastern Australia are porotic. Various factors appear to have contributed to iron
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2.5 Skeletal and dental pathological markers of deprivation 37
deficiency anemia in Australia, but parasitism is primary. The Murray Valley, southeastern
coast, and tropics provide well-suited environments for support of various intestinal parasitic
organisms, including Trichuris trichuris, Ascaris lumbricoides, Strongyloides stercoralis, and
Enterobius vermicularis. In the tropics, hookworm infection may have been a principal cause.
In living populations occupying the tropics of Australia, some 40% of children are infected
with this helminth. Although it is unknown whether hookworm parasites were present in this
region prior to contact by Europeans (in 1788), had they been, they would have caused the
same types of health problems seen in living groups today.
A limited number of settings in South America show high or temporally increased preva-
lence of porotic hyperostosis and/or cribra orbitalia (Blom et al., 2005; Hrdlička, 1914; Klaus,
Centurion et al., 2010; Klaus & Tam, 2009; Shimada et al., 2004; Ubelaker, 1981, 1992a).
Porotic lesions tend to be low in prevalence in mountainous regions, and appear to be
restricted primarily to late prehistoric and contact-era coastal or near-coastal occupations.
The penchant for coastal settings may reflect a more restricted access by native populations to
fresh, parasite-free water sources in these areas. Ubelaker (1992a) contends that the coastal
pattern of elevated porotic lesion prevalence fits the model of increased anemia due to chronic
helminth disease brought about by population crowding and reduced hygiene. In present-day
Ecuador, hookworm disease is a common and major public health problem in coastal regions.
This distribution in contemporary populations, along with the pattern of prehistoric porotic
lesions, strongly implicates parasitism as a causal factor in northwestern South America.
Moreover, in the contact period of the Lambayeque River valley of the north coast of Peru,
local circumstances during the colonial period, including increased population crowding and
generally deteriorating conditions, provided the necessary conditions leading to reduced
quality of life, including elevated levels of anemia (Klaus & Tam, 2009).
Several investigations suggest that increased prevalence of cranial porosities may be due to
highly localized factors. Lesions have been evaluated in prehistoric and contact-era popula-
tions that inhabited the southeastern US Atlantic coast of Georgia and northern Florida
(Larsen, Crosby et al., 2002; Larsen et al., 2007; Schultz et al., 2001). Maize agriculture –
introduced during the twelfth century – played an important role in changes in health in
native populations, including increased prevalence of nonspecific infections due to popula-
tion aggregation along with other evidence of increased stress (see later). However, compari-
sons of prehistoric foragers and farmers show low prevalences of porotic lesions, 6.5% and
6.2%, respectively. Marine resources contributed significantly to diets in both foragers and
farmers, which may have enhanced iron absorption in these groups. In the contact period,
both the early mission population living on St. Catherines Island, Georgia (Santa Catalina de
Guale; AD 1607 – 1680), and their descendants on Amelia Island, Florida (Santa Catalina de
Guale de Santa Maria; AD 1686 – 1702), have considerably higher prevalences of porotic
lesions, 26.5% and 27.2%, respectively. Archaeological and bone isotope evidence indicates
an increase in maize consumption in the mission groups, and reduction in marine sources of
protein.
The dramatic increase in prevalence of cranial porosities in contact-era coastal Georgia and
Florida populations may be related to similar conditions documented in the Santa Barbara
Channel Islands region. At missions in Spanish Florida, limited and easily contaminated water
sources – wells located next to the settlement – served as primary water sources. Wells in
this subtropical setting are highly susceptible to contamination by parasites and microbes that
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38 Stress and deprivation during growth and development and adulthood
cause diarrheal infections. During the mission period on St. Catherines Island, a freshwater
stream bordering the mission/village was artificially dammed and also used as a principal
water source (Larsen, Ruff et al., 1992). An abundance of archaeological refuse deposits –
mostly food remains – surrounds and intrudes into this water source today. The accumulation
of refuse during the mission period likely contributed to water contamination, thus also
providing an important source of infections potentially leading to anemia. The general
increase in concentration of population and sedentism during the mission period undoubtedly
fostered poor sanitation and living conditions (Larsen, Crosby et al., 2002). Clearly, anemia is
a likely cause of the elevated prevalence of the lesions. Indeed, histological examination of
lesions reveals morphological characteristics of the diploë consistent with this diagnosis
(Schultz et al., 2001). Moreover, these lesions may have been the result of megaloblastic
anemia caused, at least in part, by deficiencies in vitamin B12 in nursing mothers due to
reduced availability of animal food sources (the only source of vitamin B12) and increased
consumption of maize in the mission setting. Infants born to these mothers have low reserves
of the vitamin. Vitamin B12 is low in the breast milk and gastrointestinal infections and
diarrheal disease associated with poor living conditions would have exacerbated the condi-
tion, increasing the likelihood of anemia.
Data generated in the study of archaeological human skeletons worldwide indicate that the
etiology of cranial porotic lesions can only be understood in relation to multiple stressors.
Although common factors are likely present in many regions (e.g., parasitism, poor diets,
decreased sanitation), these studies also demonstrate that behavioral circumstances unique to
particular settings must be considered when interpreting porotic lesion prevalence. Much
more information is needed on details and circumstances regarding living conditions and
lifestyle (e.g., trash disposal, household and settlement size, dietary practices, food preparation
techniques). Other classes of pathological data need to be considered in understanding health
patterns potentially influencing circumstances resulting in anemia. For example, in at least
one setting, individuals having anemia-related pathology and tuberculosis showed higher
childhood mortality in circumstances where parasitism was minimal (Blom et al., 2005).
Variation in prevalence of cranial porotic lesions in human populations should inform our
understanding of the differential costs of disease stress in past societies. In some settings,
there is a consistently higher prevalence of porotic lesions in adult women than in adult men
(Dickel, 1991; Webb, 1995). For Australia, Webb (1995) suggests that a higher prevalence in
women than in men reflects the stresses of “[c]hildbearing, lactation, menstruation and the
imposition of food taboos.” Given that the pathological condition largely reflects childhood
anemia, it seems unlikely that parity status, lactation, or menstruation can explain variability
in porotic lesion prevalences. These differences suggest, however, that the growth period may
have involved greater anemia stresses in female children than in male children.
Comparison of porotic lesion prevalence across social ranks in prehistoric stratified soci-
eties suggests some differences in iron deficiency anemia. Elite individuals in several settings
in North America and South America have a lower prevalence of porotic lesions than non-
elite individuals. This difference has been reported at prehistoric Mississippian localities from
Moundville, Alabama (2.5%, elite; 9.9%, non-elite) and at Toqua, Tennessee (5%, mound
burial; 21%, village burial) (Parham & Scott, 1980; Powell, 1988, 1992a). At Mound 72 in the
late prehistoric Cahokia site, Illinois, high-status individuals lacked lesions, whereas 12.5% of
low-status individuals – female sacrificial victims – have the condition (Fowler et al., 1999).
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2.5 Skeletal and dental pathological markers of deprivation 39
Similarly, higher-status individuals (especially women) have lower prevalence than lower-
status people documented in the Middle Sicán burials in late prehistoric Peru (Shimada et al.,
2004), and most elite individuals at Copan, Honduras, have no lesions, whereas most of the
non-elite have lesions (Storey, 1998). Overall, porotic hyperostosis in this setting is nearly five
times greater in non-elite than in elite individuals. These differences suggest, therefore, that
high-status individuals may have been buffered against stressors that result in the patho-
logical condition, consistent with the notion that political and socioeconomic inequality is
linked to stress and quality of life.
In summary, a consensus has emerged that porotic hyperostosis and cribra orbitalia in
archaeological skeletal samples are due to anemia in the vast majority of cases. Genetic
anemias may have contributed to elevated porotic lesion prevalence in the past, but it is
unlikely that they would have occurred in appreciable frequencies. Moreover, significant
presence of porotic lesions in areas of the world where genetic anemias (e.g., sickle cell
anemia, thalassemia) did not occur prior to contact by Europeans – such as Australia and
the Americas – can be explained only by negative environmental factors. Moreover, only
marrow hypertrophy activated by overproduction of red blood cells, a condition associated
with megaloblastic or hemolytic anemias, will cause porotic hyperostosis and cribra orbitalia,
not iron-deficiency anemia.
Angel argued that the etiology for porotic hyperostosis and cribra orbitalia is the same,
resulting in him conflating the two terms into porotic hyperostosis (and see Blom et al., 2005;
Stuart-Macadam, 1989). However, it is becoming increasingly clear that while porotic hyper-
ostosis affecting the flat bones of the cranial vault is generally caused by anemia and is
associated with hypertrophy of the cranial vault marrow, there are a large number of cases
where cribra orbitalia is associated with subperiosteal inflammation and not marrow hyper-
trophy. For example, Wapler and coworkers (2004) found that 26% of cribra orbitalia is
associated with inflammation and not marrow expansion. That is, other pathological pro-
cesses cause circumstances resulting in inflammation, such as vitamin C and D deficiencies
(scurvy and rickets, respectively), and trauma-related hematomas. The hematomas become
plaques of new, vascular subperiosteal bone that superficially look like cribra orbitalia
associated with marrow hypertrophy (Brickley & Ives, 2006, 2008; Schultz, 2001; Sloan
et al., 1999; Walker et al., 2009). This process is common in childhood, a period of life when
the periosteum is not well attached (Ma’luf et al., 2002). These non-anemia lesions of the eye
orbits are associated with scurvy (Brickley & Ives, 2008; Halcrow et al., 2014; Klaus, 2014b;
Ortner et al., 1999), owing to the fact that vitamin C (ascorbic acid) deficiency renders the eye
orbit prone to minor trauma due to normal movement of the ocular muscles. As a result, there
is a detachment of the periosteum in the superior orbital roof and associated bleeding and
formation of new, highly vascularized bone. Similar kinds of porosities have been well
documented in other cranial areas such as the sphenoid, alveolar bone, and palatine processes
where blood vessels are close to skin surfaces and are subject to greater bleeding, especially in
association with scorbutic episodes (Figure 2.9a, b) (Geber & Murphy, 2012; Mays, 2008;
Klaus, 2014b; Ortner et al., 2001). These porotic lesions and bony plaques also co-occur on
long bones, and like the eye orbits, are bilateral (Lovász et al., 2014).
Cribra orbitalia and porotic hyperostosis lesion formation are highly age-specific
(Figure 2.10). Although the two conditions are present in older juveniles and adults in
archaeological remains, most individuals with active, unhealed lesions are young juveniles
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40 Stress and deprivation during growth and development and adulthood
(a)
(b)
Figure 2.9 Abnormal bone growth and porosity in glabellar region (a) and on greater wing
of sphenoid (b), diagnostic of scurvy; Chornancap, Peru. (Klaus, 2014b; reproduced with
permission of author and Elsevier Ltd.) (A black and white version of this figure will appear
in some formats. For the color version, please refer to the plate section.)
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2.5 Skeletal and dental pathological markers of deprivation 41
45
40 Cribra Orbitalia
Porotic Hyperostosis
35
30
% Showing Condition
25
20
15
10
0
0 10 20 30 40 50 60 70
Age (years)
Figure 2.10 Age distribution of cribra orbitalia and porotic hyperostosis, from the History of
Health in the Western Hemisphere database (Steckel & Rose, 2002). (Data from Walker
et al., 2009; reproduced with permission of John Wiley & Sons, Inc.)
(less than five years of age) for both conditions, regardless of geographic or cultural circum-
stances (Blom et al., 2005; Fairgrieve & Molto, 2000; Lallo et al., 1977; Larsen, Ruff et al.,
1992; Mensforth et al., 1978; Milner & Smith, 1990; Miritoiu, 1992; Mittler & Van Gerven,
1994; Ribot & Roberts, 1996; Salvadei et al., 2001; Stodder & Martin, 1992; Walker, 1986;
Webb, 1995; Wright, 2006; and many others). This age pattern indicates that the lesions form
during childhood (Stuart-Macadam, 1985; Walker et al., 2009). With regard to porotic
hyperostosis in particular, red blood cell production occurs in the cranial marrow in juveniles
but not the yellow marrow that characterizes adolescents and adults.
There is considerable variation of orbital versus non-orbital (cranial vault) lesions across
human populations. In prehistoric Ecuador, most lesions are associated with the cranial vault
(Ubelaker, 1992a), but in native Australians and Arctic populations, the lesions are associated
largely with the eye orbits (Dabbs, 2011; Webb, 1995). Some argue that cribra orbitalia is an
earlier manifestation of anemia than porotic hyperostosis (Blom et al., 2005; Carlson et al.,
1974; Lallo et al., 1977). Walker and coworkers (2009; Wapler et al., 2004) make the case that
at least some cribra orbitalia have little to do with iron-deficiency anemia and that common
etiology with porotic hyperostosis is not necessarily the case.
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42 Stress and deprivation during growth and development and adulthood
Figure 2.11 Harris lines on juvenile tibia (left) and femur (right); anatomical specimens. The
dashed lines indicate the contours of the growth disruption. (From Garn et al., 1968;
reproduced with permission of Eastman-Kodak Company.)
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2.5 Skeletal and dental pathological markers of deprivation 43
peaking sometime during the first five years (Clarke, 1980; Clarke & Gindhart, 1981; Dreizen
et al., 1964).
These lines are commonly referred to as growth arrest lines or growth recovery lines
representing the recovery phase following growth arrest (Garn et al., 1968). The link with
stress episodes, however, remains unclear as a number of studies have shown no association
between Harris lines and illness (Papageorgopoulou et al., 2011), and experimental studies
show that more lines are associated with rapid growth in the absence of nutritional stress
(Alfonso-Durruty, 2011). When the epiphysis commences growth following the stress event,
mineralization at the growth plate continues in the absence of epiphyseal cartilage deposition.
The bilateral nature of transverse lines and association with growth zones (e.g., same location
in distal left and right tibiae: Garn & Baby, 1969; Grolleau-Raoux et al., 1997; McHenry,
1968) provide a strong argument for their systemic origin.
Another difficulty of using Harris lines for assessing stress is the high degree of frequency
variation in relation to individual health history. The study of living individuals with known
stress histories reveals the presence of numerous lines in clinically normal children (Garn
et al., 1968), and few lines in children who are well below weight-for-age (Walimbe &
Gambhir, 1994). These findings and the lack of a close association between transverse lines
and disease episodes in archaeological populations (Mensforth, 1981) and in living popula-
tions (Marshall, 1968) suggest that this stress indicator should be interpreted cautiously in
bioarchaeological analysis, especially when considering health status and its relationship to
specific behavioral, environmental, and dietary adaptations.
Analyses of transverse lines in archaeological remains are largely built on the assumption
that they represent stress events. Comparisons of transverse lines in early and late prehistoric
foragers in central California show a general decrease in frequency, possibly indicating
improved reliability of food sources (Dickel et al., 1984; McHenry, 1968). This argument runs
counter to an abundance of evidence drawn from the study of other pathological conditions
and stature showing an increase in stress in populations in the Central Valley (Ivanhoe, 1995)
and to the south in the Santa Barbara Channel Islands region (Lambert, 1993).
Similarly, other prehistoric foragers show variable frequencies in transverse lines. High
latitude Arctic populations express elevated prevalences (>30%–50%: Buikstra, 1976b; Lob-
dell, 1984, 1988; Steffian & Simon, 1994). In several of these settings where even spacing of
lines has been observed, regularity or periodicity of stress – perhaps on a seasonal basis – is
inferred (Buikstra, 1976b; Lobdell, 1988). The common occurrence of transverse lines in these
populations indicates that metabolic stress is more characteristic of Arctic lifeways than has
often been assumed (Steffian & Simon, 1994). In addition to nutritional deficits, other
stressors associated with this setting include the constant variation in the ratio of light to
dark, placing significant demands on the body, especially in children (Condon, 1983). Sea-
sonal or other changes in the intensity and duration of daylight lead to various changes in
health and mental functioning, especially in circumstances involving depletion of light. The
study of living Inuit in the central Canadian Arctic indicates that January is a peak time of
disease susceptibility, primarily because of extremely low temperatures, low ambient humid-
ity within and outside dwellings, and lowered sunlight (Condon, 1983). This may be exacer-
bated by the effects of desynchronization of natural physiological rhythms and lack of sleep.
Temporal comparisons of transverse lines in archaeological samples presents a mixed
picture, and sometimes contrary to expectation based on other stress indicators.
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44 Stress and deprivation during growth and development and adulthood
Comparisons of three successive periods at Dickson Mounds – Late Woodland (AD 950 –
1050), Mississippian Acculturated Late Woodland (AD 1050 – 1200), and Middle Mississippian
(AD 1200 – 1300) – reveal a decrease in tibial transverse lines (Goodman & Clark, 1981).
Similarly, lines decreased in frequency in the comparison of foragers and farmers in the Ohio
River valley (Cassidy, 1984; Perzigian et al., 1984) and in the Caddo region of the southeastern
United States (adjoining the states of Texas, Oklahoma, Arkansas, Louisiana) (Rose et al.,
1984). These trends suggesting a decrease in stress are puzzling because most other indicators
of morbidity show a highly consistent pattern of increase in stress and reduced health status
(e.g., enamel defects, nonspecific infection).
The use of Harris lines for documenting stress in past populations is clouded further by the
fact that lines have a tendency to fade or vanish with advancing age due to bone remodeling.
In a study of living populations comprised of individuals of known stress history, lines
showed a decrease in width with advancing age; some lines disappeared while others were
inexplicably retained well into adulthood (Garn & Schwager, 1967; Garn et al., 1968). This
very mixed record suggests that Harris lines may not reflect stress histories, at least with any
degree of clarity. Rather, these lines correspond with normal growth and normal growth spurts
and should not be regarded as a stress indicator (Alfonso-Durruty, 2011; Papageorgopoulou
et al., 2011).
Macrodefects
Contrary to Harris lines, there is a clear link between enamel defects and stress. Indeed,
virtually any environmental factor leading to metabolic disturbance will result in visible
changes in the structure of enamel. Ameloblasts are especially sensitive to even minor
physiological disruptions. Enamel defects arising from physiological perturbation have been
documented most frequently as visible alterations of the tooth surface, especially hypoplasias,
and to a lesser extent, hypocalcifications. Hypoplasias are measurable defects characterized as
deficiencies in the amount or thickness of enamel (Goodman & Rose, 1990; Hillson, 2008,
2014; Seow, 1991; Suckling, 1989). They vary in appearance from small pits or furrows to
large, deep grooves or even large areas of missing enamel. Typically, these defects are
horizontal grooves that are called chronologic or linear enamel hypoplasias (Figure 2.13).
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2.5 Skeletal and dental pathological markers of deprivation 45
(a) (b)
Enamel
Perikymata
Striae of
Retzius
(c)
Dentin
Occlusal border
of defect
Occlusal wall
perikymata
(stress episode)
Cervical/occlusal wall junction
cervical wall perikymata
Pulp (recovery period )
Cervical border
of defect
Figure 2.12 Cross-section of tooth displaying the relationship between striae of Retzius (a) and
perikymata (b). Episodes of stress disrupt enamel growth, which are followed by recovery periods (c).
The resultant enamel surface captures this period when little or no enamel matrix was formed in
the tooth. (From Blatt, 2013; Fejerskov & Thylstrup, 1986; Guatelli-Steinberg et al., 2004; and Hubbard
et al. 2009; reproduced with permission of authors, John Wiley & Sons, Inc., and Munksgaard
International Publishers.) (A black and white version of this figure will appear in some formats. For the
color version, please refer to the plate section.)
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46 Stress and deprivation during growth and development and adulthood
Circular defects of thinned or missing enamel are also well documented, especially in
deciduous teeth (Halcrow & Tayles, 2008), but the linear form is the most common type of
hypoplasia. Regardless of form – linear or circular – the color and hardness of hypoplastic
enamel is normal.
Hypocalcifications are enamel defects involving change in color or opacity reflecting
variation in enamel quality or hardness. The enamel surface is usually smooth and appears
intact. Hypoplasias occur when ameloblasts fail to produce the normal thickness of enamel
matrix during enamel development, whereas hypocalcifications appear to result from a
disruption of the mineralization process during the maturation stage of enamel development.
However, this dichotomy may not be so clear cut as hypocalcifications have been experi-
mentally documented in the initial stage of enamel formation (Suckling, 1989).
Hypoplasias result from three potential causes, including hereditary anomalies, localized
traumas, and systemic metabolic stress, such as from illness, disease, and malnutrition (Goodman &
Rose, 1991; Hillson, 1996). Defects arising as hereditary anomalies or as localized traumas are
rare in human populations, indicating that the vast majority of hypoplasias seen in contemporary
and archaeological populations are linked with systemic physiological stress (Hillson, 2008;
Hillson & Bond, 1997; King et al., 2005). The causal stressors associated with hypoplasias are
numerous and varied. Clinical and epidemiological investigations in living populations docu-
ment associations with systemic diseases, neonatal disturbances, and nutritional deprivation
(reviewed in Hillson, 1996; Pindborg, 1982). Experimentally induced stress in laboratory animals
have also shown the direct link between enamel deficiency and stress (Kreshover, 1944;
Kreshover & Clough, 1953a, 1953b; Suckling & Thurley, 1984; Suckling et al., 1983, 1986).
Studies of non-human primates with known life histories reveal links between enamel defects,
life events, and local ecology (Dirks et al., 2002). Thus, enamel defects are a nonspecific indicator
of physiological stress (Goodman & Rose, 1990; Kreshover, 1960; Pindborg, 1982).
Ecological factors are critical for understanding the prevalence and pattern of enamel
defects in human populations. Studies of living populations with dietary deficiencies show
the primacy of nutrition in the development of normal enamel. Analysis of individuals born
during the starvation famine of 1959 – 1961 in the People’s Republic of China reveals that
enamel forming during the famine is highly defective, unlike the enamel that formed either
before or after the famine (Zhou & Corruccini, 1998). For the population as a whole, there was
a clear increase in stress as is clearly revealed in the spike in enamel defects during the famine
period (Figure 2.14). Rural individuals have more defects than urban individuals, a pattern
consistent with records documenting more stress in the rural than in the urban population.
Similarly, forced resettlement, declining nutritional quality, and poor living conditions gen-
erally of Warlpiri people in Northern Territory, Australia, and the Tupí-Mondé speakers in
Brazil in the twentieth century resulted in a profound increase in stress and a correspondingly
marked increase in prevalence of hypoplasias (Littleton, 2005; Santos & Coimbra, 1999).
Enamel hypoplasias show a predilection for anterior teeth and for the cervical and middle
thirds of tooth crowns (Barrett & Blakey, 2011; Condon & Rose, 1992; Goodman & Arme-
lagos, 1985a, 1985b; Hutchinson & Larsen, 1988, 2001; King et al., 2005; Li et al., 1995;
Pedersen & Scott, 1951; Starling & Stock, 2007; Wright, 1997; Zhou & Corruccini, 1998).
This is the case because of the layering nature of enamel deposition in forming the cuspal
component of the tooth crown (Figure 2.12). That is, enamel grows in a series of dome-like,
incremental layers, building one on top of the other, eventually forming the crown cusp(s)
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2.5 Skeletal and dental pathological markers of deprivation 47
70
Whole sample
Rural
60
Urban
Prevalence of individuals with LEH (%)
50
40
30
20
10
0
Pre-famine Famine Post-famine
Figure 2.14 Comparison of linear hypoplasia (LEH) prevalence (percentage of affected
individuals) among living rural (Qingji Township) and urban populations (Shanghai City) in
People’s Republic of China. (Adapted from Zhou & Corruccini, 1998; reproduced with
permission of authors and John Wiley & Sons, Inc.)
and occlusal surface of the tooth. Following completion of the cuspal enamel, the sides of
the tooth crown form, also in a series of discrete layers comprising the imbricational enamel
(FitzGerald & Rose, 2008; Guatelli-Steinberg, 2008; Hillson, 1996, 2008, 2014; Hillson &
Bond, 1997). The layers of cuspal and imbricational enamel are separated by incremental
lines representing the terminus of each depositional episode. These lines, or striae of Retzius,
appear on the surface of the tooth as perikymata.
The top striae of Retzius forming the cusp of the tooth are not exposed on the surface of the
tooth. Thus, except for the final dome of cuspal enamel, all earlier-formed cuspal enamel (20%
– 50% of the crown) is hidden, as are any associated enamel defects (FitzGerald & Rose, 2008;
Hillson, 2008, 2014; Reid & Dean, 2000). The perikymata along the crown sides are, however,
exposed. When insufficiencies of enamel deposition occur, it is in the imbricational enamel
where enamel defects manifest in a visible fashion. The occlusal wall of the hypoplasia
represents the period of time when the stress event occurred (Guatelli-Steinberg, 2008; Hillson
& Bond, 1997). The cervical wall of the enamel defect represents the period of recovery. Thus,
together the occlusal and cervical walls form the hypoplasia (Figure 2.12).
The record of hypoplasia defines a relationship between poor living circumstances and
increased likelihood of increased stress and poor health (see Chapter 3). Few analyses of this
compelling record address the relationship between stress and social hierarchy or inequality.
In a pioneering test case investigating the relationship between social organization and
human biology, analysis of hypoplasia reveals a striking difference in the stress experience
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48 Stress and deprivation during growth and development and adulthood
between elite and non-elite Middle Sicán society in pre-contact Peru (c. AD 1000). That is,
hypoplasia is seven times more common in the non-elite than in the elite in this setting (Klaus,
2014b). This finding is consistent with other settings (Goodman, Armelagos et al., 1984;
Swärdstedt, 1966; and see Goodman, 1998; Goodman and Martin, 2002).
Microdefects
Histological structures in dental enamel known as Wilson bands (or accentuated or patho-
logical striae of Retzius) provide a detailed record of growth disruption (FitzGerald & Rose,
2008; Goodman & Rose, 1990; Simpson, 1999, 2001). Wilson bands are thin layers of
abnormally structured enamel marking the position of the active ameloblasts at the time of
the insult (Figure 2.15). A highly common association between these incremental structures
and life history is the transition from the intra- to extra-uterine environment, resulting in a
distinctive “neonatal line” on the forming teeth, namely in the deciduous teeth and the
permanent first molars as these teeth begin to form in utero (Antoine et al., 2009; Hillson,
2014; Schour, 1936; Schwartz et al., 2010; Whittaker & Richards, 1978). Wide Wilson bands
are associated with traumatic births and perinatal strain (Eli et al., 1989), suggesting that the
width of Wilson bands may represent an indicator of stress severity. Importantly, bands that
match within a single dentition demonstrate the presence of systemic stress for the person
Figure 2.15 SEM micrograph ( 230) of a polished and acid-etched longitudinal section
from a maxillary central incisor, showing a Wilson band (arrows); Santa Catalina de Guale,
Amelia Island, Florida. (Photograph by Scott W. Simpson.)
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2.5 Skeletal and dental pathological markers of deprivation 49
represented (FitzGerald et al., 2006). Analysis in the prehistoric-colonial era series from
Spanish Florida reveals a clear association between Wilson bands and tooth size (Simpson,
1999). Individuals with at least one Wilson band have smaller tooth size (mandibular canine
cervical breadth) than individuals without Wilson bands. This finding is consistent with the
notion that stressed individuals, including short-term stress, are likely less able to resist
physiological stress and not able to achieve full growth potential.
Wilson bands are not always associated with a surface macrodefect such as a hypoplasia
(Bullion, 1986; Condon, 1981; Condon & Rose, 1992; Goodman & Rose, 1990; Hillson, 2008;
Rose, 1977; Wright, 1990). The lack of a consistent association between macro- and micro-
defects suggests that their stress etiologies may be different. Wilson bands appear to represent
brief periods of stress lasting for hours or days, whereas hypoplasias appear to represent long-
term stress lasting from weeks to months.
n = 30 / n = 39 n = 22 / n = 16 n = 22 / n = 19
4.8/5.3 4.2/5.0
4.8/5.1 3.8/4.4
4.4/4.8
4.4/4.6
4.0/4.3 3.4/3.9
3.9/4.1
3.5/3.8 2.9/3.4
3.5/3.7
3.1/3.4 2.5/2.9
3.2/3.3
2.8/3.0 2.9/2.9 2.2/2.4
2.6/2.7 1.9/2.0
2.5/2.7
2.4/2.4
2.3/2.4 1.7/1.8
2.2/2.2
2.1/2.2 1.5/1.6
2.0/2.0
1.9/1.9 1.3/1.3
1.8/1.8
1.7/1.7 1.4/1.5 1.1/1.1
1.6/1.7
1.0/1.0 0.8/1.0
1.9/2.0 1.2/1.1 1.0/1.1
1.3/1.3 1.1/1.3
2.1/2.3
1.5/1.5 1.3/1.5
2.4/2.7
1.7/1.8 1.5/1.7
2.8/3.1
1.9/2.1 1.7/2.0
n = 25 / n = 13 n = 27 / n = 13 n = 29 / n = 15
Figure 2.16 Comparison of mean chronological ages associated with enamel formation in
anterior maxillary and mandibular canines. Ages have been divided according to African
(left) and European (right) samples. (Reid & Dean, 2006; reproduced with permission of
Elsevier Ltd.)
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50 Stress and deprivation during growth and development and adulthood
40
Stillwater
35
Prevalence (%) of enamel hypoplasias
Georgia Agricultural
30 Georgia Preagricultural
25
20
15
10
0
0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 5 5.5
Age (years)
Figure 2.17 Stillwater maxillary permanent central incisor hypoplasia frequencies
(percentage per half-year age group) compared with Georgia coast foragers and farmers.
(Adapted from Hutchinson & Larsen, 1995; reproduced with permission of the American
Museum of Natural History.)
Stress chronology
Because metabolic insults leading to growth disruption affect only the part of the tooth that is
in the process of forming, location of the disturbance on the tooth crown provides a precise
chronologic indicator of stress history. Tooth enamel begins to form at about four months in
utero, beginning with the deciduous first incisors, and is complete when the crowns of the
permanent third molars are fully formed at about 12 years of age (Nanci, 2013; Smith, 1991).
The location of the position of an enamel defect (e.g., hypoplasia) relative to the cemento-
enamel junction can be used to plot the age of disturbance (Reid and Dean, 2006) (Figure 2.16).
Earlier researchers suggested that there is a preprogramed stress clock in humans (Massler
et al., 1941; Sarnat & Schour, 1942). However, the body of evidence building over the last five
decades indicates little support for a universal model of timing of growth disruption (Hillson,
2008). Rather, growth disruption is due to individual and local circumstances involving
environmental (or other) stresses. That is, even within relatively circumscribed cultural and
temporal settings, specific skeletal series show very different patterns and frequency of
enamel defects (Hutchinson & Larsen, 2001; King et al., 2005; Lieverse, Link et al., 2007;
Temple, 2007).
Although the relationship between enamel defects and age has been recognized since the
nineteenth century (Talbot, 1898), this chronological approach has been applied to archaeo-
logical remains only recently. The pioneering investigation of Medieval-era dentitions from
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2.5 Skeletal and dental pathological markers of deprivation 51
Westerhus, Sweden, by Swärdstedt (1966) revealed that hypoplasias peaked in the two-to-
four-year period, a pattern that has been identified in many other archaeological samples
(Corruccini et al., 1985; Goodman et al., 1980; Hillson, 1979; Hodges, 1989; Hutchinson &
Larsen, 1995; Martin et al., 1991; Powell, 1988; Storey, 1992a, 1992b) (Figure 2.17).
The tendency for hypoplasias to occur after the first year or so of life as documented in
archaeological samples suggests that stress may be linked to the negative effects of weaning
(Coppa et al., 1995; Corruccini et al., 1985; Lanphear, 1990; Lillie, 1996; Moggi-Cecchi et al.,
1994; Ogilvie et al., 1989; Simpson et al., 1990; Ubelaker, 1992b; Webb, 1995; and many
others). A test of the weaning hypothesis, based on the study of historical records and
archaeological dentitions from enslaved African-descent populations living in the mid-
Atlantic region of the United States (Maryland and Virginia), reveals that the peak frequencies
of hypoplasias are in the 1.5 – 4.5-year age intervals, whereas weaning took place only nine
months to one year after birth (Blakey et al., 1994). This discrepancy between age pattern of
hypoplasias and weaning led Blakey and coworkers (1994; and see Barrett & Blakey, 2011) to
conclude that weaning was not the primary causal factor leading to enamel defects. That is,
other stresses of enslavement, including nutritional problems, poor hygiene, and illness, were
likely responsible for the age pattern of physiological perturbation in this setting. Thus,
weaning as a cause for the age profile of enamel defects in archaeological settings, usually
between two and four years, is not the sole factor leading to growth arrest as documented by
enamel hypoplasias (and see Hassett, 2014; Katzenberg et al., 1996; Saunders & Keenleyside,
1999). Rather, the problems identified in African-descent populations from the mid-Atlantic
region reflect a period of weakened immunity in early life and reliance on foods that do not
include breast milk. It is this set of circumstances, of which weaning is one, that give the
infant a poor growth environment and result in growth arrest.
Insights into the timing of short-term stress from Wilson bands provide a similar conclu-
sion. Analysis of timing of Wilson bands in the first year of postnatal life in a large series of
deciduous dentitions from first- to third-century AD Rome at the cemetery of Isola Sacra
reveals peak prevalence in the second through fifth months and the sixth through nine
months. This age pattern of short-term growth arrest closely matches with historical sources
indicating two periods of infant frailty, the first occurring around three months when the
infant is first exposed to non-milk foods (e.g., bread softened with milk or wine) and a second
when the infant begins to eat solid foods (FitzGerald et al., 2006). This supports the weaning
hypothesis, but almost certainly, this setting also had other negative circumstances that
contributed to poor growth during these critical months of life.
Duration/severity of stress
Because enamel is deposited in consecutive layers from the incisal/occlusal surface to the
cementoenamel junction over some interval of time, hypoplasia width should represent a
quantification of duration of stress events (Blakey & Armelagos, 1985; Ensor & Irish, 1995;
Hassett, 2014; Hutchinson & Larsen, 1988, 1990, 1995, 2001). Based on their experimental
work with laboratory sheep, Suckling and coworkers (1986; and see Suckling, 1989) con-
cluded that the severity of the stress insult determines the magnitude of the hypoplasia. Thus,
hypoplasia size may reflect either stress duration or severity, or perhaps some unknown
combination of both.
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52 Stress and deprivation during growth and development and adulthood
On the other hand, hypoplasia size per se is not the best estimate of duration of the stress
episode, largely because enamel does not grow at a constant, linear rate (Guatelli-Steinberg,
2008; Hassett, 2014; Hillson, 2008; Hubbard et al., 2009; Reid and Dean, 2006). Simply,
enamel rates vary according to location in the tooth crown. Alternatively and far more
precisely, Hillson and Bond (1997) make the case that the duration of the episode of
developmental disruption is best estimated by counting the number of perikymata within
the margins of the defect, especially with regard to the number of perikymata in the defect’s
occlusal wall, as this is the actual period of growth disruption (Hillson & Bond, 1997)
(Figure 2.12). For example, a chronic disruption may have more than 20 perikymata, whereas
a shorter-term disruption may have fewer than five perikymata. In a Jomon series (Aichi,
Japan), Temple and coworkers (2012) documented a range from one to 20 perikymata in a
single individual, representing stress duration from 12 days to 130 days. All stress events took
place between the ages of 1.2 and 3.5 years. Precise estimates of duration of the stress episode
may be determined as each stria represents about seven to nine days of growth (FitzGerald,
1998; Guatelli-Steinberg et al., 2005; Reid & Dean, 2006). For example, an occlusal wall of a
defect containing nine perikymata represents approximately three months of growth disrup-
tion. Thus, not only do these counts provide precision with regard to timing of stress events,
they also provide an important measure of stress episode duration and inter- and intra-
population variation (Hubbard et al., 2009; Temple et al., 2012, 2013).
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2.5 Skeletal and dental pathological markers of deprivation 53
than other children (May et al. 1993). Comparable patterns of morbidity differences are
reported for Wilson band prevalence in living populations. Children with histories of chronic
systemic disease in Sheffield, England, have elevated prevalence of Wilson bands in their
deciduous teeth (Hillier & Craig, 1992; and compare with FitzGerald et al., 2006, for an
archaeological setting).
The study of dentitions from paleontological and archaeological contexts adds a great deal
to our present understanding of the history of human stress and its complex links with
environment, culture, and biology. Unlike most stress indicators, enamel defects have been
systematically investigated in a range of early hominins, including australopithecines (Tobias,
1967; White, 1978), early archaic Homo sapiens (Bermúdez de Castro & Pérez, 1995), and
Neanderthals (Guatelli-Steinberg, 2008; Guatelli-Steinberg et al., 2004; Hutchinson et al.,
1997; Molnar & Molnar, 1985a; Ogilvie et al., 1989). Early archaic H. sapiens from Atapuerca,
Spain (Bermúdez de Castro & Pérez, 1995), and Neanderthals (Ogilvie et al., 1989), possess low
to moderate prevalences of hypoplasias (permanent dentition: 12.8%, Atapuerca; 41.9%,
Krapina). Each of the two series displays similar age patterns of hypoplasias, including two
primary peaks, the first in early childhood and a second in late childhood. The late peak is
especially interesting because it represents stress events affecting posterior teeth, teeth which
rarely exhibit enamel defects in modern human populations (Ogilvie et al., 1989). Thus, the
earlier peak may reflect early childhood stress (e.g., weaning), and the later peak may
represent overall high levels of systemic stress. Infection may have been a cause of growth
disruption, but the Atapuerca and Krapina samples show very low prevalences of skeletal
infection. Instead, nutritional deficiencies – perhaps during periodic food shortages – were the
most likely causative factor (Guatelli-Steinberg et al., 2004; Hutchinson et al., 1994; Ogilvie
et al., 1989). The record suggests overall that developmental stress levels in Neanderthals by
various measures are high, including elevated prevalence of dental enamel defects and
fluctuating asymmetry (Barrett et al., 2012), but not remarkably so in comparison with
modern foragers from archaeological contexts (Guatelli-Steinberg et al., 2004; Hutchinson
et al., 1994).
Temporal comparisons of enamel defect prevalence in Holocene populations undergoing
dietary and behavioral changes show clear trends in physiological stress. Especially striking
are changes observed in human populations undergoing the shift from foraging to agriculture
or agricultural intensification and increased population density. In general, these comparisons
reveal increases in prevalence of enamel defects, especially in populations inhabiting the
Eastern Woodlands of North America (Cassidy, 1984; Cook, 1984; Goodman, Lallo et al.,
1984; Perzigian et al., 1984; Rose et al., 1984), Latin America (Márquez Morfín and Storey,
2007; Ubelaker, 1984, 1992b; but see Hodges, 1989; Lukacs & Joshi, 1992, for exceptions),
and to a lesser extent, in Asia (Rathbun, 1984; Smith, Bar-Yosef et al., 1984; but see Temple,
2010; Temple & Larsen, 2013; Yamamoto, 1992) and Africa (Starling & Stock, 2007). This is
not a universal trend, however. In comparison with late prehistoric and contact-era maize
agriculturalists from one setting in the American Southeast, there is a decrease in hypoplasia
prevalence (Hutchinson & Larsen, 1988; Larsen, Crosby et al., 2002; Larsen & Hutchinson,
1992; Larsen et al., 2007; Simpson et al., 1990). Hypoplasia widths increase in this setting.
These trends reflect a decline in the number of individuals affected, but stress episodes were of
either longer duration, greater severity, or both. Other late prehistorics or contact-period
agriculturalists in the New World express generally high frequencies of hypoplasias, which
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54 Stress and deprivation during growth and development and adulthood
reflects the deterioration of health in many of these settings (Cohen et al., 1994; Martin et al.,
1991; Milner & Smith, 1990; Pfeiffer & Fairgrieve, 1994; Stodder, 1994; Stodder & Martin,
1992).
While the record of enamel defects has been documented in populations undergoing the
transition from foraging to farming, other dietary shifts reveal a consistent set of results
linking a reduction in stress with improvement in diet. For example, in the shift from hunting
and gathering to cattle pastoralism in early Holocene North Africa, there is a decline in dental
defects in the primary dentition. This decrease reflects increased access to fat and protein in an
increasingly marginal setting of the Sahara Desert (Stojanowski & Carver, 2011). Comparison
of Jomon hunter-gatherers with Yayoi farmers in Japan also shows a decline in enamel
defects (Temple, 2010; Temple & Larsen, 2013). Temple and Larsen (2013) argue that the
introduction of a predictable and renewable food source at an advanced stage of development
resulted in a decline in physiological stress and improved health.
The impact of the adoption of agriculture on the stress experience in earlier human
populations is revealed in a number of settings where changes in health and nutrition are
documented by archaeological and osteological means. At Dickson Mounds, multiple indica-
tors show increasing levels of nutritional stress and infectious disease (see previously; and see
Chapter 3). Increase in prevalence of enamel defects – both macrodefects and microdefects – is
consistent with these findings (Goodman, 1989; Goodman & Armelagos, 1988; Goodman
et al., 1980; Goodman, Lallo et al., 1984; Lallo & Rose, 1979; Rose et al., 1978). The mean
frequency of hypoplastic defects increased from 0.9 to 1.2 to 1.6 per individual in the Late
Woodland, Mississippian Acculturated Late Woodland, and Middle Mississippian periods,
respectively. For the same three periods, the frequency of individuals affected increased from
45% to 60% to 80%. Most defects occurred in juveniles during ages two to four years for the
first two periods. Defects in the late prehistoric intensive agriculturalists (Middle Mississip-
pian) were earlier (<2 years) than in either of the two previous periods, indicating that stress
occurred in earlier childhood as nutritional quality worsened and disease intensified.
Comparisons of age-at-death for Dickson Mounds individuals with and without hypo-
plasias reveals that in the final two prehistoric periods, mean age-at-death of individuals
without hypoplasias is 36.6 years and 37.5 years, and mean age-at-death for individuals with
hypoplasias is 31.3 and 30.2 years (Goodman, Lallo et al., 1984). These results are similar to
other settings showing an inverse relationship between age and presence of enamel defects –
adults have fewer defects than juveniles, older juveniles have fewer defects than younger
juveniles, and/or older adults have fewer defects than younger adults (Boldsen, 2007; Cook,
1990; Cook & Buikstra, 1979; Duray, 1996a, 1996b; Irei et al., 2008; King et al., 2005;
Oxenham, 2006; Rose et al., 1978; Saunders & Keenleyside, 1999; Simpson et al., 1990;
Stodder, 1997; Swärdstedt, 1966; White, 1978). These findings are consistent with those in
living populations that individuals experiencing a greater number of stress episodes during
childhood are predisposed to early death, a finding consistent with the early developmental
origins hypothesis regarding the clear association between stress in early life (prenatal and
postnatal) and early mortality and compromised health in later life (and see Armelagos et al.,
2009; Beltrán-Sánchez et al., 2012; Temple, 2014).
Wilson bands in archaeological remains also provide an important avenue for investigating
other major transitions in human populations that might compromise health status and stress
levels. Wright (1990) determined prevalences of bands in precontact and contact-era
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2.5 Skeletal and dental pathological markers of deprivation 55
mandibular canines from Lamanai, Belize, a Maya center occupied from the Preclassic
through Historic periods. Following an initial period of abandonment after contact by
Europeans, the site was reoccupied as a Catholic mission by Mayan Indians until the mid-
seventeenth century. Diet changed relatively little in the prehistoric to historic transition, but
archival records indicate that other stressors (e.g., European-introduced diseases) comprom-
ised health in native populations following contact. Comparisons of Postclassic and Historic
dentitions show a dramatic increase in physiological stress: 84% of bands observed in the
samples combined are from the Historic dentitions. Historic individuals also show more bands
than seen in precontact individuals (2.4 versus 0.88 per individual, respectively). Given the
lack of major dietary changes, Wright (1990) argues that the differences in microdefect
prevalence can be attributed to changing disease patterns with contact, such as the introduc-
tion of malaria and Old World parasitic infections, diseases leading to acute health crises.
Similarly, early colonial Native Americans in Spanish Florida show considerably greater
prevalence of Wilson bands than do their prehistoric predecessors (Simpson, 2001). Although
the elevation in stress can be attributed to a variety of environmental stressors associated with
Spanish missionization in this setting, a key factor was likely dehydration owing to diarrheal
disease and the reliance on a corn-based weaning diet.
A contrasting temporal trend in microdefect prevalence in comparison with prehistoric and
contact-era native populations is identified at another mission locality in Belize. Microdefect
prevalence in mission Indians at Tipu decreases in comparison with precontact Indians (Cohen
et al., 1994; Danforth, 1997). The different temporal trends at Lamanai and Tipu may indicate
very different contact experiences within the relatively small geographical setting of Belize.
Lamanai served as an important center where populations were relocated from nearby villages
to the town, whereas Tipu was only marginally affected by population relocation and concen-
tration during the mission period. Thus, the contrasting contact experiences at Lamanai and
Tipu may explain the different patterns of morbidity.
Temporal patterns documenting increase in stress are also displayed in foraging popula-
tions undergoing significant adaptive shifts. In the Santa Barbara Channel Islands region,
there is a marked increase in frequency of hypoplasias in the Middle period when populations
underwent a transition from hunting and gathering of terrestrial foods to a heavy reliance on
marine foods, especially fish (Lambert, 1993). Hypoplasias in the mandibular left canine
increased in prevalence from 18.4% in the Late Early period to 49.2% in the Early Middle
period. This increase in stress mirrors temporal changes observed for other nonspecific
indicators (e.g., periosteal reactions), all of which appear to be associated with problems
relating to increasing sedentism, population aggregation, and declining resource availability.
Enamel defect prevalences appear to differ between class or social groupings in a number of
settings, suggesting that higher-status individuals have better diets or other positive environ-
mental factors than lower-status individuals. Prehistoric Rudston and Burton Fleming (Eng-
land) and Medieval-period Westerhus (Sweden) and Koksijde (Belgium) populations represent
unambiguous examples of status differences in enamel defect prevalence, with higher-status
individuals showing considerably fewer enamel defects than lower-status individuals (Peck,
2013; Polet et al., 2000; Swärdstedt, 1966) (Figure 2.18). Thus, in these settings, during the
years of growth and development of the dentition, higher-status juveniles may have enjoyed
better health than did lower-status juveniles. Similarly, in the Dickson Mounds series and in
the Pete Klunk Mound group in the lower Illinois River valley, high-status individuals have
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56 Stress and deprivation during growth and development and adulthood
High Status
Prevalence (%) of enamel hypoplasias
100
Intermediate Status
80 Low Status
60
40
20
0
0 0.5 1 1.5 2 2.5 3 3.5 4 4.5 5 5.5 6 6.5 7 10-17
Age at tooth mineralization (years)
Figure 2.18 Prevalence (%) of enamel hypoplasias by social groups of high, intermediate,
and low status at Westerhus, Sweden. (Adapted from Swärdstedt, 1966.)
fewer hypoplasias than low-status individuals (Cook, 1981a; Goodman & Armelagos, 1988),
suggesting a possible association between status and stress history in both settings. In
Lambayeque, Peru, and Copán, Honduras, lower-status individuals have a greater prevalence
of enamel defects than higher-status individuals (Shimada et al., 2004; Storey, 2005). The
similar pattern of age differences in vertebral neural arch size, tooth size, and enamel defect
frequency, especially in the comparison of juveniles and adults from the same population,
strongly suggests that individuals surviving to adulthood enjoyed relatively better health than
did those members of the population who expired prior to reaching adulthood.
Enamel defects are not a health risk per se, but abundant clinical evidence indicates that
enamel defects – hypoplasia and hypocalcification – predispose teeth to cariogenesis
(Baume & Meyer, 1966; Mellanby, 1934; Nikiforuk & Fraser, 1984; and see review in Duray,
1992). Called “circular caries,” the association between enamel defects and caries has been
documented in Middle Woodland and Late Woodland deciduous teeth in lower Illinois River
valley (Cook & Buikstra, 1979). Both periods contain high prevalences of circular caries, but
there is an especially high prevalence in the later period when maize agriculture intensified
(Cook and Buikstra, 1979). Follow-up study of microdefects from this setting indicates a close
association between circular caries and microdefects (Cook, 1990). Identification of age-
of-occurrence of stress indicators reveals that physiological perturbations are perinatal,
reflecting poor health of both the mother and her infant around the time of birth.
Dentitions from the Libben site, Ohio, show a gradient in caries susceptibility in comparison
with different types of gross enamel defects – teeth with gray-chalky hypocalcifications are
more carious than discolored teeth (Duray, 1990, 1992). These associations indicate that
weakened enamel structure promotes cariogenesis. In contrast to many other prehistoric
foragers, the Libben series displays high caries prevalence. Duray (1990) suggests that high
caries prevalence can be attributed to the high prevalence of hypocalcifications. In contrast to
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2.6 Adult stress 57
the association between caries and hypocalcification, teeth with linear hypoplasias appear to
be caries-resistant. Duray (1990) speculates that hypermineralization of the defect may
suppress cariogenesis.
Hypocalcifications and dental caries in deciduous teeth have also been identified in the
Classic-period Maya Indians from Copán, Honduras (Storey, 1992b, 1992c), and in native
populations from the Mariana Islands, Polynesia (Hanson, 1990). In both settings, native
twentieth-century populations have high prevalences of circular caries. In Mesoamerica, these
prevalences are linked with over-reliance on carbohydrates, poor nutrition, and the synergis-
tic relationship between diet and disease (Storey, 1992c). In the Mariana Islands, unusually
high levels of circular caries may be related to a number of local conditions, including
excessive fluoride intake by pregnant and lactating women, poor water quality, the consump-
tion of highly cariogenic starch diets with weaning, and specific nutrient deficiencies (e.g.,
protein) in mothers and their infants (Hanson, 1990).
Sex differences in prevalence of hypoplasias and other enamel defects in archaeological
series are highly variable. For example, no differences in hypoplasia prevalence are present
between adult females and males in the Dickson Mounds series (Goodman et al., 1980; and see
discussions of other North American samples by Danforth, 1997; Danforth et al., 1997;
Lanphear, 1990; Martin et al., 1991; Powell, 1988; Stodder, 1997; Wright, 1990).
Clinical, epidemiological, and bioarchaeological studies indicate that enamel defects are
more common in males than in females, more common in females than in males, and in equal
prevalences (Goodman & Rose, 1990; Goodman et al., 1991; King et al., 2005). At least one
review suggests that stress buffering, often associated with females, does not influence defect
expression to any appreciable degree (Guatelli-Steinberg & Lukacs, 1999). Immature females
have more hypoplasias than immature males in Tezonteopan, Mexico, which is consistent
with other evidence indicating worse nutrition in girls than in boys in this setting (Goodman
et al., 1991). Hypoplasia prevalence does not differ between South Asian males and females in
settings where daughter neglect results in greater malnutrition and mortality in females, at
least for some regions (Lukacs and Joshi, 1992). The differences in findings between Mexico
and South Asia suggest that differential treatment of males and females during childhood will
not necessarily be reflected in differences in enamel defect prevalences.
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58 Stress and deprivation during growth and development and adulthood
Figure 2.19 Left to right: sequential loss of adult bone with age, in second metacarpals,
Post-Medieval St. Bride’s Lower Churchyard, London. (Brickley & Ives, 2008; reproduced
with permission of the Museum of London.)
Anderson & Pollitzer, 1994; Glencross & Agarwal, 2011; Heaney, 1993; Stini, 1990, 1995).
Depending on factors relating to muscle mass, activity level, pregnancy, lactation, and diet,
bone loss after the age of 40 years is substantial, amounting to 1% – 5% per year. The
identification of variation in populations of disparate ancestry also points to genetic factors
that predispose some groups to osteoporosis (Anderson & Pollitzer, 1994; Cho et al., 2006;
Nelson et al., 2004). Two types of bone loss due to osteoporosis are identified clinically,
including that arising from reduction in estrogen levels following menopause (Type I), and
gradual age-related reduction in bone mass in adult females and males (Type II) (Drezner,
1995; Stini, 1990). Women lose relatively more bone mass than do men, due to the combined
effects of Type I and Type II osteoporosis. Estrogen is critical in bone maintenance (Drink-
water, 1994). The reduction in estrogen can even result in significant bone loss for younger
women undergoing overly vigorous exercise regimens and accompanying loss of menstru-
ation (secondary amenorrhea) (Anderson, 1995; Kreiner, 1995).
The rate of bone loss in human populations is variable, and environmental factors such as
nutritional status are significant influences (Arnaud & Sanchez, 1990; Martin et al., 1985;
Nelson et al., 2004; Pollitzer & Anderson, 1989; Schaafsma et al., 1987). Clinical evidence
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2.6 Adult stress 59
indicates that individuals with low calcium intakes are more prone to adult bone loss, and
other dietary factors such as high protein consumption are also implicated (Arnaud &
Sanchez, 1990; Nordin et al., 1993; Pfeiffer & Lazenby, 1994; Stini, 1990, 1995). Body weight,
heredity, and lactation status are also important risk factors (Agarwal & Stout, 2003; Arnaud &
Sanchez, 1990; Evers et al., 1985; Heaney, 1993; Kreiner, 1995; Pollitzer & Anderson, 1989;
Schaafsma et al., 1987; Stini, 1990, 1995). Comparisons of active versus sedentary popula-
tions or athletes versus non-athletes indicate the strong influence of physical activity on bone
maintenance: simply, active individuals have stronger, denser bone than sedentary individ-
uals (Anderson & Pollitzer, 1994; Drinkwater, 1994; Kohrt et al., 2004; Lacey et al., 1991;
Marcus et al., 1992; McMurray, 1995; Yano et al., 1984). Due to a decrease in physically
demanding lifestyles in contemporary settings (e.g., Sweden, United States, United Kingdom,
China), there appears to be a rapid secular increase in osteoporosis in general and osteoporotic
fractures in particular, matching the remarkably rapid changes in lifestyle (Allander, 1995;
Kohrt et al., 2004; Lau et al., 1990; Ruff, 2006).
Adult bone mass is documented in human remains from a variety of archaeological settings.
Much of this research shows either a general similarity or accelerated patterns of bone loss in
archaeological samples and in living populations (Carlson et al., 1976; Cho & Stout, 2003; Cook,
1984; Dewey et al., 1969; Robling & Stout, 2003; Van Gerven, 1973). Variation in relation to
differing lifestyles and subsistence strategies has been examined in some detail, using alterna-
tive data collection protocols, including raw measures of bone mass (cortical thickness [CT],
cortical area [CA], bone mineral content [BMC]), or size-standardized measures (percentage of
cortical area [%CA or PCCA] or percentage of cortical thickness [%CT or PCCT]) (Ruff, 1992).
Comparisons of femoral cortical thickness in X-group (AD 350 – 550) intensive agriculturalists
from the Wadi Halfa area of Sudanese Nubia with modern Euroamericans and Native Americans
reveal similar trends of initial gains in bone mass from the third to fourth decades, followed by
losses (Martin & Armelagos, 1979; Martin et al., 1985). In Nubian females, bone mass decreased
after 20 years of age. Martin and coworkers (1985) speculate that premature osteoporosis was
due to nutritional inadequacies associated with an over-reliance on a single dominant crop
(millet), such as protein – calorie malnutrition or imbalance of calcium – phosphorus ratios, or
the influence of disease. Perhaps the bone losses in childhood in this setting (Hummert, 1983;
and see earlier) predisposed adults, especially females, to premature bone loss.
Similarly, bone mass (percentage of cortical area from second metacarpals) in late prehis-
toric maize agriculturalists from southern Ontario is below what has been documented in
living populations (Pfeiffer and King, 1983; and compare with Garn, 1970). Although various
factors may be involved, the reliance on maize and attendant protein – calorie malnutrition
may have contributed to low bone mass.
Based on the assumption that cortical thickness and Nordin’s Index (cortical thickness/total
subperiosteal area) are useful measures of bone mass and nutritional quality, Owsley (1991)
compared femoral bone mass in a temporal series of Great Plains Arikara dating from c. AD
1600 – 1832. These comparisons reveal an increase in bone mass in the transition from the
late prehistoric to early protohistoric period in the late seventeenth century, which Owsley
regards as “a positive change in nutritional status,” perhaps relating to increased availability
of protein acquired through hunting and trade (1991:109). By 1800, bone mass declined
dramatically. Owsley suggests that these declines are due to the stresses associated with
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60 Stress and deprivation during growth and development and adulthood
biological and social disruptions of disease, warfare, and other negative environmental
circumstances in the early nineteenth century.
In contrast to the Arikara, no temporal change in bone mass (Nordin’s Index) could be
detected in a sequence of human remains from the lower Illinois River valley dating from the
Archaic to Mississippian period (Cook, 1984). This suggests that the profound change in diet
involving the shift to maize agriculture in later prehistory had no bearing on bone mainten-
ance. Some diseased individuals in the Mississippian period expressed relatively low bone
mass. Individuals with skeletal tuberculosis had remarkably low bone mass, suggesting that
non-dietary factors play an important role in bone maintenance in this region.
Most bioarchaeological research emphasizes the direct role of diet and nutrition in explain-
ing variation in bone maintenance. This perspective ignores the important influence of
mechanical loading and activity on bone mass and how it is distributed (and see Chapter 6).
Continued subperiosteal expansion in adults compensates for medullary expansion and
endosteal bone loss in order to maintain the mechanical integrity of the bone cross-section
under loading regimes. The raw measures of bone mass frequently used in studies of archaeo-
logical remains may present an incomplete picture of bone remodeling and health status (Ruff,
2008; Ruff & Larsen, 1990). Thus, low bone mass does not necessarily indicate inadequate
bone mass (Pfeiffer & Lazenby, 1994).
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2.6 Adult stress 61
(a)
Periosteum
Haversian systems
Lamella
Vein
Lacuna
Lymph vessel
Artery
(b)
Figure 2.20 Schematized (a) and actual (b) cross-section of compact bone showing major
microstructures discussed in the text. The ring-like structures (Haversian systems or
osteons) house the Haversian canals. Each lacuna contains cells that maintain the bone
tissue and are arranged in layers (lamellae). ([a] Illustration by Dennis O’Brien; Larsen,
1987. © Academic Press, Inc.; [b] photograph by Samuel Stout.) (A black and white version
of this figure will appear in some formats. For the color version, please refer to the plate
section.)
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62 Stress and deprivation during growth and development and adulthood
80
70 Isola Sacra
60 European American
50
40
30
20
10
0
Tt.Ar Ct.Ar En.Ar OPD
(mm2) (mm2) (mm2) (#/mm2)
1.4
1.2
0.8
0.6
0.4
0.2
0
r x 2 ) r) r) 2 )
t.A de m 2 /y 2 /y m
/T in m m m /m
Ar ic r( /m /m
2
C
t. bol n.
A (#
2 m
ra f m (m
O c. (m
Pa A R BFR
BF et
N
Figure 2.21 Comparison of mean histomorphometric variables: Tt.Ar, total cross-sectional
area; Ct.Ar, cortical bone area; En.Ar, endosteal area (bone marrow cavity area); OPD,
osteon population density; Ct.Ar/Tt.Ar, relative cortical area; Parabolic index, optimal
distribution of cortical bone around marrow space; On.Ar, average cross-sectional area
with minimally 25 complete osteons per section; Ac.f, mean activation frequency
associated with the number of osteons annually created per area unit; BFR, average bone
formation rate; Net BFR, new bone formation rate. (Adapted from Cho & Stout, 2003;
reproduced with permission of authors and Kluwer Academic/Plenum Publishers.)
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2.6 Adult stress 63
agriculturalists may reflect the effects of nutritional stress associated with maize-based, low-
protein diets (Pfeiffer & Lazenby, 1994; Stout, 1983), or perhaps, variation in skeletal tissue
maturation rates in different populations (Stout & Lueck, 1995).
The rate of mineralization of osteoid (unmineralized bone matrix) in the Haversian canal
can be influenced by systemic stress. Under conditions of normal bone development, osteons
mineralize uniformly, but under conditions involving slower growth – such as with nutri-
tional stress or disease – delayed osteoid mineralization results in the creation of hyperminer-
alization zones. Viewed in cross-section, one or more hypermineralized zones appear as rings
of increased density comparable to lines of growth disruption found at the ends of long bones
(Bartsiokas & Day, 1993; Martin & Armelagos, 1985; Simmons, 1985; Stout & Simmons,
1979). The presence of these “double-zone” osteons indicates the presence of physiological
stress and growth disruption. The frequency of double-zone osteons per unit area of bone is
positively correlated with the total amount (cortical area) of bone in the Nubian skeletons
(Martin & Armelagos, 1985). This may indicate that some critical threshold of metabolic
activity is essential for recovery from growth arrest (Martin & Armelagos, 1985).
Histomorphometric study of three populations with varying subsistence strategies –
primarily meat (Alaskan Eskimo), mixed foraging and farming (Arikara), and intensive maize
agriculture (Pueblo) – shows a high degree of variability in osteon structure. Osteons in a
number of individuals in these samples contain second and smaller remodeling sequences
(Eriksen, 1980; Richman et al., 1979). Called Type II osteons, these structures represent sites of
accelerated availability of calcium. Alaskan Eskimos have the highest frequency of Type II
osteons, perhaps reflecting heavy reliance on meat in comparison with more plant-oriented
Arikara and Pueblo Native Americans.
The study of histomorphometrics in adult Nubian remains provides important comple-
mentary information to the aforementioned analyses of bone mass. Nubian adult femora
possess abnormally large, active resorption spaces, which may result from nutritional stress
due to agricultural dependence (Martin and Armelagos, 1979, 1985). The overall quality of
bone is reduced in these groups due to the high degree of inadequately mineralized, porous
cortex. These findings underscore the general observation that increased porosity is due, at
least in part, to histological changes, including enlarged resorption spaces. Moreover, faster
remodeling will yield more osteons and thus more Haversian canals, contributing to increased
intracortical porosity. The role of mechanical environment is suggested strongly by variation
in microstructure, however. In this regard, adult females have larger femoral osteons than
adult males (Mulhern and Van Gerven, 1997), which reflects less mechanical demand on
women than men. This is consistent with the observation that osteon size is not different in
ribs analyzed in adult males and females because ribs are non-load bearing (Mulhern, 2000).
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64 Stress and deprivation during growth and development and adulthood
Although the reasons for this difference with modern populations are not known, the greater
mechanical demands and differences in lifestyle in general may explain the maintenance of
structural integrity in older adults at Wharram Percy in contrast to living societies where
adults are less active (Agarwal et al., 2004).
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2.7 Summary and conclusions 65
healthy. If reduced body size is adaptive, then reduced height should have no associated
functional or demographic costs. In fact, a significant body of literature from multiple
disciplines illustrates that small body size is linked with various negative factors, including
increased disease and poor nutrition. Poor growth status is associated with a range of
functional costs and consequences, including decreased activity and poorer learning (Crooks,
1995; Dasgupta, 1993; Goodman, 1991, 1994; Stinson, 1992, 2000). Although smaller body
size appears to enable individuals to perform some activities with lower energy requirements,
the efficiency is reduced (Stinson, 1992). Clearly, there are negative consequences of small
body size in disadvantaged settings, indicating that this reduction is associated with elevated
morbidity, and is in fact maladaptive.
In the study of past societies, it is possible to test the “adaptive” versus “stress” hypotheses
via observation and quantification of the growth and nutrition stress indicators discussed in
this chapter. Significantly, various indicators are linked with decreased survival as determined
by mean age-at-death of individuals with and without (or with relative differences in
prevalence of ) the indicator. Enamel defects (macro- and microdefects), vertebral neural canal
size, height, and tooth size show clear links with lifespan. Where age-at-death and stress
indicators are examined concurrently, individuals without enamel defects or anemia-related
lesions, with larger neural canal size, with greater height, and with larger tooth size died later
in life. These findings suggest that skeletal stress indicators are related to quality of life and do
not represent adaptations.
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3 Exposure to infectious pathogens
3.1 Introduction
For the entire evolution of our species, we have been exposed to a wide range of infectious
agents – parasites, bacteria, and viruses – resulting in a range of diseases. The dental and
skeletal evidence for some of these diseases, mostly chronic, is well documented (Aufderheide
& Rodríguez-Martín, 1998; Ortner, 2003; Roberts & Manchester, 2005). Current bioarchaeo-
logical inquiry emphasizes biocultural perspectives of disease in relation to social, cultural,
and environmental contexts and risks for infection, impacts on population, and implications
for pathogen – host evolution (Armelagos & Van Gerven, 2003; Buikstra, 2010; Buikstra &
Cook, 1980; Larsen & Walker, 2010). This growing record of health in past populations and
especially the emphasis on origins and evolution of infectious disease in the biocultural
context provides a powerful approach for understanding health determinants and outcomes
in the world we live in today.
Infection by a pathogen does not always result in disease. The progression from infection to
disease depends on agent pathogenicity, transmission route from agent to host, and the
strength and nature of the response of the host (Brown et al., 2011; Inhorn & Brown, 1990;
Smith & Moss, 1994). Many acute infectious diseases or epidemics result in death of the
infected individual soon after microbial attack. These infectious diseases leave no skeletal
record, clouding the full picture of disease and its relationship to mortality in past popula-
tions. A number of chronic infectious diseases affect osseous tissues in patterned ways.
Despite the interpretive drawbacks and sometimes lack of specificity, the study of bone lesions
documenting disease provides important perspectives on health in earlier societies and the
impacts of particular living circumstances on the human condition.
The frequency of members of a population affected by a disease forms the baseline of
information from which to interpret health status and factors that influence it. Various means
of data presentation are used, but incidence and prevalence are most commonly reported in
clinical and epidemiological studies (Keyserling, 1988; Waldron, 1994). Incidence is defined as
the number of new cases in a population in a given period. Because the number of new cases
cannot be identified in archaeological settings with certainty (if at all), it is not possible report
on the incidence of a particular disease or pathological conditions. Prevalence, however, can be
observed in archaeological contexts because it represents the proportion of the population
affected by the disease at a single moment or within a period. Prevalence, therefore, provides an
avenue for addressing how common pathological conditions relating to disease occur under a
given set of circumstances, including geographical and temporal variation, sex, age, status and
socioeconomic position, residence, and other key contexts. This paleoepidemiological approach
is fundamental to understanding life conditions in past populations (Boldsen & Milner, 2012).
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3.2 Dental caries 67
This chapter focuses on disease prevalence as expressed in teeth and bones. Ortner (2003)
provides details on a wide variety of infectious diseases identified in archaeological remains
from around the world. This discussion of infection pertains to dental caries, periodontal
disease, antemortem tooth loss, nonspecific periostitis, and specific infectious
diseases – treponematosis, tuberculosis, leprosy, and vectored infections, especially bubonic
plague. These conditions are among the most frequently studied, they have distinctive
pathogenesis involving bone (except for bubonic plague, which leaves no skeletal signature),
and the diagnostic tools (e.g., differential diagnosis of skeletal lesions, ancient DNA docu-
mentation of pathogens) are providing new perspectives on the occurrence and impact of
infectious disease on humankind.
Figure 3.1 Mandibular carious lesions; Ochsenfurt, Germany. Root caries have affected the
cement–enamel junction of the molar (right) and resulted in the complete destruction of the
tooth crown (left). (Photograph and copyright by Leslie L. Williams; reproduced with
permission.). (A black and white version of this figure will appear in some formats. For the
color version, please refer to the plate section.)
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68 Exposure to infectious pathogens
Several essential and modifying factors are involved in the pathogenesis of dental caries.
The essential factors include: (1) teeth the surfaces of which are exposed to the oral environ-
ment; (2) the presence of aggregates of complex indigenous oral bacterial flora (especially
including but not limited to Streptococcus mutans, Lactobacillus acidophilus), salivary glyco-
proteins, and inorganic salts adhering to the tooth surfaces (called dental plaque); and (3) diet
(Burne, 1998; Hara & Zero, 2010; Rowe, 1982; Tanner et al., 2012). Modifying factors are
those that influence the site distribution and rate of carious lesion development. Such factors
include, but are not limited to, crown size and morphology, enamel defects, occlusal surface
attrition, tooth structure, food texture, oral and plaque pH, speed of food consumption, some
systemic diseases, age, child abuse, heredity, salivary composition and flow, nutrition, perio-
dontal disease, enamel elemental composition, and presence of fluoride in drinking water, and
other geochemical factors (Bowen, 1994; Burt & Ismail, 1986; Burt & Pai, 2001; Geddes, 1994;
Greene et al., 1994; Hara & Zero, 2010; Hildebolt et al., 1988, 1989; Hillson, 2008; Hunt et al.,
1992; Leverett, 1982; Maat & Van der Velde, 1987; Meiklejohn et al., 1992; Milner, 1984;
Molnar & Hildebolt, 1987; Molnar & Molnar, 1985b; Newbrun, 1982; Powell, 1985; Rowe,
1982; Woodward & Walker, 1994). Preliminary analysis of genes involved in taste, enamel
formation, and saliva composition suggests a genetic component associated with increased
risk of contracting dental caries (Wendell et al., 2010; Wright, 2010).
Intrinsic characteristics of food, its consistency, and the manner in which it is prepared
influence cariogenesis in human populations. A range of human and animal investigations
reveal that carbohydrates – sugars and starches – are central elements promoting dental caries
(Hillson, 2008). The relative amount of carbohydrates consumed and reliance on domesticated
plants largely explains the variation in prevalence of dental caries in archaeological and other
settings. As succinctly stated by Hara and Zero (2010:459), diet is “the main driver of the
caries process.”
The degree and rate of occlusal surface wear appears to be a mitigating factor in caries rates
in archaeological settings. Maat and Van der Velde (1987) found a negative correlation
between frequency of occlusal surface caries and degree of dental wear in molars from sailors
recovered from a seventeenth- and eighteenth-century Dutch whaling station in the Spitz-
bergen Archipelago (Svalbard). In this series, increased wear appears to be associated with
fewer carious lesions. They concluded that “(t)hese findings strongly suggest a competitive
relationship between progress in caries and attrition” (Maat & Van der Velde, 1987:281). The
studies linking reduced wear to increased caries prevalence are convincing, especially because
common sites of plaque and cariogenesis are the grooves and fissures of unworn crowns (and
see Christopherson & Pedersen, 1939; Corbett & Moore, 1976; Hillson, 2008; Milner, 1984),
especially of the posterior teeth, which have a differential (higher) risk of developing carious
lesions than the anterior teeth.
The relationship between low caries rates and high occlusal-wear environments should not
be overly generalized, however. In their study of the Mesolithic dentitions from Cabeço da
Arruda and Moita do Sebastiao, Portugal, Meiklejohn and coworkers (1992) found a positive
correlation between caries and wear in molars; the most heavily worn crowns are the most
carious. In this setting, the Mesolithic Portuguese individuals consumed figs and carob, foods
high in sugar content that also produce high rates of wear. Occlusal surface wear is also
excessive in Archaic-period foragers from the lower Pecos region of southwestern Texas,
resulting in pulp cavity exposure and tooth loss (Hartnady & Rose, 1991; Sobolik, 1994).
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3.2 Dental caries 69
Caries prevalence is high (14% of teeth) and is indistinguishable from many agricultural
groups (Larsen et al., 1991).
Coprolite analysis reveals that various highly abrasive materials were included in foods
consumed, including phytoliths, seeds and small bones, and calcium-oxylate crystals from
succulents and cacti (Hartnady & Rose, 1991; Sobolik, 1994). Historic accounts also document
the introduction of abrasives to food, including ash for sotal baking and dirt to “sweeten”
meals. High-carbohydrate foods such as succulent fibers, prickly pear fruits, pecans, and
mesquite resulted in active cariogenesis. Thus, like the Portuguese Mesolithic foragers, Pecos-
region Native Americans show a positive relationship between tooth wear and caries.
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70 Exposure to infectious pathogens
25
Eastern North America
21.0
Dental Caries
20
18.0
Carious Teeth (Percentage)
14.4
15
10
5.1
5 3.3
2.1
0
Archaic
Contact
Late Woodland
Early Woodland
Late Prehistoric
Middle Woodland
Figure 3.2 Percentage of teeth affected by dental caries in eastern North America.
(Based on data from Larsen et al., 1991; Milner, 1984.)
agriculturalist periods (Late Woodland to Contact) have more than 7% of carious teeth. Within
the agriculturalist periods, there is a high degree of variability in caries prevalence. This
pattern mirrors observations of living populations with broadly similar diets (Hillson, 2008;
Walker et al., 1998). Relatively small differences in diet and food processing technology can
result in large differences in caries prevalence. Overall, there is a clear tendency for prehistoric
and early contact-era maize farmers to have higher caries prevalence than prehistoric foragers
(and see Gold, 2004; Lambert, 2000a; Larsen, 2014; Larsen et al., 1991; Milner, 1984; various
in Cohen & Armelagos, 1984; Cohen & Crane-Kramer, 2007; Steckel & Rose, 2002). The late
mission-period series from Amelia Island, Florida, shows an unusually high caries prevalence
for North America (Larsen et al., 1991, 2007). This is not unexpected given that missionization
was accompanied by considerable intensification in production and consumption of maize
during the sixteenth and seventeenth centuries in Spanish Florida (Hann, 1988; Jones, 1978;
Worth, 2001) and in other Spanish colonial settings (Bruwelheide et al., 2010; Klaus & Tam,
2010; Ubelaker, 1994).
The reasons for caries increase in Eastern Woodlands agriculturalists are varied and
complex, but the chief factor is related to the presence of sucrose in maize (Hardinge et al.,
1965). Because sucrose is a simple sugar, it is readily metabolized by oral bacteria. Declining
tooth wear in many settings was likely an important factor (see Chapter 7 for a discussion of
tooth wear change), but the common occurrence of maize consumption largely explains
these changes over this expansive area of North America. As such, the recognized
and dramatic positive association between maize consumption – in both its adoption and
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3.2 Dental caries 71
intensification – and dental caries in eastern North America is useful for tracking temporal
changes in dental health in specific settings, where social, cultural, and dietary shifts are
documented using other independent lines of archaeological evidence (Cassidy, 1984; Cucina
et al., 2011; Danforth et al., 2007; Driscoll & Weaver, 2000; Gold, 2000; Hoyme & Bass, 1962;
Larsen et al., 1991, 2007; Milner, 1984; Patterson, 1984; Perzigian et al., 1984; Sciulli, 1997;
Sciulli & Oberly, 2002; Smith, 1982; Sullivan, 1990; and others). In at least one setting where
intensive farming was replaced by mixed foraging and farming, caries prevalence shows a
decline (Lambert, 2000b). Moreover, the range of variation is strongly influenced by geog-
raphy in this and other settings. For example, in the coastal plains of North Carolina and
Georgia, variation in caries prevalence reflects differing levels of commitment to maize
farming (Driscoll & Weaver, 2000; Hutchinson, 2002; Lambert, 2000b; Larsen et al., 2007),
a pattern consistent with dietary reconstruction based on carbon stable isotope ratios
(Hutchinson, 2002; Hutchinson et al., 1998; Larsen, Griffin et al., 2001; and see Chapter 8).
For many native populations of Eastern North America, maize agriculture was the first
significant experience with plant domestication. In several regions, it was not a new experi-
ence. In the North American midcontinental region (modern states of Ohio, Kentucky, and
Illinois), at least five native starchy plants were domesticated 2000 – 4000 years ago (Gremil-
lion, 2002; Smith & Yarnell, 2009). For this period, Rose and coworkers (1991) identified an
increase in carious lesion frequency that is likely linked with this reorientation of diet well
before the adoption of maize in later prehistory (beginning c AD 800 – 900).
There is limited understanding of the relative differences of the impact of agriculture on
oral health of populations consuming various types of plant domesticates. European and
western Asian Neolithic populations have generally lower caries prevalence values than those
of prehistoric maize farmers in North America (Eshed et al., 2006; Lubell et al., 1994). This
suggests that Neolithic domesticates were either less important or less cariogenic than maize
in New World populations. Moreover, in the Levant, there is no significant difference in the
pre-farming Natufian period (6.4% carious teeth) compared to the early farming Neolithic
period (6.7%) (Eshed et al., 2006). However, early and late Natufians have 1.5% and 6.8%
carious teeth, respectively (Eshed et al., 2006), suggesting that a greater reliance on plant
domesticates began before the Neolithic period.
By the same token, some Asian populations that consumed rice have relatively low caries
prevalence compared to maize-based populations, owing to apparently lower cariogenic
properties of rice (Sheiham, 2001; and see Domett, 2001; Domett & Tayles, 2007; Oxenham,
2006; Oxenham et al., 2006; Pietrusewsky & Ikehara-Quebral, 2006; Sakashita et al., 1997;
Tayles et al., 2000; Willis & Oxenham, 2013; but see Inoue et al., 1986; Temple & Larsen,
2007, 2013). In Southeast Asia, while there are temporal increases in rice production and
consumption, there are varying associated patterns of dental caries prevalence, including
decline (Domett, 2001), very little change (Pietrusewsky & Douglas, 2002a, 2002b), and
increase (Newton et al., 2013; Oxenham et al., 2006). These findings suggest that there is a
mixed pattern of health changes with intensification of rice agriculture (and see Clark et al.,
2014).
As a general trend, however, comparisons of pre- and postagricultural populations in a
wide range of settings reveal trends involving increases in caries prevalence irrespective of the
type of cultigens consumed, including East and South Asia (Douglas & Pietrusewsky, 2007;
Fujita, 1995; Inoue et al., 1986; Lukacs, 1990, 1992; Lukacs & Minderman, 1992; Lukacs et al.,
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72 Exposure to infectious pathogens
1989; Okazaki et al., 2014; Pechenkina et al., 2013; Temple & Larsen, 2007, 2013), the Middle
East (Littleton & Frohlich, 1993; Smith, Bar-Yosef et al., 1984; Smith & Horwitz, 2007), Europe
(Bennike, 1985; Bennike & Alexandersen, 2007; Brabant, 1967; Brinch & Møller-Christensen,
1949; Brothwell, 1959; Corbett & Moore, 1976; Cunha et al., 2007; Hardwick, 1960; Meikle-
john et al., 1984; Moore & Corbett, 1971, 1973, 1975; O’Sullivan et al., 1993; Roberts & Cox,
2003; Tóth, 1970; Wells, 1975; Whittaker, 1993), northeast Africa (Armelagos, 1969; Rose
et al., 1993; Smith & Jones, 1910), and South America (Alfonso et al., 2007; Pechenkina,
Vrandenburg et al., 2007; Ubelaker, 1984, 1994; Watson et al., 2010). In the rare instances
where populations shifted back to hunting and gathering, caries prevalence declined (Lukacs,
2007a).
Measurement of carious lesion size and location on tooth crowns provides an important
means of assessing the severity of the disease process. Temporal comparisons of foragers and
farmers, or earlier and later farmers, from a number of settings indicate an increase in lesion
size and shift in the distribution on tooth crowns from mostly the occlusal surface to other
surfaces (e.g., roots) (Larsen, 1982; Pezo Lanfranco & Eggers, 2010; Smith, 1982).
Elevated prevalences or severity of dental caries are not limited to groups relying on
domesticated plants. Prehistoric foragers from the lower Pecos River valley and central Texas
have values that are well within the ranges reported for agriculturalists (Bement, 1994;
Hartnady & Rose, 1991; Marks et al., 1988; Sobolik, 1994). In these settings, the consumption
of sticky, high-carbohydrate, nondomesticated plants resulted in extensive caries.
High caries prevalence in some Mesolithic foragers from Sicily and Portugal (Borgognini
Tarli et al., 1985, 1993; Frayer, 1988a; Lubell et al., 1994; Meiklejohn et al., 1988; but see
Cunha et al., 2007) contrasts sharply with low caries rates in other Mesolithic-period European
(e.g., Scandinavia: Alexandersen, 1988; Bennike & Alexandersen, 2007; Meiklejohn &
Zvelebil, 1991); Meiklejohn et al., 1988, 1997), South Asian (e.g., North India: Lukacs & Pal,
1993, 2003), and African (e.g., Nubia: Rose et al., 1993) populations. The high prevalence of
dental caries in Sicily and Portugal has been linked with the consumption of cariogenic
nonagricultural foods (e.g., honey) or sweet, sticky fruits (e.g., dates or figs) (Lubell et al.,
1994). Comparison of foragers (Mesolithic) and later farmers (Neolithic) from Portugal
revealed an increase in caries prevalence with the adoption of agriculture (Cunha et al., 2007).
Analyses of other hunter-gatherers point to the importance of additional factors that
explain unusually high caries prevalence. In South and Southeast Asia, quite high caries
prevalence in pre-Neolithic contexts points to dietary patterns that promoted caries in either
the absence of plant domestication, or at least only limited access to consumption of crops.
Lukacs (1990) documented relatively high caries prevalence (8.0%) in a South Asian Meso-
lithic population (Langhnaj site) in comparison with other contemporary series from the
region (most are approximately or much lower than 1%). Archaeological evidence indicates
close trade relationships between Langhnaj and agricultural groups, suggesting that cario-
genic foods may have been acquired through exchanges (Lukacs, 1990). In some living
foraging groups in central Africa, although called “hunter-gatherers,” significant consump-
tion of cariogenic plant foods acquired by trade from agricultural villagers results in appre-
ciable prevalences of caries (Walker & Hewlett, 1990; and see later). At Mac Bac, Northern
Vietnam, caries prevalence of 11% in adults is considerably higher than that of other
contemporary mid-Holocene populations in the region (Oxenham & Dommett, 2011). There
is no evidence for rice consumption, suggesting that other plant sources of carbohydrates
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3.2 Dental caries 73
(e.g., taro) may have led to such a remarkably high prevalence of caries (compare with Tayles,
1999; Temple, 2007). Similarly, in other settings of late Pleistocene and early Holocene
populations where nondomesticated plant carbohydrates were consumed, dental caries preva-
lence is highly elevated, especially in comparison with most hunter-gatherers (Da-Gloria &
Larsen, 2014; Humphrey et al., 2014).
Changes in diet and subsistence technology had far-reaching implications for oral health in
some prehistoric foragers. Walker and Erlandson (1986) examined the link between dental
caries and dietary change on Santa Rosa Island, in the Santa Barbara Channel Island region of
southern California. Archaeological evidence indicates that exclusively foraging groups
occupied the island from c. 4000 to 400 yBP. For the first 1500 years of this timespan,
populations exploited predominantly terrestrial foods, primarily starchy roots and tubers.
For the remainder of the prehistoric period, diet became increasingly focused on marine
resources (see Chapter 2). A decrease in adult caries prevalence from 13.3% to 6.3% coincides
with this subsistence change, which appears to be linked to the reduction in use of plant
carbohydrates in later prehistory (Walker and Erlandson, 1986).
Other foraging groups show the opposite trend in caries prevalence. In Archaic-period
hunter-gatherers living in the Edwards Plateau of central Texas, there is an increase in
cariogenesis (Bement, 1994). This increase may reflect a reorientation of diet from a general-
ized foraging pattern involving a range of wild plants and animals to a diet focused more on
high-carbohydrate, cariogenic wild plants (Bement, 1994).
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74 Exposure to infectious pathogens
Differences in caries prevalence between males and females suggest that food consumption,
both in diet and behavior, may have been different between sexes, with males consuming
more animal sources of protein than do females and females consuming more plant carbohy-
drates than do males. This conclusion is consistent with differences documented in female and
male subsistence behavior in historic and recent agriculturalists and foragers. For example,
Southeastern North American Indian women were responsible for most plant gathering and
agricultural activities such as planting, harvesting, and food preparation. Men were respon-
sible for hunting as their primary subsistence task (Hudson, 1976; Swanton, 1942, 1946; Van
Doren, 1928). Among foragers living at Ngarulurutja, Australia, Hayden observed “that in
spite of rules about sharing, the persons who did the most hunting ate the most meat. It is
clear that the young men who actually caught the game consumed most of it” (1979:166) (see
other accounts of sex differences in diets in various traditional contexts by Hawkes et al.,
2001; Hayden, 1979; Hewlett et al., 1982; Holtzman, 2009; Lee, 1968; McArthur, 1960;
Meehan, 1977; O’Connell et al., 1992; Okazaki et al., 2013; Walker & Hewlett, 1990; Wood-
burn, 1968).
Sex differences in caries prevalence in archaeological samples are identified in populations
practicing agriculture to some degree. However, female and male foragers from the Santa
Barbara Channel Island region display differences in caries prevalence for much of prehistory
(Lambert & Walker, 1991; Walker & Erlandson, 1986). Ethnographic observations indicate a
distinctive sexual division of labor in which men hunted and fished and women collected
plants. Historic accounts indicate that men ate more of the game they hunted than did women.
Thus, it appears that greater consumption of plants by women – a factor related to their
subsistence responsibilities – is reflected in their greater caries prevalence. Similarly, con-
sumption of cariogenic plants (wild tubers and fruits) is hypothesized to explain the higher
prevalence of dental caries in females than in males in Paleoamericans from Lagoa Santa,
Brazil (Da-Gloria & Larsen, 2014). In this setting, while nearly all tooth types show greater
caries prevalence in females, the differences are especially pronounced in the second and third
molars.
Walker and Hewlett (1990) investigated dental caries in several groups of central African
pygmy foragers (Aka, Mbuti, Efe) and Bantu farmers in order to build support for a behavioral
interpretation of prevalence variation. Comparisons of subsistence patterns and food con-
sumption practices in these groups reveal clear intra- and inter-population differences in
caries prevalences that provide an important perspective on dietary behavior and cariogen-
esis. Aka and Mbuti foragers practice net-hunting, and the Efe foragers bow-hunting. The
sources of animal protein available to these groups vary, depending on the season (e.g., meat
from seasonally available animals), and is mostly obtained through hunting and collecting.
Manioc and other cultigens acquired through trade with Bantu agriculturalists form a signifi-
cant component of their diets. The relative consumption of meat to cultigens is highly
variable, ranging from a high proportion for the Mbuti (at least part of the year) to a much
lower proportion for the Efe. The Aka apparently consume similar amounts of meat as Mbuti
foragers as both are net-hunters. Based on comparisons of the amount of time spent hunting,
it appears that Mbuti and Aka foragers consume more meat than do Efe foragers. In addition
to meat and cultigens, honey is an important component of diet for part of the year. For Mbuti
foragers, this highly cariogenic food contributes nearly 80% of calories for a one-month
period (Ichikawa, 1981).
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3.2 Dental caries 75
Table 3.1 Sex differences in central African dental caries prevalences: Aka, Mbuti,
Efe, Bantu (Adapted from Walker & Hewlett, 1990: Table 2)
Teeth (n) % caries
Tribe and sex
Aka
Total (3099) 5.2
Males (1706) 4.2
Females (1393) 6.6
Mbuti
Total (1773) 6.0
Males (1048) 5.1
Females (753) 7.3
Efe
Total (277) 6.0
Males (277) 6.0
– –
Females
Bantu
Total (630) 8.1
Males (308) 9.1
Females (322) 7.1
Virtually all foods consumed by Bantu villagers are plant domesticates, including manioc,
maize, rice, peanuts, and plantains. The little amount of meat that is consumed –
approximately 2.5% of foods – is acquired from pygmy foragers. Ethnographic field observa-
tions indicate differences in food consumption between females and males in the foragers, but
not in the farmers. Diets of forager women contain more plants than those of forager men.
Aka men consume more meat than do women, which is acquired on the hunt prior to their
return to the home village; some of the choicest cuts of meat are shared among the men who
participated in the hunt.
Bantu farmers have the higher dental caries prevalence in comparison with foragers
(Table 3.1), which appears to reflect their greater consumption of plant carbohydrates.
Although foragers have relatively lower caries prevalence, the values, nevertheless, are
appreciable (5% – 6%: Table 3.1), which points to the significant component of plant carbo-
hydrates acquired from Bantu farmers via trading relationships. Caries prevalence is also
higher for pygmy women than men. These differences are related to food consumption
variation between male and female pygmy foragers. The frequency of between-meal eating
in pygmy women is an additional factor that provides partial explanation for sex differences.
In these foragers, men concentrate their eating into several large meals, and women snack
frequently during the day. Clinical evidence from Western populations indicates that snacking
between meals (especially carbohydrates) results in elevated caries rates (Burt et al., 1982,
1988; Gustafsson et al., 1954; Konig, 1970; Konig et al., 1969; Nizel, 1973; Rowe, 1982; Weiss
& Trihart, 1960; but see Rugg-Gunn et al., 1984).
Ethnographic documentation of dietary practices in native groups in South America
provides additional insight into differences between females and males in dental health
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76 Exposure to infectious pathogens
(Walker et al., 1998). In the three groups studied – the Yanomami of Venezuela, the Yora of
southeastern Peru, and the Shiwiar (Achuar) of Ecuador – meat and fish provide a significant
part of the diet, and most carbohydrates are from plant crops, such as manioc and bananas.
All three groups have significant caries rates, in part due to the consumption of cultigens, but
also due to the reduction in their isolation and greater access to processed foods. This is an
unexpected finding because Yora and Shiwiar women spend many hours processing manioc
in their mouths for production of beer (chicha). For these two groups, women display a higher
frequency of carious teeth than men. Their relatively greater exposure to cariogenic manioc
explains these higher caries rates.
Given the finding of predominantly higher prevalence of dental caries in females than in
males in archaeological contexts discussed here and in living populations (Lukacs, 2008;
Lukacs & Thompson, 2008; Sauerwein, 1974), non-behavioral reasons relating to life history
factors may also contribute in important ways to this prevalence sex dichotomy. Permanent
teeth erupt slightly earlier in females than in males, exposing their teeth at an earlier age to
caries-promoting factors (Carlos & Gittlesohn, 1965; DePaola et al., 1982; Dunbar, 1969).
However, tooth eruption differences between males and females show either weak or no
correlation with dental caries prevalence (Moorrees, 1957; Toverud et al., 1952; Ziskin, 1926)
and the absolute temporal difference in tooth eruption is not significant (12 – 16 months).
Much more likely than tooth emergence timing as a cause for sex differences in caries
prevalence are factors relating to changes in the oral environment during pregnancy owing
especially to increased production of estrogens and progesterone (Laine, 2002). In this regard,
there is considerable evidence for increasing gingival inflammation in pregnant women
(Arafat, 1974; Laine, 2002; Loe, 1965; Loe & Silness, 1963; Maier & Orban, 1949; but see
Jonsson et al., 1988). At least one meta-analysis shows, however, that inflammation is
restricted to the gingiva, does not affect the tooth supporting tissue, especially the periodontal
ligament, and generally does not result in tooth exfoliation (Laine, 2002). In addition to these
changes, bacteria are altered in the oral environment, including an increased presence of
Streptococcus mutans and other flora associated with cariogenesis, and there is a lowering of
pH. However, most investigations of caries and pregnancy are largely cross-sectional or with
limited follow-up (Laine, 2002). Moreover, because dental caries is a process that unfolds over
several years or longer, it is difficult to determine a link between caries and pregnancy. The
record is also silent on the rate of caries during pregnancy versus before or after pregnancy.
Certainly, pregnancy influences the oral environment, which could result in increased cario-
genesis. The combined effects of more cariogenic bacteria, decreased pH, and the altered
buffer effect of saliva may contribute to increased dental caries in pregnant women. On the
other hand, the research tracking change in the oral environment shows that oral buffering
capacity fluctuates during pregnancy – decreasing, increasing, then decreasing – and increas-
ing dramatically after birth (Laine et al., 1988). Therefore, the idea that the oral environment is
simply more cariogenic in pregnancy does not take into account the full pattern of oral
buffering capacity during (or after) pregnancy.
The comprehensive investigation linking increased caries to life history in women by
Lukacs and coworkers (Lukacs, 2008; Lukacs and Thompson, 2008) and potential links with
increased fertility and birthrate in the Holocene, and the additive effects of a greater number
of pregnancies in agricultural settings presents a convincing scenario that may explain, in
part, the differences in dental caries prevalence of adult males and females globally.
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3.2 Dental caries 77
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78 Exposure to infectious pathogens
to animal sources of protein and greater diversity of diet in elite individuals (Cucina & Tiesler,
2003). The greater nutritional quality, including relatively lower carbohydrate (maize) con-
sumption in elites, explains their better oral health compared to non-elites.
On the north coast of Peru, there appears to be significant status-based variation in dental
caries in the Middle Sicán theocratic state (AD 900 – 1100) (Shimada et al., 2004; Klaus,
Wilbur et al., 2010). The Middle Sicán featured rigid social stratification with multiple
archaeological reflections of immense gulfs of power between the elite Sicán and the popula-
tion of Muchik commoners they ruled over. Dental caries prevalence is significantly lower
among the Middle Sicán elite, who probably consumed more protein resources and less
carbohydrates, as opposed to the Muchik commoners, whose higher caries prevalence indicate
consumption of relatively greater proportions of starchy cultigens in their diets. This pattern
of lower caries prevalence in elites is also strongly pronounced in complex societies in the
Atacama Oases of Chile. Like the Middle Sicán elite, adult elite from Solcor 3 and Coyo 3 sites,
especially males, express considerable differences (Hubbe et al., 2012). In this setting, the
better oral health in elites appears to be related to their having relatively greater access to
animal sources of protein.
Winkler and coworkers (2012) test the hypothesis that the relative position of skeletal
remains located closest to the ritual nucleus of the church (the altar) at the seventeenth-
century mission, Santa Catalina de Guale, Georgia, reflects higher social position in the
community, and hence, greater access to resources and better living circumstances. To test
this hypothesis, they undertook a location analysis of dental caries as an indicator of dietary
quality, nutritional inference, and generalized stress. Status rank was identified on the basis of
elaborateness of grave inclusions (e.g., presence of glass beads) and burial treatment (e.g.,
inclusion in a coffin). Teeth of individuals interred closest to the altar were significantly less
carious than those of individuals interred furthest from the altar. Their study suggests that
individuals buried closer to the altar had a higher-quality diet and greater access to resources
than those buried further from the altar. These results strongly suggest clear social distinctions
in health and well-being in this colonial setting of North America.
An argument for greater access to animal protein in explaining better dental health in high-
status members of human populations is supported by ethnographic documentation of status
differences in dental health in African foragers. Walker and Hewlett (1990) found that high-
status pygmy leaders have fewer carious lesions than low-status non-leaders. They suggest
that greater access to meat by leaders (from gifts and tribute), combined with reduced
consumption of carbohyrate-rich plants, best explains the discrepancy between social ranks.
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3.3 Periodontal disease (periodontitis) and tooth loss 79
(a)
masticatory loading of the jaws and teeth. These mechanical demands are due to either
consumption of hard-textured foods or excessive extramasticatory practices, such as process-
ing of animal hides (Bement, 1994; Clarke & Hirsch, 1991; Marks et al., 1988; Molnar, 1972;
Pedersen & Jakobsen, 1989). Extreme mechanical demands on anterior teeth in northern
latitude populations result in severe tooth wear, pulp exposure, and resorption and shortening
of tooth roots. These factors contribute to tooth loss (Pedersen & Jakobsen, 1989).
Periodontal disease or periodontitis is a very common disease in industrialized countries
today (about a third of the adult population in the United States: Buhlin et al., 2003). It is
generally characterized by a loss of alveolar bone, represented either by a horizontal lowering
of the alveolar crest relative to the neck of the tooth (cementoenamel junction) or as pockets
of bone rarefaction (Hildebolt & Molnar, 1991) (Figure 3.3a). In the clinical setting, horizontal
bone loss is determined by the distance between the alveolar crest (interproximal bone) and
the cementoenamel junction. This dimension may not always represent an appropriate
indicator of periodontal disease, especially in anthropological populations where continuous
eruption during adulthood occurs in response to occlusal wear (Clarke, 1993; Clarke & Hirsch,
1991; Whittaker et al., 1985). Clarke and Hirsch suggest that periodontal disease in skeletal
remains is identified when “the crestal margin of bone undergoes loss of the surface cortical
bone, exposing the porous cancellous structure of the supporting bone, usually with an
accompanying change of the contour of the crest” (1991:241). Thus, exposure of tooth roots
as representing periodontal disease or simply compensatory continuous eruption owing to
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80 Exposure to infectious pathogens
(b)
Figure 3.3b Edentulous individual; Santa Catalina de Guale Santa Maria, Amelia Island,
Florida. (From Larsen, 1994; photograph by Mark C. Griffin.) (A black and white version of
this figure will appear in some formats. For the color version, please refer to the plate
section.)
wear and mechanical loading must be based on the quality of the alveolus – rough, irregular,
and porotic in periodontal disease, but thin, knife-edged, and dense in continuous eruption
(Ogden, 2008).
Once teeth are gone (Figure 3.3b), the soft tissue heals, and the alveolus remodels com-
pletely. Although the progression of periodontal disease is well documented in human
populations, ancient and modern, it remains an etiological conundrum. There is general
agreement that bacteria – perhaps as many as 40 different taxa – may be involved in the onset
and progression of the disease (Drake et al., 1993; Enwonwu, 1995; Li et al., 2000; Slots, 2004;
van Winkelhoff and Slots, 1999). Other important influencing factors include poor oral
hygiene, cariogenesis, malocclusion, nutritional status, and to a lesser extent, pregnancy,
puberty, and psychological stress (Hildebolt & Molnar, 1991). Some individuals appear to be
more susceptible, due in part to the presence of inflammatory gene polymorphisms (cytokines)
that determine greater presence of inflammatory serums (e.g., C-reactive protein: D’Aiuto
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3.3 Periodontal disease (periodontitis) and tooth loss 81
et al., 2004). Those individuals with genotypes specific for these cytokines appear to have
elevated levels of systemic inflammation with periodontal infection exacerbating the already
overloaded inflammatory burden. However, these genotypes are rare, and the general levels of
elevated periodontitis in human populations are largely due to environmental factors, espe-
cially diet and oral hygiene.
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82 Exposure to infectious pathogens
45
30
Prevalence AMTL (%)
25
20
15
10
0
1 2 3 4 5 6
Age Class
Figure 3.4 Comparison among age classes of posterior antemortem tooth loss (AMTL) in
late pre-Hispanic and postcontact periods, Lambayeque Valley Complex, Peru. Age classes
are defined as 0–4.9 yr (1), 5.0–14.9 yr (2), 15.0–24.9 yr (3), 25.0–34.9 yr (4), 35.0–44.9 yr
(5), and 45þ yr (6). (Adapted from Klaus & Tam, 2010; reproduced with permission of
authors and John Wiley & Sons, Inc.)
Other settings in Europe, Asia, and Africa mirror the trends documented in North America.
Seventeenth-century British mandibles exhibit lower crestal bone than Anglo-Saxon (sixth
century AD) mandibles (Lavelle and Moore, 1969). Increasing prevalence of periodontal
disease in this setting is due to the greater consumption of softer foods in the later period,
mostly arising from improvements in milling flour for bread and increased consumption of
sugar and refined carbohydrates generally. This trend closely parallels rapid increases in
dental caries in Britain for the same period (Hardwick, 1960; Moore & Corbett, 1975).
On the Indian subcontinent, Lukacs (1992) documented tooth loss in an extensive series of
human remains from various sites dating from the Mesolithic through the Iron Age, a
temporal framework representing the transition from foraging to intensive agriculture. The
later agricultural populations consumed a variety of domesticated plants (wheat, barley, peas,
sessamum). Comparisons of the earlier and later populations reveal an increase in tooth loss
resulting from an increase in consumption of plant carbohydrates (Lukacs, 1992). Similar
patterns of increased loss are found with the adoption and later intensification of agriculture
in Southeast Asia (Douglas, 2006) and South America (Alfonso et al., 2007; Klaus & Tam,
2010) (Figure 3.4).
Declining oral health has been identified in the Nile Valley in comparing dentitions from
four successive periods: Mesolithic (c. 15 000 yBP), Meroitic (350 BC – AD 350), X-group
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3.3 Periodontal disease (periodontitis) and tooth loss 83
(AD 350 – 550), and Christian (AD 550 – 1300) (Rose et al., 1993). The Mesolithic group is
characterized by a foraging economy based largely on seed gathering and hunting. Popula-
tions in the three later periods were irrigation farmers who grew cereal grains. All samples
show evidence of tooth loss. However, the Mesolithic group possessed only two teeth lost prior
to death (from 400 sockets representing 39 individuals). Later, the prevalence steadily rises
from 9.9% (Meroitic) to 34.6% (Christian). Rose and coworkers (1993) suggest that tooth loss,
although multifactorial, was due to extensive caries and excessive occlusal wear, which
predisposed teeth to decay and eventual loss. Some of the tooth loss may be due to the use
of stone grinding implements and incorporation of grit into food. The resulting wear and pulp
exposure contributes to periodontal disease and tooth loss in other areas of the Nile Valley
(Marion, 1996).
Prevalence of periodontal disease may be overestimated in archaeological samples. Hyper-
eruption of teeth (and eventual loss) may represent a normal aging process bearing little
relation to pathological processes (Clarke & Hirsch, 1991; Whittaker, 1993; Whittaker et al.,
1985). However, the important role of the dentition in food consumption argues that tooth
loss is an indicator of compromised health.
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84 Exposure to infectious pathogens
25
Male
Female
20
% Antemortem Tooth Loss
15
10
0
I1 I2 C P3 P4 M1 M2 M3 I1 I2 C P3 P4 M1 M2 M3
Maxilla Mandible
Figure 3.5 Percentage of antemortem tooth loss in males and females; Canary Island
archipelago. (Lukacs, 2007b; reproduced with permission of author and John Wiley & Sons,
Ltd.)
In the Ipiutak series, males have a very low loss of incisors (5.3%); females display a high loss
of incisors (19.4%). Hrdlička (1940a) argued that tooth loss in this region was related to the
practice of tooth ablation whereby the anterior teeth were intentionally removed (Cook,
1981b; and compare with Temple et al., 2011). Re-examination of cultural practices in these
groups suggests that greater anterior tooth loss in adult females is related to the excessive use
of front teeth in extramasticatory activities (Costa, 1980). Women in this setting engage in
behaviors that place excessive mechanical demands on the anterior teeth, such as hide
chewing. The cumulative trauma and excessive dental wear result in early loss of incisors
and canines. Thus, tooth loss differences between men and women can clearly be related to
use of the dentition in extramasticatory behaviors. The links with trauma are also well
illustrated in sex difference in antemortem tooth loss in the Canary Islands (Lukacs, 2007b),
where males generally have greater antemortem loss than females (Figure 3.5). In this setting,
antemortem losses are due to multiple factors, including accidents from walking in difficult
terrain and ritual and non-ritual forms of interpersonal violence involving mostly males. This
body of work demonstrates the complex nature of antemortem tooth loss relating to a range
of factors. The record globally, past and present, is largely explained by oral health patterns
influenced by diet, but behaviors not involving food and its mastication are important.
The influence of status and social rank on tooth loss in human populations has rarely been
systematically assessed in archaeological remains. In the Medieval series from Zalavár,
Hungary, tooth loss is considerably higher in low-status individuals buried in the chapel
(39.4%) than in high-status individuals buried in the castle (9.1%) (Frayer, 1984). The
difference between status groups is especially striking in adult males: 48.4% of teeth were
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3.3 Periodontal disease (periodontitis) and tooth loss 85
missing in the low-status group and only 5.2% were missing in the high-status group.
Females show prevalences of 32.1% and 6.7% for low- and high-status groups, respectively.
As with the dental caries prevalences from this series, sex differences in tooth loss indicate
dental health variation between men and women that is strongly influenced by sexual
dimorphism in diet and food consumption practices. In contrast, high-status persons from
the aforementioned Calakmul, Dzibanché, and Kohunlich in the northern Petén have higher
antemortem tooth loss than low-status persons (Cucina & Tiesler, 2003). In this setting, high-
status individuals express greater presence of calculus, which contributes to irritation to and
inflammation of gingival and periodontal tissues and increased risk of tooth loss. Overall, a
more refined, softer diet contributed to worse oral health in elite persons. A similar pattern is
present in Shang dynasty China where reduced tooth wear suggests a softer, more processed
diet and an oral environment more conducive to poor oral health in the comparison of
“citizens” and “slaves” (Sakashita et al., 1997).
3.3.4 Implications of poor oral health for general health: the mouth as a window
to systemic health
A wide range of clinical and epidemiological investigations have examined the relationship
between poor oral environment and general health, especially with regard to adult mortality
and chronic, systemic health conditions such as cardiovascular disease, respiratory infections,
and other diseases (Ajwani et al., 2003; Buhlin et al., 2003; DeStefano et al., 1993; Hollister
and Weintraub, 1993; Irwin et al., 2008; Johnson et al., 2006; Joshipura et al., 1996; Koren
et al., 2011; Li et al., 2000; Pihlstrom et al., 2005; Slots, 2004; Williams et al., 2008).
Periodontal disease and cardiovascular disease show the clearest association, with perio-
dontitis associated with increased risk of coronary heart disease. The mechanism is unclear
and confounding factors such as age, diet, sex, and smoking history are problematic in the
design of research and specific links. It may be the case that periodontal disease is the root
cause of systemic infection and the attendant inflammatory response, which may in turn
promote an inflammatory response in cardiovascular tissues. Nonetheless, the oral environ-
ment is a clear target for developing an understanding of adult systemic conditions and it is a
risk factor for increased mortality linked to cardiovascular conditions, pulmonary disease,
other evidence of morbidity, and increased mortality. In the last couple of decades, about 90%
of adults globally express periodontal disease. It is clear, therefore, that reconstructions of oral
health are crucial for understanding general health (DeWitte & Bekvalac, 2011; Pihlstrom
et al., 2005; Slots, 2004).
The links between inflammation associated with oral health and elevated mortality from
cardiovascular disease in living populations suggest the possibility of links between oral
health status and mortality in past populations. Indeed, the association between poor oral
health and increased risk of death (based on mortality analysis) is clearly illustrated in a large
series of remains from the medieval St. Mary Graces cemetery in London (DeWitte & Bekvalac,
2010). In this setting, there is a statistically significant relationship between oral health and
mortality: individuals with dental caries and periodontal disease are at greater risk of dying
than those without the conditions. Similarly, individuals having periodontal disease were far
more likely to have periosteal reactions than individuals not having periodontal disease
(DeWitte & Bekvalac, 2011). Historical sources show that regardless of status, individuals
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86 Exposure to infectious pathogens
living in London at the time were consuming, as the largest part of their diet, wheat, barley,
and rye in various forms (e.g., bread, ale, cereal mixtures). This, combined with poor oral
hygiene, explains the elevated levels of oral disease in this setting. Those who enjoyed better
diets (especially including animal sources of protein) almost certainly had better oral and
systemic health, and hence, lower risk of death (DeWitte & Bekvalac, 2010, 2011).
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3.4 Nonspecific infection and disruption 87
Figure 3.7 Osteomyelitis involving diaphysis of adult tibia and fibulae; King site, Georgia.
(Photograph by Clark Spencer Larsen.)
irregular elevations of bone surfaces (Figure 3.6). The skeletal tissue in the unhealed form is
loosely organized new bone. In the healed form, the skeletal tissue is incorporated into the
normal cortical bone and the surface is often smooth, undulating, and somewhat inflated. The
lesions can be highly localized, being limited to single skeletal elements, but they may also
involve multiple elements if the infection is widespread or systemic. Osteitis is usually not
identifiable without radiological observation. As most paleopathological studies do not
involve radiological analysis, osteitis is not discussed here.
Osteomyelitis involves exuberant proliferation of both endosteal and periosteal bone
surfaces (Figure 3.7), the former of which results in the restriction in diameter of the medullary
cavity. Most pyogenic (pus-producing) osteomyelitis is caused by the microorganism
Staphylococcus aureus (Aufderheide & Rodríguez-Martín, 1998; Rosenberg, 1994). The arter-
ial system of the bone is the typical route of its transport. Direct infection from a bone fracture
and break in the skin is also linked with osteomyelitis. The infection site on the bone is
sometimes associated with sinuses or holes (cloacae) for exudate or pus drainage. Chronic
osteomyelitis can occur over a period of many years due to the presence of localized infection
foci that occasionally reappear, sometimes in response to systemic or localized stress.
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88 Exposure to infectious pathogens
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3.4 Nonspecific infection and disruption 89
them more susceptible to infection (Ortner, 2003). Additionally, the anterior and lateral
aspects of the tibial diaphysis have the largest, and perhaps the most vascular and physiolo-
gically inactive surfaces in the skeleton, which may also lead to bacterial colonization and
infection (Martin et al., 1991). Circulatory flow is generally slower in the lower legs because of
gravity, enhancing the potential for bacterial colonization (Kumar et al., 2010). The anatom-
ical location – the lower end of the legs – also exposes the anterior tibia to trauma against
which little protection is offered by soft tissue. Subcutaneous and subperiosteal bruises from
trauma promote periosteal reactions. In addition, these injuries also promote bacterial prolif-
eration through release of blood and intracellular fluids from ruptured cells and vessels,
resulting in a bony reaction.
In the Eastern Woodlands of North America, several studies show important links between
subsistence, settlement pattern, and community health. In the Tennessee River valley, non-
specific periosteal lesions increase in farmers relative to earlier foragers (Danforth et al., 2007).
Similarly, in the Dickson Mounds site in the Illinois River valley, Lallo and coworkers
(Goodman, Lallo et al., 1984; Lallo et al., 1978; Lallo & Rose, 1979) compared infection
prevalence in two populations, including an earlier Woodland (called Mississippian Accultur-
ated Late Woodland) group (AD 1050 – 1200) and a later Mississippian (Middle Mississippian)
(AD 1200 – 1300) group. The later period saw an increase in consumption of maize and
reduction in animal sources of protein. There appears to have been a consolidation of
residential units into larger population aggregates at the same time. Overall, settlement
involved a marked increase in population density, decrease in mobility, and conditions that
generally provide the context for increased infection.
There is a dramatic increase in frequency and severity of infectious lesions (periostitis and
osteomyelitis) that coincides with these changes in population settlement at Dickson Mounds,
Illinois. The prevalence doubled in the later period, from 30.8% to 67.4% of individuals
affected. For the tibia, the prevalence increased from 26% to 84%, affecting all age groups
and both sexes. Comparisons of severity (from slight to severe involvement of the periosteum)
for the tibia showed that most of the infection was slight in the Woodland group whereas most
was moderate to severe in the Mississippian group.
This increase in prevalence and severity of infection could be explained in a number of
ways. The introduction of new chronic infectious disease(s) may have occurred in later
prehistory. Lallo and coworkers argue that the increase in infectious lesions in the thirteenth
century AD is possibly tied to several interlinking factors. They highlight the role of decline in
nutritional quality and sedentism in later prehistory, which likely placed populations at
increased risk of infection by decreasing the resistance to disease and by providing lower
quality of living circumstances.
Archaeological evidence suggests that there was an increase in trade networks and long-
distance social contact in the later prehistoric period. These contacts may have provided a
means for introducing new pathogens or disease vectors, or both, thus increasing the
prevalence of infectious disease. The effects of population size increase and sedentism are
well understood in infectious disease ecology and epidemiology. By increasing the size and
density of settlements, the host and pathogen are placed side-by-side in a long-term relation-
ship that may form the basis of chronic infection. The number of potential hosts is increased,
thus providing a permanent reservoir for certain infectious agents. The closer contact in a
more densely occupied settlement, coupled with the ill effects of poor sanitation resulting
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90 Exposure to infectious pathogens
from permanent occupancy of a setting, results in faster and more proficient disease trans-
mission (Armelagos, 1990; Armelagos & Dewey, 1970; Lambert, 1993; Lallo et al., 1978;
Larsen, 1982). Thus, the overall picture is one of declining community health as population
concentration increases. Moreover, the negative impact of infection on the population is
indicated by depressed survivorship for those individuals with skeletal lesions. For example,
Dickson adults (>20 years of age) with severe infections had an average age-at-death of 29.3
years, which is well below the mean age-at-death for the adult population overall (33.5 years)
(Lallo et al., 1978). This finding is consistent with the notion that individuals with elevated
stress earlier in life are predisposed to earlier death in adulthood (see Chapter 2).
Other settings from the Eastern Woodlands show results that are generally consistent with
changes observed in the Dickson Mounds populations. Comparisons of prehistoric foragers
(pre-AD 1150), prehistoric maize farmers (AD 1150 – 1550), early mission (AD 1607 – 1680),
and late mission-period intensive maize agriculturalists (AD 1686 – 1702) in the Georgia Bight
show clear temporal trends in prevalence in relation to dietary and lifeway changes (Larsen
and Harn, 1994; Larsen, Crosby et al., 2002; Larsen, unpublished; and see earlier for a more
detailed description of the samples).
Unlike the Dickson Mounds study, these analyses focus on comparisons of populations
consuming exclusively wild plants and animals with populations utilizing maize to varying
degrees. For most of the time prior to the arrival of Europeans in the sixteenth century, native
groups were nonsedentary foragers who obtained most foods from a combination of hunting,
gathering, and fishing. Archaeological and isotopic documentation of subsistence economy
indicates that marine resources from estuarine and marine contexts provided most of the
protein in native diet (Hutchinson et al., 1998; Larsen, Schoeninger et al., 1992; Reitz, 1988,
1990). During the twelfth century AD, maize rapidly took on an increasingly important
dietary role. With the arrival of Europeans and the establishment of Catholic missions, there
was a subsistence reorientation whereby maize dominated diet. Coupled with these dietary
changes were alterations in population size, density, and sedentism. Prehistoric foragers in the
region appear to have been sparsely settled and highly mobile. Prehistoric farmers, however,
were living in larger, more densely occupied villages, probably for longer periods of time
(Larsen, 1982). In the mission period, populations were coerced into living in and around
mission settlements. Although population size reduced dramatically during the contact
period, the settlements were permanent and villages were crowded.
Prevalence comparisons for periosteal reactions in adult tibiae indicate that there is an
increase in frequency prior to contact from 9.5% to 19.8%. In the early mission population,
the prevalence declined slightly to 15.4%, but then greatly increased to 59.3% in the late
mission period. These general increases affected both adult males and females. The moderate
increase in the precontact populations is well under that observed at Dickson Mounds. In all
likelihood, the relatively lower prevalence in the Georgia Bight reflects considerably smaller
population size and less of a commitment to maize agriculture in the later prehistoric period in
comparison with interior Mississippian populations (e.g., Dickson Mounds). The marked
increase in the late mission period is probably tied to the relocation and increased concen-
tration of native populations around mission centers and introduction of new diseases,
possibly including venereal syphilis. The change in settlement provided conditions conducive
to the maintenance and spread of chronic infectious diseases and other factors that lead to an
increase in bone lesions (Larsen, Crosby et al., 2002; Larsen & Harn, 1994; Larsen et al., 2007).
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3.4 Nonspecific infection and disruption 91
The effects of increased infection rates would likely have been exacerbated by the increase in
dietary emphasis on nutritionally poor foods, especially maize.
There is also a synergy between infection and malnutrition (Keusch & Farthing, 1986;
Scrimshaw et al., 1968) – malnourished individuals are less resistant to infectious pathogens
and are rendered more susceptible to infectious disease; conversely, infection worsens nutri-
tional status. In understanding the increase in infection in these archaeological (and other)
settings, the synergy with nutrition is critical. The consequences of infection and nutrition are
worse than either factor acting alone. Individuals experiencing infection exhibit higher basal
metabolic rates, which are accompanied by fever and the body’s increased demand for protein
and other nutrients necessary for the production of antibodies that fight the infection.
Therefore, in the setting of reduced nutritional quality, first in the late prehistoric and then
in the mission context, the ability to mitigate infection would likely have been hampered by a
reduction in dietary quality. Thus, infection increased in the mission period, in large part due
to the compromised health linked to poorer nutrition and population crowding.
These studies provide strong support for the epidemiological model that an increase in
population size and density often contributes to decline in community health, at least as it is
measured by prevalence of skeletal lesions. This general pattern of increase in infections with
the transition from foraging to farming or agricultural intensification is also documented in a
variety of other areas of the Eastern Woodlands undergoing the shift from foraging to farming
(Cassidy, 1984; Cook, 1984; Danforth et al., 2007; Garner, 1991; Hoyme & Bass, 1962;
Katzenberg, 1992; Pfeiffer & Fairgrieve, 1994; Rose et al., 1984, Rose, Anton et al., 1991) or
in single-component late prehistoric settings in the Eastern Woodlands (Boyd, 1986; Eisen-
berg, 1986, 1991a, 1991b; Gold, 2004; Magennis, 1986; Milner, 1982, 1991, 1992; Milner &
Smith, 1990; Powell, 1986, 1988, 1989). Increase in infection prevalence is also well docu-
mented in the American Southwest (Martin et al., 1991; Stodder, 1994; Stodder & Martin,
1992; Stodder et al., 2002), Mesoamerica (Hodges, 1989; Márquez Morfín & Storey, 2007;
Márquez Morfín et al., 2002; Norr, 1984; Saul, 1972; Storey, 1992a), South America (Klaus &
Tam, 2009; Pechenkina, Vrandenburg et al., 2007; Ubelaker, 1984, 1994), and in a few
regions of the Old World (e.g., Japan: Suzuki, 1991; Europe: Cunha et al., 2007; Grauer,
1991; Roberts & Cox, 2003, 2007).
This is not to say that increased population aggregation in farming societies invariably led
to the same levels of bone infection prevalence globally or even within broad regions. For
example, evidence of generalized infection and other health indicators are low in a number of
settings in Southeast Asia (Clark et al., 2014; Domett & Tayles, 2007; Douglas & Pietrusewsky,
2007). Community health in late prehistoric populations in eastern North America was highly
variable (Milner, 1991). Some late prehistoric agriculturalists had very high prevalences of
bone infections (Eisenberg, 1986; Lallo et al., 1978; Milner & Smith, 1990), while other groups
had somewhat lower prevalences (Larsen, 1984; Larsen, Crosby et al., 2002; Larsen & Harn,
1994; Larsen et al., 2007; Milner, 1991). Some of this variability is undoubtedly due to inter-
observer differences in recording methods. The variable pattern of infection prevalence also
points to a high degree of diversity between human groups occupying very different land-
scapes and physiographic zones, ranging from highly fertile river bottoms (Lallo et al., 1978)
to marginal uplands (Eisenberg, 1986) or coastal regions (Larsen, 1982). Detailed analysis of
population trends indicates that population histories fluctuated dramatically, with regard to
both size and distribution (Milner, 1990). Living in peripheral settings did not provide freedom
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92 Exposure to infectious pathogens
from disease – some of the highest prevalences of bone infection are in the so-called
“marginal” habitats (Eisenberg, 1986, 1991a, 1991b).
Some evidence suggests that high population density and disease burdens, in combination
with other factors (e.g., warfare), may have contributed to cultural terminations during later
prehistory well before the arrival of Europeans (Eisenberg, 1986; and see Larsen, 1994).
Improved survivorship, coupled with a decline in prevalence of periostitis and osteomyelitis
in Ontario, suggests that populations may have adjusted to high density in this setting
(Katzenberg, 1992). This decline contrasts sharply with other contact era settings where
periosteal infections have been shown to increase in a dramatic fashion (Larsen, Crosby
et al., 2002; Larsen & Harn, 1994; Larsen et al., 2007; Stodder, 1994; Ubelaker, 1994).
Timing of agricultural intensification may explain some of the variation in increasing
prevalences. In contrast to the findings from the analysis of populations from the Eastern
Woodlands, prevalence of periosteal reactions remained unchanged from earlier to later
periods in the Valley of Oaxaca, Mexico (Hodges, 1987, 1989). Unlike most settings in the
Eastern Woodlands, agricultural intensification in the Valley of Oaxaca was accompanied by
neither increased sedentism nor appreciable population growth. Unlike the Eastern Wood-
lands, agricultural development was long and gradual, taking place over several thousand
years. This contrasts with regions of secondary agricultural development, like the Eastern
Woodlands, where maize agriculture was adopted relatively rapidly. Hodges (1987) argues
that the longer period of human – plant interaction may explain some of the differences in
health declines between the Valley of Oaxaca and the Eastern Woodlands.
The shift to agriculture and increased population density in the Nile Valley of Sudanese
Nubia was also not accompanied by an elevation in frequency of bone infectious lesions. In
the X-group intensive agriculturalists (AD 350 – 550) in the Wadi Halfa area, only 12% of
individuals possess nonspecific bone infections, most of which are minor localized periosteal
reactions (Armelagos et al., 1981). This finding is especially surprising because the valley was
densely settled and populations experienced elevated stress (see Chapter 2).
Microscopic examination of femoral cortical bone from X-group individuals indicates a
pattern of fluorescence similar in many respects to tetracycline labeling in modern bone. This
analysis reveals the strong presence of tetracycline – now recognized as a broad spectrum
antibiotic – some 1400 years prior to its medical discovery in the mid-twentieth century
(Bassett et al., 1980; and see Armelagos, 2000; Bassett, 1981; Cook et al., 1989; Keith &
Armelagos, 1988; Maggiano et al., 2003, 2009; Mills, 1992; Nelson et al., 2010). Tetracycline
is highly effective against gram-negative and gram-positive bacteria as well as some other
pathogens; it may have had a highly therapeutic value for ancient Nubians. The source of
tetracycline is unclear, but Bassett and coworkers (1980) suggest that grains – wheat, barley,
and millet – stored in mud bins provided the environmental conditions and nutrients essential
for the growth of Streptomycetes, the bacteria that produce tetracyclines. Alternatively, the
pots used to store grain for brewing ale would have provided the environment suitable for
natural culturing of Streptomycetes (Armelagos, 2000). Similar patterns of bone fluorescence
have been found in skeletal remains in Kulubnarti, Sudan (Hummert & Van Gerven, 1982) and
several settings of the Dakhleh Oasis, Egypt (Cook et al., 1989; Maggiano et al., 2003, 2006,
2009), showing long-term continuance over centuries and significant regional presence. The
application of confocal laser scanning microscopy to the analysis of the Dakhleh Oasis series
in particular provides a record of fluorescence consistent with the presence of tetracycline
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3.4 Nonspecific infection and disruption 93
100 mm
Figure 3.8 Confocal laser scanning image of archaeological bone. Osteons are labeled (1)
and (2). Incorporation of tetracycline into the Haversian systems is shown in fluorescent
osteons (2) and lamellae (3). (From Maggiano et al., 2006; reproduced with permission of
authors and Elsevier Ltd.) (A black and white version of this figure will appear in some
formats. For the color version, please refer to the plate section.)
incorporated during life, likely deriving from consumption of stored grains (Figure 3.8).
However, it seems unlikely that therapeutic levels of tetracycline could have been maintained
in life, but bone infection may have been reduced owing to the presence of tetracycline in the
skeleton of affected persons (Maggiano et al., 2003).
Increased population density is not solely dependent upon the adoption of an agricultural-
based economy. A number of regions globally show an increase in sedentism and population
density in the absence of plant or animal domestication. If a chief cause for increasing skeletal
infection is related to demographic factors (i.e., population size and distribution), then
populations undergoing a shift to sedentism in foraging contexts should show similar changes
in infection prevalence as populations adopting agriculture. In order to test this hypothesis,
Lambert and Walker (1991; Lambert 1993, 1994) documented change in prevalence of
periosteal lesions in populations occupying the mainland coast and islands in the Santa
Barbara Channel Islands. Accompanying the shift toward a marine-based economy, there
was an increase in population size and density and decrease in mobility, especially in later
prehistory (Glassow, 1996). By the time of initial contact, native populations in the region had
a level of complexity of social organization and population density that rivaled many
agricultural societies in North America (Arnold, 1992).
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94 Exposure to infectious pathogens
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3.4 Nonspecific infection and disruption 95
Medieval England (Grauer, 1991), intensive agriculturalists from Mesoamerica and Peru
(Danforth et al., 1997; Hodges, 1989; Pechenkina, Vrandenburg et al., 2007), hunter-
gatherer-fishers in coastal southern California (Lambert, 1994), and colonial era African
Americans from New York (Null et al., 2009). Adult males and females show very different
temporal trends in a large series of post-Civil War African Americans in Texas (1869 – 1907;
Davidson et al., 2002). Males show a significant decline, whereas females show an increase.
Overall, males have a considerably higher prevalence than females. Both sexes, however,
display high frequencies of periosteal lesions indicating exposure to conditions conducive to
high levels of infection and infectious disease. This pattern is repeated in other samples of
African Americans, both enslaved and freed in North America (Rathbun & Steckel, 2002; and
see discussion in Blakey, 2001; Barrett & Blakey, 2011).
The tendency for greater male periostitis prevalences in these diverse settings is related to
factors that are unique to specific circumstances. In Spanish Florida, for example, resettlement
of male draft laborers in areas far from home villages may have exposed them to novel
pathogens or other infectious agents (Larsen and Harn, 1994). In the Santa Barbara Channel
Islands region, the propensity for males to participate in highly demanding physical activities
may explain a higher prevalence of periosteal reactions resulting from more blows to their
lower legs than in females (Lambert, 1994).
Several osteological samples show a tendency for adult females to have more nonspecific
lesions than adult males (Martin, 2008; Whittington, 1989). In the prehistoric series from the
La Plata Valley in northwest New Mexico, females show a much higher prevalence of lesions
than males (females¼30.7%; males¼6.2%) (Martin, 2008). At Black Mesa, Arizona, there are
no sex differences in infection prevalence, but females have more severe infections than
males. The finding of greater involvement in females, either with regard to prevalence or
severity, in these Southwestern settings may suggest a greater exposure to pathogens
resulting in infection. In the La Plata Valley, adult females have more skeletal injuries than
males, including cranial depression fractures and other broken elements. In contrast to male
burial, female burial was haphazard and devoid of grave goods. Based on this pattern, Martin
and coworkers (1991, 2001) suggest that adult females lived under suboptimal conditions, at
least in comparison with adult males. In Mórrope, Peru, postcontact females display signifi-
cantly higher periosteal lesion prevalence than their late pre-Hispanic counterparts, suggest-
ing, along with other lines of skeletal and contextual evidence, the relatively greater impact of
conquest on native women in this setting (Klaus & Tam, 2009).
Relatively little paleopathological research is devoted to the important link between status
and health, arguably an important component of complex societies (Powell, 1992a). If elite
members of a society were exempted from activities that would expose them to pathogens, they
should exhibit a relatively lower prevalence of bone infection. Comparisons of high-status with
low-status individuals at Moundville reveals no statistically significant prevalence differences
in infection, suggesting that “status differentiation at Moundville brought no substantial
biological benefits, nor levied any particularly heavy penalties” (Powell, 1992a:88). Compari-
son of skeletal elements of the high-status elites with the two non-elite groups indicates that,
with the exception of the fibula, all long bones from elite individuals have somewhat less
periostitis than those of non-elite individuals. For example, 44.8% of high-status and 51.0% of
low-status tibiae have infections (Powell, 1988: Table 35). Given the vagaries of identification
of high-status individuals as well as potential problems with sample size, these differences may
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96 Exposure to infectious pathogens
not be meaningful. Other southeastern US Mississippian centers also show no clear differences
between high and low status in infection prevalence (Blakely, 1980, 1988).
In contrast to these late prehistoric settings in the Eastern Woodlands, the picture of social
differentiation and infection prevalence is distinctive at Cahokia, a Mississippian site located
in the American Bottom region of the central Mississippi River valley. The system associated
with the site was the most organizationally complex late prehistoric Mississippian chiefdom in
the Eastern Woodlands (see discussion in Milner, 1990). By the early eleventh century AD, an
elite social stratum had emerged that had differential access to a range of prestige items, and
likely enjoyed a better quality of life than lower classes. At Cahokia Mound 72, analysis of the
remains of 261 individuals from 28 burial features revealed that only 5.3% of high-status
individuals had periostitis, whereas 25.0% of middle-status individuals had lesions (Rose &
Hartnady, 1987).
Comparison of socially elite “shamans” and commoners in the Maitas-Chiribaya culture of
northern Chile (c. AD 1000) reveals that fewer high-status individuals have bone infections
than low-status individuals (prevalence: shamans¼9%; commoner males¼20%; commoner
females¼18%) (Allison, 1984). Although preliminary, these studies indicate that high-status
individuals enjoyed a healthier lifestyle than low-status individuals, at least in these settings.
However, in the Middle Sicán culture, the elite ethnic Sicán appear to have endured signifi-
cantly less infection as well as other forms of stress than the populations that they ruled over
in the Lambayeque Valley Complex (Klaus & Toyne, in press).
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 97
the spread of infection through direct contact with infected skin abrasions and cuts. Venereal
syphilis characteristically appears in populations with higher levels of sanitation, such as in
urban settings in Western countries. These groups tend to be relatively more fully clothed,
which provides fewer opportunities for the spread of infection via skin contact in the manner
typical of nonvenereal syphilis, pinta, and yaws.
The skeletal manifestations of the treponematoses are described in the paleopathological
literature in considerable detail (Hackett, 1976; Ortner, 2003; Powell, 1988; Powell & Cook,
2005b). Yaws is most commonly represented by the inflammatory response of the periosteum
surrounding the bones of the forearm, hand, and of the lower limb bones. In the most severe
form, repeated episodes of periosteal reaction and remodeling may result in hypertrophy of
the anterior crests of the tibia and an appearance of bowing called “saber-shin.” Bone surfaces
with close proximity to skin – such as the cranial vault and the anterior tibia – also may
express active lesions or pitted defects from gummatous granulomas. Destructive nasal and
hard palate changes may occur, but are less prevalent in yaws than in venereal or endemic
syphilis.
Skeletal changes associated with endemic syphilis are similar to those that develop in yaws.
In the tertiary stage of the disease, periostitis and gummatous granulomas may develop in the
cranial flat bones and tibiae, and the tibiae take on a saber-shin appearance (Figure 3.9).
Destructive lesions of the face, especially in the nasal region, may also develop. Venereal
syphilis results in virtually the same bone lesions as endemic syphilis, including extensive
cranial vault lesions and periosteal inflammation of lower limb long bones. Other destructive
changes may be present in elbow, hip, and knee joints.
Tertiary-stage venereal syphilis may involve abundant osteosclerotic skeletal responses,
characterized by gummatous destruction of bone. The skull is frequently affected, especially
the nasal area and flat vault bones (Figure 3.10). The frontal bone typically expresses healed
star-shaped, gummatous lesions called caries sicca. Over the course of time, these lesions may
coalesce to form an ectocranial surface with a high degree of irregular topography.
Congenital syphilis results in distinctive skeletal changes, such as osteochondritis (poor
bone formation in areas of endochondral ossification), periostitis, and osteomyelitis. Dental
changes do not occur in the nonvenereal treponematoses because the teeth are fully or nearly
fully formed in the secondary and tertiary stages. In up to 30% of congenital syphilitic
children, pathognomonic modifications may occur in the forming permanent anterior teeth,
specifically the characteristic malformation of incisors in which the crown is unnaturally
constricted at the occlusal margin (called Hutchinson’s incisors, or screwdriver teeth), sharp
hypoplastic defects on canines, and the anomalous patterning of first molars involving
abnormally closely spaced cusps (Moon’s molars) or the presence of a plane-form hypoplasia
cutting into the bases of all cusps (Fournier molars) (Hillson et al., 1998) (Figure 3.11). These
lesions are extremely rare in New World archaeological remains dated before 1492, and are
more common in historic populations with documented evidence of venereal syphilis (Cook,
1994; Jacobi et al., 1992; Lambert, 2006; Mansilla & Pijoan, 1995; Nystrom, 2011; Ortner,
2003). Several possible cases from Old World contexts have been identified (Erdal, 2006;
Henneberg & Henneberg, 1994; Palfi et al., 1994), indicating that venereal syphilis in Europe
may predate the post-Columbian era.
Presence of treponematosis with special attention to venereal syphilis in past populations,
especially in the New World, has been debated for well over a century. Analysis of human
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98 Exposure to infectious pathogens
Figure 3.9 Periosteal inflammation of tibiae (treponematosis); Irene Mound, Georgia. The
tibia in the middle is nonpathological. (From Powell, 1990; reproduced with permission of
author and American Museum of Natural History.)
remains from Tennessee and Kentucky by the American Civil War surgeon Joseph Jones
revealed “the unmistakable marks of the ravages of syphilis” (1876:66). The presence of
“diseased, enlarged, and thickened” long bones convinced him that syphilis was widespread
in the region, that it had an exclusively New World origin, and that it must have been
imported to Europe via the West Indies (Jones, 1876:67). This discussion continues a debate
that is still ongoing; namely, the origin of venereal syphilis, New World or Old World (Baker &
Armelagos, 1988; Cook & Powell, 2012; Dutour et al., 1994; Harper et al., 2011; Merbs, 1992;
Powell & Cook, 2005a, 2005b). Owing to the overlapping symptoms and skeletal manifest-
ations of these three treponemal diseases and the far more extensive record of research on
North American skeletal remains than in Europe or other areas of the Old World, the issue
remains contentious.
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 99
Figure 3.10 Treponematosis (caries sicca) on cranial vault from Early Woodland (900 BC–
AD 200), Wilhoite site, Tennessee. (Photograph by Tracy K. Betsinger.)
One or more syndromes of treponematosis were present in both the Old World and the New
World well prior to AD 1492 (Baker & Armelagos, 1988; Cook & Powell, 2012; various in
Powell & Cook, 2005b; and many others). Cases of treponematosis are sparse in the Old World,
but a growing number of cases, some aided with histological evidence, are expanding that
record for Europe and East Asia (Brothwell, 2005; Cole & Waldron, 2011; Dutour et al., 1994;
Erdal, 2006; Henneberg & Henneberg, 1994; Henneberg et al., 1992; Mays et al., 2003;
Mitchell, 2003; Roberts, 1993; Roberts & Manchester, 2005; Rothschild & Rothschild, 1995;
Suzuki, 1982 – 1984; von Hunnius et al., 2006). Treponematosis is widespread in Australia
(Hackett, 1936, 1976; Webb, 1995) and in some areas of the Pacific (e.g., Marianas: Hanson,
1988; Stewart & Spoehr, 1952; Stodder et al., 1992). While these cases show the Old World
presence of some form of treponematosis, owing to the uncertainly of either clear diagnosis or
good chronological control of cases from archaeological settings, evidence supporting an Old
World origin for venereal syphilis remains elusive (Harper et al., 2011).
The evidence for treponematosis from the New World is quite abundant, having been
identified with regard to both a range of case studies and population differential diagnoses,
especially in North American contexts. In addition to earlier studies (see reviews in Baker &
Armelagos, 1988; Powell & Cook, 2005a), examples of treponematosis are associated
with highly diverse settings (Arriaza, 1995; Bruwelheide et al., 2010; Cybulski, 1980, 1990;
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100 Exposure to infectious pathogens
Lahr & Bowman, 1992; Lambert, 1994; Marden & Ortner, 2011; Owsley & Rose, 1997;
Pechenkina, Vrandenburg et al., 2007; Powell & Cook, 2005a, 2012; Sandford et al., 2002;
Schermer et al., 1994; Stodder, 1994; Stodder & Martin, 1992; Walker & Lambert, 1989;
Wright, 2006; various in Powell & Cook, 2005b). In many contexts, the skeletal manifestation
appears intermediate between those of the two modern endemic syndromes. Considering the
evolutionary nature of human infectious disease over centuries and across host populations of
differing genetic composition, these departures from the modern pattern are not unexpected.
The most extensive record of New World treponematosis data is from precontact Eastern
North America, especially in later prehistoric settings involving sedentary communities
practicing farming. As discussed earlier (see Chapter 2), it is these settings that provide ideal
conditions for the maintenance and spread of disease-causing pathogens. In the lower Illinois
River valley, a pattern of proliferative bone infection that is strongly suggestive of endemic
treponematosis rather than venereal syphilis is present (Cook, 1976, 2007). There is a high
frequency of tibial periostitis affecting adult males and females alike, progressively increasing
with age. The prevalence is especially elevated in later prehistoric, maize-dependent groups
with high population density. Although variable across populations, the pattern has been
described in prehistoric (mostly late) human remains from other groups in a wide range of
settings, but especially in the American Midwest and Southeast (Bogdan & Weaver, 1992;
Bullen, 1972, 1973; Cassidy, 1984; Cook, 2005; Danforth et al., 2007; Dickel, 1991; Eisenberg,
1986, 1991a, 1991b; Garner, 1991; Hutchinson, 1993, 2004; Hutchinson & Mitchem, 1996;
Hutchinson & Norr, 2006; Hutchinson & Richman, 2006; Hutchinson & Weaver,
1998; Hutchinson et al., 2005, 2007; Işcan & Miller-Shaivitz, 1985; Lambert, 2000b; Lewis,
1994; Miller-Shaivitz & Işcan, 1991; Milner, 1983, 1992; Milner & Smith, 1990; Monahan,
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 101
Figure 3.12 Stellate lesions on adult frontal bone (treponematosis); Tierra Verde, Florida. (From
Hutchinson, 1993; reproduced with permission of author and John Wiley & Sons, Inc.)
1995; Powell, 1986, 1988, 1989, 1990, 1991a, 1991b, 1992a, 1992b, 1994; Powell et al., 2005;
Reichs, 1989; Robbins, 1978; Ross-Stallings, 1989; Smith, 2008; Smith et al., 2011; Weaver
et al., 2005; Wilson, 2005).
A number of these settings provide details on lesion morphology and other characteristics that
strongly suggest the New World record reflects multiple forms of endemic treponematosis (Powell
& Cook, 2005a). Hutchinson (1993) found proliferative periosteal apposition on long bones
(especially tibiae) and stellate lesions in crania from late prehistoric and early contact contexts
in the central Florida Gulf coast. For example, in the sixteenth century component of the Tatham
Mound sample, three crania exhibited healed stellate lesions (Figure 3.12). Similar lesions are
present in crania from the postcontact Weeki Wachee Mound and precontact components of the
Safety Harbor and Tierra Verde sites. Although saber-shin tibiae are also present in these samples,
stellate scars appear to be the best single criterion for documenting the presence of endemic
treponematosis (Hutchinson, 1993; and see Hutchinson & Norr, 2006; Hutchinson & Richman,
2006; Hutchinson et al., 2007; Milner & Smith, 1990; Powell, 1990, 1991a) (Figures 3.10, 3.12).
How life-threatening or debilitating treponemal disease was in these earlier societies is
unknown. Owing to the apparent endemic nature of the disease, it may not have been a
primary cause of mortality. Based on the high degree of healing, Powell (1988) argues that
populations at Moundville had successfully adapted to the disease. While this may be the case,
nevertheless, the presence of the characteristic lesions indicates that the disease imposed a
significant health burden and resulted in no small amount of discomfort for those affected. In
his description of native populations in North Carolina in the early eighteenth century, the
explorer John Lawson noted that natives in the eastern part of the colony “. . .have a sort of
Rheumatism or Burning of the Limbs, which tortures them grievously, at which time their Legs
are so hot, that they employ the young People continually pour water down them. . .This not
seldom bereaves them of their Nose. I have seen three or four of them render’d most miserable
Spectacles by this Distemper. Yet, when they have been so negligent, as to let it run on so far
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102 Exposure to infectious pathogens
without curbing of it; at last, they make shift to patch themselves up, and live for many years
after. . .” (Lawson, 1967:231). These descriptions correspond well with modern clinical
descriptions of deep leg pain and orofacial lesions produced by treponemal disease (Hackett,
1951; Hudson, 1958).
Some examples of nonspecific periosteal reaction observed in archaeological skeletons
likely have a treponemal origin, especially in individuals who also display the classic path-
ognomonic lesions (e.g., stellate scars on crania and/or highly proliferative bone on tibiae
diaphyses). In cases where distinctive symptoms are absent, the delineation between non-
specific infection (periostitis) and endemic treponematosis is difficult to determine.
The well-documented paleopathological diagnoses of treponematosis in North American
archaeological settings make it clear that the disease was well established in the New World
prior to the arrival of Europeans. In the lower American Midwest and Southeast, regions that
are subtropical or that experience high seasonal humidity and temperatures, there appears to
be a cline of skeletal expression from the hotter, more humid coastal regions to the relatively
drier interior regions, resembling modern inter-regional treponemal clines spanning “classic”
yaws and “classic” endemic syphilis in central and southern Africa (Basset et al., 1994;
Froment, 1994; Grin, 1956).
Identification of Treponema DNA in archaeological skeletons is highly problematic.
T. pallidum pallidum has been successfully identified in only one setting (Kolman et al.,
1999; but see Barnes & Thomas, 2006), indicating that molecular approaches to understanding
this disease are extremely sketchy, at best (Bouwman & Brown, 2005; Buikstra, 2010; von
Hunnius et al., 2007; Wilbur & Stone, 2012). At least with respect to understanding the history
of this disease, it is, for the time being, largely dependent upon the skeletal record.
Variation by sex or status in skeletal populations is difficult to assess, especially given the
vagaries of diagnosis. Where probable cases of treponematosis have been identified, the
prevalences in adult males and females are broadly similar (Cook, 1976; Powell, 1988; Powell &
Cook, 2005a; Smith et al., 2011; but see Powell, 1990). Powell’s (1988, 1992b) investigation of
the Moundville skeletal series indicates no clear distinction between status groups. In contrast,
comparison of elite and non-elite individuals from late prehistoric (Mississippian period)
stratified societies in eastern Tennessee reveals high-status individuals as having significantly
fewer skeletal lesions associated with treponematosis than low-status individuals (Smith et al.,
2011). This clear dichotomy in disease prevalence suggests that higher-status individuals may
have enjoyed circumstances conducive to relatively better health than low-status individuals.
Alternatively, more robust health may have facilitated social advancement. Current models of
disease epidemiology suggest, however, that the former explanation is more likely.
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 103
provides a long-term context of disease history, human – animal interaction, and co-
evolution of pathogens.
Tuberculosis today is comprehensively global, infecting nearly one-third of the population,
second only to HIV as a cause of death in human populations. In 2005 alone, there were
nearly nine million new cases with most in Asia and Africa, making it a leading cause of death
(World Health Organization, 2007). Unlike treponematosis, which produces proliferative bone
apposition, tuberculosis progressively destroys bone tissue, and is commonly expressed as
erosive vertebral lesions of the lower back (lower thoracic and lumbar vertebrae) and resorp-
tive and slight proliferative changes of the pleural (internal) surfaces of ribs (Ortner, 2003;
Palfi et al., 1999; Roberts & Buikstra, 2003; Roberts et al., 1994). Tuberculosis is caused by
organisms from the Mycobacterium tuberculosis complex, here simply called “tuberculosis.”
The most common cause of tuberculosis today in humans is the bacteria M. tuberculosis and
M. bovis. The primary mode of transmission for the former is by breathing airborne microbes,
usually in droplets introduced by sneezing or coughing, and the primary mode for the latter is
via consumption of milk and meat from infected animals or by swallowing infected sputum of
the intestinal tract from animal products infected by the bacterium (Gordon & Mwandumba,
2008; Roberts, 2011).
The infection pathway is usually through the respiratory tract, resulting in a primary
infection in lung tissue and subsequent secondary infection in regional hilar lymph nodes
(Ortner, 2003). Over a period of years, the bacilli may spread to skeletal tissues via the
circulatory system, with a propensity for iron-rich hematopoietic marrow and cancellous
bone. The vertebrae, ribs, sternum, and (for subadults) long bone metaphyses are especially
favored sites of secondary infection because of the presence of a rich blood supply and the
scarcity of phagocytic cells (Hopewell, 1994; Ortner, 2003; Thijn & Steensma, 1990). The
process can result in extensive destruction of cancellous bone, most commonly in vertebral
bodies (Figure 3.13). With the loss of bone mass, vertebral bodies may collapse and fuse. The
resulting severe kyphosis is called “Pott’s disease” after the original description by Sir
Percivall Pott (1779) (Figure 3.14). However, any bone or joint can be involved in tubercular
infections (Berney et al., 1972; Ortner, 2003).
The proliferative lesions on the visceral surfaces of ribs are presumed by some to be
pathognomonic for tuberculosis, but the cause remains unclear and may reflect secondary
infection downstream from the original tubercular infection or some other infection
altogether (Kelley & Micozzi, 1984; Pfeiffer, 1991; Roberts et al., 1994, 1998; Wilbur et al.,
2008) (Figure 3.15). Examination of skeletal remains from individuals of known cause of
death shows individuals with rib lesions as also having tuberculosis (Santos & Roberts, 2001).
Individuals with non-tubercular pulmonary disease sometimes have similar rib lesions.
Therefore, all rib lesions of this type should not necessarily be interpreted as diagnostic of
tuberculosis (Roberts, 1999; Roberts et al., 1994), and may reflect other respiratory infections,
such as some form of pneumonia (Lambert, 2002b). Nevertheless, rib lesions record an
important possible signal of tuberculosis infection in past populations but the pathology data
should be interpreted cautiously (Pfeiffer, 1991; Raff et al., 2006; Roberts et al., 1998; Santos &
Roberts, 2001; and see Matos and Santos, 2006) and perhaps confirmed via analysis of aDNA
(Müller et al., 2014).
In the Old World, skeletal evidence for tuberculosis has been traced to six to seven thousand
years ago in Europe (Nicklisch et al., 2012), but with significant levels of prevalence occurring
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104 Exposure to infectious pathogens
considerably later, especially in the later Medieval period when populations began to be
concentrated in urban or otherwise settled areas (Canci et al., 1996; Gladykowska-Rzeczycka,
1999; Roberts and Buikstra, 2003; Stone et al., 2009). In many areas of the Old World, there is
limited or no record of tuberculosis. The paucity of data may simply reflect the fact that
bioarchaeological research is more comprehensive in Europe than elsewhere.
Most early authorities argued that the disease was absent in the New World prior to
European contact (Hrdlička, 1909). However, the growing skeletal record in the New World
reveals an abundance of pathology indicating that this illness appeared by the first millen-
nium AD (Drake & Oxenham, 2012; Klaus, Wilbur et al., 2010; Roberts, 2012; Roberts &
Buikstra, 2003; Stone et al., 2009). Moreover, the identification of acid-fast bacilli and soft-
tissue tubercular lesions (Allison et al., 1973, 1981), and especially M. tuberculosis DNA
in precontact Peruvian and Chilean mummies (Arriaza et al., 1995; Konomi et al., 2002;
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 105
Figure 3.14 Destructive lesions on bodies of thoracic and lumbar vertebrae, resulting in
kyphosis of the spine (Pott’s disease, tuberculosis); Norris Farms, Illinois. (Photograph by
George R. Milner and the Illinois State Museum.) (A black and white version of this figure
will appear in some formats. For the color version, please refer to the plate section.)
Figure 3.15 Tuberculosis rib lesions; Terry Collection. (Photograph by Charlotte A. Roberts.)
(A black and white version of this figure will appear in some formats. For the color version,
please refer to the plate section.)
Salo et al., 1994) and skeletons (Bos et al., 2014; Braun et al., 1998; Kaestle, 2010; Klaus,
Wilbur et al., 2010; Raff, 2008) demonstrates its presence in the New World in the pre-
Columbian era. The significant Old World distribution of the disease via detection of M.
tuberculosis DNA is also indicated in archaeological remains from a range of contexts (Baron
et al., 1996; Boros-Major et al., 2011; Crubézy et al., 2006; Fletcher et al., 2003; Haas et al.,
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106 Exposure to infectious pathogens
2000; Hershkovitz et al., 2008; Matheson et al., 2009; Mays et al., 2001; Murphy et al., 2009;
Nicklisch et al., 2012; Nuorala, 1999; Roberts, 2012; Spigelman & Lemma, 1993; Suzuki,
2013; Szelekovszky & Marcsik, 2010; Taylor et al., 1996, 2005; Zink et al., 2005; various in
Pálfi et al., 1999). The earliest purported molecular (DNA) evidence is from Neolithic settings
dating to c. 9000 years ago from Israel (Hershkovitz et al., 2008; but see Wilbur et al., 2009)
and c. 7000 years ago in Germany (Nicklisch et al., 2012).
Destructive vertebral or proliferative rib lesions are identified in archaeological skeletal
series representing a diversity of groups worldwide, including the Middle East (Baker, 1997;
Buikstra et al., 1993; Morse, 1967; Ortner, 1979; Ortner & Frohlich, 2008; Roberts & Buikstra,
2007; Strouhal, 1991; Zias, 1991), northern and central Europe (Arcini, 1999; Bennike, 1985,
1999; Blondiaux et al., 1999; Evinger et al., 2011; Formicola et al., 1987; Inglemark, 1939;
Jankauskas, 1999; Manchester, 1991; Mays & Taylor, 2003; Mays et al., 2001; Pálfi &
Marcsik, 1999; Roberts, 2011; Roberts & Buikstra, 2003; Waldron, 1993), Greece (Angel,
1984), Asia (Murphy et al., 2009; Oxenham et al., 2005; Pechenkina, Benfer et al., 2007;
Suzuki, 1991, 2013; Suzuki & Inoue, 2007; Suzuki et al., 2008; Tayles & Buckley, 2004), and
elsewhere (Ortner, 2003; Stone et al., 2009). In the New World, a spate of reports document the
presence of a disease strongly resembling tuberculosis in South America (Allison, 1984;
Allison et al., 1981; Arriaza et al., 1995; Buikstra & Williams, 1991; Klaus, Wilbur et al.,
2010; Wilbur et al., 2008). In North America, instances of tuberculosis are restricted to
primarily eastern settings (Buikstra, 1977b; Buikstra & Cook, 1978, 1981; Clabeaux, 1977;
Cook, 1984; Danforth et al., 2007; Eisenberg, 1986, 1991a, 1991b; Hartney, 1981; Katzenberg,
1977; Lambert, 2000b; Milner, 1983, 1992; Milner & Smith, 1990; Murray, 1989; Pfeiffer,
1984, 1991; Pfeiffer & Fairgrieve, 1994; Powell, 1988, 1990, 1991a, 1991b, 1992a, 1992b,
2000; Rathbun et al., 1980; Saunders et al., 1992; Widmer & Perzigian, 1981) and the
Southwest (Bruwelheide et al., 2010; El-Najjar, 1979; Fink, 1985; Hooton, 1930; Micozzi &
Kelley, 1985; Stodder, 1994; Stodder & Martin, 1992; Stodder et al., 2002; Sumner, 1985). In
addition, other cases have been documented on the Northwest Coast (Cybulski, 1978, 1990)
and Great Plains (Mann et al., 1994; Owsley & Rose, 1997; Palkovich, 1981; Williams, 1994;
Williams & Snortland-Coles, 1986). However, in Mesoamerica, a very notable lack of tuber-
culosis is well recognized (Wilbur et al., 2008).
Most paleopathological studies report on isolated cases of tuberculosis or tuberculosis-
like infections. The detailed study of prevalence, pattern, and lesion morphology in a
limited number of series provides important details on the history of the disease. In a
comprehensive study of a temporal sequence of human remains from the lower Illinois
River valley in west-central Illinois, biocultural reconstruction and interpretation of change
in pattern and prevalence of resorptive lesions indicates the presence of tuberculosis
(Buikstra and Cook, 1981; also Buikstra, 1977b; Buikstra and Cook, 1978; Cook, 1984,
2007). Over the course of the sequence, beginning in the Middle Woodland (150 BC – AD
400), followed by the Late Woodland (AD 400 – 1050) and Mississippian (AD 1050 – 1400)
periods, the region saw an increase in population density and sedentism, especially when
maize agriculture became well established during the eleventh century AD. No clear
evidence of tubercular resorptive lesions is present in the Middle or Late Woodland periods.
In the Mississippian period, however, there is a clear reorientation in disease pattern.
Human remains from the Mississippian-period Yokem and Schild sites possess destructive
vertebral body lesions consistent with an etiology of tuberculosis. The Mississippian-period
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 107
adults show a high young adult mortality and an equitable distribution among adult males
and females.
At Moundville, circumstances for the introduction and spread of tuberculosis are similar to
west-central Illinois (Powell, 1988, 1991a, 1991b, 2000). In comparison with Illinois, vertebral
lesions are rare: only three adults are affected in this manner, and only one adult displays
classic vertebral body destruction. No crania, hips, or knees possess resorptive lesions con-
sistent with tuberculosis. However, the presence of a large number of pleural rib lesions may
indicate a broader presence of the disease in this population. If so, this means that rib lesions
may be an early manifestation of tuberculosis if it first enters the lungs.
At the Irene Mound site, three individuals show osteolytic vertebral lesions. One individual
has extensive destruction of the sacroiliac joint without remodeling, two individuals have
periostitis on the anterior scapular bodies, and one individual has periostitis on the pleural
aspect of the sternum (Powell, 1990, 1991b). Additionally, eight of 10 Irene Mound individ-
uals with other tubercular lesions have subtle periosteal apposition on the pleural aspects of
the ribs. Although the skeletal changes at the Irene Mound site are not as profound as those
observed in the American Midwest (compare with Buikstra & Cook, 1981; Milner & Smith,
1990), they nevertheless fit the expected profile of tuberculosis.
Among the most comprehensive yet arguably imperfect bioarchaeological records of
tuberculosis is in the precontact Andean region, with some of the earliest cases in the
New World around 2000 years ago in the Pisco Valley (Peru) and Atacama region (Chile)
(Allison et al., 1981). These and other contexts from the region provide a number of sites
with tuberculosis, but most sites reporting disease have only one or two cases. Indeed, the
largest is represented by 10 individuals from Estuqueña (Peru) (Buikstra & Williams, 1991).
Therefore, unlike Europe where clear diachronic trends show an increase in the Medieval
period relative to earlier periods, the diachronic record in the Andean region needs to be
developed further (Klaus, Wilbur et al., 2010). Nonetheless, the skeletal record suggests that
tuberculosis peaked around AD 1000. This appears to have coincided with the appearance of
complex societies – chiefdoms and states – and associated population size increase, popula-
tion density, economic intensification, and dietary changes, promoting the infection (Klaus,
Wilbur et al., 2010). These regional analyses provide compelling evidence that, as in living
populations, the synergy of living circumstance – especially population aggregation, com-
promised health, and changed dietary foci – served to promote conditions conducive for the
pathogen to thrive in the past. This pattern is repeated worldwide where data for disease
prevalence and abundant archaeological context are available (Roberts & Buikstra, 2003;
Roberts & Cox, 2003, 2007).
Importantly, the strain or strains of M. tuberculosis in the precontact New World are
American-specific, and distinctive from the strain documented in native populations in
historic times. This finding is consistent with the hypothesis that the American-specific strains
of the tubercular pathogenic organisms were replaced by a European strain after contact (Bos
et al., 2014). One of the more provocative possibilities that Klaus, Wilbur, and coworkers
(2010) consider from their lack of identification of M. tuberculosis aDNA in a wide variety of
skeletons could indicate the infectious agent in the western hemisphere before European
contact was not tuberculosis at all, but rather, a closely related form, such as M. kansasi,
that can produce identical lesions in the human skeleton. It could also mean that the aDNA
simply is not preserved. These possibilities require further scrutiny, especially regarding the
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108 Exposure to infectious pathogens
designs of assays employed to identify tuberculosis versus other very closely related
organisms.
Distribution of tuberculosis by sex in these groups shows considerable variation (Buikstra &
Cook, 1981; Powell, 1988, 1992a). In some settings, there appears to be a male bias. In the
Andean setting, for example, the higher male prevalence may be related to exposure relating
to behaviors or dietary variation occurring prior to adulthood (Klaus, Wilbur et al., 2010). In
the Irene Mound population, however, female prevalence is twice that for males (Powell,
1991a), but this could be an artifact of the composition of the sample (Larsen, 1984, for
discussion of age bias). In terms of status differences, the only systematic analysis indicates no
apparent relationship between tuberculosis and rank in at least one setting in the American
Southeast (Powell, 1988).
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3.5 Specific infectious diseases: treponematosis, tuberculosis, and leprosy 109
2013; and others). Unlike treponematosis and tuberculosis, leprosy was not present in the New
World prior to European contact. In all likelihood, the disease was introduced to the New
World during the early colonial era (Ortner, 2003). However, no diagnoses in New World
settings have been undertaken.
Beginning largely with the work of the Danish physician and paleopathologist Vilhelm
Møller-Christensen (1961, 1978) on human remains recovered from Medieval leprosy hospital
cemeteries in Denmark (and see Andersen, 1969; Bennike, 2002; Linderholm & Kjellström,
2011), the skeletal manifestations of leprosy are well delineated. His pioneering studies
contributed to the modern diagnosis of the disease in living populations and serve as a
baseline for understanding the disease in archaeological contexts.
Excavations at St. Jorgensgard, a Danish church cemetery (c. AD 1250 – 1550) near
Næstved, resulted in the recovery of about 650 skeletons (Møller-Christensen, 1961, 1978).
This is a unique series for bioarchaeological investigation because all individuals represent
those who had been admitted into the leprosy hospital in order to isolate them from other
members of the population at large (Møller-Christensen, 1978). Møller-Christensen’s exten-
sive studies of this series revealed a distinctive “facies leprosa” or rhinomaxillary syndrome
involving atrophy of the nasal and maxillary regions, alveolar resorption, and anterior tooth
loss (Figure 3.16a). Additionally, hand and foot elements are atrophied and shortened
(Andersen et al., 1992) (Figure 3.16b). Considerable evidence from Denmark, England, and
other European settings shows that the disease first appears in the second millennium BC, has
a significant presence in the early Medieval period, and increases to its maximum in the late
(a)
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110 Exposure to infectious pathogens
(b)
Figure 3.16b Metatarsal atrophy of the feet (leprosy); Late Medieval/Early Post-Medieval,
Armoy, Ireland. (From Murphy, 2007; reproduced with permission of the author.)
Medieval period (Antunes-Ferreira et al., 2014; Arcini, 1999; Belcastro et al., 2005; Blondiaux
et al., 1994, 2002; Boldsen, 2005, 2008; Boldsen & Mollerup, 2006; Brothwell et al., 2000;
Haas et al., 2002; Judd & Roberts, 1998; Kjellström, 2012; Manchester, 1981; Mariotti et al.,
2005; Murphy, 2007; Murphy & Manchester, 2002; Pálfi, 1991; Pálfi et al., 2002; Roberts,
2002, 2011; Roberts & Cox, 2003; Rubini & Zaio, 2009; Strouhal et al., 2002). While not as
widespread as in Europe, other leprotic skeletons have been described from the Nile valley
region (Dzierzykray-Rogalski, 1980; Møller-Christensen & Hughes, 1966; Molto, 2002; Smith,
1908), Israel (Manchester, 1993; Zias, 2002; Elliot ), Turkey (Angel, 1969), Uzbekistan (Blau &
Yagodin, 2005), and India (Robbins Schug et al., 2009). The remains from Turkey date to
2700 – 2300 BC and those from India date to 2500 – 2000 BC, representing the oldest probable
cases of leprosy. As with tuberculosis, the presence of leprosy is associated with settings
involving population crowding and poor living circumstances generally. In northern Europe,
the cases are drawn largely from small villages in rural settings, similar to what is seen today.
Temporal documentation in Europe, where the data are the richest, suggests that the disease
declines after c. AD 1300 and largely disappears by the nineteenth century, perhaps due to the
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3.6 Specific infectious diseases: vectored infections 111
rise of tuberculosis and circumstances relating to cross-immunity of the two diseases (Dono-
ghue et al., 2010; Leitman et al., 1993; Manchester, 1991; Roberts & Buikstra, 2003).
Other skeletal pathology found in leprous individuals includes cribra orbitalia, periostitis on
tibiae and fibulae, and maxillary sinusitis (Andersen, 1969; Boocock et al., 1995; Møller-
Christensen, 1978; Roberts et al., 2002), although these conditions alone are not diagnostic of
leprosy. Increased prevalence of maxillary sinusitis, for example, appears to accompany
elevation in air pollution and the confines of urban living in later Medieval England (Lewis
et al., 1995) and other settings (Roberts, 2007). Some individuals display dental changes
whereby crown bases of maxillary incisors are concentrically constricted (Roberts, 1986). The
presence of malformed teeth indicates that the infection occurred early in childhood.
Additional examples of leprosy from archaeological contexts are from mostly isolated
skeletons from other Old World localities (Boocock et al., 1995; Hirata et al., 2000; Lewis
et al., 1995; Magilton et al., 2008; Manchester, 1991; Manchester & Roberts, 1989; Møller-
Christensen & Inkster, 1965; Ortner, 2003; Tayles & Buckley, 2004; Trembly, 1995, 2002; Zias,
1991), but the bioarchaeological evidence is far less profuse in these regions than in Denmark
and England. Manchester and Roberts (Manchester, 1991; Manchester & Roberts, 1989; and
see Roberts et al., 2002) assessed skeletal, archaeological, and archival evidence of leprosy in
Britain and conclude that, as in Scandinavia, the disease was endemic. They argue that
following the introduction of the endemic form of leprosy, perhaps by the late Roman period,
the disease increased in prevalence, peaking during the thirteenth century or some time later.
Leprosy then declined and disappeared by the end of the fifteenth century, but remained in
Scandinavia much later than in other regions of Europe. The general pattern is similar to
treponematosis and tuberculosis in that it increases with elevation in population size, density,
and interpersonal contact. The reasons for the disappearance of leprosy remain obscure.
Increased immunities due to centuries of exposure to the disease and improvements in
hygiene and living conditions may have contributed to its decline (Carmichael, 1993; Man-
chester, 1991).
There are no distinctive sex differences in leprosy prevalence in the Næstved adults studied
by Møller-Christensen. Status differences in leprosy prevalence are difficult to determine in
this sample. Møller-Christensen did not compare possible status differences (e.g., exterior
church versus interior church burials). However, a high-status young adult male buried in the
Næstved church choir displayed classic symptoms of leprosy. Clinical evidence indicates that
leprosy can occur in anyone, but malnutrition is a predisposing factor (Keil, 1933). The
location of leprotic skeletons in peripheral areas of cemeteries associated with churches in
Sweden points to their lower status in Medieval society in this setting (Linderholm &
Kjellström, 2011). Interestingly, stable isotope analysis reveals no evidence of a difference
in either quality or type of diet in comparison with other members of the community interred
in cemeteries in this setting.
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112 Exposure to infectious pathogens
and quality of life in past populations. These diseases, however, represent only a part of the
record. Historical sources potentially provide a more complete picture of past living circum-
stances and the larger picture of morbidity and mortality. For example, bubonic plague, which
arguably transformed the health and demographic landscape of Europe in the Middle Ages to
the end of the eighteenth century, has until recently been known from historical documenta-
tion alone as it leaves no pathognomonic signature in skeletal remains. Historians have
documented the progression in Europe of two pandemics of plague, assuming that these
plagues – the Justinian plague (AD 541 – 767) and the Black Death (AD 1347 – 1750) – were
bubonic plague, a rodent disease caused when humans are bitten by fleas carrying Yersinia
pestis (Carmichael, 1993; Haensch et al., 2010). However, some authorities have challenged
the notion that these pandemics were bubonic plague, arguing for example that mortality
profiles were unlike bubonic plague in the modern era and that anthrax or viral hemorrhagic
fever were more likely causes (Scott & Duncan, 2001; Shrewsbury, 1970).
Biomolecular analysis of bone from human remains recovered from plague cemeteries in a
variety of locations in Europe (Bianucci et al., 2008, 2009; Bos et al., 2011; Drancourt et al.,
2007; Garrelt & Wiechmann, 2003; Haensch et al., 2010; Harbeck et al., 2013; Kacki et al.,
2011; Raoult et al., 2000; Schuenemann et al., 2011; Tran et al., 2011; Wiechmann & Grupe,
2005), dating to the first and second pandemics, reveals the presence of DNA of Y. pestis (pla
and gplD genes). Moreover, Bianucci and her collaborators (2008, 2009; Kacki et al., 2011)
have developed a diagnostic immunological test for detecting the F1 antigen, which is
antigen-specific for Y. pestis. Where aDNA analysis was inconclusive, this test confirms the
presence of Y. pestis in a series of skeletons from individuals from various settings. In
particular, F1 antigen was found in remains having the pla and gplD genes, at Lambesc
(AD 1590 plague event) and Marseille (AD 1722 plague event). These details are essential for
understanding the causes, epidemiology, and evolution of disease processes specific to plague
(Dobson, 2007). Similarly, immunological analysis of bone samples in areas known to have
suffered from endemic malaria now reveal a protein signature for the presence of Plasmodium
falciparum, the parasite responsible for malaria across many areas of the world (Bianucci
et al., 2008; Fornaciari et al., 2010a, 2010b). This now makes possible the documentation of
another major disease having no skeletal signature.
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3.7 Summary and conclusions 113
new insight into the human/pathogen co-evolution and co-adaptation. Yet, DNA analysis is
not a panacea that will solve the ambiguities surrounding more traditional difficulties of
definitively diagnosing specific diseases in bone. Some investigations underscore that ancient
pathogen DNA recovery and analysis can be quite unreliable and can be fraught with
ambiguity (Roberts & Ingham, 2008; Wilbur et al., 2009). Bioarchaeological analysis of
ancient disease may indeed be strongest when multiple lines of evidence are used in concert,
such as differential diagnoses drawn from visual inspection of bone, radiographic imaging,
and DNA analysis (Klaus, Wilbur et al., 2010). Nevertheless, the so-called “Next Generation”
DNA sequencing techniques that are currently emerging may hold the promise to overcome
many of these issues and usher in a new era in pathogen DNA analysis in bioarchaeology.
In the New World, infectious disease appears to be relatively more common in late
prehistoric settings than in earlier periods. A similar pattern is expressed in Medieval Europe
in comparison with pre-Medieval Europe. In general, these increases are linked with increased
population size and aggregation, mostly in agricultural, agropastoral, and urbanized societies.
Poor dental health (based on elevated levels of dental caries, periodontal disease, wear, and
tooth loss) is related to dietary factors, and to a lesser extent, use of the masticatory complex
for mechanically demanding, nonmasticatory functions. Skeletal infections are especially
prevalent in populations living in densely settled communities. Thus, it should come as no
surprise that treponematosis, tuberculosis, and leprosy – as well as elevated levels of non-
specific bone infections – are present in these settings. In Medieval Europe, co-occurrence of
leprosy and tuberculosis reflects a similar deterioration in living standards.
These general characteristics support the contention that infectious disease as it is
expressed in osseous remains is essentially density-dependent. These diseases are opportunis-
tic in that individuals exposed to them are already stressed by poor diets and at high risk of
early death. In at least some of these settings, poor diets likely exacerbated previously
compromised health, while increased population density and poor sanitation enhanced the
burden of infectious disease and its likelihood of widespread transmission.
The presence of skeletal indicators of infection, both nonspecific and specific, is indicative
of long-term responses to pathogens. In a sense, therefore, the lesions reflect vigorous
immune response – the individual survived the initial pathogenic attack long enough to elicit
a skeletal response (Ortner, 1991; Powell, 1988). However, presence of these lesions in high
frequencies in many of the groups discussed in this chapter also reflects an elevated disease
burden and generally negative impact on adaptation and health (compare with Goodman,
1994; Goodman & Armelagos, 1989). In this regard, inflammation is traditionally thought of
as a natural outcome of infection, helping the body to heal. However, inflammatory responses
to a wide range of infections, including dental caries and periodontal disease, promote
cardiovascular disease and elevated mortality (Li et al., 2000), and thus have a detrimental
impact on overall systemic health.
The data on infectious disease, skeletal and dental, reflect social dynamics within popula-
tions. For example, caries rate differences between males and females in many societies
indicate differential access to foods and variability in dietary behavior. Differences in patterns
of health by rank or status in some groups reflect probable differences in exposure to stressors
and quality of life generally. Regardless of how infectious disease affected specific compon-
ents of the populations, the overall impact was negative – affected individuals likely had
reduced ability to acquire key resources (e.g., food) and to participate in essential work
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114 Exposure to infectious pathogens
activities, and may well have had shortened lifespans. Individuals affected by infectious
disease carried heavy social burdens. With regard to leprosy in Medieval Europe, the disease
was highly feared, and individuals with the disease were considered living dead, who were to
be isolated, forgotten, and removed from society (Moore, 1987).
Various causal factors discussed in this chapter underscore the fact that infectious disease
has a varied etiology. Undoubtedly, specific pathogens are linked with particular infectious
diseases, and are identified as their “causes.” However, even when hosts are infected by these
pathogens, actual disease transpires only when pathogen virulence coincides with host
susceptibility in a conducive environment.
The prevalence of skeletal lesions in an archaeological population does not show a direct
one-to-one correlation with actual prevalence in a living population. For example,
tuberculosis was highly prevalent in some pre-antibiotic groups, but it only rarely spreads
to the skeleton (Ortner, 2003), generally reported as 3% – 7% of cases in living populations.
Some archaeological series show comparatively higher prevalences (Buikstra & Williams,
1991; Eisenberg, 1986; Milner & Smith, 1990; Roberts and Buikstra, 2003). Additionally, it
is important to distinguish between disease and infection. Disease prevalence – whether drawn
from living or extinct populations – may represent only a small part of the total picture of
infection. Similarly, the risks outlined in this chapter for infection may differ considerably
from the factors that ultimately influence and determine if disease will develop from the
initial infection.
Diseases involving skeletal tissues must have contributed significantly to the burden of ill
health in many earlier societies, as they do today. Various segments of populations may have
been affected by infectious disease differently, but the experience was always mediated by
local ecological, cultural, social, and behavioral circumstances.
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4 Injury and violence
4.1 Introduction
Human skeletal remains from archaeological contexts provide a rich and diverse record of
past behavior, reflecting the wide array of circumstances – ecological, cultural, social,
economic, and political – that put a society and its members at risk of injury and death,
resulting from either accident or violence. Many injuries are not identifiable in human
skeletons, especially those that are limited to soft tissues. Moreover, accidental death is
virtually invisible in the archaeological record except under very special circumstances, such
as in building collapse or natural disasters (Cicchitti, 1993; Deiss, 1989; Palkovich, 1980;
Sakellarakis & Sapouna-Sakellaraki, 1991). Violent death is often invisible in the study of
skeletal remains, except in instances when bone is directly damaged and shows no evidence of
healing, such as by gunshot or a sharp-edged weapon.
Despite these limitations, osteological remains are a highly useful index for assessing
outcomes relating to unintentional (accidental) and intentional, malevolent (violent) injury
in a remarkable variety of contexts around the world (Arkush & Tung, 2013; Courville, 1962;
Domett et al., 2011; Jimenez, 1994; Judd & Redfern, 2012; Knowles, 1983; Knüsel & Smith,
2014; Lambert, 2007, 2014; Lovell, 2008; Martin & Anderson, 2014; Martin & Frayer, 1997;
Martin et al., 2012; Merbs, 1989a; Ortner, 2003; Robbins Schug et al., 2012; Roberts &
Manchester, 2005; Schulting & Fibiger, 2012; Šlaus et al., 2012; Tung, 2012a, 2012b; Walker,
2001a; Webb, 1995). This applies especially because skeletal injuries provide a snapshot of a
person’s lived experience at that particular moment when s/he sustained the injury, often
resulting in death.
There is an abundance of skeletal injury data presented in the bioarchaeological literature.
In the past, the combined sparseness of a population perspective and dominance of descrip-
tion in much of this literature, however, largely precluded the realization of the enormous
potential that these kinds of data have for drawing inferences about human behavior and
conflict situations in earlier societies. However, a richer understanding of this important
record for inferences about behavior in the past has developed, especially starting with work
on the linkages between the injured and their environmental and social contexts, the two of
which are often linked (Domett et al., 2011; Judd, 2008; Judd & Redfern, 2012; Knüsel &
Smith, 2014; Martin et al., 2012; Milner et al., 1991; Nichols & Crown, 2008; Robbins Schug
et al., 2012; Schulting & Fibiger, 2012; Tung, 2012a, 2012b; Walker, 2001a). The trauma
patterns documented in skeletal remains provide an interpretive tool for understanding the
social implications of gender and power relations. Thus, bioarchaeological study of trauma
emphasizes biocultural context and the association with person, setting, and the array of
(often complex) social and behavioral circumstances leading to injury and/or death.
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116 Injury and violence
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4.2 Skeletal injury and lifestyle 117
Figure 4.1 Perimortem cranial depressed fracture at the glabella of the frontal bone;
Phum Snay, Cambodia. (Domett et al., 2011; reproduced with permission of authors and
Antiquity.) (A black and white version of this figure will appear in some format. For the
color version, please refer to the plate section.)
elements, many past human groups show relatively high prevalence of rib, vertebral, and
radial (especially distal) fractures relating to accidental injuries and hazards of living circum-
stances (Cybulski, 1992; Dickel, 1991; Gasperetti & Sheridan, 2013; Grauer & Roberts, 1996;
Judd, 2008; Kaplan et al., 1977; Molleson, 1992; Ortner, 2003; Rose, 1985; Sandzén, 1979;
Todd, 1927; Ubelaker, 1981). Injuries to these regions provide a picture of behaviors that can
be dangerous, and they show how particular lifestyles and adaptations related to either
subsistence practices or occupation can be a risk in terms of accidental injury.
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118 Injury and violence
Figure 4.2 Antemortem fracture of rib angles from a single traumatic event; Birmingham,
United Kingdom. (Brickley, 2005; reproduced with permission of author and John Wiley &
Sons, Inc.)
AD 1850). In this context, Brickley’s (2006) analysis of an early modern (AD 1750–1850) series
from Birmingham, England, reveals that 15.6% of individuals had at least one rib fracture,
with higher prevalence in older adults, in males, and in individuals from less-wealthy graves.
The record suggests that the fractures are likely mostly due to work-related accidents
(Figure 4.2). A relatively higher prevalence in lower versus higher social classes suggests that
lower social classes were at greater risk of being injured in this way. Clinical evidence
indicates the dangers of rib fractures for health. That is, in addition to extreme levels of pain,
rib fractures are linked to increased mortality, often resulting from respiratory complications,
reduced function, and increased likelihood of infection. In the pre-antibiotic era, a rib fracture
could mean early death (Bulger et al., 2000).
The risks of urban living are also well documented in a nineteenth-century Euroamerican
series from Belleville, Ontario (Jimenez, 1994). Adult males and adult females show a very high
prevalence of rib fractures, likely arising from accidents (males¼46.6%; females¼27.9%).
As with Britain, these frequencies reflect the increased risk of injury during the period of
industrialization and the development of urban life.
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4.2 Skeletal injury and lifestyle 119
(Shahriaree et al., 1979; Snow, 1974; Wiltse, 1957; Wynne-Davies & Scott, 1979). A more
compelling explanation than genetic origin is that spondylolysis is a type of fracture: it is
absent at birth, usually absent in childhood, and increases progressively through adulthood
(Bridges, 1989a; Lester & Shapiro, 1968; Merbs, 1989b, 1995; Stewart, 1953b, 1956, 1979;
Ward & Latimer, 2005; Weiss, 2009). Moreover, there is healing in later adulthood (Merbs,
1995), the separation affects only intact bone (Wiltse et al., 1975), and it develops gradually in
response to excessive mechanical loads over a period of time (Eisenstein, 1978; Roberts, 1947;
Wiltse, 1962; Wiltse et al., 1975; and see Merbs, 1983, 1989b; Weiss, 2009).
The condition is unique to humans, suggesting that bipedality plays an important role,
largely owing to lordosis as an evolutionary adaptation in order for the body’s trunk to
maintain balance over the pelvis (Bridges, 1989a; Merbs, 1989b; Ward & Latimer, 2005). The
defect is usually bilateral (Merbs, 2002; Waldron, 1992) and almost always involves the fifth
lumbar vertebra only, although it also occurs with decreasing frequency from the inferior to
the superior lumbar spine in clinical circumstances (Moreton, 1966; Roche & Rowe, 1951) and
archaeological settings (Bridges, 1989a; Gunness-Hey, 1980; Lester & Shapiro, 1968; Lundy,
1981; Mays, 2006a; Merbs, 1983, 2002; Pálfi, 1992; Pilloud & Canzonieri, 2012; Snow, 1948;
Stewart, 1979; Waldron, 1993; Wilczak et al., 2009). The defect has also been found in
cervical and thoracic vertebrae, but these are rare occurrences.
Stewart (1953b) and Lessa (2011) reported unusually high prevalences in prehistoric Aleut-
Eskimo and coastal Brazilian populations, respectively. In the latter, for example, nearly one-
third of adults were affected, with younger adults expressing a greater frequency than older
adults. In the series studied by Stewart from north of the Yukon River, Alaska, more than 40%
of individuals exhibit separate neural arches. Inherited structural characteristics of vertebrae
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120 Injury and violence
may predispose the individual to the defect under mechanically stressful conditions. For
example, Stewart (1956) observed that spondylolysis is associated more frequently with
(1) a long “pre-arcuate” spine; (2) an acutely inclined superior sacral surface; (3) pronounced
lumbar lordosis; and (4) minimal curvature and depth of superior sacral articular facets.
Others suggest predisposing factors, including size of the articular processes (Nathan, 1959),
large pars interarticularis vascular foramina (Miles, 1975), scoliosis (McPhee & O’Brien, 1980),
presence of lumbar-sacral transition vertebrae (Merbs, 1983), spina bifida (Merbs, 1989b),
facet joint distances (Ward & Latimer, 2005), and spina bifida occulta and anterior inclination
of the sacral table (Mays, 2006a). There may well be genetically based predisposing factors for
spondylolysis, but the mechanical environment prompting its appearance is required.
High frequencies of spondylolysis in laborers (Lane, 1893) and in other individuals involved
in mechanically demanding activities, such as in college football and other athletic sports
(Hoshina, 1980; Jackson et al., 1976; McCarroll et al., 1986; Soler & Calderón, 2000), support
the mechanical stress model. The low frequency of spondylolysis in industrial populations
engaged in activities involving minimal physical exertion is consistent with this interpret-
ation. For example, one sample of twentieth-century Americans has a prevalence of only 7%
(Moreton, 1966), and the Terry and Hamann-Todd collections involve similarly low preva-
lences (Roche & Rowe, 1951; Ward & Latimer, 2005; and see Fredrickson et al., 1984).
Eighteenth- and nineteenth-century Londoners from Spitalfields also have low frequencies
that are slightly greater in males than in females (2.2% vs. 0.6%; Waldron, 1993). These values
are lower than those in earlier populations, suggesting a decrease in physical demands as it
affects the lower back in British populations (Waldron, 1991a, 1991b; and compare with
Stirland, 1996).
A range of activities involving hyperextension of the back, perhaps accompanied by jarring
and twisting, are linked with spondylolysis (Merbs, 1989b; Ward & Latimer, 2005). Given the
broad range of behaviors that are associated with these defects, no specific mechanically
related activities or stresses appear to lead to spondylolysis (Bridges, 1989a; Merbs, 1989b).
Merbs (1989b) speculates that spondylolysis may have an adaptive value in that it may
engender flexibility of the lower back (see also Snow, 1974). Bird and coworkers (1980), for
example, reported that adults with the defect considered themselves “more supple in youth”
than adults who lack the defect. The defect seems to have a minimal negative influence on
physical performance. For example, college football players with the defect appear to lose
neither practice nor playing time, and they continue to play professionally in later years (see
discussion in Merbs, 1989b).
The close association between mechanical loading of the lower back and spondylolysis
indicates that the defect should be documented in archaeological skeletal series. In
northwestern Alabama, early prehistoric hunter-gatherers have a higher prevalence of
spondylolysis than late prehistoric agriculturalists (Bridges, 1989a). Statistical analysis
of the co-occurrence of spondylolysis and osteoarthritis reveals only a weak relationship
between the two conditions. Osteoarthritis prevalence is broadly similar between individuals
with and without spondylolysis. Structural analysis of long bones reveals a general increase in
mechanical demand in the agricultural groups, which seems to contradict the findings based
on spondylolysis prevalence (see Chapter 6). Bridges (1989a) speculates that spondylolysis
may be associated with unusual postures or specific activities affecting the lower spine rather
than overall activity levels (compare with Hoshina, 1980).
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4.2 Skeletal injury and lifestyle 121
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122 Injury and violence
spinous process is fractured and displays a pseudarthrosis (Knüsel et al., 1996). Clinical
evidence suggests that the fracture is due to highly forceful muscle contraction involving
muscles of the neck or activities involving scapular retraction toward the vertebral column
during rib elevation (Knüsel et al., 1996). Virtually all individuals affected, archaeological and
contemporary, are males. Although rarely reported, avulsion (disruption) injuries involving
vertebral end plates provide a potentially important perspective on spinal injuries. Unlike
spondylolysis, which involves gradual, repeated trauma leading to fracture, vertebral evul-
sions are acute injuries, such as caused today by “whiplash” trauma in automobile rear-end
collisions (Maat & Mastwijk, 2000). In these kinds of injuries, there is abrupt hyperflexion–
hyperextension. As documented in modern Holland, skeletal changes include anteriorly
displaced annular epiphyses, compression fracture, and anterior vertebral rim fracture (Maat &
Mastwijk, 2000). As with spondylolysis, males in this series have a higher prevalence than
females, strongly suggesting greater demands on the spine in males than females, likely
relating to labor involving rapid movement.
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4.2 Skeletal injury and lifestyle 123
Figure 4.4 “Parry” fracture of right ulna. In this case, the fracture site resulted in a disunion,
or pseudoarthrosis; Lambayeque, Peru. (Photograph by Haagen Klaus.) (A black and white
version of this figure will appear in some formats. For the color version, please refer to the
plate section.)
may have been caused by accidents and not by aggressors (Smith, 1997). Alternatively, the
populations were simply successful at fending off blows, thus protecting their heads from
injury.
Conversely, crania from the Santa Barbara Channel Islands have a very high frequency of
violence-related traumatic injuries (18.3%; 138/753; and see later) and few parry fractures
(<3%) (Lambert, 1994). Forearm fractures are equally distributed throughout the temporal
sequence, whereas cranial trauma has a distinctive peak frequency in the Early Middle period
(c. 1490 BC–AD 1150). In the southern hemisphere, at a Wari-era (AD 600–1000) site in Peru,
nearly one-third of the adults have cranial trauma and 4% of the left ulnae exhibit parry
fractures (Tung, 2007). Like the series from Tennessee, these findings suggest that the
attribution of forearm fractures to parrying is better understood in relation to broad patterns
of skeletal trauma rather than a single cause such as protection of the head.
Individuals in some settings likely did use the arm to avert blows to the head, thus resulting
in forearm injury. For example, in the Honokahua precontact sample from Hawaii (Pietru-
sewsky and Douglas, 1994), and various series from Australia (Dinning, 1949; Pretty &
Kricun, 1989; Webb, 1989, 1995), Nubia (Alvrus, 1999; Judd, 2008; Smith & Jones, 1910),
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124 Injury and violence
Figure 4.5 Medial view of distal right radii: healed Colles’ fracture (left); normal (right);
anatomical specimens. (Photograph by Paul Braly.)
North America (Smith, 1996, 1997), and South America (Tung, 2007), left-side fractures have
a higher prevalence than right-side fractures, sometimes significantly so (e.g., Honokahua:
68.8% left vs. 31.3% right). Perhaps this reflects the victim holding up the left arm for
protection while holding an implement in the right hand in order to inflict injury on the
opponent.
Although behavioral circumstances associated with parry fractures cannot be known in all
circumstances, ethnographic and historic documentation provides some perspective suggesting
that the injured person was a victim of violence. For example, hand-to-hand fighting is well
documented in native Australians (Webb, 1989, 1995). Parrying shields were widely used for
protection against clubs (Massola, 1963). Kricun (1994) notes that parrying fractures could have
arisen either from direct blows to the forearm or from blows hitting the shield with the forces
transmitted to the ulna. These observations, coupled with the high levels of cranial trauma,
indicate the strong likelihood that parry fractures arose in conflict situations. On the other hand,
historic sources for ancient Nubia indicate the presence of both violence (warfare with
their Egyptian neighbors: Buzon & Richman, 2007) and sport (stick fighting: Carroll, 1988).
Interestingly, the relatively high frequency of parry fractures in Sudanese Nubia has consider-
able time depth, lasting from at least the Bronze Age (2500 BC) to sometime during the Meroitic
period (post-400 BC) (compare with Alvrus, 1999; Judd, 2008; Smith & Jones, 1910).
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4.2 Skeletal injury and lifestyle 125
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126 Injury and violence
Figure 4.6 Antemortem diaphyseal fractures of the right radius (right) and left ulna (left);
Çatalhöyük, Turkey. (Photograph by Scott Haddow.) (A black and white version of this
figure will appear in some formats. For the color version, please refer to the plate section.)
greater loss of bone in females than in males is due to hormonal changes (estrogen
reduction) at menopause. The similarity in bone mass in comparing older Çatalhöyük
females and males in their bone mass may reflect the influence of activity and gender roles
on bone metabolism. Interestingly, with the exception of two young adults with metabolic
disruption and abnormal bone, the population shows no evidence of fracture relating to
increased bone fragility. Thus, at least in this setting, age-based increases in fracture risk are
minimal, likely owing to a similarity in bone mass. Fractures certainly exist in this setting
(Figure 4.6), but the prevalence and risk are minimal.
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4.2 Skeletal injury and lifestyle 127
Overall, the study of fracture prevalence in relation to age and risk in archaeological
contexts underscores the very important point that lifestyle and the lived experience are
much more complex than the clinical record (largely based on a relatively narrow spectrum of
variation in recent humans) would indicate. Like the Çatalhöyük series from Turkey, the
Libben series from eastern North America expresses a very different pattern than in recent
urban populations (Lovejoy & Heiple, 1981). Analysis of the number of years at risk reveals
that fracture rates peaked in two age groups, adolescence/young adulthood (15–25 years) and
old adulthood (>45 years). This pattern may reflect an elevated risk of trauma due to warfare
and conflict, especially for the younger group. Lovejoy and Heiple (1981) note that women
and men were equally affected by injury, indicating that the elevated rates were due to
accidents associated with activities such as hunting forays and travel (in adults) or play (in
juveniles).
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128 Injury and violence
Figure 4.7 Atrophy and healed fracture of the right humerus; Shanidar, Iraq. The left
humerus is normal. (Photograph and copyright by Erik Trinkaus; reproduced with
permission.) (A black and white version of this figure will appear in some formats. For the
color version, please refer to the plate section.)
American rodeo athletes also have a high prevalence of head and neck injuries relative to
other regions of the body (Berger & Trinkaus, 1995). The pattern in rodeo athletes reflects the
dangers of riding highly irritated animals (e.g., Bos taurus, Equus caballus); namely, head and
neck injuries in rodeo athletes result from impacts after being thrown from these animals. By
logical extension, the high prevalence of head and neck injuries in Neanderthals relates to
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4.2 Skeletal injury and lifestyle 129
Group
Neanderthal (n¼17) 29.6 14.8 25.9 3.7 3.7 11.1 11.1
Bt-5 (n¼223) 1.8 51.1 22.4 6.3 3.1 9.0 6.3
Libben (n¼94) 6.4 21.3 29.7 0.0 0.0 39.4 3.2
Nubia (n¼160) 10.6 6.9 53.1 1.9 3.8 22.6 1.3
London (n¼1730) 6.2 7.0 31.6 24.4 0.2 23.6 7.0
New York (n¼11 959) 13.7 12.3 25.3 21.9 0.5 20.6 5.6
New Mexico (n¼792) 1.6 12.5 23.1 23.6 2.1 11.1 25.9
Rodeo (n¼181) 39.2 9.9 25.9 6.1 3.3 6.1 9.4
hunting activities, especially involving close encounters with medium-sized ungulates (Berger
and Trinkaus, 1995). Use of spears would have necessarily placed hunters in close proximity
to, and hence, bodily injury from, their enraged prey.
Angel (1974) attempted to document temporal trends by comparing post-Mesolithic arch-
aeological skeletons (n¼2125) and modern Euroamerican samples. This comparison reveals
several general characteristics of accidental injury: adult males have more injuries than adult
females; older adult females have more fractures than older adult males (Buhr & Cooke, 1959);
and recent populations have far more fractures than earlier populations. The higher frequency
of skeletal injury in adult males than females in Angel’s samples likely reflects a greater
exposure of men to trauma. The reversal in older adults likely indicates the effects of bone
loss, especially in postmenopausal women, and especially in modern industrial populations
where lifestyles are distinctively different from traditional societies sampled from the arch-
aeological past (Glencross & Agarwal, 2011). Angel (1974) contends that the higher fracture
prevalence in the modern samples results from the hazards associated with twentieth-century
technology and urban living, such as the reliance on automobiles, walking on staircases, and
urban crowding.
In addition to the anatomical region analysis discussed earlier, analysis of fracture preva-
lence for the skeleton can provide insights into local adaptation, such as those associated with
particular subsistence patterns. Contrary to Steinbock’s (1976) assertion that trauma
decreased with the foraging-to-farming transition, there is considerable variation in patterns
and prevalence of fracture that are largely driven by local circumstances (various in Cohen &
Armelagos, 1984; Cohen & Crane-Kramer, 2007; Steckel & Rose, 2002). This record suggests
that while injury prevalence owing to accidents may have decreased with the rise of farming
in some regions, there are no clear temporal generalizations that can be made about injury
relating to accidents. Nevertheless, the recent analyses of injury viewed in temporal perspec-
tive in specific communities and regions provide critical perspective on adaptation, lifestyle,
and risk.
Some North American prehistoric maize agriculturalists show generally low prevalence of
accidental injuries. At Moundville, for example, the total frequency of bones affected is only
0.7% (Powell, 1988). Many of these fractures are associated with lower-status individuals; no
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130 Injury and violence
high-status adult females have accident-related traumatic injuries (Powell, 1988). Thus, at
least in this setting, sedentary populations had a relatively accident-free lifestyle, and high-
status, elite women may have been spared altogether from activities resulting in accidental
injury (lack of elite males in the skeletal series precluded their assessment). In contrast, at the
late prehistoric Mississippian site of Chucalissa, Tennessee, high-status males have far higher
frequencies of fractures than low-status males or high-status females (Lahren & Berryman,
1984). Unlike the Moundville population, most of the injuries can be attributed to violence. In
contrast, late prehistoric intensive agriculturalists from the Dickson Mounds site show an
increase in fracture prevalence compared with earlier less-intensive agriculturalists (Good-
man, Lallo et al., 1984). This suggests that injury risk increased with the shift to more
intensive farming in this setting of the American Midwest.
In prehistoric Thailand, Domett and Tayles (2006) tested the hypothesis that more estab-
lished agriculturalists in this setting would be exposed to increased risk of injury from
extensive land clearance and other farming activities. Comparisons of Neolithic farmers with
later Bronze Age intensive farmers reveal a 10-fold temporal increase in fractures, from 0.3%
to 3.0%. Some of the fractures are parry and craniofacial, suggesting some evidence for
increased violence and competition for resources. The finding of increased skeletal injury in
the more intensive Bronze Age farmers in Thailand is consistent with results comparing urban
and rural populations in Medieval Britain. That is, the farming population from Raunds shows
four times the frequency of fracture than the urban samples (19.4% vs. <10%) (Judd &
Roberts, 1999; Roberts & Manchester, 2005). The breadth and risk of activities of farming in
Medieval England – such as plowing, tree felling, herding, transport of products – likely had
considerably more risk than village life where individuals were more tethered to the home and
craft specialization.
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4.3 Intentional injury and interpersonal violence 131
direct evidence of violent interaction. Viewed in light of living conditions and social,
economic, and political context in archaeological settings, much can be learned about the
origins and use of violence in past populations.
The skeletal and paleopathological literature on violence and injury is dominated largely by
descriptions of limited samples, such as individual instances of arrow wounds (Armendariz
et al., 1994; Bovee & Owsley, 1994; Lewis & Lewis, 1961; Pryor, 1976), decapitation (Bennike,
1985; Harman et al., 1981; Jones, 1908; McKinley, 1993; Newman & Snow, 1942; Smith,
1993; Ubelaker, 1988; Wakely & Bruce, 1989; Webb, 1974;), dismemberment (Brothwell,
1971; Smith, 1993; Snow & Fitzpatrick, 1989; Webb, 1974), sacrifice and ritual violence
(Bennike et al., 1986; Fowler, 1984; Kanz & Grossschmidt, 2006; Klaus, Centurion et al., 2010;
Pijoan & Mansilla, 1997; Stead et al., 1986), mutilation, especially scalping (Allen et al., 1985;
Hamperl & Laughlin, 1959; Hoyme & Bass, 1962; Lesley, 1995; Miller, 1994; Neumann, 1940;
Ortner, 2003; Ortner et al., 2008; O’Shea & Bridges, 1989; Smith, 1995), and cranial depressed
fractures or other forms of head injury (Haas & Creamer, 1993; Lux, 1994; Manchester &
Elmhirst, 1980; Wenham, 1989; and many others). Because many of these studies usually
involve the investigation of one or several skeletons only, they frequently provide limited
information for inferring conflict behavior in the populations from which they were drawn.
However, placement of these few skeletons within their larger, contextualized settings where
various lines of evidence about society at a particular place and time offers important insights
into circumstances and causes surrounding violent encounters (Arkush & Tung, 2013; Cook,
2012; Judd & Redfern, 2012; Klaus, 2012; Milner, 1999; Milner et al., 1991; Montgomery and
Perry, 2012; Tucker, 2014; Tung, 2012a, 2012b; Walker et al., 2011).
Contextually oriented approaches in bioarchaeology are revealing the fundamental import-
ance of skeletal data for documenting patterns of violent behavior in the past, ranging from
interpersonal conflicts between two individuals to full-scale warfare involving scores of
participants. The strength of these approaches is their reliance on context, derived from the
clinical and epidemiological literature on modern people, ethnography, climate history,
ethnohistorical accounts, and archaeology for interpreting violence, its origins, its conse-
quences, and its distinctive skeletal signatures. Indeed, skeletal trauma is among the most
common pathological conditions in the bioarchaeological record. The interpretation of the
evidence of injury requires a detailed series of interpretive pathways by the investigator
(Figure 4.8). Reflecting this level of attention paid to interpreting trauma in archaeological
contexts, there is a growing record of insight into the history of violence going back many
thousands of years.
In the following discussion, a series of bioarchaeological studies are assessed that collect-
ively illustrate the enormous variation in skeletal evidence of conflict in past populations. This
discussion is intended to be neither comprehensive nor synthetic; rather, a representative
sample of various kinds of skeletal injury useful for identification of patterns of conflict is
discussed. These studies are drawn from a diverse set of geographical and cultural settings,
including the American Midwest (Norris Farms, Riviere aux Vase), Great Plains (McCutchan-
McLaughlin, Crow Creek, Larson Village), and Southeast (Koger’s Island, Chickamauga),
southern California Pacific coast (Santa Barbara Channel islands), Arctic (Kodiak Island,
Admiralty Island, Saunaktuk), American Southwest (Anasazi), Latin America (multiple local-
ities), Easter Island, Sudan (Nile Valley), across the continent of Australia, and Southeast Asia
(Cambodia). The study of human remains from these settings provides important perspectives
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132 Injury and violence
Inference:
Post-depositional damage
during site formation?
yes
Is there active yes A perimortem injury or
new bone formation? occurred before death
no
Figure 4.8 Model for interpreting traumatic lesions on skeletons. (Walker, 2001a;
reproduced with permission of Annual Reviews Inc.)
on violence, chiefly in nonliterate tribal and state societies, mostly before contact with
Western populations.
Lastly, patterns of northern European violence, ranging from interpersonal conflict to
large-scale preindustrial warfare and execution, and the subsequent spread of European forms
of violence to the New World and military campaigns in North America, are addressed. These
investigations illustrate variability of traumatic injury in skeletal remains as well as the
impact of violence on different components of the populations involved (e.g., gender, age,
status) within a specific period or consequent to major adaptive shifts.
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4.3 Intentional injury and interpersonal violence 133
Figure 4.9 Cut marks on adult frontal bone (scalping); Norris Farms, Illinois. (From Milner &
Smith, 1990; reproduced with permission of authors and Illinois State Museum.) (A black
and white version of this figure will appear in some formats. For the color version, please
refer to the plate section.)
individual skeletons indicate that some were attacked from behind and some from frontal
assault. The former shows that some individuals attempted to flee. Evidence for mutilation is
especially abundant in the Norris Farms series. Multiple perimortem cutmarks on cranial
vaults (especially on frontal bones) produced by stone tools reveal that at least 14 individuals
had been scalped (Figure 4.9). Individuals missing their skulls and having cut cervical vertebra
show that they had been decapitated. Presence of cutmarks on multiple articular regions of
postcranial bones evinces widespread dismemberment of limbs. Many instances of punctures
and gouges in skeletal elements, mostly produced by carnivores, suggest that a significant
proportion of victims (n¼30) were exposed above ground for a period of time before
interment.
Active bone infections, articular joint dislocations, and partially healed bone fractures and
other trauma denote the presence of severe and long-standing disabilities for many individ-
uals at the time of their deaths. These conditions may well have impaired their ability to
escape confrontation, leading to early death (Milner, 1995; Milner et al., 1991).
The presence of victims – individuals who clearly exhibit evidence of violent trauma – in
individual burial pits or pits containing a few individuals indicates that violence occurred over
the entire period of the use of the cemetery, which spanned at least several decades. Over this
timespan, not all of the victims died outright as a result of violent encounters. For example, a
number of adults, both non-survivors and survivors of attack, display embedded chert
projectile points (Figure 4.10a,b).
The pattern of deadly conflict at Norris Farms is similar to that of ethnographic small-scale
societies where violence is endemic (Chagnon, 1992; Harrod et al., 2012; Heider, 1979;
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134 Injury and violence
(a)
Figure 4.10a Arrow point in adult sternum; Norris Farms, Illinois. (From Milner & Smith,
1990; reproduced with permission of authors and Illinois State Museum.) (A black and
white version of this figure will appear in some formats. For the color version, please refer
to the plate section.)
Schmidt & Schröder, 2001). Raids in these societies often involve ambush and surprise attacks,
but may also occur during chance encounters. Victims of attacks in these groups can include
individuals of all ages and both sexes. At Norris Farms, fully one-third of the adults were
victims of violent attack at one point in their lives, and they include equal numbers of adult
males and females; only two are juveniles (<15 years of age). The equal number of adult
females and males with traumatic injuries is different from ethnographically observed small-
scale societies where males are the predominant victims (Chagnon, 1992; Divale & Harris,
1976; Harrod et al., 2012; Heider, 1979; Keeley, 1996; and others). Female captives provide
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4.3 Intentional injury and interpersonal violence 135
(b)
Figure 4.10b Projectile point in left tibia; Norris Farms, Illinois. (From Ryan & Milner, 2006;
reproduced with permission of authors and Elsevier Ltd.) (A black and white version of this
figure will appear in some formats. For the color version, please refer to the plate section.)
economic return – women’s labor in many societies is essential for food collection and
preparation. Relatively, more women may have been killed at Norris Farms because of their
burden on the attacker’s resources, or it may simply have been too much trouble for the
attackers to bring captives back to their home village (Milner, 1995; Milner et al., 1991).
Warfare at Norris Farms is tied to the highly dynamic sociopolitical circumstances that
characterize this region of the American Midwest during later prehistory. The Oneota represent
an intrusion into the central Illinois River valley, replacing a somewhat more organizationally
complex system (Mississippian). Clear evidence of social tensions is indicated by fortifications
and defensible settlement locations here and throughout the late prehistoric Midwest during a
period of population expansion and growing tensions over and competition for resources
(Milner et al., 2013). Populations occupying the region may have been in competition over
productive lands and resources concentrated in river valleys (Milner et al., 1991). Violence in
this case may have been a strategy employed to gain control of these highly valued resources.
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136 Injury and violence
females (n¼14) than males (n¼5) with cranial injuries suggests that women were the preferred
target of violence. For some of these women, violent encounters may have occurred on more
than one occasion – five female crania have multiple healed depressed fractures. One of these
individuals has a severe depressed fracture and an accompanying large incision on the
occipital bone as well as multiple fractures on the left and right parietal bones.
Although other women may have been responsible for the cranial injuries in this population
(compare with Burbank, 1994), the demographic characteristics of the injured group suggest
that males were the primary aggressors. Males and females show a very different age pattern
of injury: females’ peak age of trauma is the early adult years, while male trauma is evenly
distributed across age groups. Male trauma is oriented toward the front of the vault, and
female trauma is distributed throughout the cranium. Wilkinson and Van Wagenen (1993)
suggest that violence directed at women by men or women against women or co-wives in
polygamous societies is consistent with ethnohistoric accounts of violence in American
Midwestern native populations (Wilkinson & Van Wagenen, 1993).
Crow Creek
The study of human remains from various late prehistoric sites in the Missouri River valley
located in the present states of South Dakota and North Dakota reveals evidence for violent
confrontations between tribal groups competing for overlapping resources and territory
(Bovee & Owsley, 1994; Hollimon & Owsley, 1994). The proto-Arikara (c. AD 1325) skeletal
series from the Crow Creek site supplies important details on prehistoric conflict (Gregg &
Gregg, 1987; Willey, 1990; Willey & Emerson, 1993; Zimmerman et al., 1981). Analysis of
some 500 individuals buried in a single pit indicates that nearly all members of a village were
massacred during the course of a single raid. The presence of carnivore tooth marks and
weathering reveals that following the attack, the deceased were exposed for a period of time
prior to their burial by returning survivors or allies (compare with Milner & Smith, 1989).
Analysis of the human remains suggests that although all of the deceased were victims of a
single attack, violence had a well-established history in the Crow Creek villagers; a significant
number of massacre victims had healed violence-related injuries, including scalping (Willey,
1990; Willey & Emerson, 1993).
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4.3 Intentional injury and interpersonal violence 137
Virtually all individuals in the series have unhealed cutmarks on frontal bones and other
cranial elements indicating removal of the scalp with a stone knife at or following the time of
death. Forty percent of the victims have cranial depression fractures, mostly located on the top
and sides of the vaults. In addition to scalping trauma, various other mutilations are common,
including tooth evulsions, alveolar and tooth fracture, nose removal, decapitation, and
dismemberment of both upper and lower limbs. The presence of sharp-force trauma on
mandibles – especially on ascending and/or inferior borders of rami – suggests that mutilation
also involved tongue excisions.
Demographic assessment of the Crow Creek population indicates that young adult males
(15–24 years of age) outnumber young adult females by a factor of two. Additionally, there
are twice the number of older adult females (45–59 years of age) than older adult males. The
absence of young women may reflect captive taking, escape, or actual demographic compos-
ition of the population from which victims are drawn (Willey & Emerson, 1993). Similarly, the
paucity of elderly men reflects their escape or actual demographic composition. It is unlikely
that older males were captured because they would have represented a relatively low eco-
nomic return for the raiders. Perhaps older males may be missing owing to earlier raids and
endemic warfare (Willey & Emerson, 1993).
Larson Village
During the historic period, the Arikara occupied a series of temporally successive villages as
they migrated northward up the Missouri River valley. One such village was decimated by
violence. At the seventeenth-century Larson Village site in northern South Dakota, 71 skel-
etons from house floors display evidence of violent death and mutilation (Owsley et al., 1977).
About one-third (34%) of the victims had been scalped. The mutilation patterns are similar to
those displayed in the Crow Creek population, including dismemberment, decapitation, and
facial, dental, and cranial vault trauma, as well as tongue excision. Skeletal modifications on
one young adult female run the gamut of violent injury and mutilation: “the left side of her
skull had been broken away, though cuts on the frontal, right parietal and occipital indicated
she was scalped. The distal diaphysis of her right radius and ulna are articulated in anatomical
position. . .A knife cut was made through the soft tissues in order to free the hand as a trophy
or possibly to secure a bracelet. Epiphyseal areas of both bones were broken. After severing
the muscles and tendons, the assailant simply broke the hand free. Other bones have been cut,
including five right ribs, the ventral and posterior surface of the right clavicle, the right
scapula, the right tibia and both femora. A deep cut near the distal epiphysis of the right
humerus must have resulted while separating the upper and lower arm. Cuts on the femurs are
on the neck; the objective may have been to remove the legs from the body. Bones of both legs
are associated with the skeleton though neither was articulated when excavated” (Owsley
et al., 1977:125). This analysis indicates that the Larson Village was attacked, and the villagers
attempted to defend themselves in their individual houses (Bamforth, 1994). The presence of
numerous unburied remains indicates that the Larson Village ceased to exist following the
conclusion of the attack.
Demographic composition of the historic Larson Village site and prehistoric Crow Creek
massacre victims are similar; both series contain fewer young adult females and juveniles
than young adult males. This pattern suggests that children and women may have been
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138 Injury and violence
captured rather than killed in the attack, which is documented historically in the region
(Lowie, 1954).
Analysis of scalping patterns reveals changes in warfare prior to and during the contact
period in the northern Plains (Owsley, 1994). Comparison of a large sample of late prehistoric,
protohistoric, and early historic crania (n¼751) from 15 archaeological sites indicates that
throughout the period, both males and females were at equal risk of being scalped. This risk
greatly increased for men but not for women in the early historic period; instances of scalping
tripled for males and halved for females. Most of the male victims are young adults (20–34
years of age), which likely reflects the deaths of warriors who were killed either during raids or
in defense of home villages (Owsley, 1994). Owsley’s (1994) analysis establishes that death
from violence was present throughout the late prehistoric and historic occupation of the
Plains. This violence was occasionally punctuated by eruptions of large-scale warfare
resulting in numerous deaths at one time (e.g., Crow Creek and Larson Village massacres).
Thus, contrary to earlier assertions (Newcomb, 1950), analysis of skeletal evidence of conflict
discloses that warfare and inter-group conflict did not originate during the early period of
contact with Europeans or Euroamericans, but was a well-established part of the cultural and
social behavioral repertoire of prehistoric societies living in the region. The increase in Arikara
male deaths during the historic period likely reflects elevated frequency of confrontations,
especially with the encroaching Sioux from the east (Owsley, 1994). The overall similarity
between precontact (e.g., Crow Creek) and postcontact (Larson Village) Arikara skeletal
injuries indicates an enduring pattern of conflict in this region. In the precontact Plains
setting, conflict appears to have been triggered by food shortages and stress generally, as is
suggested by the presence of stress indicators in human remains as well as paleoclimatological
evidence for periodic droughts after AD 1250 (Bamforth, 1994). Archaeological evidence also
suggests that new populations were migrating into the region during this time, resulting in
increased competition for productive lands. In view of these new developments, Bamforth
(1994) contends that the stage was set for increased violence in later prehistory, a pattern
which was exacerbated by the spread of Sioux into the Missouri River valley.
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4.3 Intentional injury and interpersonal violence 139
20
Males
15 Females
Number of Individuals
10
0
10 20 30 40 50
Age (years)
Figure 4.11 Sex-specific mortality profiles (age versus number of individuals); Koger’s
Island, Alabama. (Adapted from Bridges, 1996; reproduced with permission of
John Wiley & Sons, Ltd.)
violence in this society may have been profound – some 21% of the total number of
individuals were recovered from multiple-interment (mass) graves. The population contains
a relatively large number of infants (about 25% of the total), a pattern that is characteristic of
many preindustrial, farming-dependent populations (Bocquet-Appel et al., 2008; Lambert,
2009). Adult (>15 years of age) males and females have very different age-at-death profiles
(Figure 4.11). Female deaths peak slightly during the late teens and early twenties, progres-
sively fall to a low point in the thirties, and again rise in the forties. Male deaths are few in
number during the late teens, but high in the thirties. Presumably, many of these deaths
resulted from violent intergroup encounters. Age composition of the small burials (containing
fewer than five individuals) is different from age composition of the multiple burials (five or
more individuals). The former is dominated by infants and younger juveniles. The latter has an
unusual peak in the thirties, and most of these deaths are adult males. The pattern is
distinctive in that there are relatively few women or children in the multiple burials, suggest-
ing that women and children had either escaped or were captured; males may have died while
protecting the village from aggressors.
The loss of adults, and especially men who were responsible for protection of the group and
acquisition of resources not acquired by women (e.g., animal protein), would have had far-
reaching consequences for the population’s ability to mitigate stress in a hostile setting. In
later prehistory, political systems and population size declined in this and some other regions
of the American Southeast and Midwest (Anderson, 1994; Milner, 1990; Steponaitis, 1991).
This hostile environment may have contributed to the decreased presence of these groups in
later prehistory (Bridges, 1996).
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140 Injury and violence
Comparisons of the Koger’s Island sample with other late prehistoric populations in the
region suggest that conflict was highly localized. Less than 1% of the late prehistoric
(Mississippian period) Moundville skeletons (n¼564) from west-central Alabama have skeletal
injuries, and only a handful of these are from violence (one piercing wound and three with
cuts) (Powell, 1988). Similarly, upriver in the eastern Tennessee River valley, Mississippian-
period populations display relatively few violence-related injuries (Smith, 2003). Populations
from the preceding Late Woodland period in the Tombigbee River valley display an abun-
dance of injuries deriving from violence (Cole et al., 1982; Hill, 1981; Welch, 1990). Some of
the deceased who had died violently were buried in multiple-individual graves. Thus, in late
prehistoric west-central Alabama, there was an apparent decline in violence from the Late
Woodland to Mississippian periods, which Steponaitis (1991) argues was brought about by
increasing control of the regional population by more centralized polities, such as that at
Moundville. This pattern of declining interpersonal violence is certainly not a general out-
come, however. For example, analysis of the Cumberland River valley region in Tennessee
reveals a significant presence of injuries, including scalping, blunt-force trauma, and other
injuries in the Mississippian period (Worne et al., 2012). In general, however, there is an
overall spike in violence in later prehistory in Eastern North America (Milner et al., 2013), but
prevalence is highly variable.
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4.3 Intentional injury and interpersonal violence 141
(a)
(b) (c)
(d) (e)
Figure 4.12 Depressed fractures in adult crania: ellipsoidal parietal injury (a), deep
circular injury (b), multiple circular injuries (c), ellipsoidal occipital injury (d), circular
parietal injury (e; arrowhead points to residual fracture line); Santa Barbara Channel
Islands, California. (From Walker, 1989; reproduced with permission of John Wiley &
Sons, Inc.)
with a right-handed perpetrator striking the left side of the head of the victim (Lambert, 1994).
This patterning of nonlethal trauma is remarkably similar to trauma observed in the Yano-
mami foragers of Venezuela (Chagnon, 1992). Yanomami men attack other males with heavy
wooden clubs. Although numerous casualties result from these encounters, they are rarely
lethal.
Adult female cranial injuries are haphazardly distributed on the face and vault – only about
one-third of the injuries are on the frontal bone (Lambert, 1994). This random distribution of
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142 Injury and violence
depressed fractures indicates that although females were occasionally involved in face-to-face
attacks, most were from other directions, including from behind (e.g., while fleeing from an
attacker). Additionally, some of these nonfrontal injuries could be from accidental causes
(Walker, 1989).
The presence of projectile injuries and associated projectile points in some individuals
reveals appreciable numbers of deaths caused by violence: of 1744 individuals, 3.3% had
been killed or wounded by single or multiple projectile injuries. Unlike the cranial trauma,
the majority of projectile victims (at least 71%) died from their wounds, indicating the
lethal intentions of the attacker. The demographic composition of individuals with either
lethal (projectile wounds) or nonlethal (cranial depressed fractures) injuries is similar in at
least two respects: first, young and mature adult males are the most affected; and second,
children, older adults, and women are the least affected. Like cranial trauma, aggression
leading to injury and death involved primarily adult males under 40 years of age
(Lambert, 1997), which is consistent with patterns observed for many nonindustrial
communities globally (Chagnon, 1992; Meggitt, 1977; and see discussion in Lambert,
1994).
The temporal patterns of nonlethal and lethal skeletal injury are distinctive in prehistoric
southern California coastal populations. Nonlethal cranial depressed fractures are common
in the Early Middle period (1490 BC–AD 580), whereas lethal projectile injuries are far more
common in the Late Middle period (AD 580–1380) (Lambert, 1994, 1997). The increase in
frequency of projectile injuries may be tied to the adoption of the bow-and-arrow in
California during the sixth century AD (Lambert & Walker, 1991). Perhaps the rapid
adoption of the bow-and-arrow at this time may have been fostered by competition and
conflict between populations in North America generally (Blitz, 1988). Regardless of the
motivation for increased lethal violence, the later peak in projectile injuries signifies a shift
to more serious – and deadlier – forms of conflict in later prehistoric southern California
populations.
The availability of abundant bioarchaeological, archaeological, and climatological data
provides a comprehensive understanding of factors that may have motivated violent behav-
ior in the Santa Barbara Channel region. Analysis of tree ring and other climatological data
suggests that the Middle period saw an increase in environmental instability, periodic
droughts, and decreased terrestrial resource productivity. These changes, coupled with
warming of the Pacific Ocean during this time, reduced marine productivity (Lambert,
1994, 1997, 2007) along with increased population size and territoriality (Kennett, 2005).
In light of these changes, Lambert (1994) proposes that elevated competition from increased
resource stress may have engendered an increase in violence in these populations. This
hypothesis is consistent with other skeletal data showing a decline in quality of life and
increase in health stress (Lambert, 1993, 1994, 2002c; Walker, 1986; Walker & Lambert,
1989; and see Chapters 2 and 3). Along with the increase in trauma and disease during this
time, there is evidence for increasing social complexity. For example, burial of high-status
grave goods (e.g., beads) with infants is more common during the peak in drought in later
prehistory, suggesting ascribed rather than achieved social rank. Perhaps increased inter-
group competition – and, hence, increased violence – for scarce resources during episodes of
environmental degradation fostered more complex social organization in later prehistory
(Fischman, 1996).
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4.3 Intentional injury and interpersonal violence 143
Far from being a peaceful setting of the New World, the analysis of violence in the Santa
Barbara Channel island region reveals the presence of endemic violence, peaking during key
periods of resource stress and sociopolitical change. Similarly, analyses of a large bioarch-
aeological database of 16 820 burials from 329 archaeological series from central California
(San Francisco Bay area, Central Valley, and Sierra Nevada foothills) documents clear
patterning of three forms of violence-related trauma, including trophy-taking/perimortem-
dismemberment and sharp-force and blunt-force trauma (Pilloud et al., 2014; Schwitalla
et al., 2014). At least half of the sites represented in the central California database express
some form of violence. Evidence for all three forms of conflict-related injury is well repre-
sented in the region and in highly patterned ways. With regard to perimortem dismember-
ment, there is a low, but significant presence, in contrast to southern California where it is
completely absent (Andrushko et al., 2005, 2010; Bertelink et al., 2014). This form of violence
peaked during the Early Middle period (500 BC–AD 420) and then largely disappeared. Blunt-
force trauma is also well represented, with males and older adults having the highest
prevalence in general (Figure 4.13). This form of trauma peaked very late in the sequence,
being mostly limited to the eighteenth and nineteenth centuries when there was a four-fold
increase in males and two-fold increase in females. The highest prevalence during this period
was in regions that were contested by multiple tribes following European encroachment. The
pattern of cranial trauma shows that the peak is considerably later in central California
(historic period) than southern California (Early Middle period, 1490 BC–AD 580). Moreover,
unlike the nearly 20% prevalence in southern California (Walker, 1989), the prevalence in
central California is well below 10%.
Sharp-force trauma (from projectiles) is also well represented in the central California
region, but with a different pattern than blunt-force trauma. In this regard, sharp-force
trauma is present in 7% of individuals, and is much more common in males (11%) than in
females (5%). The relative prevalence for females is higher in adolescent and younger
individuals. Like blunt-force trauma, the peak frequency of sharp-force trauma is in the
historic period, with the peak being especially higher for males (16%). Overall, while males
are clearly more often involved in violent encounters, females were also engaged in confron-
tations, not just as recipients, but as combatants. The historical records and ethnographic
observations document a greater involvement of males in conflict, but also document the
important role of women, including female aggression directed at other females (Goldschmidt
et al., 1939).
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144 Injury and violence
0.04
0.03
Males
Trophy Taking
0.02
0.01
Females
0.00
0.16
Males
Proportion of Burials Exhibiting
0.12
Blunt Force Trauma
0.08
0.04
Females
0.15
Sharp Force Trauma
0.10
Males
0.05
Females
0.00
Figure 4.13 Graphs depicting the relative frequency of trophy taking, blunt-force trauma,
and sharp-force trauma in adult females and males from Early to Protohistoric/historic
central California. (Schwitalla et al., 2014; reproduced with permission of authors and
Elsevier Ltd.)
Saunaktuk
The study of human remains from the Saunaktuk site located east of the Mackenzie Delta in
the Canadian Northwest Territories, Canada, provides compelling evidence for violent con-
frontation between native groups (Melbye & Fairgrieve, 1994; Walker, 1990). Human remains
from this locality represent a minimum of 35 Inuit Eskimo villagers, some of whom died
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4.3 Intentional injury and interpersonal violence 145
violently in the late fourteenth century AD. Evidence for violent death and body treatment in
this Arctic setting is indicated by extensive perimortem skeletal modifications, including knife
cuts, slashing, piercing, gouging, and splitting of long bones (Melbye & Fairgrieve, 1994).
None of the remains were purposefully buried. Most of the victims are juveniles (68.6%),
suggesting that adults – particularly males – may have been away hunting, leaving a
relatively defenseless group vulnerable to attack.
Hundreds of knife cuts, especially around articular joints and the neck (e.g., occipital
condyles and upper cervical vertebrae), indicate the practice of dismemberment and decapi-
tation. Numerous cuts on facial bones on most victims identify widespread facial mutilation
or disfigurement. Many other cuts, such as on clavicles and scapulae, reflect an overall pattern
associated with purposeful dismemberment, removal of muscle and other soft tissues, and
intentional mutilation. Unique to the Saunaktuk skeletal series is the presence of gouges at the
ends of long bones. Adult distal femora from two individuals display large perimortem
mediolateral gouges passing completely through the cortical and cancellous tissue. These
modifications are consistent with oral tradition describing a type of torture whereby the
victim’s knees were pierced and the individual was dragged around the village by a cord
passed through these perforations.
With few exceptions, long bones had been split into scores of longitudinal sections. The
surfaces of split bones are smooth and display tiny step fractures identical to bone modifica-
tions in butchered animal bones. Presumably, this breakage pattern was done in order to
extract the nutritionally rich marrow for consumption (Melbye & Fairgrieve, 1994; and
compare with White, 1992). The striking similarity between butchered animal remains found
in archaeological sites and the Saunaktuk human skeletal assemblage suggests that the
deceased had been cannibalized. In summary, at least some members of this late prehistoric
Saunaktuk population had been tortured, and all were murdered, mutilated, and cannibalized.
There is a rich historical record that provides a context for understanding violence between
native groups living in this region of the Arctic. In areas where Inuit and Native Americans
came into frequent contact – such as the Mackenzie Delta region – violent interactions
between the groups were commonplace. Oral traditions and historical accounts describe the
horrific nature of inter-group violence. For example, Samuel Hearne, who explored the region
in the late eighteenth century for the Hudson Bay Company, observed the murder and
mutilation of Inuit villagers by a group of Native Americans. He noted that “the brutish
manner in which these savages used the bodies they had so cruelly bereaved of life was so
shocking, that it would be indecent to describe it. . .” (Hearne, 1971:155; quoted in Melbye &
Fairgrieve, 1994:73).
The massacre at Saunaktuk may not have been an isolated occurrence in the Arctic region.
Preliminary evidence from Admiralty Island, Alaska, indicates the presence of a small number
of broken long bones and ribs as well as perimortem cutmarks produced by stone tools (Irish
et al., 1993). Although the evidence is limited, the patterns of modification are similar to those
identified from Saunaktuk (Melbye & Fairgrieve, 1994) as well as other sites in North America
where cannibalism was probably present (American Southwest: Redmond, 1994; Turner,
1993a; White, 1992; and see the following discussion).
On Kodiak Island, Alaska, skeletal modifications suggestive of dismemberment and canni-
balism have been described. At Uyak Bay, Hrdlička (1944) briefly documented a series of
scattered human remains, some of which had been broken in a manner which he interpreted
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146 Injury and violence
as representing a practice of cannibalism. Analysis of remains from the Uyak and Crag Point
sites shows the presence of culturally modified remains of women and children, but not men
(Simon, 1992; Simon & Steffian, 1994; Steffian & Simon, 1994; and see Urcid, 1994). This
study reveals a small subsample of remains displaying cutmarks from disarticulation and
defleshing, drill holes, perimortem breakage, and longitudinal fracturing. Thus, cannibalism
was likely present in precontact Koniak Island native groups, albeit probably not at the levels
envisioned by Hrdlička.
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4.3 Intentional injury and interpersonal violence 147
Figure 4.14 Fractured and longitudinally split humeri; Mancos Canyon, Colorado. (Tim
D. White. Prehistoric Cannibalism at Mancos 5MTUMR-2346. © 1992 Princeton University
Press. Reprinted by permission of Princeton University Press.)
and longitudinal fracturing (Figure 4.14). For example, humeri shafts are highly fragmented,
which was accomplished by hammerstone percussion and anvil fracturing (White, 1992:238).
These patterns of modification are similar to fracture patterns of animal bones resulting from
removal of flesh, disarticulation, and marrow extraction (White, 1992). The strong similarity
between bone modifications associated with processing of humans and mammals in the
region indicates that cannibalism was likely practiced at Mancos Canyon and other sites in
the region.
White (1992) notes the well-documented evidence of warfare and violence in the ethno-
historical and archaeological record of the American Southwest, including defensive sites,
intentionally burned habitations, as well as remains of deceased whose deaths had been
violent (Haas & Creamer, 1993). Based on the evidence – including lack of projectile injuries,
the pattern of body modification, and the extreme reduction of skeletal materials found at
the site – he contends that it is not possible to determine the reasons for cannibalism.
Several possible scenarios emerge, such as ritualized cannibalism involving killing and
eating of enemies, or perhaps the Mancos Canyon population engaged in culinary
cannibalism – the population was so starved that they consumed friends, associates, and
relatives (White, 1992). Well-documented historical cases of starvation cannibalism provide
important support for the latter model (Bonassie, 1989). The American Southwest can be
characterized as a marginal environment where seasonal cycles and resource variability led
to frequent episodes of crop failure and famine. Paleopathological analysis provides sub-
stantial evidence for nutritional stress in the prehistoric American Southwest (Lambert,
2014; Martin et al., 1991; Stodder, 1994; Stodder et al., 2010). These data alone, however,
do not provide sufficient evidence for why cannibalism occurred. In summary, although
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148 Injury and violence
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4.3 Intentional injury and interpersonal violence 149
(a) (b)
(c)
Figure 4.15 Cut marks on the tibial and femoral shafts (a) and subtrochanteric region of left
femur (b) are indicative of tool use in processing. Perimortem fracturing of the parietal bone
(arrow) was observed on a burned cranium (c); Cowboy Wash, Colorado. (Billman et al.,
2000; reproduced with permission of authors and the Society for American Archaeology.)
Among the world’s best archaeological records of body processing is from a number of
settings in Latin America, especially in the Andean region. However, the earliest osteological
evidence is from the Lagoa Santa, Brazil, a region where a remarkable bioarchaeological
record of Paleoamerican presence has been documented (Neves et al., 2007). At the site of
Lapa do Santo (8540 yBP), an isolated skull and cervical vertebra display cutmarks consistent
with purposeful removal of the head. In particular, the right articular process of the sixth
cervical vertebra and the right posteroinferior mandibular ramus displays distinctive cutmarks
produced with a stone tool (Strauss et al., 2013). The context for decapitation is unknown,
whether as part of the mortuary ritual following death or associated with the death of the
individual.
The rise of complex societies in South America, including the iconic Andean empires, is
associated with an expansive record of body processing, best described in general as
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150 Injury and violence
Figure 4.16 Pacatnamú mass burial, second layer of human skeletal remains; Jequetepeque
Valley, Peru. (Verano, 2008; reproduced with permission of author and Springer-Verlag.)
(A black and white version of this figure will appear in some formats. For the color version,
please refer to the plate section.)
mutilation, a form of treatment occurring at and associated with death. On the northern coast
of Peru, a series of mutilated human remains and contextual data from the site of Pacatnamú
(AD 1100–1350) in the Jequetepeque River valley are well documented (Faulkner, 1986; Rea,
1986; Verano, 1986, 1995, 2007, 2008). Located outside of the entrance of the primary
ceremonial complex, the remains of 14 adolescent and young adult males (mean age¼21
years) were recovered from the bottom of a defensive trench in three superimposed groups
(Figure 4.16). The groups were separated by erosional deposits, indicating that the deceased
had been deposited in the trench in three different burial episodes. Some skeletal remains for
each of the three groups show evidence of weathering, indicating that burial did not follow
death immediately. Delayed burial is also indicated by the presence of pupal cases of muscoid
flies representing different stages of insect growth in an open environment (Faulkner, 1986).
In the top-most group, injuries include multiple stab wounds (perforations on vertebrae and
ribs) sustained from different directions. This variable pattern of wound orientation suggests
that more than one individual may have been involved in the stabbing of the victim. In the
middle group, stab wounds are not evident. The presence of cutmarks on upper cervical
vertebrae indicates throat-slashing and decapitation. The bottom-most group also shows
evidence of decapitation. Five individuals from the middle and lower groups have bisected
manubria and fractured ribs, indicating that the chest had been opened forcibly. Collectively,
these perimortem traumas present a scenario of violent death and mutilation. Many of the
victims also have healed injuries (e.g., rib fractures, depressed cranial fractures) (Verano, 1986,
2008). Based on the age distribution, sex, and pattern of healed and unhealed injuries, Verano
(1986, 2008) speculates that the victims were war prisoners. This conclusion is well supported
by iconographic depictions in art from the region (Chimú and Moche) showing ritual mutila-
tion and sacrifice of war prisoners (Sutter & Cortez, 2005; Verano, 2005, 2014).
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4.3 Intentional injury and interpersonal violence 151
35%
30%
25%
Percentage
20%
15%
10%
5%
0%
Archaic Formative Final EIP MH MH/LIP LIP LIP/LH LH
(N = 311) (N = 52) Formative (N = 377) (N = 575) transition (N = 799) (N = 555) (N = 598)
(N = 114) (N = 103)
Figure 4.17 Frequencies of adult cranial trauma throughout Andean series (EIP, Early
Intermediate Period; MH, Middle Horizon; LIP, Late Intermediate Period; LH, Late Horizon).
(Arkush & Tung, 2013; reproduced with permission of authors and Kluwer Academic/
Plenum Press.)
The links between warfare and ritual violence are well documented in the Peruvian Andes
when the Wari empire rose to its fullest influence in the region (AD 600–1000). In addition to
expansionist strategies that included engineering superiority and ideological domination, the
Wari state has long been thought to have used violence to bring populations under its control
(Lumbreras, 1974). Tung’s (2007, 2012a, 2012b) test of this hypothesis shows that imperial
core populations engaged in head-taking and the display of war trophies (see later), and that
core and peripheral populations have high rates of cranial trauma (25%–30%). For the region
overall, these high rates are part of a generally increasing frequency of violence where the
attacker targets the victim’s head (Figure 4.17). Although frequencies between the core and
periphery are similar, the lethality and head wound locations differ between the sites. At the
peripheral site of Beringa, which was located on a defensible mesa, adults have a higher
frequency of posterior cranial trauma, a pattern consistent with individuals being injured
while fleeing during a raid. They also show the greatest proportion of lethal trauma, suggest-
ing that the attackers went into the conflict with the intention of killing their opponent (Tung,
2007, 2012a, 2012b). It would appear that their concerns for defense were well founded. At the
nearby site of La Real, males show significantly more cranial trauma than females, and the
vast majority of fractures are on the anterior of men’s skulls. All cranial trauma is healed.
Importantly, 70% of those anterior wounds are on the left side, suggesting face-to-face
conflict and a hit from a right-handed attacker. This is similar to what Walker (1989) and
Lambert (1994) reported for the Santa Barbara Channel Islands populations (see earlier). The
parallels in cranial fracture patterning, combined with ethnographic evidence of ritualized
conflict resolution in the Andes and nearby Amazonia, led Tung (2007, 2012a, 2012b) to
suggest that many La Real males were injured in physical conflict resolutions, like those
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152 Injury and violence
practiced by the Yanomami, the Oro-Wari of Amazonia (Conklin, 2001), and perhaps the
Chumash of the Santa Barbara Channel Islands.
Early accounts report on the practice of removal and curation of heads in native popula-
tions in the Andes and Amazonia, Ecuador, which continued well into the twentieth century
in some Amazonian groups (Verano, 1995). The use of decapitated heads is well represented in
Andean iconography, especially among the Nasca in south-central Peru (Verano, 1985) and
the Wari in the nearby highland Andes (Tung, 2008, 2012a, 2012b). Young adult males were
the favored victims, suggesting that they represent enemy combatants (Verano, 1995) or
perhaps local warriors. The presence of cutmarks on some skulls indicates that following
death, soft tissue had been removed with a sharp implement. Although variable in their
treatment, two features characterize trophy heads from the Nasca region: the frontal bone is
perforated with a single hole and the base of the skull is damaged, ranging from enlargement
of the foramen magnum to removal of most of the cranial base. The frontal bone perforation
was used to support a rope handle. The foramen magnum may have been enlarged for removal
of brain tissue, perhaps as a part of the mortuary ritual.
At the site of Conchopata in the Wari imperial heartland, nearly half of the trophy heads
had trauma, including one individual with a perimortem fracture indicating that the victim
received a blow to the head about the time of death (Tung, 2008). Many of the skulls show
cutmarks reflecting removal of flesh in transforming the head from victim to trophy (Tung,
2008). Although there has long been debate about whether trophy heads are from local
venerated ancestors or foreign enemies, strontium isotope analysis of the Wari trophy heads
shows that the vast majority are non-local, suggesting that the trophy heads represent enemy
victims (Tung & Knudson, 2011).
Ritual and sacrifice are well represented in several settings, especially in the Middle Sicán
culture (c. 900 – 1100), the descendant culture of the earlier Moche, in the Lambayque Valley,
northern Peru (Klaus, 2014a; Klaus, Centurion et al., 2010; Shimada et al., 2004). In this
setting, a rich body of evidence for sacrifice from the capital of Sicán and the more rural
temple site of Cerro Cerrillos is revealed from the study of hundreds of skeletons. The location
of cutmarks on anterior body surfaces of cervical vertebrae reveals that the victims had their
throats slit with sharp metal (bronze) tools (Figure 4.18). Some of the victims show cutmarks
in the anterior thoracic region for opening of the chest. Comparisons with earlier sites in the
Lambayeque Valley reveal that various forms of sacrifice and body treatment were new,
especially relating to the practice of opening the chest to extract the heart and of violent child
sacrifice (Klaus, Centurion et al., 2010).
In precontact Mexico, skeletal evidence has accrued on body processing, including cut-
marks, decapitation, dismemberment, burning, and intentional bone breakage in a limited
number of different temporal and cultural contexts (Pijoan & Mansilla, 1990, 1997). As has
been discussed for the American Southwest, cutmarks and other cultural modifications of
skeletal remains, by themselves, do not provide information on the motivations for body
processing (White, 1986). However, a rich historical and iconographic record indicates that
ritual was the key motivation for cannibalism in early contact-period Mexico.
A comprehensive picture of ritualized violence and death is well documented from human
remains recovered from Teotihuacan, Mexico. Although skeletal evidence for ritual sacrifice
has been known since the early twentieth century (Hrdlička, 1910a), the number of individuals
involved were too small to develop a record of broader implications for the practice. However,
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4.3 Intentional injury and interpersonal violence 153
(a)
(b)
(c)
(d)
Figure 4.18 Composite distribution of cut marks associated with ritual killing events:
anterior aspect of cervical vertebrae (a), proximal clavicle (b), superolateral manubrium
(c), and sternal rib end (d); Cerro Cerrillos, Peru. (Klaus, Centurion et al., 2010; photographs
by Sam Scholes; reproduced with permission of the authors and Antiquity.) (A black and
white version of this figure will appear in some formats. For the color version, please refer
to the plate section.)
victims represented by single and multiple interments of mostly adults from the Temple of the
Feathered Serpent dating to the early occupation of the site and construction of the temple
(c. AD 100–250) have been analyzed (Cabrera Castro, 1993; Cabrera Castro et al., 1991; Serrano
Sánchez, 1993; Sugiyama, 1989). Sacrificial victims associated with the temple number about
120 individuals (Cabrera Castro, 1993). The skeletal remains have not been systematically
studied for cutmarks or other skeletal modifications. However, the position of the hands behind
the backs for some individuals suggests that they had been bound at the time of death.
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154 Injury and violence
The general lack of disarticulation as well as the inclusion of commingled individuals in single
burial pits (e.g., burial 190 included 18 adult males in a linear pit at the midpoint between the
southeast and southwest corners of the temple) suggests that these were simultaneous deaths
(Sugiyama, 1989). The social identity of the victims is not known. Nevertheless, in reference to
one of the multiple burials on the margins of the temple (burial 190), the inclusion of
numerous obsidian points, blades, and other offertory items suggests that they “were military
men, priestly soldiers, or men disguised as military personages” (Sugiyama, 1989:98).
Ritualized violence, sacrifice, and related behaviors are well documented in the iconog-
raphy of numerous precontact Latin American cultures. Mass burials, mutilation, decapi-
tation, and presence of trophy heads in archaeological sites in Peru and Mexico indicate that
scenes of these activities in various art forms in these regions were not just mythical events,
but rather, historically situated cultural practices.
Non-ritualized violence
Undoubtedly, the combining of ritual with violence was a highly developed social behavior in
Latin America, especially in state societies where public display of authority took on increas-
ing importance. However, like other areas discussed, the record of violence relating to
resource stress is also prominent in a number of settings. For example, temporal analysis of
remains from San Pedro de Atacama, Chile, reveals a clear pattern of elevated violence-related
cranial trauma (depressed and other types of fractures), doubling in frequency from 5%–11%
in the post-AD 600 occupation with increased sedentism and population density (Torres-Rouff
& Costa Junqueira, 2006). These increases coincided with elevated frequency of stress indica-
tors, such as enamel hypoplasia and porotic hyperostosis. Paradoxically, this period of
heightened resource stress, nutritional inadequacy, and violence was also a period of
increased prosperity, as is indicated by an increase in the quantity and quality of grave goods.
Interestingly, elite males from this setting have fewer violent-related cranial injuries than
non-elite males. These differences coincide with an apparent growth of social inequality,
particularly as it relates to the identity of injured (lower-status) victims and others (higher-
status) (Torres-Rouff, 2011). Like other settings in the New World, violence in this setting is
strongly associated with cultural and environmental circumstances, resulting in increased
competition for limited resources and social disruption generally. Moreover, level of disrup-
tion was likely partitioned along class lines.
Much of the evidence of violence in Latin America, however, relates to competition and
expansion of political control. Much of this record suggests that violence was one strategy
used by the dominant expanding polity. In the capital region of Cuzco of the Inca Empire, a
trend toward increasing cranial trauma is likely due to warfare, increasing from 2.5%
prevalence in the Middle Horizon (c. AD 600–1000) and in the Late Intermediate period
(c. AD 1000–1400) to 7.8% in the Inca Imperial period (AD 1400–1532) when the empire
reached its peak control (the area encompassing modern Peru and parts of Ecuador, Bolivia,
Chile, and Argentina) (Andrushko & Torres, 2011). Interestingly, all of the traumatic injuries
appear to be in peripheral areas and not the core region of the empire around Cuzco. The
record suggests that violence was used to subjugate and expand into peripheral areas.
Moreover, as is so well documented in historical accounts, violence did not decline in the
region during the Spanish conquest of the Inca Empire. Indeed, the study of juvenile and adult
remains shows an increase in bioarchaeological evidence of violence and traumatic injury
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4.3 Intentional injury and interpersonal violence 155
(Gaither & Murphy, 2012; Murphy et al., 2010). While some of the increase may have involved
internecine conflicts among native populations, almost certainly some originated from Span-
iards during this period of unprecedented population disruption and wide-scale suppression of
native communities.
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156 Injury and violence
of skeletal remains. Thus, folklore of Eastern Islanders overemphasizes warfare, violence, and
cannibalism.
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4.3 Intentional injury and interpersonal violence 157
respectively (Burrell et al., 1986; Kilgore et al., 1997; Van Gerven et al., 1995). Upper limb
fractures show an especially pronounced increase (30%) (Van Gerven et al., 1995).
Increase in skeletal trauma in juveniles and old adults is more pronounced than in other age
groups in the late Christian period at Kulubnarti. These age-specific increases in fracture
prevalence may be due to elevated risks of living in two-story houses in the late period versus
one-story houses in the early period (Burrell et al., 1986). Access to the living area on the
second story in late-period houses was gained by use of a retractable ladder, which may have
caused falls and other types of accidents (Burrell et al., 1986).
A significant portion of the Kulubnarti individuals possesses multiple injuries (27%). This
unusually high prevalence of accidental injuries, coupled with the presence of numerous and
severe fractures, reflects the hazards of living in a very difficult terrain. Unlike Lower Nubia to
the north, cultivated areas in Upper Nubia are highly constricted and are limited to small
pockets of flat land immediately adjacent to the Nile River. Individuals living at Kulubnarti
would have been exposed to difficult walking conditions on a daily basis. The adoption of
defensible architecture (e.g., two-story houses) later in the Christian period may also have
placed individuals at increased risk of injury.
Most Kulubnarti fractures are in the forearm (75% of fractures) (Kilgore et al., 1997).
Aggressive interactions may have contributed to some of the fractures because a push and
subsequent fall could have resulted in forearm fractures. The record suggests that the difficult
terrain and nearly continuous conflict between Egypt and Nubia resulted at various times in a
high risk of injury and death, arising from accidents and interpersonal conflict.
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158 Injury and violence
Table 4.2 Cranial trauma by region in Australia. Percentages of crania are shown (Adapted
from Webb, 1995: Table 8–2)
N One lesion (%) Two lesions (%) Three lesions (%)
Males
Central Murray (247) 13.4 3.6 0
Rufus River (122) 26.2 7.4 0.8
South Coast (138) 14.5 4.4 0.7
Desert (132) 16.7 3.8 1.5
Tropics (92) 6.6 0 0
East Coast (133) 23.3 6.8 1.5
Females
Central Murray (151) 19.9 4.0 1.3
Rufus River (83) 27.7 8.4 2.4
South Coast (123) 31.7 10.6 4.9
Desert (51) 33.3 11.8 5.9
Tropics (62) 24.2 9.7 4.8
East Coast (86) 32.6 10.5 3.5
fleeing the attacker. Adult males show a different pattern of injury location to that of women:
for the entire series, more left than right parietal bones are fractured. This pattern suggests
that male violence usually involved face-to-face confrontations.
It is not uncommon for an individual in the Australian samples to have two, three, or even
four cranial bones that display depressed fractures (Table 4.2). Consistent with the sex
differences in the prevalence of crania affected, more women than men have multiple injuries.
The general pattern, therefore, indicates that violence and aggression were directed more at
women than men in prehistoric foraging societies throughout Australia. Overall, however, the
pattern of higher injury in females than males evokes the conclusion that perhaps the record
represents continental-wide domestic violence – females being injured by male associates or
partners. Indeed, in modern societies, millions of women globally are injured by males in
domestic contexts (Novak, 2006).
In Australia, violence was not limited to prehistoric societies. Ethnographers observe that
violence is a common occurrence and a part of everyday discourse (Burbank, 1994). Unlike
Western societies, such as in the United States where fighting – and especially aggression against
women – is viewed as a deviant behavior, physical aggression in native Australians is considered
an accepted if not legitimate form of social interaction (Burbank, 1994; Myers, 1986). Burbank
(1994) provided detailed observations on physical aggression in men and women in an aborigi-
nal group living in Arnhem Land (Northern Territory). Her study shows that both men and
women were heavily involved in confrontations. However, the majority of aggressors and their
victims are adult females. These observations of both deceased and living native Australians
reveal a striking consistency in behavior between prehistoric and contemporary populations.
Women played a key role in aggressive encounters, and not simply as victims of attack. This
work underscores the important notion that violence, domestic or other, is not the sole purview
of males, certainly in this setting and likely others around the world (Guliaev, 2003).
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160 Injury and violence
well illustrated by the presence of projectile injuries, sword and axe cuts, cranial depressed
fractures, and decapitation (Bennike, 1985; Ebbesen, 1993; Kannegaard Nielsen & Brinch
Petersen, 1993). At the Mesolithic site of Bogebakken, a bone projectile was found lodged
between the second and third thoracic vertebrae of an individual. In fact, all projectile wounds
at this and many other Danish sites are found in the thoracic and head regions, revealing the
lethal intentions of the attackers.
Placement of Denmark in the larger regional context of western and central Europe reveals
that the Neolithic and the foraging-to-farming transition was a relatively turbulent period in
comparison with later times, and not only in Denmark (Fibiger et al., 2013; Smith, 2014;
various in Schulting & Fibiger, 2012). The reasons for violence in this broad expanse are
varied. For example, the co-occurrence of elevated violence-related injuries in the early
Neolithic (c. 5000 BC) and deteriorating environmental circumstances across the region
suggests the possibility of negative social, cultural, and economic factors (compare with
Orschiedt & Haidle, 2012; Teschler-Nicola, 2012; Wahl & Trautmann, 2012).
At least one Neolithic series from Talheim, Germany, represents a massacre (Wahl &
Trautmann, 2012). The posterior position of traumatic injuries for many of the 34 individuals
documented indicates that they were attacked from behind while fleeing from aggressors.
With a few exceptions, most cranial injuries (e.g., depressed fractures) throughout the region
are healed, suggesting that as in southern California coastal foragers, inflicted injuries were
not intended to have a lethal outcome (Walker, 1989; and authors cited earlier). In their
analysis of cranial trauma in Neolithic Denmark, males generally have more anterior and left-
sided trauma than females, indicating the involvement of men in face-to-face encounters
(Fibiger et al., 2013).
In interments dating to the Middle Ages, the heads of victims had been removed and placed
between their legs. The reasons for this unusual treatment are unclear, but during the Middle
Ages, the practice was associated with criminals in order to prevent their return following
death (Bennike, 1985; and see later). Decapitation and other forms of head and neck trauma
were likely more common than is indicated by the skeletal evidence alone. A number of
Neolithic and Iron Age bog corpses show evidence of decapitation and strangulation, the
latter of which may have involved hanging (Bennike, 1985; Glob, 1971). Owing to the
relatively small number of projectile- and weapon-related deaths, it is not possible to identify
a pattern of decrease or increase in violence-related death in Denmark (Bennike, 1985).
However, there is a shift from the use of projectile weapons to axes and swords in the Iron
Age (Bennike, 1985). The lethal nature of this new weaponry for the enemies of Danes is
demonstrated in at least one battle site (see later).
Violence in western Europe during the Mesolithic and Neolithic was likely highly localized,
with a relatively higher prevalence in some regions than in others (Bennike, 1985; Frayer,
1997; Jiménez-Brobeil et al., 2009; Orschiedt et al., 2003; Papathanasiou et al., 2000; various
in Schulting & Fibiger, 2012). For western Europe as a whole, violent trauma was frequent
during the Neolithic and Mesolithic (Schulting & Fibiger, 2012; but see Constandse-
Westermann & Newell, 1984; Roksandic et al., 2006; Schulting, 2006). An increase in
population density, increase in social complexity, and resource circumscription during the
period suggests the potential for an increase in hostilities. Moreover, there are clear instances
of violence predating the Neolithic, indicating possible processes underlying violence in the
region; for example, at the Mesolithic site of Ofnet (Germany) where two pits contain the
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4.3 Intentional injury and interpersonal violence 161
skulls of 32 individuals. The skulls show perimortem depressed fractures and cutmarks. Some
of the skulls have upper cervical vertebrae articulated, a number of which display cutmarks
suggesting slitting of the throat and decapitation.
Battle of Wisby
The Middle Ages in Europe involved a tremendous upsurge in armed conflict between many
warring city-states and various confederations of states. Today, across Europe, the vestiges of
walls surrounding villages and towns remain, a lasting record of violence and threat of
violence during this time. Only rarely, however, have battle sites from the conflicts resulted
in documented archaeological remains. One important exception is the battle site of Wisby,
located on the island of Gotland in the Baltic Sea. Hundreds of skeletons excavated at the
battle site present some of the grim details of preindustrial warfare in northern Europe.
The city of Wisby was invaded in 1361 by an army led by the Danish king, Waldemar
(Thordeman, 1939). Over the course of a single day, the poorly organized peasant forces
defending the city were decisively defeated and massacred by the king’s highly disciplined
army. Estimates indicate that some 1800 Gotlanders were killed in this battle (Ingelmark,
1939). Archaeological excavations of three common graves at Wisby yielded an enormous
sample of human skeletal remains (n¼1185). Analysis of these remains reveals that only
males were victims, but the age distribution was extraordinarily varied, ranging from adoles-
cents to very old adult males (Ingelmark, 1939).
Consistent with research completed on skeletons from other Middle Ages archaeological
sites (compare with Bennike, 1985), most of the injuries resulted from the use of cutting
weaponry, especially swords and axes (n¼456). A significant minority of injuries was from
projectiles (n¼126). Skeletal wounds are variable, ranging from scratches and nicks to
dismemberment. The latter, for example, is illustrated by the presence of severed hands and
feet, partial limbs, and complete limbs. One individual expresses the intensity of battle: the
lower halves of both left and right tibiae and fibulae are completely severed. The lower legs are
affected more than any other area of the body: about two-thirds (65%) of cutting trauma
involved tibiae. Ingelmark (1939) observed that the focus on the lower limbs likely reflects the
use of shields and protective clothing for the body trunk, leaving the legs especially vulner-
able to injury. Sword blows directed at the lower legs typically resulted in the slicing and
chipping of bone on the anterior crests of tibiae.
Poor protection of heads of individuals from the Gotlander army is indicated by the large
number of cranial injuries, some of which involve extremely deep cuts. Heads of some groups
of Gotlander soldiers may have been better protected than others. For example, only 5.4% of
crania are injured in common grave no. 3; this frequency contrasts with the prevalence of
cranial injuries of 42.3% and 52.3% in common graves no. 1 and no. 2, respectively. The
majority of cranial wounds are on the left side of the head, which fits the expected pattern of
injury sustained by a weapon held by a right-handed individual during a face-to-face
encounter (Courville, 1965). Some crania have injuries on posterior vaults, suggesting that
these victims were struck from behind while fleeing their attackers.
The presence of all ages of males suggests that the majority of the male population in and
around Wisby were recruited for the defense of the city. Analysis of pathological conditions
suggests that virtually anyone who could walk (and even those who could not) were drafted.
Ingelmark (1939:192) remarked on the “good many morbid processes” present in the skeletal
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162 Injury and violence
assemblage of battle victims. Many vertebrae have pronounced osteoarthritis, and at least
four individuals have extensive vertebral tuberculosis. One individual displays a completely
ankylosed (fused) knee. The angle of flexion (about 55) greatly disabled the individual:
running was impossible for this victim. A number of individuals show well-healed but poorly
aligned femoral neck fractures, which limited their ambulatory capabilities. These observa-
tions, combined with other health problems – including skeletal infections and numerous
healed, but malaligned limb fractures (n¼39) – also contributed to reduction in efficiency on
the battlefield. The defending army, therefore, was not composed of a group of robust males
who were in their peak years of fighting prowess. Like many of the skeletal samples discussed
in this chapter from both New and Old World contexts, these victims of the massacre were
members of a population not unfamiliar with violence during their lifetimes (compare with
Milner et al., 1991; Willey, 1990).
The pattern of injury in the Wisby series is similar in many respects to the pattern of injury
documented in more detail from the Towton battle site dating to AD 1461, a century after
Wisby. There, some 28 000 of an estimated 100 000 combatants died on a single day (Fiorato
et al., 2000; Knüsel, 2014). Analysis of 37 of the casualties from a mass grave revealed clear
indicators of perimortem trauma, ranging from single to multiple (up to nine) injuries (Novak,
2000). Interestingly, nine of the 29 crania examined had well-healed trauma, likely derived
from previous battle-related injuries. Most of the injuries were on the left side of the face and
vault, indicating blows struck by a right-handed person successfully landing the business end
of the weapon on the victim in a face-to-face confrontation. The injuries include sharp (blade),
blunt, and projectile wounds (Figure 4.19). The injuries provide a picture that contrasts with the
romanticized version of Medieval battle showing honor and pageantry, but show clearly that
those who had inflicted the injuries were well trained in the combat methods and practice. The
patterns provide a useful record for evaluating the consequences of conflict in this key setting
prior to the use of firearms in modern warfare (Cunha & Silva, 1997; Düring, 1997; Giuffra,
Baricco et al., 2013; Mitchell et al., 2006; Šlaus et al., 2010, 2012; Weber and Czarnetzki, 2001).
Beheading in Britain
In the upper Thames valley, a high frequency of decapitation and prone burial in Romano-
British (third to early fifth centuries AD) cemeteries (Cassington, Radley, Stanton Harcourt,
Queensford Mill, and Curbridge) has been documented (Harman et al., 1981). In total, 15.3%
show evidence of beheading. Analysis of other Romano-British cemeteries indicates that this
practice was part of a widespread behavior during this period (Anderson, 2001; Bush & Stirland,
1988; Montgomery et al., 2011; Philpott, 1991; Tucker, 2014; Wells, 1982). During the
following Anglo-Saxon period (fifth to tenth centuries AD) beheadings continued, and victims
were included in execution cemeteries (Buckberry, 2008). Reminiscent of the decapitations from
Denmark (Bennike, 1985; and see earlier), heads of a number of burials had been purposefully
placed between and associated with the legs of the deceased (Anderson, 2001; McKinley, 1993).
The simultaneous occurrence of decapitation and prone burial in Romano-British and Anglo-
Saxon cemeteries suggests a probable connection between the two practices. The demographic
composition of decapitated and prone skeletons shows a selectivity for adults, suggesting that
execution may have been the primary motive. Review of historical, archaeological, and folklore
literature indicates other possibilities, such as prevention of the deceased from walking or
communicating, sacrifice, and deprivation of the soul, for either sacrificial purposes or for
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4.3 Intentional injury and interpersonal violence 163
Figure 4.19 Sharp force trauma to the face associated with a bladed weapon; Towton, United
Kingdom. (From Novak, 2000; reproduced with permission of author and Oxbow Books.)
(A black and white version of this figure will appear in some formats. For the color version,
please refer to the plate section.)
punishment for some wrongdoing (Harman et al., 1981). Decapitation and/or prone burial, may
have been a “final form of indignity inflicted on the corpse of an individual in consequence of
particular characteristics or offenses during life. But it seems more probable that both were
believed to have some effect on the subject in an after life” (Harman et al., 1981:168).
The manner of beheading is indicated by the location and pattern of cutmarks on affected
skeletal elements. Severing of the head was usually done in the upper neck region. Damage to
anterior surfaces of cervical vertebrae in some individuals and posterior surfaces of cervical
vertebrae in others indicates that the beheading blow was delivered from in front and behind
at various times, and probably with a variety of tools (Harman et al., 1981; and see McKinley,
1993; McWhirr et al., 1982). Detailed analysis of a beheading victim from Hertfordshire shows
a series of at least six carefully placed cuts, including three cuts on the anterior odontoid
process, superior body, lower body, and right inferior articular process (McKinley, 1993). The
narrowness of the cuts indicates that the decapitation was completed with a narrow blade
administered as blows to the neck.
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164 Injury and violence
they brought with them a weapons technology that facilitated their (mostly rapid) conquest of
native populations. The early expeditions were violent affairs at times, resulting in brutal
treatment of natives (Weber, 1992). Although these tactics seem repulsive now, they were well
within the bounds of behavior regarded as fully acceptable for European males during the
Middle Ages. Historical literature and accounts of violent confrontations (e.g., Wisby) indicate
that conflict behavior between European males was often excessively cruel (Weber, 1992). The
study of native remains dating to the early period of contact with Europeans has provided a
new dimension to understanding the nature of the interactions between these groups.
La Florida
The study of hundreds of human skeletal remains from colonial-era Spanish sites in the
American Southeast (Georgia, Florida) and Southwest (New Mexico and Arizona) provides
important perspective on violent confrontation, especially during the sixteenth and seven-
teenth centuries. In Spanish Florida (present northern Florida and coastal Georgia), the region
named La Florida by Juan Ponce de León in 1513, short-term encounters between native
populations and Spaniards occurred during the exploration period (c. 1513–1565), followed
by long-term, sustained contact during the mission period (1565–1704) (McEwan, 1993). The
earliest contacts frequently resulted in hostile interactions and deaths of both Europeans and
natives (Varner & Varner, 1980). The later mission period was relatively peaceful; long periods
of calm were occasionally punctuated by native revolts violently put down by Spanish
military forces (Hann, 1988).
Analysis of skeletal remains from both periods of Spanish occupation in the region
produced only limited evidence of violent interactions. In skeletons from the Tatham Mound
site on the Gulf coast of western Florida – the probable location of Hernando de Soto’s visit
in 1539 – perimortem trauma caused by the impact of metal weapons is present in 17 skeletal
elements (Hutchinson, 1991, 1996, 2006). The most dramatic examples of cut bones include
the severed acromion process of a right scapula and a left humerus diaphysis cut through
60% of the midshaft (Figure 4.20). Neither bone showed evidence of healing. Other long
bones in the sample have multiple cutting injuries around diaphyseal perimeters. In total, the
pattern of damage appears to be associated with purposeful dismemberment (Hutchinson,
1996). It is possible that Indians inflicted the injuries using captured Spanish weapons. The
early dates of the site (early sixteenth century) suggest that this is an unlikely possibility.
The injuries were more likely inflicted by Spaniards (Hutchinson, 1996). The pattern of
injury is also consistent with European battle tactics. For example, the orientation in the
scapular cut is consistent with the practice of removing the fighting arm of the opponent
(Hutchinson, 2006).
Only one other skeleton in Spanish Florida shows evidence of violent confrontation.
A single high-status male from Mission San Luis de Talimali (AD 1656–1704) likely died
from a gunshot wound, but it is not possible to identify the perpetrator, Indian or Spaniard
(Larsen, Huynh et al., 1996). Thus, based on the study of violent trauma in skeletal remains,
the legends suggesting an unusually cruel treatment of natives – at least as it is indicated by
metal-edged or firearms weaponry – are not substantiated. Rather, the form of violence that
has been well documented in this setting is what Klaus (2014a) described as bioarchaeological
evidence of structural violence (see later).
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4.3 Intentional injury and interpersonal violence 165
(a)
(b)
Figure 4.20 Cut adult humerus (a) and scapula (b); Tatham Mound, Florida.
(From Hutchinson & Norr, 1994; reproduced with permission of authors and
John Wiley & Sons, Inc.) (A black and white version of these figures will appear in some
formats. For the color version, please refer to the plate section.)
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166 Injury and violence
American Southwest
There is a highly visible record of violence in precontact Southwestern populations, much of
which highlights episodes of stress and (possibly motivated by starvation and heightened
resource stress generally) cannibalism. This record is closely linked with periods of violence as
shown by an abundant skeletal record of injury and violent trauma, sometimes in association
with processed remains (Turner & Turner, 1999; White, 1992). The presence of fortifications
and other defensive architecture and high frequencies of traumatic injury (including injury
resulting from violence) in some later prehistoric sites suggests that confrontations were
frequent (Stodder & Martin, 1992). Cranial trauma in the American Southwest shows an
increase in prevalence during the late prehistoric period, which continued into the historic,
mission period (Bruwelheide et al., 2010; Stodder, 1990, 1994; Stodder & Martin, 1992).
Archaeological and historical evidence shows that the high frequency of cranial trauma in
the historic period can be attributed to confrontation between Spaniards and Indians as well
as among Pueblos, and between Pueblos and non-Pueblo native groups. The study of skeletal
remains reveals that most cranial injuries are in adult males (Stodder, 1994). At San Cristobal
and Hawikku, sites with significant contact-period skeletal assemblages, very high frequen-
cies of cranial injuries have been reported (Stodder, 1994). Twenty percent and 17% of males
have cranial trauma from the San Cristobal and Hawikku sites, respectively. Paleopathological
markers of stress (e.g., dental defects) indicate that nutritional and other health disruptions
were generally widespread during the late prehistoric and contact periods, which may have
contributed to fostering intra- and inter-group hostilities during this time (Stodder, 1994;
Stodder et al., 2002). At Pecos Pueblo, the prevalence of cranial trauma, largely in the form of
depressed fractures, in the mission population (42.1%) is considerably higher than the
prehistoric ancestral population and most other samples in North America where data have
been collected systematically (Bruwelheide et al., 2010).
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4.3 Intentional injury and interpersonal violence 167
Analysis of the remains of five adult males buried in a mass grave within the fort indicate
clear evidence of violence-related injuries, bearing testimony to the historical and fictional-
ized (Cooper, 1919) accounts of the battle (Liston & Baker, 1996). Four of the five show
perimortem trauma to the legs reflecting injuries received during the siege of the fort. None of
the injuries are healed, suggesting that they died prior to or during the siege. The trauma was
not lethal, but serious enough so as to prevent their departure from the fort. These skeletons
display a range of perimortem trauma that likely represents injuries resulting in death. One
individual shows a series of four cutmarks on the posterior surface of the odontoid process of
the second cervical vertebra. Another individual expresses a series of radiating fractures
through the face and frontal bone, indicating crushing of the skull with a blunt object. The
pattern of trauma suggests that the soldier had been beheaded from behind. All five individ-
uals show notches, slices, and gashes in skeletal elements of the anterior and posterior trunk
(e.g., scapula, ribs) and pubic region. The morphology of cutmarks evinces the use of both
knives and axes in the mutilation of victims.
Snake Hill
Some of the most intense fighting between the British and Americans during the War of
1812 took place in the frontier region between Lake Ontario and Lake Erie along the Niagara
River (Whitehorne, 1991). During the four-month period in 1814 when American troops
successfully captured and held Fort Erie on the Canadian side of the river, heavy siege and
combat resulted in the deaths of hundreds of soldiers from both the British and American
armies. Archaeological excavations at the battle site of Snake Hill, Ontario, resulted in the
recovery of the skeletal remains of American soldiers from burial and medical waste pits
(Owsley et al., 1991; Thomas & Williamson, 1991). Demographic assessment of the complete
or nearly complete skeletons indicates that they were young adult males, aged 15 to 29 years;
seven soldiers were older than 30 years at death. Half (50%) of the individuals in the sample
had fractures caused by damage from firearms projectiles. The general lack of healing in most
cases indicates that these wounds were usually fatal. The highest percentage of fractures were
ribs (28%; 7/25), followed by femora (25%; 7/28), and crania (9.1%; 2/22).
Locational assessment of skeletal wounds indicates that most injuries (69.8%) were above
the waist. With regards to the total number of noncranial and nonvertebral trauma (n¼53),
twice the number of fractures occurred on the left side (54.7%; 30/53) than on the right side
(26.4%; 14/53) of battle victims. This pattern may reflect handedness or body postures during
the battle (Owsley et al., 1991). Cause of death is especially apparent for several victims. For
example, a young adult died of a massive head injury whereby a firearm’s projectile had
passed through the left and then right parietal bones. This individual also had a large,
completely healed cranial depressed fracture from earlier injury. Other individuals had frac-
tured facial bones from a firearms projectile and shattered long bones.
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168 Injury and violence
battle is part of an overall pattern of political domination and control of lands and resources
in the Great Plains by opposing groups.
Within two days of the battle, eyewitness accounts described mutilation (including
scalping) and dismemberment not unlike patterns observed in other Plains samples (e.g.,
Crow Creek, Larson Village site; see earlier) (Scott et al., 2002; Snow & Fitzpatrick, 1989).
Temporary graves were hastily prepared at the locations where individuals were killed. Some
of the bodies were identified, but owing to decomposition and mutilation, many were not.
Skeletal fragments of battle victims from erosion and limited test excavations provide the
basis for detailed study of battle injuries (Snow & Fitzpatrick, 1989). Analysis of 375 partial
and complete bones and 36 teeth from a minimum of 34 individuals indicates three primary
types of perimortem trauma, including blunt-force trauma, cutmarks, and bullet wounds
(Scott et al., 2002; Snow & Fitzpatrick, 1989; Willey & Scott, 1996). Blunt-force trauma
involved catastrophic fragmentation of crania, and to a lesser extent, postcranial elements.
All 14 partial crania showed massive injuries due to heavy blows. Additionally, presence of
cutmarks on various skeletal elements indicates widespread perimortem mutilation. Several
different forms of cutmarks, ranging from fine cuts to pronounced incisions, reflect the use of
metal arrows or knives.
Cutmarks on a variety of skeletal elements indicate the high degree of mutilation of battle
victims. One individual, for example, has cutmarks on a humerus head and sternum. The use
of heavy metal-edged weapons (e.g., hatchets) is clearly indicated in several instances.
Elsewhere, a completely transected cervical vertebra indicates decapitation by a single blow
to the neck. Another individual shows distinctive sets of chopping blows to the proximal ends
of the left and right femora indicating purposeful dismemberment.
In addition to traditional native weaponry, the presence of gunshot wounds in six individ-
uals indicates the use of firearms by Native Americans at the battle site (Scott et al., 2002;
Willey & Scott, 1996). Individual M85, for example, had at least two upper body gunshot
wounds, including an entrance wound on a rib margin and shattered ribs from another
wound. A third wound is represented by a bullet or bullet fragment embedded in the distal
left radius. This individual also displays cutmarks on his clavicle. Three gunshot wounds are
located in the crania, one entering from the back and two entering from the right side and
exiting from the left.
In summary, based on the study of human remains from the battle site, a sequence of events
can be reconstructed: namely, soldiers were wounded, killed (frequently with blunt-force
trauma to the skull), and mutilated (Snow and Fitzpatrick, 1989). A consideration of direction
of entry wounds is consistent with historic records indicating that the battle was chaotic
(Willey & Scott, 1996). As would be expected, except for the use of firearms, the pattern of
killing and mutilation of victims is strikingly similar to that observed in other North American
native populations from the Great Plains and Midwest discussed in this chapter (e.g., Crow
Creek, Larson Village site, Norris Farms).
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4.4 Medical care and surgical intervention 169
social group to ensure that the injured person returns to a functional state of health and well-
being. Ethnographic and historic accounts of nonindustrial societies report tremendous
variation in the care of traumatic injuries, ranging across alignment of fractures and use of
splints, immobilization, oral medicines, and other treatments (Ackerknecht, 1967; Ortner
2003; Roberts, 1991). For example, lack of angulation or significant difference in length of
long bones in the fractured versus the normal side has been documented in the Libben
population from the American Midwest (Lovejoy & Heiple, 1981) and in Medieval populations
from England (Grauer & Roberts, 1996).
Similarly, in prehistoric Australian skeletal series, most fractures show proper unification
and alignment (Kricun, 1994; Webb, 1989, 1995), which Webb regards as evidence for “a firm
commitment to care and concern for the injured patient” (1995:200). The presence of well-
healed amputations in two individuals from the central Murray Valley region suggests
knowledge of this type of surgical procedure before the arrival of Europeans (Webb, 1995).
In five to six thousand Nubian skeletons, some 160 fractures are present, most of which are
well-healed and aligned, with little evidence of infection (Smith & Jones, 1910). Bark splints
were found associated with limb fractures in a couple of instances (Smith, 1908).
Some earlier societies appear to have lacked either the ability or interest in treatment of
fractures. For example, the fourteenth-century Wisby battle victims had a high degree of
angulation of healed fractures, suggesting only minimal levels of treatment (Ingelmark, 1939).
The battle victims are largely drawn from the peasant class, and may not have had access to
the same medical care afforded the nobility.
Temporal assessment of a large sample of Roman, Anglo-Saxon, and Medieval long bone
fractures reveals changes in injury management in these populations (Roberts, 1991). During
the earlier Roman and Anglo-Saxon periods, healing of fractures was generally good,
suggesting that fracture sites were correctly reduced and aligned, probably with some type
of a support (Roberts, 1991). Treatment was so widespread in the Roman period that
deformities from poorly healed or misaligned fractures were no more common than they
are in living populations. Medieval management of fractures appears to have been less
efficient than that in earlier periods. There is a generally higher prevalence of deformation
and angulation of long bones (Roberts, 1991). Additionally, many fractures (35/59) from this
period had associated periosteal infections. If this analysis and the findings based on the
study of the Wisby sample are representative, it appears that northern European populations
living during the Middle Ages were far less knowledgeable about fracture management than
their forebears.
Treatment of head injuries is suggested by the association between trephination and cranial
trauma in a number of settings. In Denmark, trephinations frequently accompany cranial
trauma, including fractures and sword or axe cuts (Bennike, 1985). Most of the Danish
trephinations are on the left side of the cranial vault, which coincides with the location of
skull injuries received in battle. This pattern is also consistent with the predominance of
trephinations in males, the primary participants in battle and interpersonal conflict. Similarly,
over 50% of recorded trephinations from Anatolia documented by Erdal and Erdal (2011) were
associated with cranial trauma.
Precontact sites in Andean South America also show an abundance of trephination,
especially in Peru and Bolivia where scores of cases are reported. Analysis of crania from
central and south coastal Peru and regions of the highlands of the Central Andes reveals that
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170 Injury and violence
Figure 4.21 Incomplete trephination and depressed skull fracture on right frontal bone;
Cinco Cerros, Peru. (Verano, 2007; reproduced with permission of author and University of
Arizona Press.) (A black and white version of this figure will appear in some formats. For
the color version, please refer to the plate section.)
adult males comprise the majority of trephined individuals, but adult females and some
juveniles are also included (Verano, 2003). The association between cranial injuries and
trephination indicates that this form of surgery was likely performed as a treatment for head
trauma (Figure 4.21). Diachronic comparisons indicate that the frequency of well-healed
trephinations increased over a 2000-year period (400 BC–AD 1532). The highest rate of
success was from the latest precontact period (Inca), including some individuals with as many
as five to seven healed trephinations. The apparent increase in survival may have been due to
the reduction in size of the trephination opening as well as to the greater use of the circular
grooving technique of excision, thus reducing the risk of dura mater penetration and neuro-
logical damage (Verano, 2003).
The lack of an association between trephination and head injury in other settings suggests
that there may have been other motivations, including treatment of real or imagined ills. For
example, none of the few trephined crania from North America are associated with cranial
injury (Ortner, 2003; Stone & Miles, 1990).
Evidence for treatment of dental disease has been identified in the form of alterations of
teeth, namely drilling in tooth roots (Schwartz et al., 1995) and crowns (Bennike, 1985;
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4.4 Medical care and surgical intervention 171
Figure 4.22 Anteromedial (above) and scanning electron microscopic (below) perspective
of drilled right mandibular canine; Sky Aerie Overlook, Colorado. (White et al., 1997;
reproduced with permission of authors and John Wiley & Sons, Inc.)
Koritzer, 1968; Turner, 2004; White et al., 1997). These holes are usually found in association
with carious or otherwise diseased teeth, indicating a therapeutic intention (Figure 4.22).
In summary, the study of samples worldwide reveals that injuries sustained either by
accidental means or during violent confrontations were treated in some earlier societies.
These findings indicate that many past societies were aware that proper restoration of
function could only be brought about by appropriate treatment protocols (Roberts, 1991).
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172 Injury and violence
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4.5 Interpreting skeletal trauma 173
injury occurs in individuals and populations who were the victims of conflict. Thus, cemetery
assemblages representing groups who were the winners would not be expected to exhibit the
frequency of injury seen on the losing end of violent encounters.
Elevated prevalence of violence and injury mortality in some prehistoric and historic
settings and their possible relationship with increased population density and/or resource
circumscription is similar to that of recent, twentieth-century populations. For example,
Relethford and Mahoney (1991) documented markedly higher rates of injury mortality in
the most densely populated areas of New York State (excluding New York City). These
similarities may reflect common themes between past and recent humans, such as high
density of population and social inequalities that serve to promote violence. Population
density is a complex composite of a number of factors, such as the physical and sociocultural
environments, demographic, cultural, and social influences, and individual behavior. There-
fore, although these apparent similarities are informative, it is important to identify specific
causal factors in specific settings before making conclusions generally regarding the relation-
ship between population distribution and injury mortality in humans.
In some regions, clear patterns of violence have begun to emerge. For example, an increas-
ingly hostile landscape in the millennium preceding European contact in the Eastern Wood-
lands generally is corroborated by the archaeological evidence of an increase in defensive
construction in later prehistory (Keener, 1999; Milner et al., 1991). It is during this time that
there is the rise of chiefdoms, population increase, and competition between neighboring
villages. Similar patterns of increasing conflict and aggression in the final centuries before
European conquest are indicated in the southern California Santa Barbara Channel islands,
the American Southwest, Southeast, Great Plains, and Arctic (Lambert, 2007, 2014). Regional
investigations suggest that resource productivity and climatic instability may have had a
strong influence on the presence or degree of conflict in the past. This hypothesis is supported
by the increasing evidence for broad patterns of violence in association with climatic instabil-
ity and drought in the later prehistory of North America. In particular, some of the most severe
violence occurs in settings and periods where climate is unstable and prone to drought. This is
not to say that climate causes violence. Rather, periods of drought were triggering events,
setting off a stream of events resulting in violent encounters between groups competing for
the same limited resources. Similar patterns of the increase in evidence of violence are also
documented in a wide range of Old World settings (Kennedy, 1994; various in Knüsel & Smith,
2014; Schulting & Fibiger, 2012).
Overall, bioarchaeological studies indicate that violence and conflict are not random
events, but are strongly influenced by extrinsic factors, such as resource depletion and
competition for important resources. Dietary deprivation may have been a motivation for
cannibalism. Historical records in a number of settings are informative. For example, canni-
balism may have been symptomatic of a larger pattern of animosity and aggression between
groups (e.g., Saunaktuk) or a key part of ritualized violence involving sacrifice and
cannibalism (e.g., Teotihuacan).
The study of trauma in skeletal remains reveals that, on comparing different societies, areas
of the body attacked are highly patterned. Walker (1997, 2001a) observed that in modern
industrial Western societies (e.g., United States), the head and neck are highly favored targets
of attack, probably for both strategic and symbolic reasons. He argues that the face is an
appealing target because the injuries are especially painful. The face and head generally bleed
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174 Injury and violence
profusely and bruise easily, which may symbolize the aggressor’s dominance (Walker, 2001a).
This probably explains why the most highly traumatized focal point of the body in recent
urban populations is the face (Allen et al., 2007; Hussain et al., 1994).
Many past societies show a penchant for head injury, but these injuries are usually directed
at the vault and not the face or dentition. Dental fractures are present in archaeological
remains, but they are relatively rare (Alexandersen, 1967; and see Leigh, 1929; Lukacs, 2007b;
Lukacs & Hemphill, 1990, for regional studies documenting dental trauma). The location of
the injury on the body provides insight into some of the details of conflict between the
individuals involved. For example, many cranial injuries are found on the left side of the
frontal bone or other anterior elements, indicating that a right-handed attacker successfully
engaged his/her weapon while facing the victim (e.g., native Australian males). A more
haphazard pattern of cranial injury (e.g., prehistoric Michigan) or higher frequency of trauma
on the right side or posterior vault indicate that injuries were sustained while the victim was
fleeing their attacker or perhaps while lying prone (e.g., Wisby). This pattern is more common
in women than in men in some settings (e.g., Australia, Michigan, Peru), suggesting that
aggression was also directed at women. Ethnographic evidence reveals that although the
aggressor was often an adult male, attack by adult females on other females (and on males)
occurs in no small number in some settings (Burbank, 1994).
Historical documentation indicates that children have long been a target of violent injury
and death. DeMause (1974), for example, regarded child abuse as widespread in Europe prior
to the eighteenth century. Yet, examination of thousands of archaeological skeletons reveals
remarkably little evidence of skeletal trauma – localized trauma-induced subperiosteal lesions
in multiple stages of healing, and perimortem fracture – associated with battered-child
syndrome (Fibiger, 2014; Lewis, 2014; Walker, 2001b; Walker et al., 1996; Wheeler et al.,
2013). Certainly, juveniles in earlier societies were victims of homicide and violence (e.g.,
Ofnet, Crow Creek, Norris Farms). Juvenile skeletons, however, lack the injuries associated
with long-term abuse. This suggests that, as with the pattern of facial and other injuries in
twentieth-century Western societies (Love et al., 2011), child abuse resulting in severe skeletal
trauma is primarily a modern phenomenon. Walker (2001b) suggests that the rise of childhood
abuse is due to the loss of social controls over behavior in largely recent urban settings in
comparison with controls present in earlier and traditional societies.
Technological factors are important in interpreting patterns and types of skeletal injuries.
The introduction of the bow-and-arrow is linked with an increase in lethal conflict (e.g.,
southern California). Prior to the invention of firearms, violence-related injuries were caused
primarily by projectiles, cutting, and blunt force.
The skeletal record shows the presence of both lethal and nonlethal forms of trauma, thus
providing essential insight into the previous history of interpersonal aggression both within
and between past societies. The Wisby, Towton, Crow Creek, and Norris Farms victims display
numerous healed injuries (e.g., cranial depressed fractures) that reflect a long and well-
established history of conflict well prior to the event or events resulting in widespread death.
In a sense, then, these injuries foreshadow a future act (e.g., a major battle) that later resulted
in more widespread violence.
The study of human remains from these sites suggests that debilitating injuries, poor health,
or generally high levels of physiological stress may have increased the susceptibility of a
population to attack and defeat. The Norris Farms and Crow Creek skeletons display numerous
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4.5 Interpreting skeletal trauma 175
pathological indicators of stress, including iron deficiency anemia, dental defects, tubercu-
losis, and generalized infection, that reflect compromised health and the reduced ability to
perform subsistence and other arduous tasks (Bamforth, 1994; Milner et al., 1991). Although
this pattern of poor health does not explain the demise of the population, it suggests that they
may have had a reduced ability to successfully mitigate hostile social environments. Some
populations display fractures and other debilitating conditions that limited their ability to
protect themselves or even flee a more powerful adversary (e.g., Wisby).
Although both nonlethal and lethal forms of violent injury are highly prevalent in many of
the populations discussed in this chapter, the dominance of one category over the other
informs our understanding of the intentions of the attacker. For example, the higher preva-
lence of nonlethal than lethal injury in a number of settings – Australia, Santa Barbara
Channel, Peru, and Easter Island – indicates that injury was meant to maim and not kill the
victim. Death of the opponent was clearly the preferred outcome of attack in the Middle Ages
of Europe (e.g., Wisby), late prehistoric Great Plains and Midwest (e.g., Crow Creek, Norris
Farms), in the Arctic (e.g., Saunaktuk), and in historic North America (e.g., Snake Hill). In
prehistoric settings, it usually is not possible to determine the reasons for preference of lethal
or nonlethal forms of violence. In California, the shift to lethal forms of injury from projectiles
in later prehistory may have been influenced by the change in weapons technology coupled
with increasing resource stress.
Clear patterns of mutilation of victims are well documented in a number of prehistoric and
other New World settings. In North America, the evidence for removal of soft tissue and skin
from the cranial region – especially scalping – is abundant. Typically, the scalp was removed
by first slicing skin along the frontal and parietal bones and peeling back of the skin (e.g.,
Norris Farms, Koger’s Island), but other approaches involved removal of facial and other
tissues (e.g., Saunaktuk). Mutilation was a highly visible behavior. In addition to scalps,
tongues, noses, limbs, and heads were removed from the near-dead or deceased (e.g., Great
Plains). A more profound mutilation to the head than scalping was decapitation. Decapitation
was practiced in the New World and Old World, and for a variety of reasons. In Roman Britain
(and throughout history), this was a preferred form of execution in some groups. In northern
Europe, the head of the victim was sometimes placed between the legs (Denmark, England),
perhaps as the ultimate insult. Other unique and highly localized forms of body treatment of
the living victim were likely practiced. For example, gouging of knees at Saunaktuk in the
Arctic may have been associated with a practice of dragging the victim through the village
prior to his or her death.
Trauma data are important for dispelling prejudices and assumptions about past societies.
For example, hunter-gatherer societies around the world are often characterized as peaceful
inhabitants of stress-free environments living in a state of blissful repose (Lee & DeVore,
1968; Service, 1966; and see discussions by Burbank, 1994; Fienup-Riordan, 1994; Walker,
2001a). This characterization may reflect the fact that anthropologists doing fieldwork among
these societies are guests – after all, what guest is going to go back home and write about the
unpleasant things they observed, especially with regard to violent encounters between
individuals (Erchak, 1996; Keeley, 1996)? Many anthropologists underplay the negative or
offensive, avoiding realism in social beliefs. In fact, a number of cultures described as
nonviolent or peaceful have homicide rates far exceeding those of some Western nations
(Knauft, 1987).
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176 Injury and violence
Ethnographic research has undergone a dramatic change, with key developments showing
the social, political, and economic contexts for a variety of settings and the relevance of
violence to social life (Burbank, 1994; Kapferer, 2012; Stewart & Strathern, 2002). The point is
not to replace peaceful characterizations of earlier societies with violent ones. Rather, these
findings underscore the importance of substituting an incorrect image with one that fits the
evidence, past and present. This approach is critical for informing our perspective on past
groups as functioning societies rather than as images of what earlier social behavior must
have been like. This newfound precision adds a more complete historical context for the study
of recent human behavior. Anthropologists and others seem to employ – either consciously or
unconsciously – their own cultural and social assumptions about earlier societies in order to
“remember” the past (Keeley, 1996). We project these assumptions into a past that seems to
reflect current and highly biased perspectives on the condition of humankind, be they
peaceful or violent. These skeletal data help us to reconstruct and interpret trauma and
violence in a more comprehensive and accurate manner.
This chapter has discussed the obvious skeletal correlates of violence and trauma in a
wide range of societies around the world, namely injuries received in interpersonal
conflict, ritual, warfare, and other settings where individuals are intentionally injured or
killed. Yet, there is another kind of violence that places people at risk of harm or death,
perhaps not immediately, but rather the kind that inhibits a person from maintaining
homeostasis owing to social circumstances. This structural violence pertains to members of
societies where needs are not met or where they are exploited for economic, political, or
other reasons (Farmer, 1996). Borrowing from developments in other social sciences, Klaus
(2012, 2014a) makes the case that broken bones and cutmarks are not the only reflection
of violence in human remains. Specifically, he documents evidence of the impairment of
human needs caused by social structures that prevent persons from reaching biological,
ocial, and economic potentials in prehistoric societies involving social inequality. Focus-
sing on the colonial-era Lambayeque Valley, Peru, he draws on ethnohistoric and arch-
aeological context in the newly established Spanish colonial state. This hierarchical
structure contained strong elements of violent repression for members of the native
populations in the region. The dominant European power viewed native peoples as a
source ripe for exploitation and ruled by a set of repressive laws where their labor was
bought, sold, and (usually) abused. This kind of exploitative system denied certain foods
and other resources, yet at the same time, demanded heavy physical labor. The predictable
outcome, of course, would be evidence of poor health, disease, physiological stress, and
biomechanical demand. In these circumstances, this outcome in health is best described as
structural violence.
In order to test the hypothesis that structural forms of violence were in place in the
Lambayeque Valley, a region colonized by Spain beginning in 1534, Klaus (2014a) compared
the record of health and diet of precolonial populations with colonial-era populations
represented by remains recovered from the Chapel of San Pedro de Mórrope dating from the
period of 1536–1750. This record reveals a compelling picture of outcomes of repression,
including disrupted homeostasis and decline, a remarkable increase in porotic hyperostosis
(154% increase), enamel hypoplasia (184% increase), and periosteal reaction owing to infec-
tion (471% increase). Along with the bioarchaeological record of violence and traumatic
injury associated with the Spanish conquest elsewhere in the Inca Empire (Gaither & Murphy,
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4.6 Summary and conclusions 177
2012; Murphy et al., 2010), the record of structural violence provides a window onto a rapidly
changing political landscape involving conquest and population collapse.
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Activity patterns: 1. Articular
degenerative conditions and
5 musculoskeletal modifications
5.1 Introduction
Physical activity is a defining characteristic of human adaptations. Hunter-gatherers, for
example, are often characterized as highly mobile, hard-working, and physically active. In
contrast, agriculturalists are sometimes seen as having an easier life – they are settled in one
place and have a lighter workload than hunter-gatherers. In his popular and influential
archaeology textbook, Robert Braidwood (1967:113) distinguished hunter-gatherers as leading
“a savage’s existence, and a very tough one. . .following animals just to kill them to eat, or
moving from one berry patch to another (and) living just like an animal.” Ethnographic and
other research calls into question these simplistic portrayals of economic systems. Following
the publication of Lee and DeVore’s (1968) Man the Hunter conference volume, and especially
Lee’s (1979) provocative findings regarding work behavior and resource acquisition among the
!Kung in northern Botswana, a consensus emerged that, contrary to the traditional Hobbesian
depiction of hunter-gatherer lifeways as “nasty, brutish, and short,” prehistoric foragers were
not subject to overbearing amounts of work, and life overall for them was leisurely, plentiful,
and confident (Sahlins, 1972). More importantly, these developments fostered a wider discus-
sion by anthropologists and other social scientists of activity, behavior, and lifestyle in both
present and past hunter-gatherers (Kelly, 2013). These discussions led to the conclusion that
human adaptive systems are highly variable. As a result, it is now clear that it is not possible to
make blanket statements about the nature of workloads or other aspects of lifestyle in foragers
and farmers (Kelly, 1992, 2013; Larsen, 1995). Rather, workload and lifestyle are highly
influenced by the kinds of resources exploited, climate, and sometimes highly localized
circumstances. Nevertheless, there are some general patterns that emerge via bioarchaeological
study of past human populations, which this chapter will discuss, in part.
Workload and activity have enormous implications for demographic history of a popula-
tion. The study of living humans indicates, for example, that demanding physical activity in
reproductively aged females results in reduced ovarian function and fecundity (Dufour, 2010;
Ellison, 1994; Jasienska, 2010). Thus, the identification of workload and patterns of physical
activity from the study of human remains may provide indirect reflections of variation in
birthrates and fertility in some past populations.
Of course, we cannot observe and document levels of ovarian function in past populations
directly. However, human remains offer a fund of data for documenting and inferring patterns
and level of workload and other aspects of lifestyle that involve physical activity. Specifically,
the study of pathological and nonpathological changes of articular joints and behaviorally
related modifications of nonarticular regions offers important insights that are not available
from any other record derived from archaeological settings.
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180 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
by ship builders and others results in similar modifications (Lawrence, 1955, 1961); lifting of
long tongs to move hot metals by foundry workers results in degenerative changes in the
elbow (Hough & Sokoloff, 1989); and repetitive activity involving the hands in cotton mill
workers results in different patterns of osteoarthritis (Hadler, 1977; Hadler et al., 1978).
Other findings for manual laborers, farmers, ballet dancers, various types of athletes, and
those who engage in rigorous exercise generally support these observations (Coggon et al.,
1998; Cooper et al., 1994; Croft, Coggon et al., 1992; Croft, Cooper et al., 1992; Felson
et al., 1988; Forsberg & Nilsson, 1992; Lawrence, 1977; McKeag, 1992; Nakamura et al.,
1993; Stenlund, 1993).
Epidemiological findings are providing important corroboration for conclusions linking
mechanical demand and osteoarthritis. Comparisons reveal markedly higher prevalence of
knee and hip osteoarthritis in North Carolina rural populations than in the United States
(primarily urban) population as a whole (hip: 25.1% vs. 2.7% in the 55–64 age cohort:
Jordan et al., 1995). These differences suggest the greater physical demands of the rural
lifestyle in the modern United States. This same pattern of variation is expressed in at least
one archaeological context. In this regard, the comparison of rural and urban populations
from ancient Corinthian Greece reveals a generally higher prevalence in the former than the
latter, reflecting more strenuous physical labor (Rife, 2012). Rife (2012) suggests that the
greater prevalence of shoulder and vertebral osteoarthritis in the rural individuals reflects
their exposure to work involving agricultural labor, such as field labor, tending livestock,
and other behaviors.
The links between physical activity and osteoarthritis are not straightforward, however.
The hand bones of weavers from the Spitalfields, London skeletal series have no more
osteoarthritis than hand bones from the general sample (Waldron, 1994; and see various
citations in Jurmain, 1999). Manual laborers in this series have no more or less osteoarth-
ritis than the population as a whole. These findings and a survey of inconsistencies found
in the epidemiological literature led Waldron to conclude “that there is no convincing
evidence of a consistent relationship between a particular occupation and a particular form
of osteoarthritis” (1994:94). On the other hand, in some unusual circumstances, there
appears to be a pattern of articular modifications that link with known and highly specific
physical activities (Ciranni & Fornaciari, 2003) or age-related degeneration of a particular
joint or joints of the skeleton in relation to a particular activity regimen (Stirland, 2002;
and see Waldron, 2012). Thus, while articular pathology relating to activity offers an
important insight into behavioral characteristics of human populations in a general sense,
the identification of specific activities or occupations from individual remains is the rare
exception to the general rule.
Like other chronic diseases or disorders, the influence of environmental conditions in the
prenatal environment, infancy, and childhood (low birth weight, poor nutritional status) has
been implicated (Jordan et al., 2005; Melanson, 2007; Peterson et al., 2010). Epidemiologists
and anthropologists observe a great deal of worldwide variation in osteoarthritis in relation to
age (Corti & Rigon, 2003). For example, young adults and older juveniles in some human
populations express a relatively high frequency of osteoarthritis (Chapman, 1972; Chester-
man, 1983; Larsen, Ruff et al., 1995; Rojas-Sepúlveda et al., 2008). In urbanized industrial
societies, osteoarthritis rarely occurs before the age of 40 years (Arden & Nevitt, 2006), a
pattern that is clearly different from the archaeological record showing a much earlier age of
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5.3 Articular joint pathology: osteoarthritis 181
onset (Larsen & Hutchinson, 2010). Regardless of cause, the record shows consensus on one
key factor. That is, owing to disability, the economic, social, and behavioral costs of osteo-
arthritis are substantial in light of the cascade of negative health outcomes from loss of
movement (Corti & Rigon, 2003; Sharma, 2001).
In some respects, epidemiological studies provide an important baseline for interpreting
osteoarthritis in past human populations. For example, the very large samples present a
compelling picture of a huge amount of variation. However, much of the literature is
hampered by the focus on the comparison of folk racial groups (African American vs. White
vs. Asian), having little to do with population biology or modern concepts of human variation
(Allen, 2010). Moreover, bioarchaeological and epidemiological studies are not strictly com-
parable in that the latter are almost always based on clinical contexts, either radiological
examinations or patient interviews, which do not identify subtle degenerative changes seen in
the actual skeletal specimens that bioarchaeologists study. Moreover, clinical evaluations
include factors such as joint capsule spacing. Thus, hard tissue changes observed in the
clinical setting are not strictly comparable to those observed in archaeological or other types
of skeletal collections.
The pathophysiology of osteoarthritis is complex and incompletely understood, especially
regarding the relationship between hyaline cartilage and bone changes. Some have argued
that changes in cartilage – including fibrillation or tearing – precede bony responses; others
contend that minute changes in subchondral bone precede cartilaginous changes (Radin,
1982). For archaeological remains, the exact order of tissue response to mechanical stress is
immaterial because regardless of the order of events, the skeletal changes arising from
osteoarthritis are universal, including proliferative exophytic growths of new bone on joint
margins (“osteophytes” or “lipping”) and/or erosion of bone on joint surfaces (Figure 5.1). In
some joints, the cartilaginous tissue covering the articular surface has failed, resulting in
pitting or rarefaction of the surface. In instances where the cartilage has disintegrated
altogether, the articular surface becomes polished due to direct bone-on-bone contact
(Figure 5.2). Because the surface has a glistening appearance reminiscent of ivory, the
polished area is called eburnation. In the hinge joints of the knee and elbow, deep, parallel
grooves may be present on the eburnated surface (Ortner, 2003; Rogers & Waldron, 1995). The
presence of eburnation indicates that although the articular cartilage is missing, the joint was
still actively used at the time of death (Rogers & Dieppe, 2003).
Osteophytes vary from fine tuft-like, barely perceptible protrusions to large projections of
spiculated bone. Even in the extreme, mobile diarthrodial joints do not usually fuse. In spinal
osteoarthritis, the marginal osteophytes of two adjacent vertebrae may unite, thus forming
a bridge of continuous bone. This change (ankylosis) is also accompanied by reduction in
disk space separating the two vertebral bodies, and hence, marked reduction in mobility of
the spine.
Compression or crush fracture of anterior vertebral bodies – an occasional concomitant
of spinal osteoarthritis – gives them a wedge-shaped appearance (Figure 5.3). Additionally,
herniation of the intervertebral disk results in irregular depressions on intervertebral body
surfaces called Schmorl’s depressions (Ortner, 2003; Schmorl & Junghanns, 1971)
(Figure 5.4).
Biological anthropologists, anatomists, and others have systematically collected data on
osteoarthritis for more than a century. Wells referred to osteoarthritis as “the most useful of all
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182 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
(a)
(b)
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5.3 Articular joint pathology: osteoarthritis 183
(c)
(d)
diseases for reconstructing the life style of early populations” (Wells, 1982:152). Osteoarthritis
is present in all human populations, and regardless of etiology, the patterns documented and
interpreted by bioarchaeologists provide a picture of the cumulative effects of mechanical
stress and age on the body in different human groups. Owing to the lengthy history of study
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184 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
(e)
(f)
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5.3 Articular joint pathology: osteoarthritis 185
Figure 5.2 Distal right humerus showing eburnation (osteoarthritis); anatomical specimen.
(From Larsen, 1987; photograph by Barry Stark; reproduced with permission of
Academic Press, Inc.) (A black and white version of this figure will appear in some formats.
For the color version, please refer to the plate section.)
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186 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
Figure 5.3 Collapsed thoracic vertebrae; Cochiti, New Mexico. (Photograph by Clark
Spencer Larsen.) (A black and white version of this figure will appear in some formats. For
the color version, please refer to the plate section.)
particular lifestyles. For example, it is highly unlikely that sedentary United States popula-
tions in the twenty-first century exhibit the same profile of osteoarthritis as Great Basin
foragers.
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5.3 Articular joint pathology: osteoarthritis 187
Figure 5.4 Schmorl’s depression on the superior surface of the thoracic body;
Lambayaque, Peru. (Photograph by Sam Scholes.) (A black and white version of this figure
will appear in some formats. For the color version, please refer to the plate section.)
lifting and carrying. The conspicuous anterioposterior vertebral body elongation may be
caused by various activities that involve extreme ventral flexion of the body trunk.
A number of Neanderthal skeletons have distinctive patterns of osteoarthritis that are
useful for reconstructing posture and activity in the late Pleistocene, providing a context
for interpreting behavioral antecedents to modern humans. Based on his study of the La
Chapelle-aux-Saints skeleton, Boule reconstructed the individual “as an almost hunchbacked
creature with head thrust forward, knees habitually bent, and flat, inverted feet, moving along
with a shuffling, uncertain gait” (Straus and Cave, 1957:348). This image of Neanderthal
locomotion served as a model for behavioral reconstruction, and it reinforced the popular
image of Neanderthals as less than human. Straus and Cave (1957) suggested that Boule
misinterpreted key aspects of the anatomy of the skeleton and overlooked the possibility that
severe osteoarthritis may have prevented the individual from normal perambulation. Analysis
of the La Chapelle skeleton reveals the presence of widespread degenerative pathology
(especially marginal lipping) involving the temporomandibular joint, the occipital condyles,
lower cervical vertebrae, and thoracic vertebrae (the T1–T2 exhibits eburnation, and the T6,
T10, and T11 have possible eburnation) (Trinkaus, 1985). The left acetabulum shows extreme
lipping and eburnation. Although the right acetabulum is missing, the head of the right femur
is normal, suggesting that the hip osteoarthritis is unilateral. The severe osteoarthritis in the
left hip suggests that it would have been painful for the individual to walk or run. The overall
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188 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
pattern of degenerative pathology indicates that locomotor abilities may have been somewhat
limited, but certainly not in the manner imagined by Boule (Trinkaus, 1985).
Osteoarthritis is also extensive in the Neanderthal adults (n¼6) from Shanidar, Iraq (Trin-
kaus, 1983). The widespread nature of articular pathology in these individuals reflects a highly
physically demanding lifeway for these archaic Homo sapiens. This conclusion is confirmed
by other lines of evidence, such as the high overall robusticity and bone strength in these
hominins (Lovejoy & Trinkaus, 1980; Ruff et al., 1993; Trinkaus, 1984; Trinkaus & Ruff, 1999,
2012; and see Chapter 6).
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5.3 Articular joint pathology: osteoarthritis 189
supplies and long-distance travel was physically demanding, requiring heightened strength
and endurance of long distance travel (Larsen, Ruff et al., 1995).
In order to determine which of the two models best characterizes adaptive strategies of
native populations in the Great Basin, Larsen, Ruff, and coworkers (1995, 2008) assessed
pattern and prevalence of osteoarthritis in prehistoric human remains recovered from the
Stillwater Marsh region, a large wetlands area in western Nevada. Analysis of these remains
revealed an abundance of osteoarthritis. Most adults, including all individuals over the age of
30 years, have osteoarthritis in at least one, and usually multiple, articular joints. Articular
pathology for older adults involves severe proliferative lipping on joint margins, eburnation,
vertebral compression fractures, and Schmorl’s nodes. Contrary to expectations of the limno-
sedentary model, these findings suggest that hunter-gatherers in this setting led extremely
demanding lives. The high prevalence of osteoarthritis suggests elevated mechanical demand,
such as in heavy lifting and carrying. These findings also imply that prehistoric groups may
not have been tethered to the marsh, but they exploited a wide range of resources from both
the marsh and the surrounding uplands. Beyond concluding that the Great Basin lifeway was
physically demanding, however, it is not possible to state whether these populations were
sedentary or mobile from osteoarthritis evidence alone. Analysis of long bone structural
morphology is more informative on this point (Larsen et al., 2008; Ruff, 2010b; and see
Chapter 6).
Similar assessment of osteoarthritis in the Malheur wetlands in the northern Great Basin
(Oregon) reveals that, like the Stillwater series, there is a strikingly high level of articular joint
pathology, especially in comparison with foragers and later farmers from the Georgia Bight
(Hemphill, 2010) (Figure 5.5). Interestingly, the prevalences for each articular joint, comparing
the two Great Basin series, are statistically indistinguishable (chi-square; P>0.05). The high
values for both series speak to the likelihood that both settings – Stillwater in the western
Great Basin and Malheur in the northern Great Basin – were engaged in very similar adaptive
strategies focusing on largely the same resources, manner of acquiring them, and level
of activity and workload that make the acquisition process economically successful
(Hemphill, 2010).
Population comparisons
These studies of archaic hominins and modern foragers underscore the highly variable nature
of osteoarthritis. In order to assess general patterns of and variation in past physical activities
on a broad basis, comparisons of many different skeletal samples are necessary. Bridges
(1992) attempted a broad-scale analysis by reviewing published studies on appendicular
(shoulder, elbow, hip, knee) and axillary (vertebrae) osteoarthritis in native populations from
North America. In the 25 skeletal samples included in her review, osteoarthritis shows the
highest prevalence in the knee for 17 samples; elbow osteoarthritis is either the most or next
most prevalent for 15 samples. No clear association between osteoarthritis and subsistence
mode emerges in comparing hunter-gatherers and agriculturalists. However, agriculturalists
tend to have a low prevalence in the wrists and hands, but not all foraging groups have high
levels in these joints. For nearly all populations reviewed, ankle or foot arthritis is less
common than hand osteoarthritis.
The comparison of different populations in published findings (Bridges, 1992) contributes
to an understanding of variation in work burdens and activity. However, these comparisons
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190 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
Males
100 Malheur
Stillwater
Percentage of Individuals Affected
80 Georgia Preagricultural
Georgia Agricultural
60
40
20
–20
Cer Tho Lum Sac Sho Elb Wri Han Hip Kne Ank Ft
Joint Region
Females
90 Malheur
80 Stillwater
Percentage of Individuals Affected
Georgia Preagricultural
70
Georgia Agricultural
60
50
40
30
20
10
–10
Cer Tho Lum Sac Sho Elb Wri Han Hip Kne Ank Ft
Joint Region
Figure 5.5 Frequency and distribution of osteoarthritis among males and females in
the Great Basin (Malheur Lake and Stillwater Marsh) and Georgia coast preagricultural
and agricultural populations.
are limited by the variable nature of the methods of data collection used by the different
researchers (see discussions by Bridges, 1993; Lovell, 1994; Waldron & Rogers, 1991). This
factor alone may prevent investigators from presenting clear diachronic trends or population
differences in osteoarthritis prevalence when comparing findings reported by different
researchers (Cohen, 1989). Data collection and population comparisons by the same researcher
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5.3 Articular joint pathology: osteoarthritis 191
or by researchers sharing the same methods circumvent this problem. These types of compari-
sons provide an important perspective on general characteristics of different lifestyles,
especially with regard to workload and level of mechanical demand on the body.
In a classic investigation of variation in degenerative articular pathology, Jurmain (1977a,
1977b, 1978, 1980) assessed osteoarthritis patterns in the appendicular skeleton (shoulder,
elbow, hip, and knee) in a range of populations, including American Whites and Blacks (Terry
Collection), Eskimos (Alaska), and Native Americans (Pecos Pueblo, New Mexico). Eskimos
have a higher prevalence and severity of osteoarthritis than do American Whites and Blacks
or Pecos Pueblo Native Americans; Pecos Pueblo adults have the least prevalence and severity
of osteoarthritis among the four groups (Table 5.1). These population differences reflect the
highly variable mechanical demands associated with contrasting lifestyles and subsistence
strategies. For example, mechanical demands for the Pecos Pueblo agriculturalists may be
mostly limited to the growing season, whereas Eskimos are subjected to high levels of activity
throughout the year (Jurmain, 1977a; and see Merbs, 1983).
The impact of specific lifestyles and occupations on patterns of degenerative articular
pathology in various colonial and postcolonial North American populations has received
considerable attention by biological anthropologists. These studies reveal that for many
settings, physical activities were highly demanding. African Americans from a range of
circumstances provide a growing record of lifestyle and activity in urban and rural settings.
For example, individuals in the First African Baptist Church (urban Philadelphia) cemetery
display extensive degenerative spinal pathology, including osteoarthritis (males, 69%;
females, 39%) and Schmorl’s nodes (males, 31%; females, 13%) (Parrington & Roberts,
1990; and see Angel et al., 1987; Davidson et al., 2002). Similarly, in the African Burial
Ground (urban New York), osteoarthritis is highly prevalent, with the lumbar vertebrae being
especially highly affected (45% of individuals aged 15–24 years; Wilczak et al., 2009). These
prevalences are higher than those of a contemporary African American population from a
rural setting in Cedar Grove, Arkansas (compare with Rose, 1985) and such differences
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192 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
suggest that the urban lifestyle was far more mechanically demanding than the rural lifestyle.
The differences in degenerative joint pathology between the urban and rural settings may
be due to specific differences in habitual activities. For example, historical records indicate
that individuals interred in the Philadelphia cemetery held unskilled, physically demanding
jobs (see also other settings of African Americans in relation to mechanical environment:
Davidson et al., 2002; Kelley & Angel, 1987; Owsley et al., 1987; Rathbun, 1987; Rathbun &
Steckel, 2002; Thomas et al., 1977).
Highly demanding circumstances are also inferred from the study of osteoarthritis in
pioneer Euroamericans living in the rural frontier of the American Midwest and Great
Plains. Euroamerican adults from Illinois and Texas have remarkably elevated prevalences
of osteoarthritis and highly developed muscle attachment sites on limb bones (Larsen,
Craig et al., 1995; Winchell et al., 1995). Articular degenerative pathology includes
extensive marginal lipping on weight-bearing and nonweight-bearing joints, eburnation,
and extensions of articular surfaces (e.g., anterior femoral head and neck). High prevalence
of nonspecific physiological stress indicators (e.g., enamel defects) and historical evidence
indicate that life on the early American frontier was generally unhealthy and physically
demanding. Numerous historical accounts from the early to mid-nineteenth century
discuss the extremely hard physical labor that pioneer families endured, especially in
preparation of fields and tending and harvesting crops (see discussion in Larsen, Craig
et al., 1995).
Degenerative joint pathology among war casualties is especially revealing about phys-
ical activity in military contexts, mostly drawn from rural, frontier circumstances. Many
of the Euroamerican skeletal remains from the War of 1812 Snake Hill cemetery near Fort
Erie, New York, and the Little Bighorn, South Dakota battlefield display Schmorl’s nodes
on vertebral bodies, with an unusually high prevalence, including the majority of individ-
uals (Owsley, 1991; Scott & Willey, 1997). Some individuals from Snake Hill, for example,
have multiple Schmorl’s nodes: six individuals have five or more vertebrae with nodes,
and one soldier has pronounced nodes in 11 vertebrae. In addition, several individuals
have vertebral compression fractures resulting from excessive mechanical loading of the
back. Similarly, military recruits serving on the mid-sixteenth-century warship, Mary Rose
display a high prevalence of Schmorl’s depressions (thoracic vertebrae: 26.7%) (Stirland,
2002; and compare with Knüsel, 2000). These recruits, composed mostly of adolescents
and young adults, were subjected to heavy mechanical loading in general, and of the
back specifically, in a range of activities, including lifting in quite confined spaces. In
these military settings, the elevated level of vertebral pathology indicate that pre-modern
military recruits were subjected to excessive loading of their spines, such as from lifting
heavy military hardware, carrying heavy loads, and participation in rigorous activity
regimens.
In sharp contrast to these settings, the largely sedentary urban population from
nineteenth-century Belleville, Ontario, presents a remarkably low prevalence of osteoarth-
ritis, including in the spine (Saunders et al., 2002). This speaks to the relatively low amount
of physical activity in comparison with the rural and military individuals described earlier.
In addition, a poorhouse population from Rochester (New York) displays a relatively low
prevalence and severity of osteoarthritis, although not as low as that of Belleville (Higgins
et al., 2002).
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5.3 Articular joint pathology: osteoarthritis 193
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194 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
Horseback riding
The horse was an important mode of transport for many Holocene societies, in the Old World
and later in the New World following European contact, and into the early twentieth century.
Some populations show articular degenerative sequelae of an equestrian lifestyle in the
limited number of settings studied by bioarchaeologists (Dutour & Buzhilova, 2014; Edynak,
1976; Larsen, Craig et al., 1995; Owsley et al., 2006; Pálfi, 1992; Reinhard et al., 1994).
Following the introduction of the horse to the American Great Plains by Europeans, native
populations relied on this animal as the key element in the acquisition of resources. Patterns
of osteoarthritis attributed to horseback riding include a high frequency of degenerative
changes in the vertebrae and pelves of adult males in historic-era Omaha and Ponca from
northeastern Nebraska, along with other skeletal features that are best explained by mechan-
ical loading of specific joints during horseback riding (Reinhard et al., 1994). Features
associated with horseback riding are especially diagnostic in the hip joint (innominates,
proximal femora). These features include superior elongation of the acetabulum, extension
of the femoral head articular surface onto the anterior femoral neck (Poirier’s facets), and
hypertrophy of muscle attachment sites for the muscles: adductor magnus, adductor brevis,
vastus lateralis, and gastrocnemius (medial head) (Dutour & Buzhilova, 2014; Erikson et al.,
2000; Reinhard et al., 1994). The development of these hip (and knee) muscles reflects the
emphasis on stabilizing the hip and keeping the rider upright. Extensive osteoarthritis of the
first metatarsals is suggestive of mechanical stresses associated with the placement of the first
toe into a leather thong stirrup (Reinhard et al., 1994). In all settings studied, more males than
females have pathological changes associated with horseback riding, thus indicating that men
were habitually engaged in behaviors involving the use of the horse, more so than women.
Vertebral osteoarthritis
The vertebral column has been studied in a large number of settings in the Americas
(summarized in Bridges, 1992) and elsewhere. For prehistoric North America, these compari-
sons reveal a number of tendencies. First, prevalence is always greatest in the articular region
between the fifth and sixth cervical vertebrae; second, there is a tendency for the lower
thoracic to be affected more than the upper thoracic vertebrae; third, the second to fourth
lumbar vertebrae usually show the greatest degree of marginal lipping in comparison with
other vertebrae; and finally, the region encompassing the seventh cervical vertebra to the
upper thoracic vertebrae (to about the third thoracic vertebra) is always least affected by the
disorder (Bridges, 1992). The relatively minimal amount of osteoarthritis in the thoracic
vertebrae is due to the lower degree of movement in this region of the back (Waldron, 1993).
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5.3 Articular joint pathology: osteoarthritis 195
For a wide range of populations globally, the highest prevalence of vertebral osteoarthritis
is in the lumbar spine, followed by the cervical spine (Bennike, 1985; Bridges, 1994; Gunness-
Hey, 1980; Jurmain, 1990; Klaus et al., 2009; Merbs, 1983; Snow, 1974). Some human
populations show relatively higher levels of osteoarthritis in the cervical vertebrae. For
example, cervical vertebral osteoarthritis is relatively elevated in the Spitalfields, London
industrial urban group (Waldron, 1993). Similarly, Harappan populations from the Indus
Valley display higher frequencies of osteophytes and articular surface pitting of cervical
vertebral bodies than in either the lumbar or thoracic spine (Lovell, 1994). This pattern
suggests an activity-related cause, such as carrying of heavy loads on the head. Individuals
in traditional agricultural communities and from lower socioeconomic groups from urban
settings in South Asia habitually carry loads on their heads (Lovell, 1994). These loads include
laundry bundles, water jars, firewood, and dirt-filled containers at construction sites. Clinical
and observational studies confirm that the upper (cervical) spine is susceptible to injury and
cumulative degenerative changes by persons carrying heavy loads on their heads (Allison,
1984; Levy, 1968; Lovell, 1994). The greater severity of osteoarthritis in the cervical spine in
women than in men suggests that the practice of burden-carrying with the use of the head is
gender specific. For example, severity of cervical osteoarthritis is greater in adult females than
adult males in the Romano-British Bath Gate populations from Cirencester, England (Wells,
1982; and see Lovell, 1994).
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196 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
Averbuch males have significantly greater frequencies of osteoarthritis for the shoulder and
hip, but not the knee. This pattern is suggestive of change in workload and activity with the
adoption of agriculture.
Unlike males, agriculturalist females from the lower Illinois River valley have a higher
prevalence of vertebral osteoarthritis than forager females from the same region (Pickering,
1984). These differences are especially pronounced in cervical vertebrae, which may be related
to an increase in mechanical demand in this region of the skeleton with the shift to agriculture
(Pickering, 1984).
Fahlström (1981) identified an unusually high prevalence and severity of shoulder osteo-
arthritis in adult males in the Medieval skeletal series from Westerhüs us, Sweden. Historical
analysis of this population suggests that the high frequency in males reflects work and
activity practices that are exclusive to men, including parrying in sword fighting, spear
throwing, timber cutting, and other activities associated with repetitive, heavy loading of
the shoulder joint (Fahlström, 1981).
Some analyses reveal no appreciable differences between males and females. For example,
males and females in the Dickson Mounds, Illinois series show no differences in prevalence of
appendicular osteoarthritis (Goodman, Lallo et al., 1984; Lallo, 1973). The similarity between
sexes infers that mechanical loading of most articular joints in this setting was broadly the
same in adults regardless of sex, in contrast to most other prehistoric Eastern Woodlands
populations (compare with Bridges, 1992). Similarly, documentation of prevalence of
osteoarthritis in two series of African American adults from nineteenth- and twentieth-
century Washington, DC shows no appreciable differences between adult males and females,
suggesting that labor demands were similar for men and women in this setting (Watkins, 2012).
Two clear trends emerge when examining sex differences (Bridges, 1992). First, where there
are statistically significant differences between males and females, males nearly universally
show a higher prevalence of osteoarthritis than females. Second, when looking at specific
regions of the New World, maize agriculturalists tend to display more sexual dimorphism
in degenerative pathology than foragers. This suggests a difference in behavior leading to
degeneration of articular joints in agriculturalists but not in earlier foragers. The change
in pattern of sexual dimorphism suggests that there was a fundamental shift in the division of
labor once agriculture was adopted (Bridges, 1992).
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5.3 Articular joint pathology: osteoarthritis 197
remains from coastal British Columbia (Cybulski, 1992). Adult females are older than the adult
males in the assemblage. Thus, an unusually high prevalence in females relative to males is
likely due to the difference in age composition rather than variation in mechanical environment.
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198 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
different settings (Jurmain, 1977a, 1977b, 1978, 1980) indicate differences in osteoarthritis –
and presumably workload and activity – in relation to subsistence. Regionally based temporal
studies of osteoarthritis give additional perspective on change in functional demand as
populations underwent adaptive shifts in the past. Based on comparisons of earlier and later
societies from the same region, it has become possible to assess the relative labor costs of
change in economic focus, at least as these costs are measured by mechanical stress.
The most extensive temporal studies of osteoarthritis have been completed for several
settings in North America. The study of osteoarthritis prevalence in Archaic-period hunter-
gatherers and later Mississippian-period maize agriculturalists from northwestern Alabama
suggests changes in activity and workload, especially when viewed in the context of diet
and lifeway (Bridges, 1991a). Archaic-period populations exploited a range of terrestrial
and riverine animals and plants, including deer, raccoon, beaver, fish and shellfish, wild
plants, and limited cultivation of sunflower, sumpweed, chenopod, squash, and bottle gourd
(Dye, 1977). Populations moved seasonally from river valleys to nearby uplands. Later
prehistoric groups were intensive maize agriculturalists, but also exploited a limited number
of species of nondomesticated plants and animals (Smith, 1986). These later groups were
largely sedentary and lived primarily in villages on river floodplains, although smaller
temporary upland habitations were utilized for hunting deer and other animals on a
seasonal basis (e.g., small mammals, turkey, waterfowl). In summary, although sharing
some features, the subsistence strategies and settlement patterns in the earlier and later
periods were very different. Because foraging and farming involved very different kinds of
physical activity, the respective populations should display different prevalence and pat-
terns of osteoarthritis.
Comparisons of shoulder, elbow, wrist, hip, knee, and ankle osteoarthritis show a number of
important temporal trends in the Alabama series (Bridges, 1991a). For individuals 30–49 years
of age-at-death, the Archaic group generally has more osteoarthritis than the Mississippian
group, and these differences are especially consistent for males (Table 5.2). Statistically
significant differences between periods are present in only a few of the joints. However, the
overall greater prevalence in the Archaic sample is clear. The severity of osteoarthritis tells the
same story: Archaic populations generally have greater severity of the disorder than Mis-
sissippian populations. The pattern of degenerative pathology is remarkably similar in the
prehistoric foragers and farmers and in the males and females within each group – for all
samples, osteoarthritis is most common in the elbow, shoulder, and knee, and it is least
common in the hip, ankle, and wrist.
Prehistoric and contact-period human remains representing a temporal succession of
Native American populations living in the Georgia Bight have been the focus of research on
physical activity and behavioral changes by Larsen and coworkers (Fresia et al., 1990; Larsen,
1982, 1984, 1998; Larsen & Ruff, 1991, 1994, 2011; Larsen et al., 2007; Ruff & Larsen, 1990;
Ruff et al., 1984). Temporal comparison of osteoarthritis prevalence shows a distinctive
decline in prehistoric farmers relative to earlier foragers. For the series as a whole (sexes
combined), statistically significant reductions occur for the lumbar vertebrae (26.2%), elbow
(6.8%), wrist (4.5%), hip (3.8%), knee (7.2%), and ankle (4.0%) joints. Frequency of osteoarth-
ritis either reduces or does not change in all other joints. Both adult females and adult males
show the same trend of reduction; more significant reductions occur in females than in males
(six joints versus three joints; and see earlier). The pattern of osteoarthritis prevalence in the
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5.3 Articular joint pathology: osteoarthritis 199
Table 5.2 Percentage of individuals with moderate to severe osteoarthritis, aged 30–49 years
(Adapted from Bridges, 1991a: Table 2)
Males Females
Joint
Shoulder 36.8 42.1 30.0b 30.4 7.7 28.6 10.0 17.6
(n) (19) (19) (20) (23) (13) (14) (20) (17)
Elbow 27.3 40.9 28.0b 24.0 26.4 37.6 15.8 20.0
(n) (22) (22) (25) (25) (19) (16) (19) (20)
Wrist 9.5 15.8 0.0 17.4 0.0 6.7 5.6 0.0
(n) (21) (19) (23) (23) (13) (15) (18) (14)
Hip 5.0 5.0 0.0 0.0 0.0 0.0 7.1 0.0
(n) (20) (20) (21) (21) (13) (10) (14) (17)
Knee 27.3 31.8 21.7 8.6 15.8 22.3 21.1 23.5
(n) (22) (22) (23) (23) (19) (18) (19) (17)
Ankle 23.8a–c 0.0 0.0b 4.8b 0.0d 5.9 0.0 0.0
(n) (21) (22) (24) (21) (18) (17) (16) (19)
a
Frequency significantly greater in males than in females (chi-square: P<0.05).
b
Severity significantly greater in males than in females (Mann-Whitney rank sum: P<0.05).
c
Frequency significantly greater in Archaic group than in Mississippian group (chi-square: P<0.05).
d
Severity significantly greater in Archaic group than in Mississippian group (Mann-Whitney rank
sum: P<0.05).
skeleton is similar in both the preagricultural and agricultural groups. In both series the
lumbar and cervical vertebral, elbow, and knee joints show the highest prevalences.
Comparison of the postcontact (seventeenth-century) group with late prehistoric agricul-
turalists reveals a striking increase in osteoarthritis prevalence for most articular joints. Some
of these increases are extraordinary; for example, from 16.3% to 52.9% for the male lumbar
vertebrae and 1.1% to 41.6% for the male foot. Overall, these findings suggest two significant
trends: a decrease in mechanical demand with the introduction of maize agriculture, followed
by a marked increase in mechanical demand following the arrival of Europeans. Foot
osteoarthritis shows the most pronounced increase in comparison with the other articular
joints, especially in males. The increase in degenerative pathology in comparing different joint
types is suggestive of a general and pronounced increase in workload after contact. The very
high increase for the foot suggests that these populations – especially the males – were
engaged in a type or range of activities involving pronounced mechanical demands on the
lower limb. Because the lower limb and foot function primarily in ambulatory activities (i.e.,
walking and running), the increase in contact-era foot osteoarthritis suggests that adult males
were engaged in a great deal of walking.
The changes in osteoarthritis in contact-era populations are consistent with behavioral
characteristics historically documented for the mission period in Spanish Florida. Namely,
native males were drafted into work service under the repartimiento labor system and forced
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200 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
to make long-distance trips to various localities in the province (Hann, 1988; Worth, 1995).
These trips involved carrying heavy burdens over great distances (Hann, 1988), which placed
demands on the lower limbs in walking, and on upper limbs and trunks. The general increase
in mechanical demands on native populations in the seventeenth century is also well
documented in the historical literature. The Spanish viewed native populations as an inexpen-
sive labor source. Native labor was a central element in their economic and political success in
the area. Indian laborers were used for cargo bearing, agricultural production, construction
projects, woodcutting, and a variety of other physically demanding activities (Hann, 1988;
and see Larsen, 1990a; Worth, 1995). For example, Governor Canzo remarked in his report to
the Crown in 1602–1603 that:
. . .but with all this and the grain from maize, the labor that they endure in the many cultivations that are
given is great, and, if it were not for the help of the Indians that I make them give, and they come from
the province of Guale, Antonico, and from other caciques, it would not be possible to be able to sow
any grain. . . (Unpublished translation provided by J. H. Hann; cited in Larsen, 1990a:16.)
These historical accounts strongly suggest that the increase in osteoarthritis prevalence during
the mission period was due, at least in part, to the increasing labor demands placed on native
groups at that time (see Chapter 6). Similar dramatic increases in oetoarthritis prevalence in
other regions of the New World coming under Spain’s control have been well documented
(Klaus et al., 2009; Ubelaker & Newson, 2002), changes almost certainly due to labor
exploitation (Figure 5.6).
In prehistoric New World populations that underwent the transition from foraging to
farming, there are some trends that point to different lifestyles associated with each adaptive
regime. Both in the Georgia Bight and the upper Midwest (Ohio), there is a decline in prevalence
of osteoarthritis, suggesting a change in lifestyle (Larsen & Ruff, 2011; Larsen, Crosby et al.,
2002; Sciulli & Oberly, 2002). The relatively low prevalence of osteoarthritis in the Georgia
Bight may be influenced by terrain. That is, the comparison of late prehistoric maize farmers
from upland, interior Georgia populations with coastal Georgia populations reveals higher
prevalence of osteoarthritis in the uplands groups (Larsen et al., 2007; Williamson, 2000).
In summary, based on the osteoarthritis evidence, there was an apparent decline in physical
demands with the transition to agriculture in northwestern Alabama and coastal Georgia. It is
important to point out that the Alabama and Georgia populations may not be directly
comparable. There is a large temporal gap between the early (foragers) and late (farmers)
prehistoric groups in northwestern Alabama. The skeletal series used for making comparisons
are separated in time by about 2200 years (1000 BC–AD 1200). The osteoarthritis profile
during the period of time immediately prior to the Mississippian agriculturalists is unknown. It
is possible that a period of less intensive maize agriculture prior to AD 1200 produced levels of
osteoarthritis similar to either the foragers or the farmers. Larsen’s (1982) study of osteoarth-
ritis in the prehistoric Georgia Bight also involved the analysis of skeletal remains from
populations spanning a lengthy period; the precontact preagricultural group is comprised of
all human remains predating AD 1150. However, most prehistoric forager remains are drawn
from the 450-year period immediately preceding the adoption of maize agriculture (c. AD
700–1150). Unlike the Alabama series, there is a cultural and biological continuum with little
or no time gap separating the population groupings. The temporal differences between the
Alabama and Georgia skeletal assemblages may not be significant because subsistence
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5.3 Articular joint pathology: osteoarthritis 201
(a)
80
Foragers
70
Farmers
60
Osteoarthritis prevalence (%)
Late Mission
50
40
30
20
10
0
C-spine
T-spine
L-spine
Shoulder
Hip
Wrist
Hand
C-spine
T-spine
L-spine
Shoulder
Hip
Wrist
Hand
Males Females
(b)
80 Late pre-Hispanic
70 Postcontact
OA Prevalence (%)
60
50
40
30
20
10
0
C-spine
T-spine
L-spine
Shoulder
Hip
Wrist
Hand
Elbow
Schmorl’s nodes
Knee
Ankle
Foot
C-spine
T-spine
L-spine
Shoulder
Hip
Wrist
Hand
Elbow
Schmorl’s nodes
Knee
Ankle
Foot
Males Females
Figure 5.6 Prevalence and distribution of osteoarthritis in the Georgia Bight (a) and
Lambayaque Peruvian (b) populations, demonstrating the physiological effect of colonial
rule on local inhabitants. Note the peak in osteoarthritis in late mission versus preceding
forager and farming populations, as well as the increase from late pre-Hispanic to
postcontact populations in Peru.
reconstruction based on stable carbon isotopes for the Eastern Woodlands indicates that the
shift to maize agriculture was widespread in the region after about AD 900 (Ambrose, 1987;
Smith, 1989). Thus, the two regions are broadly comparable, at least with respect to the timing
of the introduction of maize agriculture and underlying socioeconomic factors.
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202 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
The adaptive systems represented in the Alabama and Georgia populations are different in
some other important aspects. The Alabama skeletal series represents populations who were
intensive terrestrial maize agriculturalists, whereas the Georgia series represents populations
who were maize agriculturalists, but also engaged in fishing and collecting marine resources
from local estuarine and ocean contexts. Activity differences reflecting these adaptive con-
trasts are suggested by long bone structural analysis (see Chapter 6).
It would be overly simplistic to say that the shift to or intensification of agriculture in
prehistoric North America involved a reduction in workload in native populations. For
example, comparisons of less intensive with more intensive agriculturalists from the
Dickson Mounds site, Illinois show a general increase in prevalence and severity of vertebral
osteoarthritis in the more intensive agriculturalists (Lallo, 1973). In adults (sexes combined),
the frequency increases from 39.7% in the Late Woodland period to 65.8% in the Middle
Mississippian period. Similarly, there is a much greater frequency of osteoarthritis in
Mississippian-period agriculturalists from the Averbuch site compared to foragers from the
Indian Knoll (Pierce, 1987; and see Hodges, 1989).
Much of the focus on the temporal comparisons of osteoarthritis deals with the shift from
foraging to farming. The study of osteoarthritis in relation to other changes in economic
systems has also proven highly informative. In the Santa Barbara Channel islands and
mainland Pacific coast, the focus on hunting and gathering of terrestrial resources was
replaced by intensive fishing in later prehistory (Walker & Hollimon, 1989). The latter
adaptation is especially well documented by early explorers and others who first arrived in
the region. These observations provide important perspective on the types of activities
undertaken by native populations that may have potential influence on articular pathology.
Early accounts of native groups note the presence of an elaborate fishing technology and
material culture, including such items as harpoons, fish traps, nets, and fishhooks. In addition
to fishing, shellfish were collected from rocks by the use of prybars constructed from wood or
bone. Boats made from carved planks were used for travel between islands and between
islands and the mainland. Plant foods (e.g., acorns, chia, and other seeds) were collected in
large quantities, especially on the mainland. Various roots and bulbs were extracted from the
ground with digging-sticks. Economic tasks followed a strict division of labor by gender –
men hunted and fished, and women collected plant foods and shellfish. These differences in
work activities are reflected in dietary differences between adult females and males (Walker &
Erlandson, 1986). For example, early prehistoric women have higher caries rates than early
prehistoric men, which reflects the greater consumption of cariogenic plant carbohydrates by
women (see Chapter 3).
Comparison of osteoarthritis in early and late prehistoric Indians from this setting reveals
temporal changes that are suggestive of alterations in activity and workload with the transi-
tion to a marine focused economy (Walker & Hollimon, 1989). Severity of osteoarthritis,
ranging from slight articular surface porosity to extensive marginal lipping and eburnation,
increases in the late prehistoric period, especially in the lower limb (Walker & Hollimon,
1989). Based on archaeological evidence, Walker and Hollimon (1989) speculate that the
increase in osteoarthritis may be due to increased work that involved trade, exchange, and
more pedestrian travel.
Severity of elbow and wrist osteoarthritis increased in Late period adult males, but not in
adult females in the Santa Barbara Channel Islands region. Osteoarthritis severity declined in
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5.3 Articular joint pathology: osteoarthritis 203
the shoulder and hand. These changes may be linked to an alteration in weaponry (replace-
ment of the atlatl by the bow-and-arrow) and fishing equipment (shift from harpoons to nets).
The temporal increase in forelimb osteoarthritis is perhaps related to the increased use of
canoes and fishing nets in the Late period (Walker & Hollimon, 1989). Overall, the increase in
osteoarthritis was greater for males than females. Although there are several possible explan-
ations for this trend, the greater role of men in fishing suggests that the workload increase was
greater for males than females. The net result was a decrease in difference in osteoarthritis
prevalence between men and women in the Late period (Hollimon, 1991).
Viewed in a regional perspective, these North American studies show a tremendous range of
variation in mechanical stress loads. This high degree of variability from one setting to
another suggests that at least some factors that influence osteoarthritis are dependent on
local circumstances. These regional studies serve to underscore the point that what often are
perceived as uniform adaptations – even over very large areas – are, in fact, highly variable.
Comparative analysis of osteoarthritis prevalence reveals very different patterns and mech-
anical stress levels. For example, in Australia, elbow osteoarthritis is virtually nonexistent in
east coast populations, whereas it is commonplace in the Murray River valley (Webb, 1989,
1995). This variability reflects a remarkable degree of diversity in use of the upper limb.
The results of analyses of skeletal series representing populations that shifted economic
focus from foraging to farming are available for a limited number of regions in the Old World
(e.g., Europe: Meiklejohn et al., 1984; South Asia: Kennedy, 1984). These studies indicate a
reduction in osteoarthritis prevalence, and suggest a decline in mechanical loading with the
adoption of agriculture, patterns similar to some regions of North America (e.g., coastal
Georgia).
Temporal comparison of earlier and later hunter-gatherers reveals considerable variation,
likely reflecting changes in resource utilization and lifestyle practices. Comparisons of a series
of foragers from the Lake Baikal region of eastern Siberia from the Bronze Age to the Early
Neolithic (c. 6000–1000 BC) reveal key trends, interpreted within the context of a highly
detailed archaeological record of ecology, diet, resource acquisition, and lifestyle (Lieverse,
Weber et al., 2007, 2011; Weber et al., 2010). In particular, analysis of osteoarthritis preva-
lence and patterning provides important insights into activity, workload, and mobility of
foraging populations from a climate having extensive, cold winters and short, mild summers
combined with a wide spectrum of plants and animals. For the 6000-year period, climate and
biota appear to have changed very little. On the other hand, the earlier populations consumed
more fish than the post-hiatus populations (Katzenberg et al., 2012; Lieverse et al., 2011).
Until the final period, there is a slight increase, although not statistically significant, in the
number of individuals affected by osteoarthritis, especially in males. Similarly, the prevalence
and distribution of osteoarthritis in the skeleton is broadly similar for all periods and between
sexes within each period. However, earlier males have a significantly greater prevalence of
knee osteoarthritis than earlier females and later males. These differences may reflect mobility
change over time but with no real change in level of activity, a finding consistent with
archaeological and isotopic reconstructions of mobility in this setting (Lieverse et al., 2011).
One of the most abundant skeletal records for any period and any place is the Medieval
period of the United Kingdom, representing 2635 skeletons dating to three periods, pre-
Medieval (AD 400–1050), Medieval (AD 1050–1550), and post-Medieval (AD 1550–1850)
(Roberts & Cox, 2003). No change in the distribution of osteoarthritis is found in the
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204 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
pre-Medieval and post-Medieval period (Waldron, 1995). However, there is a clear change in
the post-Medieval period relative to the Medieval period. That is, there is an increase in the
proportion of knee osteoarthritis (4.4%) and a decrease in that of hip osteoarthritis (2.9%).
Waldron (1995) suggests that the changes are so rapid that they are probably behavioral in
origin. The pattern of greater knee osteoarthritis continues to the present, suggesting
that the increased incidence of greater body weight – a hugely important predisposing
factor, at least for the knees – may have had its origin centuries ago (see discussion in Roberts
& Cox, 2003).
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5.4 Nonpathological articular modifications 205
Trinkaus’s (1975) study reminds us that it is difficult to sort out general activity levels and
specific articular joint modifications in interpreting skeletal morphology. The study of dis-
tinctive metatarsophalangeal joint articular modifications in several different contexts reveals
that habitual kneeling – in food preparation tasks or other occupational activities – can be
identified. When walking or running, a high degree of dorsiflexion at the metatarsophalan-
geal joint of the toes is sustained only momentarily. In kneeling postures where the hyper-
dorsiflexed position of the toes is sustained for extended periods, the joints develop articular
modifications reflecting these behaviors. Metatarsophalangeal alterations have been identi-
fied in late prehistoric (AD 700–1550) human remains from the Ayalan site, Ecuador (Ubela-
ker, 1979). The articulations are characterized as small extensions or facets, or both, on the
distal ends of metatarsals and the proximal ends of proximal foot phalanges (Figure 5.7). The
facets are flat with clearly demarcated proximal borders and an accompanying superior
surface bony extension extending distally from the proximal articular surface. These alter-
ations are present in about 20% of Ayalan foot bones; they are bilaterally distributed among
the first three metatarsals and first phalanx, and are far less common in the fourth and fifth
metatarsals.
Comparisons of the Ayalan series with other archaeological series (Eskimos, Hawikuh site
Zuni, Mobridge site Arikara, Late Woodland Nanjemoy, and Terry Collection Blacks and
Whites) show a great deal of variation in metatarsophalangeal articular facets (Ubelaker,
1979). The alterations occur in all of these samples, but they are most common in the Ayalan
group. The prevalence in the Hawikuh and Nanjemoy samples is 5%; the other groups have
prevalences of 2% or less. The distribution among the digits of the foot also varies between
samples. In the Ayalan series, the alterations are present on the first through fourth metatar-
sals and first proximal phalanges, whereas in the Hawikuh and Eskimo series, they are on the
second through fourth metatarsals.
Although little research has been conducted on these articular modifications, they are
found in a wide diversity of human populations. In addition to those analyzed by Ubelaker
(1979), Mesolithic and Neolithic skeletal series from the early agricultural settlement at Tell
Abu Hureyra, Syria, display metatarsophalangeal alterations in the first metatarsals (Molle-
son, 1989, 1994). In older adults, the margins of these facets are associated with degenerative
changes (osteoarthritis). At Bronze-Age Bab edh-Dhra’12, Jordan, adults display more alter-
ations in the earlier than later occupation, suggesting alteration in activity involving a change
in repetitive activity and posture (Ullinger et al., 2012). Proximal extensions of the distal
articular surfaces in many of the metatarsals are also identified in prehistoric samples from
the north coast of Rota, Mariana Islands (Hanson, 1988). The association with significant
mechanical stresses of the foot is also indicated by the co-occurrence of osteoarthritis in this
setting. Similar joint modifications of the first metatarsals are present in nineteenth-century
fur-trappers from Alberta, who in life spent long hours canoeing from one location to the
next, primarily in kneeling postures with their toes dorsiflexed (Lai & Lovell, 1992; Lovell &
Lai, 1994). Likewise, the characteristic morphology, location, and association with osteoarth-
ritis in older adults in Syria and Ecuador indicate that the metatarsophalangeal joint alter-
ations were probably produced by prolonged hyperdorsiflexion of the toes while kneeling
(Molleson, 1989, 1994; Ubelaker, 1979). With regard to the Ecuador series especially, Ubelaker
(1979) speculates that kneeling while grinding maize with stone metates was the most likely
cause. Molleson (1989) provides confirmation of a similar posture in regard to cereal grinding
depicted in Assyrian and Egyptian dynastic tomb art.
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206 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
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5.5 Nonarticular pathological conditions relating to activity 207
(a)
Figure 5.8 Entheseal changes on the medial epicondyle of the humerus (a), radial
tuberosity of the radius (b), and attachments for infraspinatous and teres major muscles.
(Photographs by Sébastien Villotte.) (A black and white version of these figures will
appear in some formats. For the color version, please refer to the plate section.)
2006). Various skeletal elements are affected, but enthesial changes are especially common-
place for the humerus, radius, ulna, tibia, femur, metacarpals, metatarsals, and terminal
phalanges (Bufkin, 1971; Edgerton et al., 1990; Owsley et al., 1991).
Although referred to in the literature as musculoskeletal stress markers, cortical defects, or
other synonyms (Brower, 1977; Bufkin, 1971; Edgerton et al., 1990; Owsley et al., 1991;
Resnick & Greenway, 1982), the irregularities on the outer margin of the joint and/or other
characteristics (e.g., foramina) are called enthesial changes (Villotte & Knüsel, 2013)
(Figure 5.8). Although the link between form and function is debated, it is clear that enthesial
changes provide a potentially important source for inferring patterns of activity (Dutour,
1986; Hawkey, 1998; Hawkey & Merbs, 1995; Ibanez-Gimeno et al., 2013; B. Kennedy, 1989,
1998; Niinimäki, 2012; Ruff et al., 2006; Schlecht, 2012; Stirland, 2002; Villotte et al., 2010a,
2010b; Weiss, 2007; various in Henderson & Cardoso, 2013).
This body of work was motivated by a set of investigations undertaken by Angel and his
collaborators, beginning in the 1950s, who were interested in reconstruction of physical
activity from analysis of skeletal morphology (Angel, 1952; Angel et al., 1987; Kelley &
Angel, 1987). There is now an extensive record of research focusing on enthesial changes,
much of which is built on the influential study of Hudson Bay Inuit skeletal remains where
robusticity markers (degree of muscle markings), stress lesions (pitting or furrows), and
ossification exostoses (bony spurs) for each muscle attachment site were documented by four
well-defined grades (Hawkey & Merbs, 1995). The results of this and other investigations
suggested patterns of variation that reflected muscle use and activity involving specific joints.
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208 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
(b)
Most investigations following Hawkey and Merbs’s (1995) study use this method or its
derivations (Eshed et al., 2004; Hawkey, 1998; Lieverse et al., 2009, 2011; Molnar, 2006, 2008;
Molnar et al., 2011; Perry, 2008; Peterson, 1998, 2002; Steen & Lanes, 1998; Weiss, 2007).
Approaches to data collection based on understanding of the types of entheses – fibrous and
fibrocartilaginous – provide a more biologically meaningful assessment of enthesial changes
in relation to function and activity (Schlecht, 2012; Villotte & Knüsel, 2013; Villotte et al.,
2010a, 2010b). These two types of entheses have biomechanical and histological differences, a
factor not recognized in most earlier studies using form to infer function. Villotte and
coworkers (2010a, 2010b) propose focusing on fibrocartilaginous entheses, both in order to
understand the underlying etiology to better inform behavioral inference and to avoid
conflating different tissue types and drawing generalizations about activity.
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5.5 Nonarticular pathological conditions relating to activity 209
(c)
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210 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
example, an increase in markers for pectoralis major in comparing Early and Late Thule males
suggests an increase in downward thrusting, such as that associated with harpooning
seals. Ethnographically, this thrusting motion targets the seal as it comes to its breathing
hole in the ice.
Musculoskeletal functional analysis of entheses in other settings where archaeological,
ethnographic, historical, and other contextual evidence is available reveals important behav-
ioral and lifestyle interpretations. On the island of Gotland (Sweden; see Chapter 4), where
Neolithic (3400–2300 BC) coastal populations focused their diets on fish and seals from the
Baltic Sea, markers are higher in males than females and in older adults compared to younger
adults (Molnar, 2006, 2008). The common trend for many of the investigations of enthesial
changes suggests that much of the variation can be explained by accumulation of modifica-
tions with advancing age. This makes the strong case that age must be accounted for when
documenting enthesial changes for reconstructing and interpreting activity patterns (Weiss,
2007).
In contrast to the clear age progression of enthesial changes in the series from Gotland,
Sweden, there are clear age changes, but for females only, in the later post-hiatus series from
Lake Baikal, Russia (Lieverse et al., 2009, 2011). Moreover, there is an increase in the severity
of upper limb changes in the post-hiatus males. These patterns of general increase, especially
in males, are consistent with the notion that later-period populations were involved in
hunting forays, in particular hunting large terrestrial mammals (e.g., ruminants). Interest-
ingly, in both males and females, markers for conoid and costoclavicular ligaments and
pectoralis major, deltoid, pronator quadratus, and brachioradialis were quite pronounced,
regardless of the period.
There is considerable debate about the foraging-to-farming transition in the Levant,
especially in the region encompassing the modern country of Israel (Hershkovitz & Galili,
1990). In order to address the question, did the transition to farming involve reduced
workload? Eshed and collaborators analyzed upper limb markers for four pre-farming (Natu-
fian) and four early farming (Neolithic) skeletal series. Based on the consistent increase in
scores, the results of their study suggest a clear increase in mechanical stress and loading of
the upper limbs. Males generally show greater scores than females in both periods. However,
Natufian females express higher scores than males for the deltoid attachment and show a clear
dominance of right over left side. Eshed and coworkers (2004) suggest that this reflects the
elevated use of the forearm muscles, perhaps in pounding of grain with a stone or wood pestle,
but likely other activities requiring heavy use of the arm in a range of work activities. In her
analysis of changes in the foraging-to-farming transition in the Levant, Peterson (2002)
found a distinctive Natufian pattern of high scores in adult male upper limbs, especially for
the right triceps brachii and anconeus, muscles involved in forearm extension. This kind of
movement is associated with activities in overhand throwing motions, such as for atlatl use
and spear throwing. Some of the individuals were young adult males, suggesting that the
pattern is not just an age-related phenomenon. On the other hand, both males and females
show similarities in scores, such as for pectoralis. This suggests overlapping patterns of upper
limb use for males and females. In sharp contrast, the Neolithic adults show a very different
pattern than the Natufian adults. In particular, Neolithic adults show a distinctive symmetry
in scores comparing right and left sides for both males and females, suggesting bilateral tasks
in the use of the upper limbs. In addition, the scores for the brachialis – a forearm flexor – are
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5.5 Nonarticular pathological conditions relating to activity 211
much more pronounced in the Neolithic males and Natufian males. Overall, the pattern of
changes shows fewer sex differences and more emphasis on bilateral activities with the shift
to and intensification of farming. The archaeological context shows emphasis on farming
activities, including field preparation, harvesting, and food preparation.
In contrast to the symmetry of enthesial changes in the upper limbs in early farmers in the
Levant, prehistoric males from Europe express greater frequency of enthesial changes in the
right elbow (humeral epicondyles) than in the left elbow (Villotte & Knüsel, 2014). This strong
degree of asymmetry is consistent with an extensive record of articular injuries resulting from
overhead throwing using the right arm, a largely male-based activity pertaining to use of the
atlatl and other projectile technology.
Analysis of a large agricultural-based population from Grasshopper Pueblo, Arizona
(Pueblo IV period: AD 1275–1600) expresses a very different kind of pattern in comparison
with the agriculturalists from the Levant and some other contexts (Perry, 2008). In this setting,
males show strong unilateral emphasis on muscle groups that are involved in movement of
the arm and shoulder, stabilization of the chest and shoulder, and flexion of the arm. The
right–left asymmetry with right dominance is especially pronounced for the right deltoid and
costoclavicular ligaments in males, and at considerably higher levels than for females. There
are various activities that could explain this highly distinctive pattern in the late prehistoric
Southwest. However, in the context of behavioral reconstruction based on archaeological,
ethnohistorical, and ethnographic evidence, the most likely male activity includes use of the
bow-and-arrow in large game hunting and in its masculine identity. The distinctively bilateral
development of changes in the upper limb in females likely reflects corn grinding and other
domestic activities associated with food preparation. On the other hand, females show a
distinctive unilateral development of the transverse head of the adductor pollicus, a muscle
that brings the thumb to the center of the palm of the hand, such as that associated with
scraping ceramics, using a scraping tool held in the right hand during production. Similarly,
analysis of War of 1812 battle casualties from Snake Hill, Ontario reveals an unusually high
frequency (40%) of enthesial changes on proximal humeri, significant higher than in other
series, such as Civil War casualities and Native American hunter-gatherers and agricultural-
ists (Owsley et al., 1991). Moreover, there is significant right dominance (62.5%) in the Snake
Hill series; if bilaterally present, the defect on the left side is small and shallow compared to
the much larger defect on the right side. This pattern of variation likely reflects right hand
dominance.
Overall, therefore, sex and age patterns of enthesial changes in archaeological settings are
highly consistent: males generally express higher prevalence, severity, and right-dominated
upper limb articular injury than females, which can broadly be interpreted to mean that males
have greater and more intensive physical activity and muscle use than females. Similarly,
with few exceptions, severity increases with age, and the most severe forms of enthesial
changes are associated with elderly adults (Villotte et al., 2010a, 2010b).
One would expect to find evidence for a relatively greater workload and demanding
lifestyle in low-status individuals who had been exploited as a labor source. Indeed, this
expectation is met with regard to exploitation of native populations by Spaniards during the
colonial era in the American Southwest (Chapman, 1997) and in a Medieval-era setting in
east-central Europe (Havelková et al., 2011). In both settings, individuals exposed to a high-
level mechanical demand express elevated prevalence of enthesial changes.
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212 Activity patterns: 1. Articular degenerative conditions and musculoskeletal modifications
Although analyses of enthesial variation from bioarchaeological settings are highly sug-
gestive, there remains very little direct evidence for a relationship between entheses variation
and activity, either specifically involving a particular muscle group or general level of degree.
Most of the aforementioned studies assume that activity is responsible for increasing perios-
teal apposition in those areas of the skeleton associated with muscle tendon insertions
(Schlecht, 2012). That is, bioarchaeologists have extrapolated from the findings of experi-
mental skeletal biologists (Chamay and Tschantz, 1972; Woo et al., 1981) who have docu-
mented the direct link between activity increase and periosteal apposition in tendinous
regions of the skeleton. Applying these findings, bioarchaeologists argue that the periosteum
associated with muscle sees increases in periosteal capillary volume with an increase in
muscle contraction. This causes the periosteum in the region associated with the muscle to
hypertrophy, thereby increasing the strength of the tendon-anchoring sites (Hawkey & Merbs,
1995; Torg et al., 1972; Zumwalt, 2006). Schlecht (2012) notes, however, that while the work
of experimental skeletal biologists has documented some key insights, they may not be
applicable to understanding the etiology and derivation of entheses. Importantly, little work
has been done on enthesial changes, documenting variation in loading in the regions
involved. Analysis of enthesis morphology of exercised sheep compared with non-exercised
sheep in an experimental context revealed no linkage between enthesis morphology and
exercise (Zumwalt, 2006). On the other hand, histomorphological analysis of the proximal
radius in a human cadaver sample of known age and sex revealed evidence of increased bone
turnover, suggesting a potential relationship between enthesial changes and mechanical
strain (Schlecht, 2012). This record – bioarchaeological, experimental, and anatomical –
indicates that enthesis morphology, while likely linked to activity, is complex and incom-
pletely understood.
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5.6 Summary and conclusions 213
The study of enthesial changes indicates that they are likely related in some manner to
activity and lifestyle, providing an important avenue of inquiry for interpreting behavior as it
relates to activity and lifestyle. Additional exploration, especially through histological analy-
sis, experimental investigation, and covariance with other indicators of activity (e.g., cross-
sectional geometry), have the potential to provide a much clearer understanding of the
adaptive responses to a diversity of activity regimens and variation by sex, age, body size,
and labor intensity (Ibanez-Gimeno et al., 2013; Jurmain, 1999; Niinimäki, 2009; Pearson &
Buikstra, 2006; Schlecht, 2012; Stirland, 1998; Villotte et al., 2010a, 2010b; Weiss, 2007;
Zumwalt, 2006, Zumwalt et al., 2000).
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6 Activity patterns: 2. Structural adaptation
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6.2 Cross-sectional geometry 215
exercise tend to have higher bone mass and size than adults who are relatively sedentary,
especially in males (Lorentzon et al., 2005).
Loss of normal functional loading, especially involving extended periods of bed rest,
weightlessness in spaceflight and absence of gravitational loading, or from partial or complete
immobilization of limbs, results in decreased bone mass (Abram et al., 1988; Holick, 1998;
Jenkins & Cochran, 1969; Kazarian & Von Gierke, 1969; Kiratli, 1992; Lanyon & Rubin, 1984;
Lazenby & Pfeiffer, 1993; Meade, 1989; Miyamoto et al., 1998; Morey & Baylink, 1978; Peck &
Stout, 2009; Prince et al., 1988; Sevastikoglou et al., 1969; Todd & Barber, 1934; Whalen
et al., 1988; see discussion in Trinkaus et al., 1994). The loss in mass is largely due to profound
reduction in remodeling rates of bone whereby the normal linkages between osteoclasts and
osteoblasts are decoupled (Alexandre & Vico, 2011; Schlecht et al., 2012). This dysfunctional
replacement process leads to more removal, especially endosteally, than replacement of bone,
and ultimately, considerable reduction in bone density and mass.
Bone is anisotropic in that it is characterized by different material properties depending
upon the direction of loading (Cowin, 1989; Currey, 2002; Nordin & Frankel, 2001;
Wainwright, 1988). Long bones, for example, are stronger in the longitudinal direction than
in any other plane (Wainwright, 1988). The primary loading forces affecting bone include
tension, compression, shearing, bending, and torsion (Figure 6.1). These can be best understood
by considering a thin section, or slice of bone. Tensile loading occurs when equal and opposite
forces are applied outwardly from the surface of the slice. Compression, the opposite of tension,
occurs when equal and opposite loads are directed toward the two surfaces. Shearing loads
involve the application of forces parallel to the surfaces under consideration. Bending forces
produce two types of stresses, tension on the convex side and compression on the concave side.
Torsional loading is the twisting of the skeletal element about an axis and results in a
combination of tension, compression, and shear forces. Individual skeletal elements have an
irregular geometric structure, and a range of forces usually acts on bone in normal physio-
logical activities, such as running and walking. Therefore, loading almost always involves a
combination of these modes, but the largest and most common loading parameters affecting
the human skeleton are bending and torsion, especially for the long bones.
This extensive body of research makes clear that there is strong support for inferring patterns
of activity from skeletal morphology. However, it must be kept in mind that there are other
factors that influence morphology. There are genetic influences, and skeletal form represents
compromises between mechanical and other influences, such as nutrition, hormones, and age
(Gosman et al., 2011; Nelson et al., 2004; Ruff, 2008; and see Chapter 2). This chapter takes the
position that mechanical loading, as it relates to lifestyle and habitual activity, plays the leading
role in explaining robusticity and its variation (Daly et al., 2004; Martin, 2007; Robling et al.,
2002; Stock, 2006). In the larger picture, this record reveals the fundamental importance of
adaptive plasticity, or the ability to respond to environmental circumstances during the years of
growth and development (Gotthard & Nylin, 1995; Rovosa et al., 2008; and see Chapter 7).
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216 Activity patterns: 2. Structural adaptation
Currey, 2002; Ruff, 2008). Unlike straight mechanical analysis of building materials, biomech-
anics deals with dynamic tissues that modify themselves continuously in relation to loading
modes and activity levels. That is, building materials do not adapt and alter their form in
response to the mechanical environment. Only dynamic, living tissues adapt and alter their
form in response to the mechanical environment.
Density of bone tissue differs within the skeleton and within individual bones in response to
varying mechanical demands. As such, mineral content is a component of bone strength
(Burr, 1980; Martin et al., 1998). However, the response to increased loading is primarily in the
distribution of bone rather than in its density or any other intrinsic material property (Beck
et al., 1990; various in Burr & Allen, 2013; Burr et al., 1989; Gosman & Ketcham, 2009;
Grynpas, 2003; Ruff, 2008).
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6.2 Cross-sectional geometry 217
Borrowing the simple beam model used by civil and mechanical engineers to analyze
buildings and design structures (Huiskes, 1982; Timoshenko & Gere, 1972; Turner & Burr,
1993), biological anthropologists and others have analyzed stresses – the pressures on the
inside of materials – that result from loads applied externally. These stresses have been
calculated via analysis of cross-sectional geometric properties in various settings, including
archaeological (Bridges, 1989b; Bridges, Blitz et al., 2000; Larsen & Ruff, 1994; Lieverse et al.,
2011; Maggiano et al., 2008; Ruff & Hayes, 1983a, 1983b; Ruff & Larsen, 2001; Stock &
Pfeiffer, 2001; Stock et al., 2011), fossil hominins (Churchill & Smith, 2000; Holt, 2003; Holt &
Formicola, 2008; Trinkaus & Ruff, 1999; Trinkaus et al., 1994), and non-human primates (Burr
et al., 1989; Kimura, 2002; Polk et al., 2000; Ruff & Runestad, 1992). These investigations, as
well as experimental evidence based on laboratory animals (Abram et al., 1988; Simkin et al.,
1989), indicate the value of structural analysis in drawing inferences about physical activity
and behavior patterns. However, experimental research also suggests that mechanical loading
of long bones is an approximation of the loading of simple beams (Lieberman, Polk et al.,
2004), but the large and growing body of work on humans and animals indicates that cross-
sectional geometric analysis is the best available source for understanding variation in
mechanical loading and integrity (Ruff, 2008; Ruff et al., 2006; Turner & Burr, 1993).
Stresses are expressed as two specific characteristics: rigidity (the ability to resist bone
deformation during loading) and strength (the ability to resist structural failure or fracturing
during loading). Both characteristics are central to understanding skeletal adaptation. In
bending and torsion of a hollow beam, such as a long bone, the magnitude of stresses is
proportional to the distance from the central or “neutral” axis of the bone (Figure 6.2). The
neutral axis is the plane (bending) or axis (torsion) where stress is zero. Thus, all else being
equal, the cross-section that is strongest is that in which the material is oriented furthest from
the neutral axis (Currey, 2002; Nordin and Frankel, 2001; Ruff, 2008). Additionally, by
inference, the greater the distance from the neutral axis, the greater the magnitude of stresses
to which the bone has adapted over time (Nordin & Frankel, 2001; Ruff, 2008; Wainwright,
1988). In long bones and in some other elements, especially tubular-shaped bones (e.g.,
metacarpals), the cross-sectional area and the manner in which the bone is distributed about
its long axis reflect mechanical/functional behavior.
Bending a ruler is a good analogy for demonstrating the principles associated with beam
analysis. If one attempts to bend a ruler against its narrow edges, there is little or no give. In
contrast, if forces are applied virtually anywhere along its flat surface, especially toward the
middle, the ruler bends readily. From a mechanical perspective, the small amount of give
when applying bending forces to the narrow edges occurs because the materials in this axis
are distributed relatively far from a central, neutral axis. Thus, in this plane of bending, the
ruler has a great deal of resistance. Conversely, the ease of deformation when applying
bending forces to the flat surface is made possible by the lack of material far from the neutral
axis; thus there is very little resistance to bending of material when the ruler is subjected to
bending forces in this direction. A ruler is structured so as to resist bending from one direction
only. Given the tubular shape of long bones, they are able to efficiently withstand mechanical
demands associated with bending and torsion from various directions, although as noted
later, some bone sections are more adapted to loads applied from particular directions.
Beam analysis involves the measurement of geometric properties from cross-sections taken
perpendicular to the long axis of a skeletal element. As demonstrated by the ruler analogy, these
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218 Activity patterns: 2. Structural adaptation
(a)
NEUTRAL AXIS
(b)
NEUT
RAL
AXIS
Figure 6.2 Cross-section of a bone undergoing bending (a) and torsion (b), and showing
stress distribution around the neutral plane and axis, respectively. Note that the magnitude
of forces (indicated by heavier arrows) is greatest at the periphery of the bone and least
nearest the neutral plane or axis. (From Nordin & Frankel, 2001; reproduced with
permission of authors and Williams & Wilkins.)
properties are based on both the amount and the distribution of bone tissue in the cross-section.
As such, they are direct measures of the aforementioned rigidity and strength of the bone cross-
section. Therefore, the properties measured in archaeological human bones should represent a
measure of the cumulative forces operating on the skeletons of individuals during their lifetimes.
Cross-sectional geometric properties measure the amount and the distribution of skeletal
tissue in a section. For measuring rigidity, these properties include section “areas” and
“second moments of area” (or “area moments of inertia”) (Figure 6.3). Areas include total
subperiosteal area (TA), endosteal or medullary area (MA), and cortical area (CA). Measure-
ment of mediolateral (ml) and anteroposterior (ap) breadths of long bone diaphyses can be
used to calculate areas using formulae for a cylinder (Ruff & Jones, 1981):
TA ¼ π(Tap/2)(Tml/2)
MA ¼ π(Map/2)(Mml/2)
CA ¼ TA – MA
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6.2 Cross-sectional geometry 219
Medial-lateral
bending
Ix
Anterior-posterior
bending
Iy
Femur midshaft
Figure 6.3 Cross-section of the femur midshaft and associated geometric properties.
Second moments of area about the x- and y-axes (Ix and Iy) are associated, respectively,
with anterioposterior bending and mediolateral bending of the femoral midshaft.
where T is the total diameter of the cross-section and M is the medullary diameter. However,
this technique only works in bone regions with relatively regular, circular cross-sections, and
even there it is subject to error compared to more direct measurements (O’Neill & Ruff, 2004).
Cortical area is a measure of the amount of cortical bone in a cross-section and is also an
indicator of strength of the long bone diaphysis under pure axial loading (loading that is
simultaneously applied to both ends of the bone along its long axis). The percentage of
cortical area (PCCA) where PCCA ¼ (CA/TA) 100 is an alternative indication of the amount
of compact cortical bone, but it has no direct mechanical significance (Ruff, 2008). Cortical
area and percentage of cortical area provide very different representations of bone mass
owing to the fact that the latter, but not the former, measures cortical bone relative to total
subperiostial area (Ruff, 2008). Total subperiostial area and medullary area are measurements
of the two major surfaces of the bone cortex, including the outer periosteal and inner
endosteal surfaces, respectively (Ruff, 1991). Expansion in total subperiostial area and medul-
lary area indicates a greater distribution of skeletal tissue further from the neutral axis of
the bone.
Bone areas, especially cortical area, are proportional to strength in compression and tension
when the forces are applied non-eccentrically along the central longitudinal axis of the bone
diaphysis (axial loading). Because long bones are curved and are affected by muscular forces
applied off-center and at angles to the longitudinal axis, pure axial loadings in either
compression or tension are rare; most forces involving long bone diaphyses are eccentrically
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220 Activity patterns: 2. Structural adaptation
applied, thus create bending and/or torsion. Second moments of area have been shown
generally to be more accurate indicators of bone rigidity and mechanical function than areas
alone. For example, analysis of second moments of area in second metacarpals from
the dominant hand in a large sample of urban Americans (n¼992) reveals that in both left-
handed and right-handed individuals the metacarpals in the dominant hand have signifi-
cantly greater second moments of area than metacarpals in the nondominant hand (Roy et al.,
1994). Increased bone strength is not the result in this case of greater cortical thickness.
This finding underscores the point that cortical thickness by itself is not an appropriate
indicator of functional/mechanical demand (Ruff, 1992; Ruff & Larsen, 1990; Trinkaus &
Ruff, 2012).
Second moments of area are geometric properties that are used to measure bending and
torsional rigidity. Bending rigidity, values of which are called “I” with a subscript that
references the specific axis running through the cross section (e.g., Ix), is calculated in relation
to the neutral axis. Typically, diaphyseal geometry is assessed at specific percentages of length
of the bone. For example, the 50% section for the femur refers to the diaphyseal geometry at
the midshaft (Figure 6.3).
The general formula for calculating I in relation to a particular axis is:
X
I¼ ai d i 2
i
where ai is the unit area and di is the perpendicular distance from the center of the unit to the
neutral axis. Ix refers to the bending rigidity in the anteroposterior plane and Iy to bending
rigidity in the mediolateral plane (Ruff, 2008). Other values of I expressing the maximum and
minimum bending rigidity in a cross-section are referred to as Imax and Imin, respectively,
where Imax measures the maximum rigidity in resistance of bone to bending and Imin measures
the minimum rigidity in resistance of bone to bending. Torsional rigidity is calculated with
reference to the neutral center or “centroid” of the cross-section and is called the polar second
moment of area or “J.” J is equal to the sum of the values of Imax and Imin (or Ix and Iy), which
are always perpendicular to each other (Ruff, 2008). J is proportional to torsional rigidity and
the average bending rigidity in any two perpendicular planes. Therefore, it represents a good
measure of overall rigidity (Ruff, 2008).
Values of bending rigidity (I) and second moment of area (J) are calculated as products of
very small unit areas in the cross-section and squared distances of the unit areas relative to
the neutral axis (for values of I) or the neutral center of the section (for values of J). Therefore,
second moments of area are presented as linear dimensions to the fourth power. Because of
variability in body size or length of long bones in comparison with different population
samples or temporal series within a particular setting, properties should be size-standardized
when making comparisons between or within groups (Ruff, 2008). Earlier work suggested that
for the femur, dividing areas by length2 and second moments of area by length4 was an
appropriate means of size standardizing (Ruff, 1984; Ruff & Larsen, 1990). More recent
analyses utilizing additional and more extensive samples of extinct and recent humans
suggest that more appropriate powers for size standardization of the femur are bone length3
and bone length5.33 for areas and second moments of area, respectively (Ruff et al., 1993).
Lastly, development of estimates of body mass via femoral head breadth provides another
method of size standardization (Trinkaus & Ruff, 2012).
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6.2 Cross-sectional geometry 221
Cross-sectional geometric properties used to measure bending and torsional strengths are
slightly different from those used to measure rigidities. Maximum stresses occur on the outer
surface of the bone cross-section. Thus, strength cross-sectional properties (or section moduli)
are calculated by dividing the second moments of area described for rigidity by the distance
from the bone surface to the neutral axis. Section modulus values can also be estimated by
taking the rigidity values to the power of 0.73 (Ruff, 2008). These section moduli properties for
measuring strength are designated as values of “Z” with the subscript describing the plane
(e.g., Zx), with torsional strength expressed as the value Zp, representing the sum of values
perpendicular to each other and representing a measure of torsional loading. Regardless of
which are analyzed, rigidities or strengths, both variables present a record for interpreting the
same basic components of mechanical loading, with each reflecting relative patterns of
mechanical demand in past populations. Indeed, some refer to strength and rigidity under
the general term of strength, even though technically they relate to the specific meanings
described here.
Cross-sectional geometric analysis was first applied to samples of greater than a single bone
in a variety of settings involving both human and non-human primates (Jungers & Minns,
1979; Kimura, 1971, 1974; Klenerman et al., 1967; Lovejoy et al., 1976; Martin & Atkinson,
1977; Miller & Piotrowski, 1977; Minns et al., 1975; Piziali et al., 1976). These studies were
generally limited to fewer than 10 individuals because of the lengthy and tedious process
involved in manually calculating geometric properties. Specifically, two problems with
calculating section properties are the determination of endosteal and periosteal boundaries
and the mathematical integration of areas that are required for the calculations (Ruff, 2008).
The development of automated protocols for computer analysis of large numbers of cross-
sections (Nagurka & Hayes, 1980) and new technologies have made it possible to carry out
studies involving more extensive samples. These developments have fostered a more compre-
hensive understanding of variability both within and between populations from archaeo-
logical and paleontological contexts (Bridges, 1989b, 1991a, 1995; Bridges, Blitz et al., 2000;
Brock, 1985; Brock & Ruff, 1988; Churchill, 1994; Churchill & Smith, 2000; Holt, 2003; Holt &
Formicola, 2008; Kimura & Takahashi, 1982; Larsen & Ruff, 1991, 1994; Larsen, Ruff et al.,
1995; Larsen et al., 2007, 2013; Lieverse et al., 2011; Maggiano et al., 2008; Rhodes & Knüsel,
2005; Robbins et al., 1989; Ruff, 1987, 1989, 1991, 1994b, 2010a, 2010b; Ruff & Hayes, 1982,
1983a, 1983b; Ruff & Larsen, 1990, 2001; Ruff et al., 1984, 1991, 1999; Stock et al., 2011;
Stock & Pfeiffer, 2001; Sumner et al., 1985; Trinkaus et al., 1994; Van Gerven et al., 1985).
The method of geometric analysis involves the preparation of an image of a section made
perpendicular to the longitudinal axis of the bone. This image is obtained through several
alternative means, from existing breaks on long bone diaphyses (Lovejoy & Trinkaus, 1980),
direct cutting (Larsen & Ruff, 1994; Ruff & Hayes, 1983a, 1983b; Ruff et al., 1984), or
noninvasive imaging, especially computed tomography (CT) (Brock & Ruff, 1988; Bridges,
1989b; Larsen et al., 2013; Larsen, Ruff et al., 1995; Ruff, 1987, 1999), or a combination of
methods (Larsen et al., 2013; Ruff & Larsen, 2001). Other useful noninvasive techniques
include biplanar radiography (Biknevicius & Ruff, 1992; Fresia et al., 1990; O’Neill & Ruff,
2004; Runestad et al., 1993; Trinkaus & Ruff, 1989) and photon absorptiometry (Martin &
Burr, 1984; Van Gerven et al., 1985; see discussion of imaging techniques by Ruff, 1989,
2008; Ruff & Leo, 1986). Noninvasive imaging is useful in situations where cutting of bone
specimens is not possible (e.g., fossil hominins). Unlike invasive cutting, the advantage of
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222 Activity patterns: 2. Structural adaptation
noninvasive analysis is that section properties can be determined directly from the images
(e.g., CT scan) using a variety of software. More importantly, noninvasive methods leave the
bone intact.
Accuracy of geometric analysis is dependent upon the availability of fully intact periosteal
and endosteal section contours. In addition to well-preserved periosteal and endosteal sur-
faces, the accuracy of analyses – including size (length) standardization as well as precise
location of sections – requires the presence of largely intact ends of long bones. Therefore,
even though large numbers of skeletons may be present in an archaeological series, if
differential preservation exists, then only subsamples composed of well-preserved specimens
can be analyzed (Larsen et al., 2013; Ruff, 2010a, 2010b; Ruff et al., 1984).
There are various ways of obtaining landmark data for section analysis. Once the section
image is obtained, a magnified photographic slide image can be projected onto a digitizer
screen, and the periosteal and endosteal borders then manually digitized or scanned using an
optical scanner and directly input to a desktop or laptop computer. Automated computer
programs using engineering principles based on standard algorithms are widely available
(Nagurka & Hayes, 1980; Sylvester et al., 2010) for calculating section properties.
In summary, studies of cross-sectional geometric properties address a range of long-
standing issues in anthropology dealing with activity and physical behavior, especially in
regard to subsistence strategies and tool and weapon use, the relationship of sex differences to
dietary and subsistence adaptation, and variability in skeletal growth and development in
response to environmental and dietary change.
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6.2 Cross-sectional geometry 223
CA CA
150 150 150 CA
50 50 50
Georgia Ag.
Georgia Ag.
Georgia Ag.
Pecos
Pecos
Georgia Preag.
Georgia Preag.
Georgia Preag.
Stillwater
Stillwater
Stillwater
Gr. Pl. Coal
Gr. Pl. Precoal
Figure 6.4a Comparisons of cross-sectional areas – total subperiosteal area (TA), cortical
area (CA), and percentage of cortical area (PCCA) – for three long bone sections. Note that
for each graph the foragers are at the left (Stillwater at far left) and farmers are at the right.
Areas are standardized over bone length2 and multiplied by 105. (From Larsen, Ruff et al.,
1995; reproduced with permission of American Museum of Natural History.) Cultural units:
Stillwater (1300 BC–AD 1300); Georgia Preagricultural (500 BC–AD 1150); Great Plains
Precoalescent (AD 400–1600); Pecos Pueblo (AD 1300–1650); Great Plains Coalescent (AD
1600–1850); and Georgia Agricultural (AD 1150–1550). Filled squares, males; open
squares, females.
in upper and lower limbs in a harsh desert setting (Larsen, Craig et al., 1995, Larsen, Ruff
et al., 1995, 2008; Ruff, 1999, 2010a, 2010b). Comparisons of percentage of cortical area
(PCCA), a relative measure of bone mass, with other North American archaeological series
reveal that the Stillwater group (but not Malheur or Great Salt Lake) is on the low end of a
continuum (Figure 6.4a). In sharp contrast, total subperiosteal area and J are remarkably
high in the Great Basin generally. These findings indicate that there is a relatively low
amount of femoral cortical bone for the Stillwater population but the bone that is present is
distributed far from the neutral axes relative to that for many other populations, indicating
very high bending and torsional strength. Humeral cortical area is generally reduced in all
three Great Basin samples, possibly a reflection of dietary insufficiency. As a whole, the Great
Basin samples have relatively low percentage of cortical area, that is, thin cortices, compared
to Native Americans from other regions (Ruff, 1999, 2010a), which may reflect the combined
effects of systemic stress (relatively poor diet) and a high level of physical activity.
Females and males from these various North American settings show somewhat different
patterns of skeletal morphology (Larsen, Craig et al., 1995, Larsen, Ruff et al., 1995, Larsen
et al., 2008; Ruff, 1999, 2010a). In males, torsional rigidity (J) in the femoral midshaft closely
parallels the subsistence strategy – hunter-gatherers tend to be high (e.g., Great Basin, Georgia
preagricultural), and conversely, agriculturalists tend to be low (e.g., Pecos Pueblo). This trend
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224 Activity patterns: 2. Structural adaptation
Georgia Ag.
Georgia Ag.
Georgia Preag.
Georgia Preag.
Georgia Preag.
Pecos
Pecos
Stillwater
Stillwater
Stillwater
Gr. Pl. Coal
Gr. Pl. Precoal
follows the rationale for a decline in mechanical loading of the femur in agriculturalists in
comparison with hunter-gatherers, as a more sedentary lifeway associated with farming
emerged and intensified. This is not to say that foragers are subjected to high levels of
mechanical loading and agriculturalists are not. Rather, these biomechanical studies suggest
tendencies in patterns of mechanical loading from high levels in foragers to relatively lower
levels in agriculturalists, likely associated with different degrees of mobility.
Torsional loading bears no apparent relationship to subsistence strategy in females in these
North American groups. Rather, female torsional loading corresponds with degree of rugged-
ness of terrain – mountainous populations (Great Basin, Pecos Pueblo) have the highest values
of J, coastal populations have the lowest values (Georgia coast), and Great Plains populations
are intermediate (Figure 6.5).
Humeri show a somewhat different pattern of area and second moments of area than
femora. Especially striking are the low values of humeral cortical area and second moment of
area (J), notably in the Stillwater Great Basin series (Larsen, Ruff et al., 1995; Ruff, 2010a).
This finding is consistent with the hypothesis that mechanical loading is localized in the
skeleton (Lanyon & Skerry, 2001; Macdonald et al., 2009; Peck and Stout, 2009).
Thus, the generally high level of robusticity in Great Basin adults – especially in femora – is
consistent with findings based on osteoarthritis prevalence (and supporting a limnomobile
subsistence strategy; see Chapter 5). Behaviors causing osteoarthritis and elevated robusticity
result in respective high frequencies of joint degeneration and high second moments of area.
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6.2 Cross-sectional geometry 225
550
500
450
400
350
Mountains Plains Coastal
Terrain
Figure 6.5 Torsional rigidity correlates with terrain, with mountainous populations (Great
Basin and Pecos Pueblo) showing markedly higher relative femoral strength (J) than coastal
populations (Georgia Bight). (Ruff, 2008; reproduced with permission of author and John
Wiley & Sons, Inc.)
Although Great Basin foragers from all three settings show quite high levels of skeletal
robusticity, for the lower limbs, some populations from the region of the Great Basin pursued
a maize farming strategy. That is, stable carbon isotope analysis of the Great Salt Lake later
prehistoric Fremont (AD 400–1150) shows an increasing reliance on maize (Coltrain &
Stafford, 1999; and see Chapter 8). Interestingly, individuals having a strong isotopic signa-
ture for maize consumption have generally lower skeletal robusticity (J) for both femora and
humeri compared to individuals who have a weak isotopic signature for maize consumption,
with the pattern more pronounced for males (Ruff, 1999) (Figure 6.6). These findings show a
clear link between dietary regime, behavior, and their combined influence on skeletal
morphology.
Subarctic hunter-gatherers
Populations inhabiting the Aleutian Islands, Alaska, from the mid-eighteenth century to the
present, also led physically demanding lifestyles. An especially demanding task was kayaking
on the open ocean (Laughlin et al., 1991, 1992; and see Chapter 5). This activity underlies the
extreme external robusticity of Aleut humeri (Hrdlička, 1945) and high levels of osteoarthritis
and musculoskeletal stress markers in these populations (Hawkey & Street, 1992; and see
Chapter 5). Comparisons of Aleuts with other populations in humerus torsional rigidity reveal
that these subarctic peoples surpass values derived from a range of modern groups (Churchill,
1994; Ruff et al., 1993) (Table 6.1). These elevated levels of bone rigidity in adult humeri
reflect intensive mechanical loading of the upper limb, especially in use of the arms in
paddling of watercraft on the open ocean. Moreover, the findings are consistent with the
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226 Activity patterns: 2. Structural adaptation
(a)
800
Females
700 Males
Femoral Size-Standardized J
600
500
400
300
200
100
0
-20 -18 -16 -14 -12 -10
d13C (ppm)
(b)
1100
Females
1000
Males
Humeral Size-Standardized J
900
800
700
600
500
400
300
-20 -18 -16 -14 -12 -10
d13C (ppm)
Figure 6.6 Femoral (a) and humeral (b) relative strength (J) are plotted relative to stable
carbon isotope ratios. Stable carbon ratios reflecting maize-rich diets (less negative
δ13C values) are associated with lower femoral and humeral robusticity. (Ruff, 1999;
reproduced with permission of author and University of Utah Press.)
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6.2 Cross-sectional geometry 227
Table 6.1 Humerus polar moments of area (35%, 50%, and 65% sections) in selected fossil
and recent human adults (Adapted from Churchill, 1994: Tables 16 and 17)
Females Males
analysis of cross-sectional geometry of rowers whereby rowers tend to have high levels of
robusticity, but especially in populations rowing in open ocean settings (Ruff, 2006; Weiss,
2003).
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228 Activity patterns: 2. Structural adaptation
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6.2 Cross-sectional geometry 229
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230 Activity patterns: 2. Structural adaptation
Variability between status groups in bilateral asymmetry may reveal behavioral differences
within ranked societies. Degree of left–right dominance in external measurements of Meso-
lithic limb bones from Western Europe shows some difference in work and activity between
upper- and lower-status groups (Constandse-Westermann & Newell, 1989). High-status
females have reduced right dominance when compared with low-status females.
Constandse-Westermann and Newell (1989) contend that the greater upper limb lateralization
in low-status women reflects their heavier work demands than those of high-status women.
Higher-status males possess greater right dominance than lower-status males. Although
reasons for the contrasting pattern in males and females are unclear, they speculate that in
order for males to achieve high status, they were required to perform more demanding and
more differentiated tasks.
Analysis of individuals engaged in highly specific, highly demanding activity that required
the use of a dominant arm provides important perspective on the interrelationship between
form, function, and behavioral inference. Examination of cross-sectional geometry of the
humeri of battle victims in Medieval contexts from Towton (see Chapter 4) and Fishergate,
England, shows striking differences in diaphyseal robusticity and shape of the left and right
humeri (Rhodes & Knüsel, 2005). Comparison of blade-injured and non-blade-injured males
reveals distinctive patterns of variation in areas and second moments of area (CA, TA, J, and
Ix/Iy). Blade-injured males display consistently higher values for these properties compared to
non-blade-injured males from the same period. In particular, values of J, a good overall
indicator of resistance to bending/torsion, are remarkably high in the Towton battle victims.
With regard to diaphyseal shape asymmetry, values of Ix/Iy present distinctive left–right
differences in the Towton series. This pattern of bilateral shape asymmetry represents signifi-
cant differences in loading, movement patterns, and use of the arms generally, in these
individuals. These right–left side differences reflect years of training and the adaptation of
bone to profoundly demanding activities. Rhodes and Knüsel (2005) present the case that the
upper left limb in these individuals was adapted to activities that increase bone deposition,
especially in the mediolateral dimension, such as movement patterns involving abduction,
adduction, and rotation of the arm, possibly using a longbow. In contrast, Fishergate males
show right-dominance in cross-sectional properties, suggesting long-term training in use of
the right arm, which is likely related to use of a right-handed unimanual weapon, such as a
sword or lance.
While the bioarchaeological record shows highly specific patterns of humeral strength (or
rigidity) that reflect local adaptations, there is a general pattern of reduction in upper limb
asymmetry over the course of human evolution. In particular, early hominins are highly
asymmetrical, Holocene foragers are moderately asymmetrical, and Holocene farmers and
modern humans having the least degree of asymmetry in the upper limb (Figure 6.7).
Asymmetry of the lower limb is poorly known in archaeological or other population
settings. Lateralization of lower limb bones has been studied from California (Ruff & Jones,
1981), Pecos Pueblo (Ruff & Hayes, 1983b), and Mesolithic Western European (Constandse-
Westermann & Newell, 1989, 1990). These investigations reveal that the left side tends to have
slight size dominance and greater mechanical strength, especially in anteroposterior bending
forces. Generally, lower limb asymmetry in structure and overall size is either considerably
less than upper limb asymmetry or shows no consistent pattern (Auerbach & Ruff, 2006;
Borgognini Tarli & Repetto, 1986; Constandse-Westermann & Newell, 1989, 1990;
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6.2 Cross-sectional geometry 231
40
Humeral Bilateral Asymmetry in Z p (%)
30
20
10
0
Georgia Mission
Jomon
Euroamerican
Calif. Amerindian
SW Amerindian
Aleut
Neanderthal
Tennis Players
Figure 6.7 Humeral bilateral asymmetry in Zp among Holocene and modern human
populations. Paleolithic and Neanderthal populations demonstrate humeral bilateral
asymmetry that equated with modern day tennis players. This similarity in asymmetry
indicates that early hominins engaged in activities where one arm was consistently used
more than the other. In contrast, modern humans from temperate coastal (Georgia Mission)
to Arctic (Aleut) environments show significantly less humeral bilateral asymmetry. These
results indicate generally equal distribution of mechanical loading on both left and
right arms. (Ruff, 2008; reproduced with permission of author and John Wiley & Sons, Inc.)
Dangerfield, 1994; Ruff & Hayes, 1983b). The differences in asymmetry in upper and lower
limbs in humans reflect the fact that the upper limbs are used in a wide variety of non-
ambulatory activities, whereas lower limbs are primarily used in bipedal locomotion, a super-
dominant function requiring equal use of left and right sides.
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232 Activity patterns: 2. Structural adaptation
population in comparison with the Archaic population for both adult males and females, thus
indicating greater bone rigidity in the agriculturalists than in the foragers. Structural analysis
of male humeri shows that the two temporal series are virtually indistinguishable. Thus, for
males, activity levels increased, but primarily in relation to lower limb (ambulatory) functions.
In females, there are significant increases in humeral and femoral rigidity. Bridges suggests
that the increase in bone rigidity in female femora and humeri reflects a relatively greater
range of activity changes in them than in males. Thus, as revealed by the osteoarthritis
analysis, the shift to food production may have had a relatively greater impact on women
than men in this setting. Some of these activities are likely related to who is responsible for
food processing; increases in female humeral bone rigidity may be associated with maize
processing. Historic-era Southeastern native women used wooden mortars and pestles
that required very demanding physical labor involving the upper limbs. One nineteenth-
century observer equated the task of maize pounding to blacksmithing (Current-Garcia &
Hatfield, 1973).
These findings from northwestern Alabama suggest that the adoption of an agricultural
lifeway involved more strenuous physical activity than in earlier populations. The decline in
osteoarthritis prevalence for these populations suggested a decrease in mechanical loading
(see Chapter 5). Thus, these two indicators of mechanical stress seem to yield contradictory
results. Bridges (1991b) suggests that osteoarthritis and long bone geometry are due to
different types of activities. For example, citing findings from sports medicine and other
research, she notes that some normal activities – such as running – do not contribute to
osteoarthritis (Eichner, 1989; Panush & Brown, 1987). Less frequent movements that lead to
microtrauma and injury of joints may be important factors in the explanation of osteoarth-
ritis. It is virtually impossible to distinguish traumatic from daily “wear-and-tear” osteoarth-
ritis, thus making it difficult to identify specific causes where behaviors are unknown, such as
in archaeological settings. Skeletal structural change has been tied to long-term repetitive
forces (Lanyon & Rubin, 1984; Shaw et al., 1987). Thus, osteoarthritis and diaphyseal cross-
sectional geometry should not necessarily be concordant because osteoarthritis would be
expected to develop in older adults, whereas diaphyseal remodeling is a lifelong response to
mechanical stimuli (Ruff, 1992).
A similar pattern of change in cross-sectional geometry is documented in the lower Illinois
River valley, a region that saw a transition from mixed foraging/hunting and consumption of
domesticated plants (native starchy seed crops such as Chenopodium) during the Middle
Woodland (50 BC–AD 200) and Late Woodland (AD 600–1050), to the introduction of maize
in the later Late Woodland, and intensive maize production and consumption in the Mis-
sissippian period (AD 1050–1250). Analysis of skeletal morphology (cortical area, CA, and
second moment of area, J) in light of economic intensification associated with more intensive
plant production and especially increased maize production in the Mississippian period
revealed little evidence of change in males (Bridges, Blitz et al., 2000). In contrast, females
show a distinctive pattern of increase in mechanical loading, as it is represented in second
moment of area (J) and cortical area (CA), through the later part of the Late Woodland period.
In females, however, there is a decline in cross-sectional geometric properties in the Mis-
sissippian period. This pattern of change reveals that the economic shifts more profoundly
affected women than men, perhaps indicating that women were more heavily involved in
agricultural production and food preparation than men; men continued similar kinds of
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6.2 Cross-sectional geometry 233
behaviors, such as those associated with hunting, throughout all four periods. The decline in
activities affecting especially the upper limbs of females in the Mississippian period may be
related to innovations in how maize was prepared in later prehistory in the lower Illinois River
valley. For example, innovations in ceramic technology that allowed increased boiling of
maize and other food may have resulted in reduced workload involving pounding and
processing, hence explaining the clear evidence for reduction in mechanical demand on
women’s upper limbs (Bridges, Blitz et al., 2000).
Analysis of femoral and humeral diaphyses from preagricultural and agricultural Georgia
Bight populations yields an unambiguous decline in second moments of area for both males
and females, reflecting a decline in workload in this setting (Larsen & Ruff, 1991, 1994, 2011;
Ruff & Larsen, 1990; Ruff et al., 1984) (Figure 6.8).
The results reported from the three regions of the southeastern United States – Pickwick
Basin, lower Illinois River valley, and Georgia Bight – are different, especially in comparison
with second moments of area (Larsen & Ruff, 2011) (Figure 6.8). However, the comparisons of
periods within each of the three settings described here are informative about workload and
activity and how they vary across landscapes and adaptive shifts. Comparison of temporal
trends for J for the femur and humerus reveals significant differences between the three
groups. For males, femoral midshaft rigidity increases in Alabama, shows no change in the
lower Illinois River valley, and decreases substantially in the Georgia Bight. In females,
femoral midshaft rigidity increases in Alabama, increases in the lower Illinois River valley
in the first three periods, but then decreases in maize farmers, and decreases in the Georgia
Bight. Comparisons of cross-sectional geometry in the humerus show broadly similar patterns
across the three settings. However, male humeri show no change in Alabama and the lower
Illinois River valley, and a decrease in the Georgia Bight. Female humeri show an increase in
Alabama and the lower Illinois River valley, followed by quite a substantial decrease in the
Mississippian period and a decrease in the Georgia Bight.
While these changes are somewhat complex in viewing the region as a whole, several
general trends are clear. First, the comparisons reveal different patterns of temporal change
in males and females in the foraging-to-farming transition. The degree of change is greater
in females in Alabama and the lower Illinois River valley, but greater in males in the
Georgia Bight. This suggests that the transition had a great impact on women in the
terrestrial, inland setting and on men in the marine, coastal setting. In two of the three
regions (Alabama, Georgia Bight), there was only one agricultural period (introduction of
maize), whereas in the third region (lower Illinois River valley), there were two agricultural
periods (local domestication of weedy plants; introduction of maize). The latter involved
changes in activity in both periods, but with different outcomes – an increase in activity in
the shift to exploitation of oily starches, followed by a decrease in activity subsequent to
the adoption of maize. The implications of the biomechanical analyses are that the transi-
tion had behavioral significance well before maize was adopted and intensified. Import-
antly, these findings strongly suggest that the agricultural transition was not a monolithic
event having the same outcome across the region of the American Southeast, or even
within a single region (e.g., lower Illinois River valley). In fact, factors of terrain and
ecology – markedly different when comparing inland and coastal settings – almost cer-
tainly influenced the kinds and levels of activity that affected skeletal growth and adult
diaphyseal morphology.
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234 Activity patterns: 2. Structural adaptation
(a)
Pickwick Basin
450 Foragers
Farmers
400
Femur JSTD
350
300
250
200
Males Females
350
300
250
200
Males Females
Georgia Bight
600
Foragers
550
Farmers
500
Femur JSTD
450
400
350
300
250
200
Males Females
Figure 6.8 Second moments of area (JSTD) in femora (a) and humeri (b) from Lower Illinois
Valley and Georgia Bight populations decline in late prehistory. These results suggest a
decline in upper- and lower-body workload. In contrast, the Pickwick Basin populations
increase in JSTD values. This difference suggests that variation in mobility and activity
exists within subsistence strategies between periods and regions. (Larsen & Ruff, 2011;
reproduced with permission of John Wiley & Sons, Inc.)
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6.2 Cross-sectional geometry 235
(b)
Pickwick Basin
Foragers
Farmers
600
500
Humerus JSTD
400
300
200
Males Females
Mississippian Farmers
500
400
300
200
Males Females
Georgia Bight
800 Foragers
Farmers
700
Humerus JSTD
600
500
400
300
200
Males Females
The study of structural morphology in Georgia Bight limb bones has been expanded to
include the descendant, contact-era populations, including the descendant Guale population
on St. Catherines Island, Georgia (Santa Catalina de Guale), the later descendant Guale on
Amelia Island, Florida (Santa Catalina de Guale de Santa Maria), and an early contact series
preceding full missionization from Amelia Island representing Timucua (Larsen & Ruff, 1994,
2011; Ruff & Larsen, 1990, 2001). Cross-sectional geometric analysis for the femur reveals
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236 Activity patterns: 2. Structural adaptation
(a)
600
Males
Females
Femoral Midshaft Bending/Torsional Robusticity (JSTD)
500
400
300
200
100
0
EPG LPG EMG LMG EMT
Group
Figure 6.9 Torsional robusticity (JSTD) decreases in femoral (a) and humeral (b) midshafts
from precontact to postcontact populations on Amelia Island: EPG (Early Preagricultural
Guale), LPG (Late Preagricultural Guale), EMG (Early Mission Guale), LMG (Late Mission
Guale), and EMT (Early Mission Timucua). (Ruff & Larsen, 2001; reproduced with
permission of University Press of Florida.)
that beginning with the early contact period on St. Catherines Island, there is a reversal of the
decrease in bone rigidity that had been documented in the late prehistoric populations from
the region (Figure 6.9). Geometric properties increase in the early contact period, but decline
in the late contact period in the Guale series. The very early Timucua have values of J more
like those of the precontact agricultural Guale from St. Catherines Island for the femur.
In males, humeral second moments of area increase successively in the early and late
contact periods. In females, there is a continued decline in humeral values in the early contact
period, but this reverses in the late contact period. Thus, both females and males in the late
contact period experience a marked increase in humeral rigidity. In general, the changes
observed in bone rigidity in the contact period indicate that native populations experienced
increases in mechanical loads, probably due to increased manual labor and physical demands
placed on them by the Spanish (see Chapter 5). The early contact-era Timucua, however, are
more similar in these values compared to later contact populations. This finding is consistent
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6.2 Cross-sectional geometry 237
(b)
Males
800 Females
Mid-Distal Humeral Bending/Torsional Robusticity (JSTD)
600
400
200
0
EPG LPG EMG LMG EMT
Group
with the notion that the earliest contact populations that preceded full missionization and its
associated exploitation of native labor would be more similar to precontact populations than
to later contact populations in workload and lifestyle generally (Ruff & Larsen, 2001).
Because nutritional quality also influences skeletal size to a degree (see Chapter 2), the
possibility remains that the mechanical environment may not be the sole factor that explains
the structural modifications documented in the Georgia Bight. Standardized values of cortical
area show very little change in the comparison of the four periods, thus suggesting that bone
mass remains essentially unchanged through time. In contrast, the distribution of bone tissue
alters dramatically, which is consistent with skeletal adaptations to localized (mechanical)
factors rather than systemic (nutritional) stress (Larsen & Ruff, 1994; Ruff, 1999; Ruff &
Larsen, 2001; Ruff et al., 1984).
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238 Activity patterns: 2. Structural adaptation
new arrivals prior to the European invasion. With the arrival of Europeans in the seventeenth
century came increased reliance on domesticated plants (maize) and farming overall, intensi-
fied long-distance hunting of bison, and intensified trade and warfare between native
populations and Euroamericans (Blakeslee, 1994; Owsley & Jantz, 1994). Much of this record
pertains to the late prehistoric Plains Village period (post-AD 950–1600) and Historic period
(post-AD 1600) and involved dramatic lifestyle and activity changes. Biomechanical analysis
contributes important insight into these developments, sometimes rapidly occurring and
limited to the region (e.g., intensified hunting with reliance on the horse) but also in
comparison, more broadly, with North America generally (maize farming). The adoption of
the horse had a profound impact on cultures in the region. In addition to altering subsistence
practices, the adoption of the horse facilitated inter-tribal trade and increased the ability to
travel long distances. In comparing cortical area (CA) and second moment of area (J) for
earlier and later Historic populations, no differences in mechanical loading of femora were
found (Ruff, 1994b). However, there is a clear decline in sexual dimorphism in these properties
in comparing males and females, suggesting that the transition involved increasingly similar
activities between the sexes, a pattern much like settings in the American Southeast (e.g.,
Georgia Bight). With regard to specific tribes, biomechanical comparisons of late prehistoric
and historic Arikara reveal that with intensified farming, a development related to increasing
economic focus on creating surpluses of crops for trade, mechanical loading increased to some
extent but primarily in females (Wescott & Cunningham, 2006). The greater work change in
women than men suggests that women may have been the primary source of labor for
increasing crop production for trade.
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6.2 Cross-sectional geometry 239
86
50% Section Femur
84
50% Section Tibia
Mean Cortical Percentage (%CA)
82
80
78
76
74
72
70
68
Early Mid Neanderthals MPMH EUP/MUP
Pleistocene Pleistocene
Pleistocene Homo
Figure 6.10 Temporal reduction in femoral midshaft robusticity in Homo from the Early
Pleistocene to Upper-Middle Paleolithic. (Adapted from Trinkaus & Ruff, 2012; reproduced
with permission of authors and PaleoAnthropology Society.)
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240 Activity patterns: 2. Structural adaptation
Late Pleistocene and early Holocene – between the Upper Paleolithic and Neolithic – there is a
significant reduction in lower limb rigidity and mobility (Holt et al., 2012; Shaw & Stock,
2013), indicating major alterations in mechanical loading in general and mobility in
particular.
Femoral head size relative to body size also remains relatively constant throughout most of
the Pleistocene (Ruff et al., 1993). These findings lend support to the hypothesis that joint
morphology is more genetically canalized or less developmentally plastic than long bone
diaphysis morphology. In adults, articular joint size and shape does not alter in response to
mechanical loading, unlike diaphyseal structure (Rafferty & Ruff, 1994; Ruff & Runestad,
1992; Ruff et al., 1991, 1994). Changes in articular loading in adults can have profound effects
on subchondral and trabecular bone structure organization underlying the joint surface
(Pauwels, 1976; Poss, 1984; Radin et al., 1982, 1984; Rafferty & Ruff, 1994; Ruff, 1992,
2008), but these changes are not manifested in external articular size. Thus, internal and
external joint structures are independent and represent contrasting expressions of the mech-
anical environment.
The lack of influence of mechanical loading on joint size has important implications for
functional studies of archaeological remains, especially where joint size differences are used
to infer mechanical differences between groups. For example, increase in femoral head size in
comparing prehistoric hunter-gatherers and agriculturalists in the Caddo region of the Ameri-
can Southeast was interpreted to reflect an increase in protein consumption and increase in
mechanical loading (Rose et al., 1984). These new findings comparing articular and diaphy-
seal structure indicate that mechanical loading as a causal factor in explaining temporal
change in joint size is highly unlikely.
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6.2 Cross-sectional geometry 241
250
Male
Female
200
Standardized Humerus J
150
100
50
0
Jebel Sahaba el-Badari Hierakonpolis Kerma
Site
Figure 6.11 Decline in male humeral robusticity (J) from preagricultural (Jebel Sahaba) to
agricultural (el-Badari, Hierakonpolis, and Kerma) Egyptian populations. Females show no
significant difference in humeral robusticity. (Burger & Thomas, 2011; reproduced with
permission of author and John Wiley & Sons, Inc.)
males, femoral robusticity also shows a sharp decline after the Late Paleolithic, and then
remains essentially unchanged in the sequence of the three agricultural periods. In females, no
change occurs in femoral robusticity immediately following the Late Paleolithic, but there is a
dramatic decline in the final two periods of agricultural intensification. Overall, there are
significant declines in levels of habitual activity, but with different timing between adult men
and women. This pattern fits the general finding in other regions where there is evidence for a
decline in sexual dimorphism. On the other hand, this pattern is overlain by other factors
affecting workload and activity differently for men and women that are specific to the Nile
Valley.
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242 Activity patterns: 2. Structural adaptation
Table 6.2 Percentage change with age in femoral and tibial cross-sectional geometric
properties. Calculated by the formula: {[(40 þ years) – (20 to 39 years)] (20 to 39 years)}
100 (Adapted from Ruff & Hayes, 1982: Table 1)
CA MA TA Imax Imin
Bone
(section location)
Males
Tibia (20%) −1.1 13.6a 7.3a 9.4 7.8
Tibia (50%) 0.2 35.7c 12.4c 14.6a 19.9a
Femur (50%) 0.1 25.4c 6.7b 12.3b 6.8
Femur (80%) −0.5 13.1b 4.7a 7.5 4.8
Females
Tibia (20%) −14.5b 9.1a 9.1a −1.5 −3.0
Tibia (50%) −13.1b 62.4c 12.9b 10.3 4.1
Femur (50%) −6.6 66.6c 11.7c −9.6 15.6b
Femur (80%) −2.4 41.2c 13.6c 16.5b 16.0a
a
Statistically significant between age groups (Student’s t-test: P<0.05)
b
Statistically significant between age groups (Student’s t-test: P<0.01)
c
Statistically significant between age groups (Student’s t-test: P<0.001)
CA, cortical area; Imax and Imax, second moments of area/bending rigidity; MA, medullary area;
TA, total subperiosteal area.
1967, 1992; Heaney et al., 1997; Russo et al., 2006) and archaeological series (Carlson et al.,
1976; Pfeiffer, 1980; Ruff, 2010b; Ruff & Hayes, 1983b; Stirland, 1993). Some contend that
periosteal expansion represents a compensatory response to endosteal bone loss and thinning
of the cortex with advancing age (Garn et al., 1967; Ruff & Hayes, 1982; Smith & Walker,
1964; see discussion in Martin et al., 1998). Until recently, this hypothesis was difficult to test
because of the imprecision of radiographic measures of cortical bone remodeling (Ruff &
Hayes, 1982). In order to examine the issue of age changes and periosteal expansion in more
detail, Ruff and Hayes (1982, 1983b; Ruff, 2010b) analyzed section properties – areas and
second moments of area – from the late prehistoric/protohistoric Pecos Pueblo site, New
Mexico. Analysis of femoral and tibial diaphyseal sections reveals that both sexes saw
increases in medullary area and total subperiosteal area and decreases in cortical area with
advancing age (Table 6.2). Second moments of area (Imax and Imin) increase in older adults.
Thus, in support of the compensatory hypothesis, continuous periosteal expansion in older
adults appears to maintain the mechanical integrity of the long bone despite overall decline in
bone mass.
Variation in different sections along femoral and tibial diaphyses also reveals that skeletal
remodeling with age is less pronounced in the most distal and proximal sections of the
diaphyses. The greater remodeling in tibial and femoral midshaft and adjacent sections than
at the distal and proximal ends is likely due to the relatively greater mechanical loads –
especially bending (Ruff, 1992; Ruff & Hayes, 1982).
While the analysis of cross-sectional geometry and behavioral inference is richly repre-
sented in adults, it is the record of growth and development in juveniles that is important for
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6.2 Cross-sectional geometry 243
comprehending the acquisition of adult robusticity, the timing of its development, and the
overall record of ontogeny that informs our understanding of robusticity (Ruff, 2003).
Analysis of Medieval-period juvenile (<16 year of age) tibiae from Kulubnarti, Sudanese
Nubia, shows increases in percentage of cortical area and second moments of area (Ix, Iy) from
3 to 12 years of age (Van Gerven et al., 1985). After the age of 12 years, areas decline, while
second moments of area continue to increase dramatically. Thus, despite decline in bone area,
continued increase in second moments of area in the later juvenile years appears to preserve
mechanical integrity throughout the years of growth and development.
Similarly, analysis of various populations from North America, Europe, and the Middle East
representing a variety of behavioral, ecological, and dietary circumstances, reveals common
patterns of age changes from birth through adolescence, yet important differences reflect
significant regional variation (compare with Cowgill, 2010; Cowgill & Hager, 2007; Garofalo,
2013; Gosman et al., 2013; Larsen et al., 2013; Van Gerven et al., 1985). These comparisons
show developmental trajectories of long bone rigidity and strength. For example, in the first
year of life, Medieval Nubians and twentieth-century Portuguese show relatively low robus-
ticity, whereas Medieval Bosnians and prehistoric Inupiat show elevated robusticity (Cowgill,
2010). Sully (Arikara), Point Hope (Inuit), Çatalhöyük (Anatolian Neolithic), and Barton-
upon-Humber (Medieval-Industrial United Kingdom) express relatively high robusticity
throughout the juvenile years (Garofalo, 2013) (Figure 6.12). High levels of robusticity early
in ontogeny predict, therefore, the high levels documented for adults (Çatalhöyük;
Figure 6.13).
In order to document ontogenetic age patterns of diaphyseal remodeling in more precise
fashion, Ruff and coworkers (1994) examined areas and second moments of area of humeri in
professional tennis players aged 14–39 years. Both males and females show a pattern of
endosteal contraction and periosteal expansion resulting in large increases in bone area (CA)
and torsional rigidity (J) in the dominant playing arm. The resulting robusticity is primarily
due to greater periosteal expansion and not endosteal contraction. The degree of humeral
robusticity has a strong association with age: individuals who began playing tennis earlier
have greater robusticity than individuals who began later. The increased mechanical loading
in children and young adolescents has a more pronounced effect on the periosteal surface;
after mid-adolescence, loading appears to have a more pronounced effect on the endosteal
surface. These findings indicate an age pattern of sensitivity in the bone forming surfaces –
periosteal versus endosteal – in response to increases in mechanical stimuli. The cause of the
shift in focus of sensitivity is unknown, but may be related to changes in hormonal levels and
their different effects on the two bone surfaces (Ruff et al., 1994).
Relatively little is known about the variation in cross-sectional morphology along the
diaphyses of limb bones, especially with regard to ontogenetic changes in response to activity.
In order to draw a more precise understanding of behavioral and developmental variation,
Gosman and collaborators (2013) scanned (high-resolution X-ray CT) five sections (20%,
35%, 50%, 65%, 80%) of femoral and tibial diaphyses of juveniles ranging in age from
neonate to young adults from the Norris Farms, Illinois skeletal series. As would be predicted,
their analysis revealed that cross-sectional geometric shape (Imax/Imin) changes throughout the
life course (Figure 6.14). However, these changes are highly patterned. That is, there are peak
levels of changing morphology during two key periods of life; first, during early childhood
(attainment of walking); second, during early adolescence (hormonal and body-mass
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244 Activity patterns: 2. Structural adaptation
(a)
10 000
9000
8000
7000
Humerus Rigidity (J)
6000
5000
4000
3000
2000
1000
0
Tigara
Tigara
Tigara
Tigara
Barton
Barton
Barton
Barton
Arikara
Arikara
Arikara
Arikara
Çatalhöyük
Çatalhöyük
Çatalhöyük
Çatalhöyük
Infant Young Child Middle Child Old Child/
Adolescent
(b)
35 000
30 000
25 000
Femur Rigidity (J)
20 000
15 000
10 000
5000
0
Tigara
Tigara
Tigara
Tigara
Barton
Barton
Barton
Barton
Arikara
Arikara
Arikara
Arikara
Çatalhöyük
Çatalhöyük
Çatalhöyük
Çatalhöyük
Figure 6.12 Developmental differences in humeral (a) and femoral (b) rigidity (J) according to
age groups and variable populations. Rigidity increases significantly from middle child to
old child/adolescent years among Neolithic Çatalhöyük, Protohistoric Arikara, and Medieval-
Industrial Barton-upon-Humber samples. Prehistoric Tigara (Point Hope) individuals do not
show this markedly increased rigidity at adolescence, indicating inter-regional variability in
long bone ontogeny. (Adapted from Garofalo, 2013; produced with permission of author.)
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6.2 Cross-sectional geometry 245
120
Males
Females
110
100
Relative Femoral Strength
90
80
70
60
ze
c
ic
.
.
hi
l
l
Pa
Pa
ith
on
lit
ol
eo
Br
p.
p.
es
U
U
N
M
te
rly
La
Ea
Figure 6.13 Femoral midshaft strength (Zp) in Çatalhöyük and comparative samples. (Adapted
from Larsen et al., 2013; reproduced with permission of Cotsen Institute of Archaeology.)
Figure 6.14 Cross-sections of the femoral (above) and tibial (below) midshafts among five age
categories: Age Group 1 (0–1.9 years), Age Group 2 (2–4.9 years), Age Group 3 (5–8.9 years),
Age Group 4 (9–13.9 years), and Age Group 5 (14–17.9 years). Cross-sectional geometric
morphology (maximum diameter/minimum diameter, Imax/Imin) of the femur is generally
conserved during development while tibial midshaft becomes anterioposteriorly
strengthened. (Adapted from Gosman et al., 2013; reproduced with permission of authors
and John Wiley & Sons, Inc.)
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246 Activity patterns: 2. Structural adaptation
increase). Moreover, for the tibia, the round diaphysis in earlier childhood becomes increas-
ingly anterioposteriorly strengthened. The femur, on the other hand, has more region-specific
changes, with the midshaft (50% section) being much more conserved than the distal and
proximal diaphyses. All statistically significant changes are associated with the ends of the
diaphyses, suggesting that these areas are more subject to morphological change in response
to mechanical loading during the years of growth and development. Overall, the diaphyseal
cross-sections change from relatively circular early in life to more asymmetrical in late
adolescence. This finding underscores the critical importance of understanding ontogeny as
a background to interpreting adult variation. Virtually all of the records described in this
chapter focus on adults and especially the midshaft of adult long bones. The study of Norris
Farms femoral development in particular reveals that the record of ontogeny and biomechan-
ical adaptation may be best explored and interpreted by taking a whole bone approach to
include the midshaft as well as other diaphyseal regions. Clearly, the midshaft is an important
part of the behavioral/adaptive story, but only part of the story, especially when attempting to
reconstruct key events in the lifecourse.
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6.4 Behavioral inference from external measurements 247
other modern humans, and hence, a greater volume of bone formed per unit area. This
interpretation is in accordance with the findings based on structural and geometric analysis
of femur cross-sections in Pecos Pueblo adults (Ruff, 1991; Ruff & Hayes, 1983a, 1983b).
This reasoning is in line with the very significant positive correlations found between
osteon density and anteroposterior and mediolateral second and polar moments of inertia
in a Euroamerican cadaver sample consisting of older adults (>50 years of age); individuals
with high levels of mechanical loadings have high osteonal densities (Walker et al. 1994).
Similarly, comparisons of femoral osteon density between adult males and females from
late Christian-period Nubia (AD 550–1450) show that males have more osteons than
females, which may represent greater bone turnover from higher activity regimens in males
than in females (Mulhern, 1996). As with experimental evidence (Lanyon & Baggott, 1976),
these studies indicate that activity has a strong influence on histological variation in
cortical bone.
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248 Activity patterns: 2. Structural adaptation
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6.4 Behavioral inference from external measurements 249
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250 Activity patterns: 2. Structural adaptation
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6.4 Behavioral inference from external measurements 251
Ix
Ix
Iy Iy
Ix/Iy = 1.0
Ix/Iy = 1.5
in the anteroposterior plane (i.e., ratio is >1.0) represents relatively greater bone rigidity and
functional demand in the anteroposterior direction than in the mediolateral direction. Ruff
(1987) has shown a temporal decline in the ratio in recent human groups, which he interpreted
to reflect a general reduction in the amount of anteroposterior bending forces as populations
have become increasingly sedentary (compare with Lovejoy et al., 1976). Therefore, the Ix/Iy
ratio represents a mobility index, herein referred to as Ruff’s mobility index.
Analysis of the generally high mobility index in hunter-gatherers from the American Great
Basin, especially from Stillwater and the Great Salt Lake regions, reveals the dominance of a
highly mobile lifestyle, especially in males (Ruff, 1999). Their values are among the highest in
comparison with other North American populations, and like data presented for second
moments of area (J), the mobility index values fit into a continuum from generally high in
hunter-gatherers to relatively lower in agriculturalists. Based on this index, Great Basin
females appear to be less mobile than males, which is true for all other North American series
(Ruff, 2010b). Comparisons of females and males in the Ix/Iy ratio show that Great Basin
foragers are highly sexually dimorphic (Figure 6.17), which is a paramount characteristic of
hunter-gatherers generally, and is related to sexual division of labor and a strong male–
female dichotomy in activity patterns (Larsen, Ruff et al., 1995; Ruff, 1987, 2010a; Ruff &
Larsen, 2014; Stock et al. 2011).
In the Georgia Bight, early contact-period femora show a general reduction in the femoral
midshaft Ix/I y ratio relative to earlier prehistoric populations. Historic sources indicate that
populations during the historic period became generally less mobile as they were either forced
or coerced to live in and around mission centers (e.g., Santa Catalina de Guale, Georgia). This
skeletal indicator of mobility is, therefore, in accordance with other sources describing
substantial population sedentism during the historic period. These structural modifications
suggest that mission Indians in Spanish Florida had a more physically demanding lifestyle,
but within the relative confines of the mission setting (Larsen & Ruff, 1994).
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252 Activity patterns: 2. Structural adaptation
30
Alabama
Georgia-Florida
20
15
10
Hunter-Gatherers Agriculturalists
0
Pecos
Malheur
Stillwater
Delaware
Guale (HG)
Guale (AG)
Alabama (HG)
Alabama (AG)
E. Gr. Salt Lake
Figure 6.17 Mobility indices for multiple preagricultural and agricultural populations
indicate an appreciable percentage of sexual dimorphism in comparison to hunter-
gatherers and horticulturalists in North America. However, intensive labor exploitation
among later Spanish mission Indians in Guale indicate great differences between males and
females. (From Larsen & Ruff, 2011; © 2011 John Wiley & Sons, Ltd.)
Similarly, temporal comparisons of Early Classic (AD 250–550) and Late Classic
(AD 550–750) populations from Xcambó on the northern coast of the Yucatán peninsula,
Mexico revealed a significant decline in the Ix/Iy ratio in males (but not females), indicating
considerable decline in mobility (Maggiano et al., 2008; and see Wanner et al., 2007). In this
setting, there was no clear change in diet (Maggiano et al., 2008). Rather, the change
in activity – also accompanied by a general decline in robusticity – appears to be related to
improvements in general living standards. In particular, evidence for improvements in
material wealth (e.g., architecture, grave inclusions) suggests that increasing economic
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6.4 Behavioral inference from external measurements 253
complexity associated with the administrative function of the community resulted in reduced
mobility and workload in general, especially in males. These findings are consistent with other
circumstances showing a negative association between wealth and workload in stratified
societies (Robb et al., 2001).
An important European record of diaphyseal shape ratios using cross-sectional shape
reveals similar trends in activity. In a series of femora from multiple localities representing
the Late Upper Paleolithic, Mesolithic, Neolithic, Eneolithic, and Iron Age (c. 19 000–4000
yBP) in northwestern Italy, analysis of the Imax/Imin ratio (the ratio of plane of maximum
bending strength to plane of minimum bending strength) shows a continuous decline in
females, reflecting an increasingly round shape in cross-section of the femoral diaphysis
(Marchi, 2008; Marchi et al., 2011; Sparacello et al., 2011). This expected finding contrasts
with the record of the Imax/Imin ratio for males. That is, following the Mesolithic, Neolithic
males show a dramatic rise in the ratio, approaching the mean for males of the Late Upper
Paleolithic. This suggests that the Neolithic males, but not females, were likely as mobile as
Late Upper Paleolithic males, certainly more so than the Mesolithic and all post-Neolithic
populations in the region (Marchi et al., 2011; and compare with Sládek et al., 2006a, 2006b
for central Europe). Overall, therefore, these findings point to an exception to the general
finding of declining mobility from the late Pleistocene through recent times. What is it that
Neolithic males from northwest Italy are doing that requires elevated mobility? Marchi and
coworkers (2011) suggest that the spike in mobility in Neolithic males reflects the emphasis on
pastoral activities, especially animal husbandry, and not agricultural/farming activities. For
example, Robb (1994) makes the argument that these Neolithic males were involved in
herding over rough and uneven terrain. Indeed, the biomechanical evidence strongly supports
this explanation. This pattern of elevated mobility contrasts with patterns of the probability
ratio for Neolithic Çatalhöyük (Larsen et al., 2013) (Figure 6.18). That is, while the ratio
documenting the anterioposterior to mediolateral bending strength is relatively high (>1.0), it
is considerably lower than samples documenting the European Upper Paleolithic, Mesolithic,
Neolithic, and Bronze Age. Thus, while physically active, the relatively lower mobility index
value for Çatalhöyük is more consistent with a population that is relatively sedentary. On the
other hand, within the 1400-year period of occupation, there is a trend showing an increase in
the index, suggesting that the population became increasingly mobile, perhaps in response to
a decrease in food and other resources immediately adjacent to the community, in combin-
ation with increased aridity prior to the abandonment of the community, as well as a range of
other factors (Larsen et al., 2013) (Figure 6.19).
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254 Activity patterns: 2. Structural adaptation
1.6
Males
Femoral A-P/M-L Bending Strength
Females
1.4
1.2
1.0
0.8
.
l.
ic
ze
l
Pa
Pa
hi
ith
on
lit
ol
p.
p.
eo
Br
es
U
N
M
rly
te
La
Ea
Figure 6.18 Mobility indices (femoral A-P/M-L bending strength) of Neolithic Çatalhöyük
samples compared with Early Upper Paleolithic, Late Upper Paleolithic, Mesolithic,
Neolithic, and Bronze Age European populations. Çatalhöyük males and females have
mobility indices close to 1.0, suggesting a more sedentary lifestyle relative to these
European sites. (Reproduced with permission of Cotsen Institute of Archaeology.)
1.3
Males
1.2
Femoral A-P/M-L Bending Strength
Females
1.1
1.0
0.9
0.8
0.7
Early Middle Late
Figure 6.19 Mobility indices (femoral A-P/M-L bending strength) for Neolithic Çatalhöyük
indicate an increase in mobility from Early to Late periods in both males and females.
(Reproduced with permission of Cotsen Institute of Archaeology.)
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6.5 Summary and conclusions 255
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Masticatory and nonmasticatory
functions: craniofacial adaptation to
7 mechanical loading
7.1 Introduction
The influence of environment and behavior on skull morphology was first discussed centuries
ago. In the fifth century BC, Herodotus remarked on apparent differences in cranial robusticity
between Persians and Egyptians: “The skulls of Persians are so weak that if you so much as
throw a pebble at one of them, you will pierce it; but the Egyptian skulls are so strong that a
blow with a large stone will hardly break them.” He interpreted the strength of Egyptian skulls
to the lifelong exposure of their heads to the sun. Since the time of Herodotus, the influence of
environment and behavior, no matter how specious the interpretation, has been only minim-
ally considered in discussions of cranial morphology in archaeological remains. Beginning in
the eighteenth century, osteologists relied on craniofacial variation for determining popula-
tion history and classification, with little attention given to the biological significance of
observed patterns, especially from a population perspective (Armelagos et al., 1982; Carlson &
Van Gerven, 1979; Cook, 2006; Larsen & Walker, 2010; Powell, 2005). As with the study of
long bone morphology discussed in Chapter 6, there has been a gradual reorientation from
typological/historical to population variation analyses. This emphasis focuses on processes
and functions that influence cranial morphology, revealing the adaptive and behavioral
significance of variation from a population perspective.
This chapter’s central discussion is built on the premise that craniofacial morphology is a
biological ensemble, influenced by processes involving the central role of the masticatory
complex – composed of the jaws, teeth, and associated muscle – bone interactions – during
growth and development and throughout adulthood (Daegling, 2010; Lieberman, 2011). Both
intrinsic and extrinsic factors influence craniofacial growth, just as with any other skeletal
tissue in the body. Throughout life, the cranium grows and adapts in an epigenetic context
where environmental and functional factors operate on an underlying genetic infrastructure
(Carlson, 2005).
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7.2 Cranial form and functional adaptation 257
observational studies on animals and humans (Daegling, 2010; Herring, 1993; Kiliaridis,
1995; Lieberman, 2011; Raadsheer et al., 1999; Ravosa et al., 2008) show the considerable
influence of environment, especially in relation to the cumulative effects of mastication and
mechanical loading of the face and jaws during the years of growth and development and
adulthood. Cranial form cannot be regarded as a single unit with similar genetic and environ-
mental influences. That is, regions of the cranium that are subjected to masticatory loading
(e.g., attachment sites for masticatory muscles in the face and jaws) show more variation than
other regions of the cranium (Lieberman, 1995; von Cramon-Taubadel, 2009b).
Experimental studies involving extirpations of masticatory muscles in laboratory animals
show associated craniofacial skeletal modifications, especially with regard to size and robus-
ticity (Avis, 1959, 1961; Horowitz & Shapiro, 1951, 1955; Moore, 1967, 1973; Pratt, 1943;
Schumacher et al., 1979; Washburn, 1947a, 1947b). Craniofacial skeletons of animals fed soft
foods tend to be smaller and less robust than animals fed hard foods (Beecher & Corruccini,
1981, 1983; Bouvier & Hylander, 1981, 1982, 1984; Bouvier & Zimny, 1987; Corruccini &
Beecher, 1982, 1984; Hinton, 1990; Lieberman, Krovitz et al., 2004; Moore, 1965; Tuominen
et al., 1993; Watt & Williams, 1951; Whitley et al., 1966; and see reviews by Herring, 1993;
Kiliaridis, 1995; Lieberman, 2011). The profound effects of alteration of masticatory loading
are also demonstrated in the experimental transpositions of masseter and temporalis muscles
in laboratory monkeys (Hohl, 1983). The anterior relocation of these muscles leads to a
number of changes, including superior facial tilting. These extirpation and translocation
studies show that alterations in mechanical loading produce shifts in masticatory behavior
that result in distinctive craniofacial morphological changes. Collectively, this body of work
underscores the centrality of adaptive plasticity, or the organism’s ability to respond to
environmental circumstances during their ontogeny (Gotthard & Nylin, 1995; Ravosa et al.,
2008; and see Chapter 6).
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258 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
White males
144
White females
Black males
142
Black females
140
Basion-Bregma Height
138
136
134
132
130
128
126
124
122
in width (Buretic-Tomljanovic et al., 2006; Cameron et al., 1990; Jantz, 2001; Jantz & Logan,
2010; Jantz & Meadows Jantz, 2000; Weisensee & Jantz, 2011) (Figure 7.1). Simultaneous
increases in head length and long bone length documented in North Americans of European
and African ancestry suggest that these are responses to similar forces affecting growth (Jantz
& Meadows Jantz, 2000).
An independent approach to understanding plasticity of cranial shape involved efforts to
document the influence of culture and behavior in past populations. In sharp contrast to
ancient British populations, Keith observed that in modern Britons “many persons have small,
contracted palates. . .Their noses are narrow; so are their faces” (1950:402). Late Celtic faces
became smaller, reflecting in part the “change in dietetic which has occurred since the early
years of the Christian era, cooked food and soft cereals replacing tough meats and imperfectly
ground corns” (Keith, 1916:198). Keith (1916) noted concomitant changes in the occlusal
surfaces of teeth, especially with regard to a reduction in tooth wear in later populations.
Keith’s observations regarding the influence of dietary consistency on facial robusticity were
prescient of a large body of work showing the effects of cooking and reduction of food and
links with reduction of growth of the masticatory apparatus both in Britain (Frake & Goose,
1977; Goose, 1962, 1981; Goose & Parry, 1974; Lavelle, 1972; Moore et al., 1968) and
elsewhere.
Following Keith’s efforts to relate temporal trends in craniofacial morphology to shifts in
masticatory behavior and diet, various researchers have documented other trends in archaeo-
logical populations, both regionally and globally. Weidenreich (1945) recognized the
inappropriateness of using cranial shape for identifying racial groups, instead observing a
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7.2 Cranial form and functional adaptation 259
trend in human evolution – and especially during the Holocene – for increasingly shorter
crania, a process he called brachycephalization. Most workers argued that long-headed
(“dolichocephalic”) populations had been replaced by alien short-headed (“brachycephalic”)
populations, thus explaining the trend (Retzius, 1900; and see review in Weidenreich, 1945).
Contrary to this consensus, Weidenreich (1945) argued against invasion and replacement
models by showing the widespread trend of cranial vault shortening, taking place in earlier
populations throughout Europe (Sokal & Uytterschaut, 1987; Sokal et al., 1987), the Middle
East, and South and Central Asia (Vladescu, 1992).
As with other regions of the world, studies of native New World groups in the first half of
the twentieth century emphasized diffusionistic interpretations of cranial shape variation,
especially arguing that earlier long-headed “dolichocephals” were replaced by later “brachy-
cephals” in native New World groups (Dixon, 1923; Hooton, 1930, 1933; Hrdlička, 1922a;
Hulse, 1941; Newman & Snow, 1942).
Coinciding with and following the publication of Weidenreich’s (1945) classic article, other
researchers recorded temporal changes in cranial form, especially involving reductions in
robusticity and/or increasing vault roundness and brachycephalization, in the comparison of
earlier and later prehistoric North American Indians (Anderson, 1967; Boyd, 1988; El-Najjar,
1981; Guagliardo, 1982b; Hoyme & Bass, 1962; Ivanhoe, 1995; Newman, 1962; Newman &
Snow, 1942; Steele & Powell, 1992; Webb & Snow, 1945) and elsewhere around the globe
(Abdushelishvili, 1984; Henke, 1984; Kaifu, 1997; Mushrif-Tripathy & Walimbe, 2006; Naka-
hashi, 1993; Newman, 1951; Okazaki, 2004; Rightmire, 1984; Rösing & Schwidetzky, 1984;
Sagne, 1976; Sardi et al., 2004; Smith, Bar-Yosef et al., 1984; Smith & Horwitz, 2007; Suzuki,
1969; Walimbe & Gambhir, 1994; Walimbe & Kulkarni, 1993; Walimbe & Tavares, 2002;
Wu & Zhang, 1985; Wu et al., 2012; and references cited later). In some settings, cranial vault
shortening can also be attributed to artificial deformation in some cultures, a practice largely
limited to late prehistoric populations. For much of this record, there is a general pattern of
gracilization of the face and jaws in comparing hunter-gatherers and early farmers in various
settings (Bulbeck & Lauer, 2006; Larsen, 1982, 1995; Sardi & Béguelin, 2011; Sardi et al.,
2004; von Cramon-Taubadel, 2011).
The Nile Valley: invasion and replacement or subsistence change and craniofacial adaptation?
Beginning in the nineteenth century, various workers speculated on the origins of human
groups occupying the Nile Valley (Morant, 1925; Morton, 1844; Smith, 1910). Following
Morton’s (1844) highly influential study of archaeological crania from Egypt and Nubia, the
prevailing notion was that two biologically distinct groups occupied the Nile Valley in
temporal succession. In Lower Nubia, Morant (1925) identified an earlier “Upper Nile type,”
with predominantly “Negroid” features, and a later “Lower Nile type,” which lacked “Negroid”
features. The changes were viewed in a diffusionistic paradigm: simply, the disappearance of
“Negroid” features resulted from an invasion and subsequent replacement by alien “Caucas-
oid” (Egyptian) peoples from the north (Calcagno, 1986a, 1986b; Carlson, 1976; Carlson &
Van Gerven, 1977, 1979; Van Gerven et al., 1973, 1977).
Recent analyses of crania and dentitions from Lower Nubia indicate that the evidence for
the diffusionist model of biological change is less than compelling. Independent analyses of
skeletal and dental discrete and metric variables and other lines of evidence suggest that the
earlier and later Nubian populations largely represent a biological continuum (Batrawi, 1946;
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260 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
Berry & Berry, 1972; Calcagno, 1986a, 1986b; Franceschi et al., 1994; Greene, 1972; Mukher-
jee et al., 1955; Nielsen, 1970; Van Gerven et al., 1977). Certainly, there was considerable
contact and likely gene flow between Nubia and other settings in northeast Africa, as is
clearly illustrated in the long history of contact between Nubia and Egypt (see Chapter 4).
However, there is no evidence of population replacement. Therefore, the differences in cranial
morphology between earlier and later populations observed by Smith and Jones (1910),
Morant (1925), and others are best understood in relation to factors not involving population
replacement.
In order to understand these factors better, especially those related to dietary and
technological change, Carlson and Van Gerven and their coworkers (Armelagos et al.,
1984; Carlson, 1976; Carlson & Van Gerven, 1977, 1979; Hinton & Carlson, 1979; Van
Gerven et al., 1973, 1977) compared craniofacial morphology in a Nubian-based temporal
sequence, including foragers from the Mesolithic (c. 12 000 yBP), initial agriculturalists
from the combined A- and C-groups (3400–1200 BC), and intensive agriculturalists from
the combined Meroitic, X-group, and Christian periods (AD 0–1500). These comparisons
reveal that Nubian foragers and incipient farmers have flat and elongated vaults with
well-developed, protruding supraorbital tori and occipitals. In contrast, later intensive
agriculturalists have rounded vaults with small and more posteriorly positioned faces and
masticatory muscle (temporalis and masseter) attachment sites and reduced temporoman-
dibular joint size (Figure 7.2).
Carlson and coworkers posit a masticatory-functional hypothesis for explaining cranio-
facial changes in Nubia (Figure 7.3). They argue that the primary factor influencing Nubian
craniofacial anatomy was the change in subsistence economy, from foraging to food produc-
tion and the shift to consumption of softer foods. These changes resulted in a reduction in
loading of the masticatory muscles. Alteration in masticatory function led to alteration in
craniofacial growth in two ways: (1) decreased stimulation of bone growth leading to a
reduction in facial robusticity, and (2) progressive alteration of the overall growth of the face
and vault, resulting in a smaller and more inferoposteriorly oriented face relative to the
cranial vault.
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7.2 Cranial form and functional adaptation 261
softer foods (Sully Arikara; AD 1650–1750) (Holmes & Ruff, 2011). In this regard, the Point
Hope corpi are considerably more buttressed mediolaterally than Sully site corpi. Moreover,
second moments of area (Ix) of mandibular corpi are significantly greater in the Point Hope
series than in the Sully series overall (see Chapter 6 for discussion of cross-sectional geometry).
Interestingly, the differences emerge in young juveniles (aged 1–6 years), and become statistic-
ally significant in later childhood (6–12 years old), adolescence (12–18 years old), and adult-
hood (post-18 years of age). Similarly, comparisons of mandibular robusticity in prehistoric
Japan reveal differences in comparison with modern Japanese (Fukase & Suwa, 2008; Kaifu,
1997; Okazaki, 2004), patterns that emerge early in childhood. These examples show that
differences in robusticity that develop through ontogeny meet expectations of reduced masti-
catory loading. The evidence for either development of robusticity or reduced robusticity during
the growth years also lends considerable support for adaptive plasticity in explaining these
patterns of variation.
Temporal trends in skeletal series from the Eastern Woodlands of North America are
consistent with the masticatory-functional paradigm. Crania of late prehistoric (Mississippian
period) farmers from Tennessee show a general decrease in robusticity and a reduction in size
and more posterior orientation of the masticatory muscles in comparison with crania of early
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262 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
Foraging to farming
Altered chewing
Reduced masticatory
muscle activity
Reduced robusticity of
face and jaws
prehistoric (Archaic period) foragers (Boyd, 1988). In this setting, size reduction is especially
pronounced in the mandible and lower face, suggesting that the change in morphology
resulted from decreased mechanical loading of the face and jaws brought about by consump-
tion of softer-textured foods in later prehistory (Boyd, 1988). Accompanying these changes is
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7.2 Cranial form and functional adaptation 263
a marked reduction in size of the temporomandibular joint (Hinton, 1981a, 1983). Experi-
mental studies show that this joint is highly sensitive to alterations in mechanical loading
(Bouvier & Hylander, 1981, 1982, 1984; Tuominen et al., 1993), and the joint tends to be
largest in human populations with high masticatory stresses (Corruccini & Handler, 1980;
Hinton, 1981a, 1981b, 1981c, 1983; Hinton & Carlson, 1979; Wedel et al., 1978). Like the
assessments of robusticity and vault shape generally, these findings denote the primacy of
mechanical factors in interpreting craniofacial morphology, including when population
continuity can be established.
Temporal analysis of crania from a very similar setting from the Ohio River valley reveals
complementary evidence of masticatory function and craniofacial robusticity (Paschetta
et al., 2010). Comparison of foragers (Archaic-period Indian Knoll), incipient farmers
(Woodland-period sites, Kentucky), and intensive farmers (Mississippian-period sites) demon-
strates clear differences in craniofacial robusticity, with the temporal changes of reduction in
robusticity in the face (zygoma and alveolus reduction), and especially reduced projection of
the face, and reduced size of the temporomandibular joint. These changes link directly with
reduced masticatory loading of the temporalis muscles and increased consumption of softer
foods.
Similarly, comparisons of prehistoric Georgia coastal forager-fishers (pre-AD 1150) and
farmers (AD 1150–1550) reveal that, like the Tennessee and Ohio River valley populations,
there is a general decrease in craniofacial robusticity. Like these settings, the reductions
in facial and mandibular dimensions and attachment sites for masticatory muscles (tem-
poralis and masseter) were more pronounced than reduction in nonfacial dimensions
(Larsen, 1982, 1984). These changes in craniofacial size and robusticity in prehistoric
Georgia Indians appear to be due to increased consumption of soft, maize-based foods
in later prehistory (Larsen, 1982). These findings, therefore, are strongly suggestive of
responses of the craniofacial skeleton to change in subsistence and the manner in which
food is prepared.
Strikingly similar diachronic patterns of craniofacial robusticity are revealed in a suite of
analyses from South America in the comparison of earlier foragers and farmers/pastoralists
(González-José, Ramírez-Rozzi et al., 2005; Sardi & Béguelin, 2011; Sardi et al., 2006). In the
Argentine Center-West, the southernmost region of the Andes where the foraging-to-farming
transition took place, beginning c. 3000 BC and heavily emphasized by AD 600, the dietary
transition involved the adoption of beans, maize, potato, and manioc. These domesticated
plants were ground and boiled to a soft consistency. The differences in the comparison of the
earlier foragers and later farmers are profound, especially in cranial morphology and the
remarkable reduction in the masticatory apparatus in the farmers (Sardi & Béguelin, 2011;
Sardi et al., 2006). Like the transitions in North America, this regional change in South
America appears to represent a continent-wide trend in craniofacial modifications concomi-
tant with masticatory loading and differences associated with hunter-gatherers and
agriculturalists (González-José, Ramírez-Rozzi et al., 2005).
Thus, the record of evidence indicates that the worldwide trend of cranial shortening and
gracilization is best understood in relation to masticatory, dietary, and technological changes,
especially those associated with the shift from foraging to food production and consumption
of softer foods by later populations. These changes in subsistence practices and their influence
on craniofacial anatomy are global in scope (Larsen, 1995; von Cramon-Taubadel, 2011).
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264 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
While the general trends of reduction in craniofacial robusticity – especially in the mastica-
tory complex – are clear, it is important to emphasize that there is significant variation. For
example, in the southern Levant (Israel), the transition from foraging to farming had consider-
able effects on tooth size (reduction) but only modest effects on mandibular size (Pinhasi
et al., 2008). In this regard, analysis of a series of populations representing foragers and
farmers reveals a statistically significant reduction in ramus breadth and symphysis height
only. Although these changes likely represent responses to changing loading patterns, the
modest reductions reflect local circumstances in food preparation technology and the kinds of
foods consumed. Moreover, in South Africa, there is a reduction in craniofacial size and
robusticity (c. 2000–1000 BC) coincident with increased intensification and breadth of foods
consumed (Stynder et al., 2007). However, unlike other settings described earlier where this
kind of adaptive intensification led to domestication and farming (e.g., North America,
Europe, Levant), in this setting agriculture did not arise. Rather, the craniofacial changes in
South African foragers were due to some other adaptive shift, perhaps related to the economic
intensification unique to this particular setting. Finally, it is important to emphasize that
other, non-environmental factors play an important role in understanding craniofacial
changes. In the Great Plains region of North America, for example, morphometric analyses
implicate gene flow between neighboring populations rather than masticatory loading (Jantz,
1973; Key, 1983, 1994; Ubelaker & Jantz, 1986). This underscores the important point that
cranial morphology – as with all other phenotypic variation – reflects the interaction between
genes and environment.
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7.2 Cranial form and functional adaptation 265
(AD 610–1033) with mixed forager/agriculturalists from the Middle Missouri (AD 900–1675)
and Coalescent (AD 1600–1832) traditions reveals a general gracilization of the supraorbital
torus from the earlier to later periods (Cole & Cole, 1994).
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266 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
Other high latitude foragers display pronounced craniofacial robusticity. Like Eskimos,
crania from Tierra del Fuego and Patagonia, South America, bear robust supraorbital tori
and anteriorly placed zygomas, sagittal keeling, occipital tori, and pronounced attachment
sites for the temporalis muscle (Lahr, 1995). The functional-masticatory paradigm (Hylander,
1977; Spencer & Demes, 1993) best explains the similarities between Eskimos and Fueguian/
Patagonians, especially with regard to the common skeletal responses to highly demanding
masticatory regimes in these two different settings.
Some workers have argued that the round head of high latitude populations would be best
suited for cold adaptation, indicating that a sphere maximizes volume for heat retention and
minimizes surface area for heat loss prevention (Beals, 1972; Beals et al., 1984; Crognier,
1981). However, the aforementioned studies linking craniofacial morphology with trends in
masticatory loading and dietary change in the Holocene and changes in head shape in the
absence of climatic change makes thermoregulatory models or similar types of ecological
arguments less than compelling (Armstrong, 1984; Gibson, 1984; Henke, 1984; Henneberg,
1984; Lahr, 1995; Morimoto, 1984). In North America, analysis of cranial dimensions based
on thousands of native individuals measured by Boas in the late nineteenth century shows no
relationship between climate and head shape (Jantz et al., 1992; and see De Azevedo et al.,
2011). Finally, the cold stress model has limited explanatory power because a wide range of
other human populations with round heads live in warm climates, and various populations,
living and extinct, having robust, forwardly placed molars are associated with hot, dry
climates (Hylander, 1977). Moreover, recent global analyses show that climatic adaptation
does not explain cranial shape differences at a global level but that it might explain thermo-
regulatory adaptation in high-latitude populations (Harvati & Weaver, 2006; Hubbe et al.,
2009; Noback et al., 2011; von Cramon-Taubadel, 2009a, 2011).
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7.2 Cranial form and functional adaptation 267
Figure 7.4 Superior (top) view of large mandibular tori: Kodiak Island, Larsen Bay, Alaska,
pre-AD 1000. (Photograph by G. Richard Scott.) (A black and white version of this figure
will appear in some formats. For the color version, please refer to the plate section.)
There are relatively high frequencies of palatine tori in far northern and circumpolar
populations, including Icelanders (Hooton, 1918), Lapplanders (Schreinder, 1935), and Eskimos
(Hylander, 1977; and prior discussion). Torus prevalence is also high in Medieval Norse living in
Iceland and Greenland in comparison with Europeans generally (Mellquist & Sandberg, 1939;
Pedersen, 1944). In order to address the question of why Medieval Norse and indigenous Arctic
groups have developed a convergence in torus expression with Eskimos and other circumpolar
groups, Halffman and coworkers (1992; Scott et al., 1992) studied torus frequency and size in a
series of Medieval Norse skeletal remains from Norway, Greenland, and Iceland dating to the
eleventh through fourteenth centuries. Temporal comparisons reveal that later Norse from
Norway, Greenland, and Iceland have a significantly higher prevalence of tori than do early
Norse from Greenland. Tori prevalence approaches 100% in later populations, which is among
the highest in the world (compare with Halffman et al., 1992:156; Hauser & DeStefano, 1989).
Tori increase in size in older adults, suggesting that the trait is strongly influenced by age. The
pattern of increase in tori frequency and size within a relatively short temporal span (several
hundred years) suggests that environment in general and masticatory loading in particular
exert more influence on torus expression than does inheritance.
Changes in subsistence and food preparation techniques provide some insight into possible
reasons for the secular increase in tori in Greenland. Archaeological evidence indicates that
the subsistence economy of later Greenlandic Norse became increasingly focused on wild
game (e.g., seals and caribou), rather than domestic animals (cattle, sheep, and goats). Easily
chewed foods such as grains and breads became increasingly scarce in the later Norse in
Greenland. Archaeological evidence indicates a decrease in cooking of food due to declines in
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268 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
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7.2 Cranial form and functional adaptation 269
vault circumference, length, bizygomatic breadth, and head breadth. One of the most dynamic
areas of size increase is height of the lower face, including the mandible, with relatively little
change in the upper face (Forsberg et al., 1991). Interestingly, this is also the most dynamic
area of adaptation in masticatory stress as demonstrated by experimental and population
studies (see prior discussion). The results of this process are documented in cross-sectional
studies involving comparisons of individuals of different ages within the same population
(Baer, 1956; Goldstein, 1936; Hooton & Dupertuis, 1951; Howells & Bleibtreu, 1970; Hrdlička,
1936; Lasker, 1953; Todd, 1924; Walker, 1995; and see Albert et al., 2007), and in longitudinal
studies comparing sequential observations of the same individual at different ages (Behrents,
1985; Israel, 1968, 1973, 1977; but see Tallgren, 1974). Overall, these investigations show the
dynamic nature of craniofacial architecture during adulthood, especially with regard to
increases in craniofacial dimensions, even in later years when muscle mass reduces (Albert
et al., 2007; Garn, 1985).
Most investigations of bone apposition are based on industrialized urban populations, and
thus represent a narrow perspective on human variation. Studies of archaeological samples
are important in that they provide a wider context for assessing the roles of genes and
environment. In this regard, Ruff (1980) compared a large sample (n¼136) of adult males
from the Archaic-period Indian Knoll, Kentucky series. Statistical comparisons of younger
(20–34 years of age) with older (35–50 years of age) males revealed that virtually all
dimensions (e.g., face width, face height) were larger in the older adults, six of which reached
significance (P<0.05). Comparisons of craniofacial dimensions in a series of adult females and
males from Late Woodland and Mississippian-period sites in west-central Illinois (Droessler,
1981) revealed similar trends to the Indian Knoll series. Comparisons of young adults (20–34
years old), middle-aged adults (aged 35–49 years), and old adults (50þ years of age) indicate
that cranial size tends to increase with age, but to a relatively greater extent in the face, and
more so in females than in males (Table 7.1). Age comparisons of crania from Europe,
Melanesia, and the American Great Plains also revealed significant increases in adult cranio-
facial size (Guagliardo, 1982b; Sejrsen et al., 1996). Increases in facial projection, interorbital
width, orbit and mastoid size (Guagliardo, 1982b), and palate width (Sejrsen et al., 1996) are
especially prominent.
Although these studies show generally increasing size with advancing age, there is some
intra-population variability, such as changes in females but not in males (Albert et al., 2007;
Droessler, 1981), earlier changes in adult males than in adult females (Walker, 1995), or lack
of age changes entirely (Lynnerup, 2001; Sjøvold, 1995). This variability, especially between
males and females, may reflect differences in tooth loss or some unknown factors (Droessler,
1981). The congruence of results from studies of populations representing very diverse
lifestyles and ecological settings suggests some common factor or factors – genetic and/or
environmental – that influence bone apposition during adulthood.
Baer (1956) suggested three possible explanations: secular change, selective survival, and
true ontogenetic development. In working with archaeological samples, special problems
unique to these kinds of data sets emerge. For example, archaeological series usually include
multiple generations spanning many years and differing adaptive regimes. The west-central
Illinois series, for example, encompasses about 600 years of occupation with a major dietary
shift from foraging to farming (Droessler, 1981). The consistency of craniofacial changes with
advancing age in investigations of modern (clinical) and archaeological populations is
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270 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
Table 7.1 Age comparisons of west-central Illinois adult cranial measurements: young (20–34
years), middle (35–49 years), old (50þ years) individuals (Adapted from Droessler, 1981. All
measurements in millimeters)
Young (n¼54)a Middle (n¼58)a Old (n¼39)a
striking, suggesting that the trends are more real than apparent. As discussed previously,
biomechanical factors likely influence craniofacial expansion, including changes with age
due to the cumulative forces associated with mastication.
Regardless of cause, the increasing expansion of the craniofacial complex with advancing
age underscores the importance of considering the demographic composition of skeletal and
clinical series when making comparisons and drawing inferences about craniofacial morph-
ology. Comparison of a predominantly young adult series from one population with an old
adult series to another might give the impression of differences in robusticity. Consideration
of age structure, therefore, represents a vital part of skeletal analysis. These craniofacial size
differences are most apparent in the comparison between very young and very old adults
(Behrents, 1985). Most archaeological samples contain relatively few older adults. Thus,
differences in craniofacial robusticity between different archaeological series are likely due
to factors other than age.
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7.3 Dental and alveolar changes 271
space for all teeth (e.g., tooth impaction), and (2) underdevelopment or overdevelopment of
the maxillary dentition relative to the mandibular dentition or vice versa (e.g., overbite and
overjet) (Corruccini, 1991; Proffit et al., 2013; Scott & Turner, 1988).
The causes of malocclusion have been debated for years, some workers invoking genetic
explanations (Hrdlička, 1922b; Lundström, 1948; Smith & Bailit, 1977) and others offering
environmental explanations (Begg, 1954; Hunt, 1960, 1961; Sagne, 1976). Comparisons of
monkeys (squirrel monkeys, macaques, and baboons) fed hard and soft diets reveal significant
differences in occlusion; animals fed soft diets tend to have narrower maxillary arches, higher
frequencies of tooth impactions, rotations, and crowding than do animals fed hard diets
(Beecher & Corruccini, 1981; Corruccini and Beecher, 1982, 1984). Human studies are consist-
ent with these findings. In Western societies where highly processed foods make up a consider-
able part of diet, the prevalence of malocclusions exceeds half of the population (Evensen &
Øgaard, 2007; and see Proffit et al., 2013). In contrast, non-Western societies consuming
traditional diets composed of hard-textured foods have a very low prevalence of occlusal
abnormalities (Corruccini et al., 1981, 1983; Hunt, 1961; Lombardi & Bailit, 1972; Lu, 1977;
Moorrees, 1957; Price, 1936). In addition, Corruccini and Whitley (1981) showed that older
Euroamericans in rural Kentucky who had been raised on traditional diets (e.g., dried pork,
heavy corn-bread, wild garden foods) have a low prevalence of malocclusion. In contrast, high-
status South Asians who have greater access to soft, refined foods than low-status South Asians
have an elevated prevalence of occlusal abnormalities (Corruccini et al., 1983). Similarly, urban
Greek populations consuming nontraditional diets dominated by soft, processed foods have
more dental crowding and non-edge-to-edge bite than rural Greek populations consuming
traditional diets (Angel, 1944). These changes reflect the secular trend of increased malocclu-
sions (Corruccini, 1984). Similarly, comparisons of Medieval Norwegians with modern
Norwegians reveal a relatively low prevalence of malocclusions (35%) in the former compared
to the latter (65%), with females showing greater prevalence than males in the Medieval series
but no difference in modern Norwegians (Eversen & Øgaard, 2007).
Populations shifting to highly processed, nontraditional diets show an increase in overbite,
overjet, impactions, crowding (Brown, 1992; Corruccini, 1984; Corruccini & Choudhury,
1985; Corruccini et al., 1981, 1990; Moorrees, 1957; Price, 1936; Waugh, 1937; Wood,
1971), and a general narrowing of the arches (Corruccini & Whitley, 1981; Goose, 1962,
1972; Lundström & Lysell, 1953; Lysell, 1958). The latter findings suggest that the
narrower faces of American-born children in comparison with their European-born parents
(Boas, 1912, 1916; Hrdlička, cited in Boas, 1916) resulted from the shift to softer, more
processed foods.
The association between malocclusion and consumption of soft-textured foods in animal
and human studies supports the “disuse” hypothesis; namely, reduction in masticatory
mechanical loading has led to a reduction in growth of bone tissue supporting the teeth.
Certainly, some populations appear to have a genetic predisposition for occlusal abnormalities
(Corruccini, 1991), and there is definitely a genetic component to dentofacial abnormalities in
general (Carlson, 2005). However, overwhelming evidence indicates that reduction in masti-
catory stress has engendered an elevation in occlusal abnormalities in recent human history.
From this, it can be implied that tooth size has reduced over the course of human evolution
(and see later) at a relatively slower rate than that of the supporting bony structures of the
maxilla and mandible. Differential reduction in tooth and jaw size has been documented in
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272 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
recent human populations. Comparisons of British orofacial and dental dimensions spanning
the Neolithic to the nineteenth century show a distinctive reduction in bone dimensions (e.g.,
mandible, palate width) but no or little reduction in tooth size (Goose, 1963; Keith, 1924;
Moore et al., 1968). Similarly, comparisons of Medieval and recent Swedish skulls show
reduction in mandible size but not in tooth size (Sagne, 1976). These findings indicate that
temporal increases in crowding are essentially reductions in bone size without corresponding
decreases in tooth size. This differential reduction is due to the fact that, unlike bone, teeth
cannot respond to changes in use via differential growth. Thus, while jaw size reduces in
response to dietary changes, tooth size is not able to do so.
Findings based on temporal comparisons from several archaeological contexts are consist-
ent with the disuse hypothesis concerning the replacement of hard-textured by soft-textured
foods in the foraging-to-farming transition or within increasing urbanization. Comparison of
Archaic hunter-gatherers with Mississippian agriculturalists from Koger’s Island in the Pick-
wick Basin, Tennessee, reveals an increase in occlusal abnormalities in the agriculturalists
relative to the hunter-gatherers (Newman & Snow, 1942). Because there was also a decline in
tooth wear in this setting, Newman and Snow (1942) concluded that dietary consistency
declined due to the use of wooden mortars and pestles in food preparation in later prehistory,
thus indirectly implicating decreased masticatory stress as a factor contributing to increased
prevalence of malocclusion.
In the eastern Mediterranean, decrease in edge-to-edge bite from the Neolithic to the mid-
twentieth century also coincides with a reduction in tooth wear, which led Angel (1944) to
infer a decrease in mechanical demand on the faces and jaws. The pattern was also linked to a
decline in craniofacial robusticity, accelerating dramatically in recent Greek populations.
The trend of increasing prevalence of occlusal abnormalities has been documented in a
variety of settings and interpreted in various ways (compare with Davies, 1972; Dickson,
1970). This trend is well documented in archaeological remains from Japan, especially in view
of changes in diet, food preparation technology, and orofacial adaptation (Hanihara et al.,
1981; Inoue et al., 1986; Ito et al., 1983; Kaifu, 1999, 2000; Kamegai et al., 1982). Compari-
sons of adult skulls from the Jomon (1000–500 BC), Yayoi, Kofun (AD 50–350), Medieval
(AD 1300–1600), Yedo (AD 1600–1900), and modern (AD 1964–1966) periods reveal clear
trends in orofacial adaptation (Hanihara et al., 1981; Kaifu, 1999, 2000). Frequency of
malocclusion increased from 20.0% in Jomon hunter-gatherers to 45.5% in Kofun incipient
agriculturalists, to 76.2% in modern Japanese. Building on these findings, Inoue and
coworkers examined the relationship between food consistency and occlusal abnormalities
in Japan, succinctly arguing that “soft and highly nutritious food reduces the functional
activity of the human masticating system, because it requires less chewing force and less
chewing time. Thus reduction of the jaw bone has progressed through the course of human
micro-evolution, resulting in disharmony between the sizes of teeth and jaws” (Inoue et al.,
1986:164; and see Suzuki, 1969). To test this hypothesis further, malocclusion prevalences of
Jomon-period foragers (divided into an early and late sample), Yayoi-period early farmers,
and Kofun-period protohistoric farmers mostly from southern Japan were compared. Factor,
principal components, and cluster statistical analyses indicated a grouping of occlusal pat-
terns based on subsistence categories. This analysis revealed a low prevalence of malocclu-
sions in the foragers followed by increases in the later, agricultural populations with increased
use of soft foods. Among the most striking temporal changes in this complex of craniofacial
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7.3 Dental and alveolar changes 273
changes is protrusion of the anterior dentition, overbite, and increasing alveolar prognathism
in later Japanese relative to earlier populations (Kaifu, 1999, 2000). Almost certainly, highly
distinctive changes are related to decreasing masticatory loading, perhaps relating to the
greater reduction of supporting bony architecture relative to large anterior teeth. However, the
role of anterior tooth wear – especially its considerable reduction and the associated conse-
quences for increasing malocclusion – is an essential component for understanding these very
significant craniofacial changes in East Asia and elsewhere.
Collectively, the record of skeletal, tooth size, and tooth wear research reveals a complex
interaction of bone, tooth size, and rate of dental attrition. It may be the case that a number of
epigenetic mechanisms are responsible for the integration of craniofacial growth, tooth
growth, and tooth wear (Lieberman, 2011). While it is the case that tooth size has a higher
genetic component than the masticatory skeletal apparatus, permanent tooth size nonetheless
has a significant environmental component (Harris & Rathbun, 1991; and see discussion in
Lieberman, 2011). Tooth size has decreased over the last 1000 years (Brace et al., 1991).
Moreover, in China, posterior tooth size has increased in just the last several generations
relating to improved nutrition and increased body size (Harris et al., 2001). Lieberman (2011)
speculates that just as chewing of hard food stimulates jaws and tooth roots to become larger,
such a stimulus may also act on tooth crowns during their development to become larger.
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274 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
(a)
1600
1500
Summed Posterior Areas (mm2)
1400
1300
1200
1100
1000
900
800
al . .
lis us od od
abi ect r th m m
.h . er n de rly te
H ea la
H ea
N
Hominid
Figure 7.5a Reduction in summed posterior tooth size in Homo from early species to
anatomically modern humans. (Data from Calcagno & Gibson, 1991.)
breadth is consistent with other analyses of Holocene dental size reduction in populations
undergoing the shift from foraging to farming in Europe (Brace et al., 1987; Pinhasi &
Mieklejohn, 2011) and northeast Africa (Calcagno, 1989).
Yet, mechanisms for tooth size reduction remain elusive (see Kieser, 1990, and Pinhasi
et al., 2008 for review of alternative models explaining tooth size reduction). Tooth size
appears to be highly heritable, suggesting that size reduction may be largely an evolutionary
(genetic) change. The rapid reduction in tooth size accompanying shifts in dietary focus and
other circumstances (Harris et al., 2001; Hinton et al., 1980; Larsen, 1982, 1983; Sciulli, 1979)
or the presence of small teeth in physiologically stressed individuals (Garn & Burdi, 1971;
Garn, Osborne et al., 1979; Kieser, 1990; Townsend & Brown, 1978) underscores the influence
of environment (e.g., nutrition, maternal health). The greater heritability of tooth size than
bone size and the fact that teeth do not remodel once formed underlies the potential imbal-
ance between teeth and supporting bone, as is so well illustrated in the aforementioned studies
of occlusal abnormalities.
Large teeth located in proportionately small alveolar tissue also represent a potential health
risk, predisposing an individual to impactions, dental caries, periodontal disease, and tooth
loss (Calcagno & Gibson, 1991; Inoue et al., 1986). Caries and dental impactions can also lead
to abscesses and localized infections, which can progress to more severe types of systemic
infections (e.g., gangrene, septicemia, osteomyelitis; Calcagno & Gibson, 1991). Because
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7.3 Dental and alveolar changes 275
(b)
UP3BL
12.5
UM2BL
12 LP4BL
11.5
11
Tooth length (mm)
10.5
10
9.5
8.5
7.5
Natufian
Natufian
Natufian
PPNA
PPNB
PPNC
Middle
Early
Final
Period
Figure 7.5b Reduction in upper premolar (UP3BL), upper second molar (UM2BL), and lower
premolar (LP4BL) lengths from Early Natufian (preagricultural) to Pre-pottery Neolithic C
(PPNC) periods (agricultural). (Data from Pinhasi et al., 2008.)
(c)
14
14
12 Male mandibular teeth Male maxillary teeth
12
10
Breadth (mm)
Breadth (mm)
10
8
8
6 6
Final Paleolithic Final Paleolithic
4 Agriculturalist Agriculturalist
4
Intensive Agriculturalist Intensive Agriculturalist
2 2
0 0
I1 I2 C P3 P4 M1 M2 M3 I1 I2 C P3 P4 M1 M2 M3
12 14
Female mandibular teeth 12 Female maxillary teeth
10
Breadth (mm)
10
Breadth (mm)
8
8
6
6
4 Final Paleolithic Final Paleolithic
Agriculturalist 4 Agriculturalist
2 Intensive Agriculturalist
2 Intensive Agriculturalist
0 0
I1 I2 C P3 P4 M1 M2 M3 I1 I2 C P3 P4 M1 M2 M3
Figure 7.5c Significant reduction in breadth of all teeth between Nubian Final Paleolithic
(preagriculturist) and agriculturalist populations. (Data from Calcagno, 1986a.)
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276 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
(d) 140
EUP
120 LUP
MESO
100
Vexin Medieval
Tooth area (mm2)
80
60
40
20
0
C P3 P4 M1 M2 M3 C P3 P4 M1 M2 M3
Mandible Maxilla
Teeth
Figure 7.5d Overall decreases in mandibular and maxillary tooth areas of populations from
Early Upper (EUP) and Late Upper (LUP) Paleolithic periods to Mesolithic (MESO) and
Merovigian (Vexin Medieval) periods. (Data from Frayer, 1977.)
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7.4 Dental wear and function 277
with grit inclusions within the food. Wear can also occur on adjacent surfaces between teeth as
well as food movement against the cheeks and tongue, which causes a relatively small amount
of wear, enough to obliterate perikymata, for example (Lucas, 2004). Over the course of human
evolution, there has been a shift away from the use of the teeth toward the use of the hands in
the manipulation of the environment and a reduction in the importance of mastication in the
consumption of processed foods. It is also true, however, that the presence of significant
amounts and variable patterns of occlusal surface wear, especially, in past and living human
populations, indicates the continued importance of teeth in survival and adaptation.
Two forms of wear, abrasion and attrition, are most commonly discussed (Kaidonis et al.,
1993; Lucas, 2004; Townsend et al., 1994). Abrasion is caused by contact between the tooth
and food or other solid exogenous materials, especially as food is forced over occlusal
surfaces. Attrition is caused by tooth-on-tooth contact in the absence of food or various
other abrasives. Additionally, erosion – the loss of tooth surfaces due to chemical dissolution –
is sometimes considered as a form of wear (Davis & Winter, 1980; Linkosalo & Markkanen,
1985). Because of the difficulty of distinguishing between abrasion, attrition, and erosion, this
discussion regards “wear” as any combination of abrasion, attrition, and erosion.
Tooth wear shows considerable variation between human populations. Owing to localized
behavioral characteristics, and differences in cultural practices, age, sex, diet, and orofacial
morphology, it provides enormously important information on earlier foodways and masti-
catory behavior (Benfer & Edwards, 1991; Clement & Hillson, 2012; Hillson, 2005; Molnar,
1971, 1972; Molnar & Molnar, 1990; Molnar et al., 1983; Powell, 1985; Richards, 1990;
Richards & Miller, 1991; Rose & Ungar, 1998; Walker et al., 1991). The significance of dental
wear in relation to diet was succinctly summarized by Walker, who noted that “[f]rom an
archaeological standpoint, dietary information based on the analysis of [wear] is of consider-
able value since it offers an independent check against reconstruction of prehistoric subsist-
ence based on the analysis of floral, faunal and artifactual evidence” 61978:101).
Dental wear variation is largely described and analyzed according to its severity or form. In
addition to occlusal surface wear, wear can involve substantial losses in the regions of contact
between teeth (Begg, 1954; van Reenen & Reinach, 1988; Wolpoff, 1971), resulting in
reduction in length (mesiodistal). Although severe wear can predispose a tooth to disease
(e.g., pulpitis, caries) and loss, tooth wear is a normal physiological process showing predict-
able modifications rather than a disease or pathological state. Viewed from this perspective,
tooth wear is a functional adaptation to stresses placed on teeth during mastication.
The physiological process of tooth wear is well understood. With regard to the most
common type of wear, occlusal surface wear, it commences with loss of occlusal enamel
and often the loss of cusps and crown height. If dentin is exposed, there is deposition of
secondary dentin serving as a protective zone overlying the pulp chamber. In extreme forms
of wear, all enamel is removed from the occlusal surface, leaving a nearly continuous surface
of dentin surrounded by a partial or complete rim of enamel. As the process continues in older
adults, the nerve supply in the pulp chamber withdraws toward the root tip and is replaced in
the pulp chamber by dentin. This replacement process is essential because it accommodates
extreme wear – in some populations, involving complete loss of crown height (Hartnady &
Rose, 1991; Larsen & Kelly, 1995). Moderate to severe wear results in a reduction in tooth size
due to the combined effects of occlusal and interproximal wear (van Reenen, 1982). In
especially severe wear environments, loss of crown size in molars appears to be compensated
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278 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
Figure 7.6 Lingual tilting of mandibular first molar; Golovin Bay, Alaska, protohistoric.
(Photograph by G. Richard Scott.) (A black and white version of this figure will appear in
some formats. For the color version, please refer to the plate section.)
for by a combination of extra-eruption and lingual tilting (Clarke & Hirsch, 1991; Comuzzie &
Steele, 1989; Reinhardt, 1983; Taylor, 1963, 1986a, 1986b) (Figure 7.6). The added chewing
surface on the buccal aspects of the molar roots in these teeth likely represents an adaptive
response to severe wear by maintaining occlusal surface area commensurate with the dietary
or masticatory needs of the individual.
Severity of wear is highly influenced by the consistency and texture of food, which is
determined by either the characteristics of the food (e.g., presence of phytoliths or cellulose in
plants), the manner of its preparation, or some combination of these. In order to facilitate the
ready availability of nutrients to the digestive enzymes or to remove undesirable constituents
(e.g., fiber, toxins), humans process plants in a variety of ways, including grinding, pounding,
grating, soaking, leaching, drying, heating, and fermenting (Stahl, 1989). Some of these
processing techniques may involve the introduction of abrasive elements that promote tooth
wear (e.g., use of grinding stones for making flour from cereal grains), or they may involve the
removal of abrasives as in highly processed foods consumed by Western industrialized
populations. The following sections discuss wear as documented by two primary means,
namely through visual inspection of gross or macrowear and observation of microwear.
7.4.1 Macrowear
Some of the most comprehensive information on the behavioral significance of tooth wear is
from temporally successive series of human remains representing populations that had
undergone shifts in diet. Investigation of tooth wear and the mechanical environment reveals
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7.4 Dental wear and function 279
a high degree of consistency with skeletal indicators of masticatory stress. Populations with
high levels of mechanical demand and/or reliance on abrasive diets have relatively advanced
wear (Barondess & Sauer, 1985; Clement & Hillson, 2012; Hansen et al., 1991; Hartnady &
Rose, 1991; Hemphill, 1992; Marks et al., 1988; Molnar et al., 1983; Powell & Steele, 1994;
Scott et al., 1992; Walker, 1978). Traditional populations undergoing shifts from diets con-
taining tough foods to diets dominated by processed foods during the twentieth century show
reduction in wear (Davies & Pedersen, 1955; Staz, 1938). Important differences in interprox-
imal wear and specific patterns of wear also help inform our understanding of masticatory
behavior in human populations.
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7.4 Dental wear and function 281
Neolithic
7
Natufian
6
Attrition Level (1–10)
0
I1 I2 C P3 P4 M1 M2 M3 I1 I2 C P3 P4 M1 M2 M3
Mandible Maxilla
Figure 7.7 Changes in dental attrition comparing Natufian (preagricultural) and Neolithic
(agricultural) populations in the southern Levant. The transition to an agricultural
subsistence refocuses attritional wear from anterior to posterior dentition. However, a
general reduction in attritional levels can be observed from Natufian to Neolithic periods.
(Data from Eshed et al., 2006.)
Elsewhere in Asia, limited evidence from a handful of settings shows a general pattern of
decreasing severity of wear with the foraging-to-rice farming transition or rice intensifica-
tion, for example in Japan (Inoue et al., 1986; Kaifu, 1999) and China (Liu et al., 2010;
Pechenkina et al., 2002). Similarly, in Neolithic Thailand, there is a decline in severity of wear
inferred from the prevalence of individuals with advanced occlusal surface wear (Douglas,
2006), and in later contexts where rice cultivation and consumption intensified (Domett &
Tayles, 2007). Throughout East Asia, rice was the dominant domesticated plant, taking on a
central role of diet in later prehistory and the historic period. As with other settings globally,
the general reduction in tooth wear reflects the co-occurrence of two factors, namely the
consistency and texture of the food consumed and the manner in which it is processed for
consumption. With regard to rice and its preparation in Japan, considerable evidence reveals
that rice, when boiled, is soft textured, and other treatments that developed over the course of
time served to reduce the texture to being remarkably soft. Moreover, experimental evidence
of food preparation practices that were reconstructed from ancient Japanese periods and the
food fed to students revealed decreased chewing time and masticatory cycles (Kaifu, 1999).
Moreover, Kaifu (1999) suggests that the adoption of chopsticks first invented in China and
then adopted in Japan sometime between the third and seventh centuries AD resulted in
reduced initial food preparation by the anterior teeth, thus explaining, for this setting, the
reduced anterior tooth wear in agricultural-intensive populations.
A general decline in masticatory demands with agriculture and its intensification has also
been documented via analysis of occlusal wear in northeastern Africa (Greene et al., 1967).
Mesolithic foragers in Nubia have severe occlusal wear, which reduces considerably in the
following agricultural-dependent populations (Meroitic, X-group, Christian group). This
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282 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
decline in wear had important implications for changing patterns of oral health, as the
reduction of wear coincides with an increase in dental caries and other dental pathological
conditions. Perhaps, then, the continued presence of caries-prone grooves and fissures from
reduced wear in the agricultural groups may have predisposed their teeth to increased decay
with the shift to an emphasis on plant carbohydrates (Greene et al., 1967).
Collectively, these studies underscore the very strong influence of dietary change on the
wear environment, especially in relation to the worldwide transition from food collection to
food production and the later intensification of food production with the focus on domesti-
cated plants. Analysis of dental wear in settings involving other kinds of dietary shifts, within
both hunter-gatherers (Lieverse, Link et al., 2007; Walker, 1978) and agriculturalists (Gua-
landi, 1992), demonstrates appreciable changes in masticatory behavior not involving the
foraging-to-farming transition. Early and late period populations from the Santa Barbara
Channel Islands region saw a dramatic shift in dietary focus – from an emphasis on terrestrial
foods to a reliance on marine foods after AD 1150 (Glassow, 1996; Walker, 1978). In the early
period, terrestrial foods included seeds, prepared with grinding stones, and significant quan-
tities of shellfish (Erlandson, 1994; Glassow, 1996). In the later period, a much greater
emphasis was placed on fishing and hunting of marine mammals (Glassow, 1996; Walker,
1991–1992). The changes in tooth wear in these populations are profound, including a
decrease in wear as measured by dentin exposure area on occlusal surfaces. This trend is
likely related to the change in dietary emphasis and food preparation technology. The use of
milling equipment by earlier populations contributed to relatively greater wear than in the late
prehistoric populations. Similarly, in the Lake Baikal setting of central Asia, there was an
expansion of types of foods eaten in comparison of Kitoi culture (late Mesolithic–early
Neolithic; c. 6800–4900 BC) and later Isakovo-Serovo-Glaskovo culture (middle Neolithic–
early Bronze Age; c. 4200–1000 BC). Analysis reveals consistently high levels of occlusal wear
throughout the temporal sequence (i.e., there was no change in wear over time). The lack of
change is interesting because there had been significant behavioral (increased mobility) and
dietary (increased breadth, but a relatively reduced focus on fish) change in the comparison of
the two periods (Lieverse, Link et al., 2007). On the other hand, individuals associated with
lake shore sites, in both earlier and later periods, had a consistently lower severity of wear,
suggesting circumstances involving that particular context.
Interproximal wear
Bioarchaeologists have given considerably less attention to the study of interproximal wear,
which involves flattened facets at the areas of contact between adjacent teeth. Interproximal
wear results from differential movement of teeth during chewing; the greater the mechanical
forces placed on the teeth during mastication, the greater the amount of interproximal wear
(Hinton, 1982; Wolpoff, 1971). For example, Australian Aborigines eating tough or abrasive
foods have far more interproximal wear than Europeans eating soft, processed foods (Wolpoff,
1971). Hinton (1982) documented a significant reduction in interproximal wear as one moves
in temporal succession from prehistoric Archaic- to Woodland- to Mississippian-period
premolars and molars in the Tennessee River valley (Table 7.2, Table 7.3). This trend suggests
a decrease in masticatory loading of the dentition, a finding consistent with declining
craniofacial robusticity in these populations (Hinton, 1981a). For example, temporomandib-
ular joint size reduces from the earliest to latest periods. This record reveals that although the
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7.4 Dental wear and function 283
Table 7.2 Interproximal wear-facet breadth, PM2/M1, stratified by level of occlusal wear;
prehistoric Tennessee Native Americans (Adapted from Hinton, 1982. All measurements in
millimeters)
Occlusal wear level on lower M1
3 4 5 6 7 8
Period
Archaic 4.9 5.4 5.3 6.1 6.4 7.1
(n) (8) (6) (6) (18) (31) (22)
__
Woodland 4.8 5.3 5.4 5.9 6.0
__
(n) (33) (11) (9) (22) (8)
__ __
Mississippian 4.3 4.6 4.9 4.7
__ __
(n) (43) (22) (10) (6)
Table 7.3 Interproximal wear-facet breadth, M2/M1, stratified by level of occlusal wear;
prehistoric Tennessee Native Americans (Adapted from Hinton, 1982. All measurements in
millimeters)
Occlusal wear level on lower M2
2 3 4 5 6 7 8
Period
Archaic 4.8 5.5 6.0 6.3 6.7 6.7 6.9
(n) (7) (16) (7) (5) (18) (30) (8)
__ __
Woodland 4.2 5.0 5.4 5.7 6.2
__ __
(n) (14) (32) (13) (9) (8)
__ __ __
Mississippian 4.0 4.3 5.1 4.5
__ __ __
(n) (28) (42) (3) (3)
dietary change likely had an important influence on both interproximal wear and craniofacial
robusticity, the more important change may have been related to the manner in which food
was prepared. At the time of contact, native populations were consuming various soft foods,
such as mushes, stews, and puddings (Hudson, 1976; Swanton, 1946). Archaeological and
ethnobotanical evidence of foods and food processing in the earlier periods indicates a greater
coarseness of diet requiring more mechanical force in chewing than in the later periods.
Comparisons of a temporal series spanning the shift from foraging to farming in southern
Ontario likewise show a decrease in interproximal wear (Patterson, 1984). This change is
consistent with Hinton’s model of reduction in masticatory loading in relation to consump-
tion of soft, agricultural-based foods during later prehistory in the Eastern Woodlands.
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284 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
conditions present in human remains recovered in Nubia, Jones (1910:279, 282) compared
occlusal wear patterns in Predynastic dentitions with later samples (e.g., Christian and Byzan-
tine periods), noting a clear tendency for flat (“level and even”) wear on tooth crowns in the
former, contrasting with “a marked hollowing-out of the centre of the crowns” in the latter.
In order to test the hypothesis that systematic differences in human populations in tooth
wear patterns are related to subsistence and food preparation, Smith (1984) examined flatness
of occlusal wear in hunter-gatherers versus agriculturalists on a worldwide scale. Foragers
included European and Near Eastern Middle and Upper Paleolithic, French Mesolithic, pre-
contact Australian, precontact Eskimo, and Archaic-period Native Americans from Alabama;
agriculturalists from the British and French Neolithic, Nubia, historic Britain (Iron Age,
Anglo-Saxon, Medieval), Mississippian-period Alabama, and late prehistoric American
Southwest Pueblo (Smith, 1984:42). Although the general severity of wear was similar within
the two groups, and foragers overall had relatively more severe wear than agriculturalists;
these differences were not uniform. For example, tooth wear in the Nubian agriculturalists
was as severe as tooth wear in Australian Aborigine hunter-gatherers.
In spite of a great deal of variation in diet and food preparation technology across the
samples, highly consistent differences in the angle of the occlusal wear plane on the man-
dibular first molars were documented. Agriculturalists show higher angles of occlusal surface
wear than hunter-gatherers. For the more advanced wear stages (see Figure 7.8), these
differences approach 10. Overall, hunter-gatherers have flatter wear than agriculturalists.
Moreover, wear on first molars differs by subsistence strategy: wear is cupped in agricultur-
alists and flat in hunter-gatherers (Figure 7.9; and see Eshed et al., 2006; Kasai & Kawmura,
2001; Pinhasi et al., 2008). This compelling analysis shows that wear plane angles and form
reflect the kinds of foods being eaten as well as the manner in which they are prepared. Smith
drew the general conclusion that these differences were related to greater “toughness or
fibrousness” of the diets in foragers than in farmers.
Thus, the pattern of flat and cupped wear that Jones (1910) first observed in his compari-
sons of Predynastic and Christian-era populations in Nubia reflects the distinctions between
the respective hunter-gatherer and agriculturalist groups documented in Smith’s study.
Additional confirmation of Smith’s findings has been provided in other settings undergoing
the shift from food collection to food production. In South Asia, Pastor (1992) compared gross
wear and microwear (discussed later) in a series of prehistoric hunter-gatherers’ lower first and
second molars from the Mesolithic site of Mahadaha in the Ganga River valley and incipient
agriculturalists from the Chalcolithic site of Mehrgarh in the Indus River valley. Diet in the
Mesolithic was comprised of tough, fibrous foods, including nondomestic mammals and wild
grains, roots, and other plants that were processed in stone querns. Chalcolithic populations
consumed domestic animals (cattle, sheep, goats) and several varieties of wheat and barley. In
addition to showing a marked decline in wear severity, the pattern of wear is strikingly similar
to Smith’s hunter-gatherer and agriculturalist samples for the Mesolithic and Chalcolithic,
respectively: Mesolithic teeth are worn flat at slight angles, and Chalcolithic teeth are worn at
a greater angle with distinctive cupping.
In the previously mentioned Portuguese populations, molar occlusal surfaces also show a
temporal change from flat to cupped wear as one moves from the Mesolithic to Neolithic sites
(Lubell et al., 1994). This study also serves to illustrate the potential variation of wear patterns
in human populations as wear angles in Mesolithic foragers are much greater than in later
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7.4 Dental wear and function 285
(a)
(b)
Figure 7.8 Lateral views of tooth wear in Nubian A-group agriculturalist (a) and Eskimo
hunter-gatherer (b), showing greater angle of wear plane in the former than the latter.
(From Smith, 1984; reproduced with permission of author and John Wiley & Sons, Inc.)
(A black and white version of these figures will appear in some formats. For the color
version, please refer to the plate section.)
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286 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
(a)
(b)
Figure 7.9 Occlusal view of tooth wear in Nubian X-group agriculturalist (a) and Mesolithic
hunter-gatherer (b) showing cupped occlusal wear on first molars in the former but not the
latter. (From Smith, 1984; reproduced with permission of author and John Wiley & Sons,
Inc.) (A black and white version of these figures will appear in some formats. For the color
version, please refer to the plate section.)
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7.4 Dental wear and function 287
Neolithic farmers. The reasons for the differences from Smith’s findings are largely unknown.
Lubell and coworkers (1994) speculate that differences between the Portuguese populations
and the samples studied by Smith, especially in age composition and molar crown morph-
ology, may be important considerations. Thus, while a common pattern includes flat wear in
hunter-gatherers and oblique wear in agriculturalists, variability is present in some groups
(Schmucker, 1985).
Other types of wear demonstrate significant variation in masticatory behavior in past
human populations. Analysis of prehistoric foragers and farmers from a wide range of
contexts reveals important differences in forms and severity of anterior tooth wear compared
to posterior tooth wear. In general, in hunter-gatherers, severity of anterior tooth wear is
greater than, or equal to, severity of wear on posterior teeth. In farmers, anterior wear is
appreciably less severe than posterior wear (Bennike & Alexandersen, 2007; Brown & Molnar,
1990; Dahlberg, 1963; Deter 2009; Duray, 1996a, 1996b; Eshed et al., 2006; Hinton, 1981d,
1982; Kaifu, 1999; Molnar, 1971, 1972; Schmucker, 1985; Sciulli, 1997; Tomenchuk &
Mayhall, 1979; Turner & Cadien, 1969; van Reenen, 1992; Walker & Erlandson, 1986). The
form of wear on the anterior teeth is distinctive: prehistoric foragers show a characteristic
rounded wear, whereas the agriculturalists have cupped wear (Hinton, 1981d). The consist-
ency of these findings indicates that tooth wear is mediated by masticatory behaviors. The
relatively heavier use of the anterior teeth in hunter-gatherers is in line with ethnographic
reports of tooth use in Australians and Eskimos, in both dietary and non-dietary functions.
The distinctive labially oriented rounded wear in hunter-gatherer dentitions is likely a
function of the use of the incisors and canines in various extramasticatory activities (e.g.,
hide preparation; and see following discussion). The distinctive cupping wear on agricultur-
alist incisors and canines appears to be especially prominent in individuals who have lost
posterior teeth prior to death. Thus, this anterior wear pattern may represent an alteration of
tooth use arising from the loss of posterior teeth, namely from the use of the incisors and
canines for a combination of food preparation and mastication.
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288 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
shows differences between the sexes, suggesting especially gendered patterns of wear. For
example, Euston females have a flat wear plane on the anterior teeth and maxillary premolars
(Littleton et al., 2013). Similarly, Gillman females lack heavy anterior and premolar wear, but
some males do. In contrast, in Kaurna foragers, females, but not males, expressed heavy wear
on incisors and canines (Littleton et al., 2013; Richards, 1984). These findings strongly suggest
gendered roles as they pertain to extramasticatory tooth use. Although the reasons for this
variation are unknown, they attest to the presence of highly variable wear patterns between
closely related populations in what is often characterized as a homogeneous region (Littleton
et al., 2013; Molnar et al., 1989).
Analyses of tooth wear patterning in other settings generally show more wear on occlusal
surfaces in males than females (Da-Gloria & Larsen, 2014; Douglas, 2006; Esclassan et al.,
2008; Fujita & Ogura, 2009; Kaifu, 1998, 1999; Meng et al., 2011; Pechenkina et al., 2002).
Regardless of the differences – male wear greater than female wear, or female wear greater
than male wear – the record shows evidence for clear sex differences in the wear environment,
suggesting differences in either diet or use of teeth, or both, are a part of gender identity as it
relates to masticatory and extramasticatory function.
Gross wear differences by social group and rank have been identified in past societies. In
the Medieval Edo period of Japan, members of the elite Shogun class had virtually no occlusal
surface wear, unlike lower status, non-Shogun individuals (Suzuki, 1969). This indicates a less
mechanically demanding, less abrasive diet in the elite than in the remainder of Japanese
society. The presence of narrow faces, reduced size of the maxillae and mandibles, and gracile
masticatory muscle attachment sites in Shogun individuals corroborates a reconstruction of
the consumption of soft, processed foods by the elite.
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7.4 Dental wear and function 289
Figure 7.10 Adult mandibular dentition showing occlusal surface grooves; Great Basin,
Nevada. (Reproduced with permission of John Wiley & Sons, Inc.) (A black and white
version of this figure will appear in some formats. For the color version, please refer to
the plate section.)
indicate that, as in the Great Basin setting, some type of flexible material had been passed
transversely over the anterior teeth in a repetitive and habitual fashion, especially in process-
ing materials such as sinews for bow strings or plant fibers for cordage or basketry.
Ethnographic and historical evidence provides corroborative support for possible uses of the
dentition that may have resulted in grooves. In the southwestern margin of the American Great
Basin (Death Valley), Coville (1892) described native Panamint women preparing sumac and
willow for materials to be used in making “wicker-work utensils”: A woman “selects a fresh
shoot. . .bites one end so that it starts to split into three nearly equal parts. Holding one of these
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290 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
parts in her teeth and one in either hand, she pulls them apart, guiding the split with her
fingers. . .Taking one of these, by a similar process she splits off the pith. . .leaving a pliant, strong,
flat strip of young willow or sumac wood” (Coville, 1892:358). Greenland Eskimo women pull thin
cords of animal tendon across the clenched anterior teeth in order to moisten and soften the sinew
(Pedersen and Jakobsen, 1989). Exclusively older women (>40 years of age) are responsible for
the preparation of sinew in this manner. Interestingly, in archaeological dentitions, only older
adult female dentitions display these “sinew grooves” (Pedersen & Jakobsen, 1989). The presence
of grooves in females only in Eskimos and in males only in Great Basin Native Americans suggests
that activities causing these grooves were gender-based (but see Schulz, 1977).
While a range of settings show the presence of occlusal surface grooves in anterior teeth in
New World foragers, these dental modifications are increasingly becoming recognized in Old
World foragers and farmers. Early Holocene foragers from north Africa (Uan Muhuggiag,
Libya: Minozzi et al., 2003), pre-Neolithic (Natufian) incipient farmers from Israel (Bocquentin,
2007; Eshed et al., 2006) and Jordan (Webb & Edwards, 2002), Neolithic farmers from Syria
(Abu Hureyra: Molleson, 1994) and Poland (Lorkiewicz, 2011), and Byzantine dentitions from
western Asia (Kovuklukaya, Turkey: Erdal, 2007) also display distinctive patterns of occlusal
surface grooving. In these settings, grooves may have been produced by dragging plant fibers
across the surface of the teeth as part of the process of preparing materials for basket production
(Eshed et al., 2006; Lorkiewicz, 2011; Minozzi et al., 2003; Molleson, 1994) or yarn for making
clothing (Erdal, 2007). Preparation of tough plant materials for utilitarian purposes also may
explain the presence of occlusal surface grooves in prehistoric manioc agriculturalists in
St. Thomas, US Virgin Islands (Larsen et al., 1998; Larsen, Teaford et al., 2002).
Alterations of anterior teeth involving notching of the mesial or distal occlusal margins
have been observed in southeastern US samples from Tennessee (Blakely & Beck, 1984) and
the Georgia coast (Larsen & Thomas, 1982). In the latter case, an individual shows wear on the
mesial corner of a maxillary right first incisor that resulted from extramasticatory use, such as
from clamping fishing nets or from processing plant materials. The presence of cumulative
wear indicates that the teeth were not intentionally altered for personal ornamentation, such
as with purposeful drilling.
Lingual surface wear on maxillary incisors and canines is another highly distinctive pattern
of tooth wear involving extramasticatory behaviors (Figure 7.11). Most (39/46) adult crania
from the preceramic (4200–3000 yBP) site of Corondó, Rio de Janeiro State, Brazil, display
pronounced lingually oriented flat wear (Turner and Machado, 1983). There is no corresponding
wear on the lower teeth. Called lingual surface attrition of the maxillary anterior teeth
(LSAMAT), the orientation of wear striations strongly suggests that some kind of extramasti-
catory use of the teeth caused the wear pattern. Its presence in adult females and males suggests
that it is not associated with a gender-specific activity. The presence of moderate lingual
wear in anterior teeth of older children (beginning at about 10–11 years of age) indicates the
time in life when the behaviors that cause the wear commence. Given the high prevalence of
caries in this group (10.7% of teeth) and the known heavy reliance on manioc – a cariogenic
carbohydrate requiring extensive preparation prior to consumption – the teeth appear to have
been used to process plants for consumption (Turner & Machado, 1983). Support for this
argument has been provided by the study of additional prehistoric dentitions from other New
World settings, including Panama (Irish & Turner, 1987), Belize (Saul & Saul, 1991), Texas
(Hartnady & Rose, 1991), and the US Virgin Islands (Larsen et al., 1998; Larsen, Teaford et al.,
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7.4 Dental wear and function 291
2002). In these settings, the form of wear, reliance on manioc (or other fibrous plants), and
high prevalence of dental caries argue that these modifications are due to the habitual use of the
upper teeth in preparation of abrasive plant material. Because these studies indicate that wear
occurs mostly on the upper teeth, the material being manipulated must have been placed between
the upper teeth and tongue and drawn across the upper teeth in a back-to-front motion. With
regard to the Brazilian populations, Turner and Machado (1983) suggest that the upper teeth were
used to shred or peel tuberous plants (e.g., manioc) “comparable to the modern way we eat
artichokes – by pulling and planing the edible petals across the occlusal surfaces of our anterior
teeth” (1983:128; and see Irish & Turner, 1987). Microwear analysis of incisors from the Tutu
population from the Virgin Islands shows that minute striations caused by the passing of material
across the tooth surface involved a back-to-front motion (Larsen et al., 1998), thus adding
additional confirmation of Turner and Machado’s artichoke hypothesis.
A similar pattern of lingual wear in maxillary anterior teeth has been identified in a range
of Old World settings, including eastern Europe (Lorkiewicz, 2011), the Mediterranean basin
(Harper & Fox, 2008), Africa (Irish & Turner, 1997), western Asia (Lukacs & Pastor, 1988), and
northern China (Liu et al., 2010; Meng et al., 2011; Miao et al., 2013; Pechenkina et al., 2002).
Each of these settings has very different dietary and subsistence technology, suggesting
alternative uses of the lingual surfaces of teeth in extramasticatory functions. For example,
Lukacs and Pastor (1988) suggest that lingual wear in a Neolithic dentition from Mehrgarh,
Baluchistan, resulted from the use of teeth for the processing of animal skins. However, the
strong association between caries and lingual wear in at least one setting in historic-period
West Africa suggests a link between high carbohydrate diet (manioc) and dental processing
(Irish & Turner, 1997). On the other hand, the significant presence in low-carbohydrate
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292 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
contexts reveals the varied associations of this behavioral marker (Miao et al., 2013; Pechen-
kina et al., 2002). It is likely that this form of wear relates to preparation of fibers generally,
for either dietary purposes (the dominant pattern in the New World) or material culture.
7.4.3 Microwear
Microwear, or microscopic damage on teeth by materials processed in the oral cavity, provides
essential insight into dietary adaptation and masticatory function. The development of a number
of tools for accessing intricacies of dietary adaptation, tooth use, and masticatory behavior make
possible more effective ways of understanding use wear (Teaford, 1991; Ungar et al., 2008). The
depth of focus and highly detailed resolution in SEM analysis (Teaford, 1991), and more recently,
microwear texture analysis (Ungar et al., 2008) have expanded our understanding of diet and
tooth use for a wide range of primates, including humans (Figure 7.12). Because the analysis of
microwear features (e.g., pits and scratches) requires high magnification (e.g., 500), only small
areas can be assessed at any one time. Texture analysis, on the other hand, provides a more
(a)
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7.4 Dental wear and function 293
(b)
(c)
Figure 7.12c Confocal microscope 3D image ( 100) of same location on same tooth. (Image
courtesy of Melissa Zolnierz and Christopher Schmidt.) (A black and white version of this
figure will appear in some formats. For the color version, please refer to the plate section.)
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294 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
species. The key limitation of microwear analysis is that it provides a record of tooth use that
documents masticatory or dietary behavior for the time immediately prior to death, from
hours to a few days, otherwise known as the “Last Supper” phenomenon (Grine, 1986).
However, the generally high degree of consistency in various analyses indicates the import-
ance of this kind of study in assessing patterns of tooth use and in drawing inferences about
masticatory behavior (Teaford, 1991; Ungar et al., 2008).
Experimental research indicates that teeth of animals and humans fed soft or nonabrasive diets
have fewer microwear features than animals and humans fed hard or abrasive diets (Teaford,
1991; Teaford & Lytle, 1996). Like macrowear, these differences are influenced by the nature of
the food itself and by extraneous elements, such as soil particles, which may be incorporated in
the food prior to its consumption. Microwear variation in archaeologically recovered humans
indicates differences related to food texture and diet within specific groups (Danielson &
Reinhard, 1998; Fine & Craig, 1981; Ma & Teaford, 2010; Mahoney, 2007; Marks et al., 1988;
Organ et al., 2005; Puech et al., 1983) or in relation to subsistence change (Bullington, 1991; Hojo,
1989; Homes Hogue & Melsheimer, 2008; Mahoney, 2005, 2006, 2007; Rose, Anton et al., 1991;
Schmidt, 2001; Teaford, 1991; Teaford et al., 2001; Ungar & Spencer, 1999).
Occlusal microwear on molars from Early Mississippian-period pre-maize populations and
Middle Mississippian-period (Zebree site: AD 900–1200) maize-dependent populations in the
central Mississippi River valley underwent a change from the presence of numerous large
striations to few large striations; occlusal surfaces go from being rough to smooth (Rose,
Anton et al., 1991). Rose and coworkers (1991:17) interpret the striking change in surface
topography as indicating “a radical reduction in abrasive particles associated with changes in
both food content and preparation.” They note increased prevalence of dental caries, from 2.6
to 3.5 lesions per dentition, and less negative δ13C values, indicating that the microwear
alterations resulted from a change in food consumption.
Similarly, an association between dietary change and microwear has been documented in a
temporal series from the southeastern US Atlantic coast and interior northern Florida,
including precontact hunter-gatherers, later precontact farmers, and early and late mission-
era populations living in Catholic missions (Teaford, 1991; Teaford et al., 2001). The pre-
contact populations underwent a transition from foraging to maize farming in late prehistory,
and the later contact populations were largely intensive maize agriculturalists. Overall,
precontact molars display narrower scratches on the occlusal surfaces than do the mission
molars. These changes in microwear reflect a reduction in abrasiveness of diet during the
contact period, suggesting that maize-based diets were of a softer-textured nature than the
foraging-based diets. On the other hand, the analysis revealed considerable variation in
microwear features in the region, reflecting the highly localized aspects of diet and habitat,
patterns which reflect similar conclusions drawn in the analysis of carbon and nitrogen stable
isotopes (Hutchinson et al., 1998; Larsen, Griffin et al., 2001). At San Luis de Apalachee (AD
1656–1704), an economically important center at the western end of the mission system in
Spanish Florida, molars express an unusually high volume of surface pits and scratching
(Organ et al., 2005). Historical sources point to the importance of cattle in this setting and of
maize farming. However, the low prevalence of dental caries and preliminary isotopic
evidence for lower meat consumption than at other missions suggests that the unusual
microwear reflects a diet relatively higher in meat. If meat played an important role in diet,
then it would have been relatively tough, perhaps with soil and/or grit inclusions.
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7.4 Dental wear and function 295
The number of microwear features on occlusal surfaces of deciduous incisors and molars
from Middle Woodland (50 BC–AD 250) incipient pre-maize agriculturalists and Mississippian
(AD 1000–1350) maize agriculturalists in the lower Illinois River valley of western Illinois
increased with age during both periods (Bullington, 1991). Reflecting the difference in use of
anterior and posterior teeth, the incisors have relatively more scratches than pits and the
molars have more pits than scratches. These differences indicate the use of the incisors in
initial processing and the molars in crushing hard objects. The earlier group focused on wild
plants and animals in combination with a reliance on various starchy seeds and those with
hard seed coats. The later group continued to utilize these food items, but with their partial
replacement by maize. In addition, these foods were likely boiled in ceramic vessels for long
periods of time, thus reducing the toughness of foods consumed by prehistoric populations
(Bullington, 1991). Unlike the changes observed in the Mississippi River valley and the
Georgia Bight, comparisons of the Middle Woodland and Mississippian groups revealed no
differences in microwear. Bullington (1991) suggests that the lack of temporal change may
reflect the fact that her study focuses exclusively on young juveniles; other investigations
focus on adult microwear. There is a strong similarity in microwear between the foragers and
farmers overall, but the two groups are distinctive in the youngest cohort (c. 6–12 months).
Mississippian teeth have lower feature frequencies than Middle Woodland teeth. This suggests
that very young children in the later period were consuming softer foods than very young
children in the earlier period.
Similar changes in microwear in relation to the shift from hard- to soft-textured diets have
been identified in a number of settings in the Old World. Comparison of canines and molars
from two Neolithic (c. 2000 BC) and two historic (AD 1750–1800) dentitions from western
coastal Kyushu, Japan shows a change from generally long, wide striations to thin, narrow
striations (Hojo, 1989). Hojo (1989) suggests that the later diet included smaller abrasives than
did the earlier diet.
Change in occlusal microwear in relation to the shift from foraging to dependence on cereal
grains in western Asia has been investigated via comparison of dentitions from the previously
discussed Neolithic settlement at Abu Hureyra in the Euphrates River valley in northern Syria
(Molleson, 1994; Molleson & Jones, 1991; Molleson et al., 1993). Microscopic analysis of
molars provides corroborative evidence for the study of gross wear. Comparison of pit
diameter, feature frequency, pit density (percentage determined from number and size of
pits), and ratio of striations and other linear features to pits shows a clear shift in microwear in
deciduous and permanent teeth through time. The change in wear is especially conspicuous in
the shift from foraging in the Mesolithic to early agriculture in the pre-pottery Neolithic,
being accompanied by a dramatic increase in microwear feature density. Molleson and
coworkers (1993) attribute this trend to a shift from a relatively soft diet, based on the
consumption of roots and small, wild grains, to a few cereal types prepared with grinding
stone equipment. In the later Neolithic, there is a reduction in feature density and pit diameter,
reflecting a return to softer diets, namely involving cooking of food with ceramic vessels and
the increased consumption of meat. Clearly, the adoption of pottery and its use in cooking
food to a soft consistency had great implications for the manner in which teeth were worn.
The trends identified in Syria are broadly similar to the temporal trends in microwear in the
southern Levant. In this setting, analysis of a small sample of Upper Paleolithic individuals
from Ohalo II (23 500–22 500 yBP) reveals the presence and high frequency of long, narrow
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296 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
scratches characteristic of populations eating tough, abrasive diets (Mahoney, 2007). Maho-
ney (2007) speculates that this represents consumption of items requiring shearing, such as
wild plants. The infrequency of pits suggests a lack of a hard, brittle diet that would have
required strong compressive forces in the jaws and teeth. In the late Pleistocene and early
Holocene Natufian foragers, however, the molars have various distinctive large pits and wide
scratches, reflecting consumption of hard foods requiring compressive forces in chewing.
Various factors likely explain this temporal change, but the use of grinding stones and the
transfer of quartz particles to the food is the likely cause of this change from earlier popula-
tions (Mahoney, 2005, 2006, 2007). In the early Neolithic populations, the pits became smaller
and the scratches narrower than in the preceding Natufian populations, reflecting a decline in
texture and toughness of foods consumed, similar in many respects to the Upper Paleolithic
individuals from Ohalo II. However, in the later Neolithic period, the pits and scratches
became distinctively larger, a pattern suggesting an increase in toughness of foods consumed,
as in the earlier Natufian population. This pattern indicates an increase in hardness of diet in
the Levantine Neolithic. Overall, there were trends in dietary adaptation, not representing an
increase or decrease in toughness and abrasiveness, but rather, a mosaic of modifications
reflecting both types of foods consumed and how they were prepared for consumption.
A parallel development for a shift from foraging to farming has been documented via
microwear analysis in South Asia, based on the analysis of dentitions from the Mesolithic
(Mahadaha site; 8000–1000 BC), the pre-pottery Neolithic (Mehrgarh site; 7000–6000 BC),
incipient agricultural Chalcolithic (Mehrgarh site; 4500 BC), and urban agriculturalist Har-
appa culture (Harappa site; 2500–2000 BC) (Pastor, 1992, 1993). Comparative analysis of
microwear in permanent first and second molars indicates a generally similar pattern of
increase in features (pits, scratches) as Abu Hureyra. In general, trends in microwear indicate
the higher prevalence of finer scratches in the hunter-gatherers and incipient agriculturalists,
whereas microwear features become coarser in more intensive agriculturalists. The latter
characteristic reflects an increased consumption of cereal grains and use of grinding equip-
ment in the Neolithic and Chalcolithic. The use of stone grinding equipment and its influence
on microwear is illustrated by comparisons of microwear in a living American fed a typical
soft diet with the effects of sandstone-ground maize (Teaford & Lytle, 1996). These compari-
sons reveal that microwear on a maxillary molar increased by 30 times in the consumption of
ground maize over a period of a week. This study supports the observation that food
preparation technology has a profound effect on tooth wear.
Another implication for the research on economic transitions from microwear analysis and
other approaches to functional analysis of the craniofacial complex is the trend for reduction
in hardness and texture of diet, albeit with some variation (Mahoney, 2006, 2007). Presum-
ably, this tendency would be especially pronounced in those members of populations in
higher socioeconomic groups. In late eighteenth- and early nineteenth-century Baltimore,
MD, comparison of occlusal microwear from the Christ Church cemetery and Potter’s Field
East provides an opportunity to characterize and interpret patterns of tooth use in wealthy
and poor persons living in the same habitat, but with different diets (Ma & Teaford, 2010).
Individuals interred in both cemeteries would have consumed seafood, pork, and maize,
among other foods, but the foods consumed by individuals associated with higher-status
Christ Church would have been of a higher quality, prepared in sauces and wine, and they
would have had access to various fruits and vegetables, certainly more frequently than those
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7.4 Dental wear and function 297
individuals associated with Potter’s Field East (Carson, 1985; Steckel, 1999). Analysis of
microwear, however, reveals no clear differences in how this culinary variation would have
impacted tooth surfaces. That is, analysis of microwear features – pits and scratches – reveals
that while diets were free of abrasives, there are no discernible differences between the two
socioeconomic groups. It may be that traditional microwear analysis based on scanning
electron microscopy is not robust enough to provide the kind of detail on diet and tooth use
needed to discern subtle differences in foodways between the two groups. Importantly,
however, the approach clearly reveals the nature of diets in the nineteenth-century urban
setting of the mid-Atlantic: rich and poor alike enjoyed an abrasive-free diet.
Virtually all of the aforementioned microwear analyses focus on the occlusal surfaces of
teeth. Buccal surfaces have the potential to provide important perspective on tooth use because
wear on them is not influenced by contact between opposing teeth (Ungar & Teaford, 1996). The
importance of buccal surfaces is emphasized in research by Lalueza Fox and coworkers (1996)
on a wide range of past and recent human populations, including hunter-gatherers, pastoral-
ists, agriculturalists, and fossil hominins. Analysis of buccal surfaces of premolars and molars
in these samples reveals that foragers with high-meat diets have fewer and more vertically
oriented striations than agriculturalists. Greater frequency of buccal striations in plant-
consumers may be due to the presence of abrasive phytoliths (Lalueza Fox et al., 1996).
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298 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
Figure 7.13 Chipped mandibular incisors from the Cathedral of Santa Maria in Vitoria,
Spain, c. AD 1400–1500. (Photograph courtesy of G. Richard Scott.) (A black and white
version of this figure will appear in some formats. For the color version, please refer to the
plate section.)
settings is associated with oral processing and consuming frozen or otherwise tough foods
requiring substantial reduction. Unlike these groups expressing chipping mostly in the poster-
ior dentition, populations from Medieval Norway and Medieval and post-Medieval Spain
express damage but mostly in the anterior dentition (Scott & Winn, 2011) (Figure 7.14). These
differences highlight significant and important variation in how these modern populations
process food and in use of the teeth as an important tool.
As with the Inuit, damaged teeth are also relatively commonplace in other settings. For
example, at Taforalt, Morocco, chipping of teeth is present in 94% (31/33) of individuals, both
in anterior teeth (21.1%) and posterior teeth (32.8%) (Bonfiglioli et al., 2004; and compare
with Milner & Larsen, 1991). While the chipping is likely related to the use of teeth for
processing tough, perhaps uncooked foods, these terminal Paleolithic foragers also practiced
tooth avulsion involving purposeful extraction of anterior teeth, especially permanent inci-
sors. The consequent reliance on the posterior teeth for processing normally involving the
anterior teeth may have contributed to an elevated prevalence of damaged molars (Bonfiglioli
et al., 2004). The considerably greater prevalence of dental chipping in males than in females
in this and most other settings where the condition has been documented suggests a division
of labor and/or difference in diet between sexes (Scott & Winn, 2011). Similarly, prehistoric
Hawaiians display strong differences in dental chipping, but with greater frequency for
women than men (18% vs. 6%). In this setting, the higher prevalence in women is consistent
with ethnographic accounts of women habitually using their teeth for various extramastica-
tory tasks, such as cord production, leaf cutting, and nut cracking. In these populations,
maxillary and mandibular tori are also more prevalent in females than males, thus supporting
a functional interpretation of these skeletal features.
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7.4 Dental wear and function 299
90 Mandible
80
70
Chipped teeth (%)
60
50
40
30
20
10
0
LM3 LM2 LM1 LP4 LP3 LC LI2 LI1 RI1 RI2 RC RP3 RP4 RM1 RM2 RM3
80
70
Chipped teeth (%)
60
50
40
30
20
10
0
LM3 LM2 LM1 LP4 LP3 LC LI2 LI1 RI1 RI2 RC RP3 RP4 RM1 RM2 RM3
80
70
Chipped teeth (%)
60
50
40
30
20
10
0
LM3 LM2 LM1 LP4 LP3 LC LI2 LI1 RI1 RI2 RC RP3 RP4 RM1 RM2 RM3
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300 Masticatory and nonmasticatory functions: craniofacial adaptation to mechanical loading
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Isotopic and elemental signatures of diet,
8 nutrition, and life history
8.1 Introduction
Documentation of past foodways is not only intrinsically valuable, but it also provides the
requisite resource context for evaluating effects of nutrition on growth and development,
assessing stress and disease from paleopathological indicators, and interpreting skeletal
adaptation, among other topics discussed in the foregoing chapters. There is a range of
conventional approaches for characterizing past diets and inferring nutritional outcomes,
including analysis of plant and animal remains, coprolites, pottery residues, and tools used for
food production and consumption. These approaches provide a record of what foods were
consumed, but do not give a representation of the proportions of foods or food classes
consumed by past populations. The poor preservation of plants in many archaeological
contexts, for example, inhibits documentation of their role in diets in many settings. Food
refuse is often subject to preservation-related biases that prevent accurate or otherwise
meaningful nutritional interpretation. Moreover, food refuse deriving from ritual uses may
or may not have been eaten.
Beginning in the late 1970s, a collaboration between an archaeologist and a geochemist on
the analysis of stable isotope composition of carbon in archaeological human remains (van
der Merwe & Vogel, 1978; Vogel & van der Merwe, 1977) started a revolution in dietary
reconstruction having implications for issues of interest in the study of past populations, all
linked either directly or indirectly to diet. Following this pioneering research, stable isotopes
have become a standard data set for addressing fundamental questions about diet and dietary
adaptation in past populations, enhancing our ability to characterize past human diets and
their variation in general (Burton, 2008; Katzenberg, 2008; Lee-Thorp, 2008; Schoeninger,
2010). Simply, the documentation of isotope signatures passed from the foods being eaten to
the consumer allows the identification of diet with considerably greater precision than with
conventional archaeological data involving recovery of plant and animal remains alone.
These biochemical signatures do not represent a “reconstruction” of diet; rather, they facilitate
the identification of consumption profiles of different foods eaten by past populations. If
consumption profiles are identified accurately, then it becomes possible to get at the real
“meat” of the matter, namely, nutritional inferences and broader implications for understand-
ing past human adaptation in a broad biocultural context.
Regional and methodological perspectives on isotope and elemental analysis in archaeo-
logical remains and identification of dietary signatures are presented in a number of compre-
hensive reviews of the older literature (Katzenberg & Harrison, 1997; Pate, 1994; Price et al.,
1985; Sandford, 1992, 1993a; Schoeninger, 1989, 1995a; Schoeninger & Moore, 1992;
Schwarcz & Schoeninger, 1991) and newer developments building on earlier advances
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302 Isotopic and elemental signatures of diet, nutrition, and life history
(Ambrose & Krigbaum, 2003; Brown & Brown, 2011; Burton, 2008; Katzenberg, 2008; Lee-
Thorp, 2008; Schoeninger, 2010; Schwarcz & Schoeninger, 2011). This chapter discusses
isotopic and elemental analyses of archaeological human remains and the ways in which
these analyses contribute to our understanding of dietary behavior, nutritional ecology, and
milestone events in early life (e.g., weaning) and in later life (e.g., residential change). The
record is based largely on bone collagen, the structural protein that makes up most of the
organic component of bone, but other sources such as bone mineral, tooth enamel, and
calculus serve as isotope-based dietary proxies (Austin et al., 2013; Schoeninger, 2010; Scott &
Poulson, 2012). Importantly, because enamel does not remodel, the chemical signature
remains largely unchanged once the enamel is fully formed. In contrast, because bone does
remodel over the life course, its signature reflects diet and other aspects of biochemistry for
well over a decade (Hedges & Reynard, 2007; Hedges et al., 2007). Thus, an individual aged
45 years at death has a hard-tissue record of diet from both the juvenile years (teeth) and at
least the last 10 years of his/her life (bone).
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8.2 Isotopic analysis 303
18
16
Marine
14
d15NAIR ( ‰ )
12 Preagricultural
Agricultural
C4-No marine
10
8
C3-No marine
δ13C values averaging −26‰, with a range of −22‰ to −38‰ (Tieszen, 1991). In C4 plants,
plants that are typically adapted to hot and dry climates (e.g., tropical grasses like maize,
sugar cane, some amaranths, chenopods, setarias), the reduced discrimination against the
13
C isotope results in values averaging −12.5‰, with a range from about −9‰ to −21‰
(Tieszen, 1991). On average, C4 plants have δ13C values that are about 14‰ less negative than
C3 plants and their consumers (Figure 8.1). Plants with CAM photosynthesis pathways (many
cacti and succulents) have δ13C values that overlap the values in C3 and C4 plants, because
they use either a C3 or C4 pathway as determined by environmental circumstances.
The human consumers of these plants retain the differences in δ13C values, but with values
shifting approximately 1‰–4‰ in experimental settings (DeNiro & Epstein, 1978; Hare et al.,
1991; Schoeninger, 2010; Tieszen & Boutton, 1989; Tieszen et al., 1983) and 5‰ in field
investigations (Lee-Thorp et al., 1989; Schoeninger & Moore, 1992; van der Merwe, 1982)
from the food to what is observed in their bone collagen. Sometimes called the “fractionation”
factor, this offset value represents what is better termed the “diet-to-tissue spacing” or
“trophic effect,” because fractionation technically refers to specific equations that measure
the relative amount of two stable isotopes before and after a particular chemical reaction
(Katzenberg, personal communication; and see Fry, 2006). Photosynthesis is a well-known
chemical reaction, but there are a number of reactions that occur between consumption of a
particular food and the formation of bone collagen (or apatite). Moreover, fractionation in
specific reactions is a constant and it is known that diet-to-tissue spacing is not a constant.
Collagen requires essential amino acids for its formation, and therefore reflects the majority
(~60%) of the δ13C protein component of diet. On the other hand, the mineral carbonate from
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304 Isotopic and elemental signatures of diet, nutrition, and life history
bone apatite represents more of an approximation of the whole diet, which may include
carbohydrates and fats, in addition to protein not used for protein tissue synthesis (Krueger &
Sullivan, 1984; and see Ambrose & Norr, 1993; Ambrose et al., 1997; Cooke et al., 1996;
Harrison & Katzenberg, 2003; Tieszen & Fagre, 1993).
Marine plants have δ13C values that are between the values of C3 and C4 terrestrial plants
due to variation in their carbon sources, ranging from detritus of terrestrial origin (mix of
local terrestrial plants), dissolved CO2 (−7.0‰), and dissolved carbonic acid (0‰) (Schoenin-
ger & Moore, 1992). Thus, marine organisms consuming these plants have values ranging
from closer to C3 plants at one extreme to C4 plants at the other. Marine fishes and mammals
have δ13C values that are less negative (by about 6‰) than do animals feeding on C3-based
foods, and values that are more negative (by about 7‰) than do animals feeding on C4-based
foods (Schoeninger & DeNiro, 1984).
-5
-10
Mean d13 C‰ (±1 s.d.)
-15
-20
-25
-4000 -3000 -2000 -1000 0 1000
Age (BC/AD)
Figure 8.2 Temporal change in stable carbon isotope ratios in eastern North America. Error
bars indicate one standard deviation from the mean where N>1. (Adapted from Ambrose,
1987; courtesy of the Center for Archaeological Investigations. © 1987 by the Board of
Trustees, Southern Illinois University.)
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8.2 Isotopic analysis 305
-5
-10
d13 C‰
-15
-20
-25
2500 BC 1000 BC 500 BC AD 1 AD 500 AD 1000 AD 1500
Date
Figure 8.3 δ13C values from bone collagen for each sample (circles) documents an increase
in C4 plant consumption after AD 1000 in Ontario, Canada. Mean values from the same date
are indicated by filled circles and mean values are indicated by squares. (From Harrison &
Katzenberg, 2003; reproduced with permission of authors and Elsevier Ltd.)
1995; Schwarcz et al., 1985; van der Merwe et al., 2003). After AD 1000, however, isotope
values rapidly become considerably less negative (Figure 8.3). The presence of relatively less
negative δ13C values in later prehistoric and historic sites in these settings is generally
consistent with findings from other localities in the interior of eastern North America
(compare with Ambrose, 1987; Boutton et al., 1991; Buikstra, 1992; Buikstra & Milner,
1991; Schurr, 1992; Schurr & Schoeninger, 1995; and see following discussion). Analysis of
apatite values shows somewhat earlier use of maize in the diet beginning before AD 1000
(Harrison & Katzenberg, 2003; Katzenberg, 2006). These values suggest that maize was likely
eaten, but not in appreciable amounts. Overall, this record shows that while maize began to
enter diets of native populations in Ontario by AD 500, it did not become a staple until after
AD 1000, when there was a clear and quick shift from C3- to C4-dominated subsistence.
For the broad region of midwestern and interior northeastern North America, the period AD
500–1300 evinces a trend of increase in δ13C values from the earliest to latest periods, but with
notable variation, related to geography, climate, and perhaps cultural preference. In the
southern region of the eastern United States (states of Missouri, Arkansas, Tennessee), the
Ohio River valley, and the American Bottom of the Mississippi River valley, isotope analysis
indicates appreciable increases in maize consumption that occurred rapidly but late in
prehistory (Boutton et al., 1991; Buikstra et al., 1988; Greenlee, 2006; Reber, 2006). In
contrast, populations living in some areas (e.g., Illinois) have gradually higher δ13C values
that suggest a slower rate of adoption of maize (Buikstra et al., 1987).
Temporal comparisons of Ontario populations show a trend that is intermediate between
the southeastern US and Illinois samples (Katzenberg et al., 1995). Katzenberg and coworkers
(1995) suggest that the variation between northern and southern regions of eastern North
America reflects a shorter growing season in the former than the latter, and hence, a relatively
reduced use of maize. They speculate that the later full-blown adoption of maize in the
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306 Isotopic and elemental signatures of diet, nutrition, and life history
southern area may reflect the existence of an established agricultural lifeway at the time of the
introduction of maize (e.g., squash and other indigenous plants). As is characteristic of other
cultural phenomena, dietary practices may have been adopted quickly by some groups and
not by others for reasons that remain unclear. Additionally, other factors influencing stable
isotope values include regional variation in δ13C values of maize, consumption of C3 versus C4
plants by economically important animals (e.g., deer), fish consumption practices, and the
presence of other C4 plants in the diet (e.g., amaranth in the middle Mississippi and Ohio River
valleys) (Katzenberg et al., 1995).
Even within relatively restricted geographic regions, there are apparent spatial differences
in maize consumption. For example, maize appears to have been a greater part of diet in the
outlying areas around the late prehistoric Mississippian center of Cahokia than in Cahokia
itself (Buikstra & Milner, 1991). These differences may represent social distinctions between
the core (elite) and hinterlands (sub-elite) of the Cahokia chiefdom. Moreover, the somewhat
more negative δ13C values in high-status individuals than in low-status individuals within the
Cahokia center point to slightly lower consumption of maize in those elite that have greater
access to a more diverse diet and animal sources of protein (Ambrose et al., 2003).
Analysis of δ13C values in the late prehistoric Ohio River valley drainage similarly
suggests a link between social complexity and maize production and consumption (Schurr &
Schoeninger, 1995). Comparison of populations from late prehistoric Fort Ancient sites
(Baum, Gartner, Feurt, SunWatch) with contemporary populations from organizationally
more complex Mississippian sites (Angel, Wickliffe Mounds) reveals very different isotopic
compositions. Average δ13C values are generally less than −11% for the Fort Ancient samples,
whereas the values are generally more than −10% for the Mississippian samples. Additionally,
there is a strong correlation between site size and isotope values in both this series of sites and
other Fort Ancient and Mississippian period sites. These and other observations (e.g., location
of Mississippian sites in areas with higher agricultural productivity [compare with Ward,
1965]) suggest that, on average, maize played a more important role in subsistence strategies
of Mississippian chiefdoms than of Fort Ancient groups. Schurr and Schoeninger (1995)
speculate that Fort Ancient groups had a subsistence system that was more stable than that
practiced by Mississippian groups, as indicated by the persistence of Fort Ancient but not
Mississippian societies into the historic period. This persistence may reflect the fact that Fort
Ancient populations developed in situ and were well adapted to the limitations of the Ohio
Valley ecosystem. Mississippian societies may represent a cultural intrusion into the region
and practiced an agricultural and subsistence system that may not have been as well suited to
this setting. These complex societies did not share the relatively greater long-term success of
the Fort Ancient cultures – they effectively migrated out of this area and/or became more
similar in structure to the Fort Ancient type of adaptation well before the arrival of Europeans.
Overall, this setting shows a confluence of complex interactions and movements between
regions.
In the late prehistoric central Mississippi River valley in the American Bottom and north of
the American Bottom, spatial analysis of δ13C values shows considerable variation, even
within small regions. Some individuals had C3 diets, whereas others had C4 diets to varying
degrees (Hedman et al., 2002; Rose, 2008; Schoeninger, 2009). The variation does not relate to
sex, age, or status, and appears to reflect resource extraction that is highly geographically
localized, suggesting that food consumption practices, while having broad temporal patterns
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8.2 Isotopic analysis 307
(i.e., maize as a staple in the last several centuries prior to European contact), are immediate –
simply, travel distance to acquire food is limited. Similar patterns of localized dietary
regimens are also well documented in the Georgia Bight region (Hutchinson, 2002; Hutch-
inson et al., 1998; Larsen, Griffin et al., 2001; Norr, 2002; and see later). These findings
suggest the possibility that multiple subsistence strategies were practiced, perhaps reflecting
group or individual decisions pertaining to risk management (Schoeninger, 2009; Winter-
halder & Kennett, 2009).
On the other hand, the local variation described here may represent cultural differences in
foodways. For example, in the rich agricultural region represented by the Ohio River valley
region, analysis of stable carbon isotope ratios reveals key differences between the Fort
Ancient and Mississippian period cultures. The traditional perspective has been that they
are two distinctive cultural traditions (Griffin, 1943). However, in order to explain the sharing
of attributes, some argued that this was due to migration of groups of Mississippians and
interregional contact (Griffin, 1967; but see Pollack & Henderson, 2000). Cook and Schurr
(2009) have revisited these differences in cultural, social, and dietary attributes of Fort
Ancient and Mississippian traditions, especially in spatial and temporal contexts. Rather than
showing a dichotomy in diet in the comparison of Fort Ancient and Mississippian sites,
comparisons of carbon isotope ratios reveal increasing similarity of diets over time. During
the later years, the Fort Ancient and Mississippian stable isotope ratios are statistically
indistinguishable. This confluence of diet in these late prehistoric societies occurs simultan-
eously with the presence of Mississippian architecture and ceramics in the Fort Ancient region
(Cook & Schurr, 2009). Cook and Schurr (2009) make a strong case that the growing similarity
of diet represents the migration of Mississippians into the Fort Ancient region. Thus, at least
with regard to this regional pattern, when viewed in temporal perspective along with a rich
archaeological context, dietary variation may be due to social dynamics involving migration
and interaction.
In addition to documenting variable patterns of maize consumption in the broad region of
the Eastern Woodlands, these studies point to another important finding. Pollen and other
non-isotopic evidence indicate that maize was likely eaten by native populations quite early
in some areas of the Eastern Woodlands (e.g., 3500 yBP at Lake Shelby, Alabama [Fearn & Liu,
1995]). Isotopic analysis of a wide array of prehistoric native populations indicates, however,
that maize did not become economically important and serve as a key source of energy until
very late in prehistory, certainly after AD 900 in eastern North America (Figure 8.2). As
discussed earlier, in the Great Lakes region, analysis of apatite suggests that maize was
consumed well prior to late prehistory. However, populations were not committed to maize
production as a dominant component of diet. This temporal pattern documenting the increas-
ing focus on maize coincides with other evidence of increased morbidity, increased warfare,
and declining skeletal robusticity in later prehistory (Cohen & Armelagos, 1984; Larsen,
1990b; Larsen & Milner, 1994; Steckel & Rose, 2002).
Stable isotope analysis presents a less clear picture of maize consumption in western North
America, primarily because native populations consumed significant amounts of various
foods with δ13C values similar to maize, such as bison meat, amaranth, and cacti (Schoeninger &
Moore, 1992). For example, less negative δ13C values (−8‰) were identified in skeletal series
from northern Texas and Oklahoma (Habicht-Mauche et al., 1994). These values reflect the
consumption of cactus and other nondomesticated C4 plants, bison, and perhaps some grown
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308 Isotopic and elemental signatures of diet, nutrition, and life history
or traded maize. In other late prehistoric Great Plains settings (e.g., Crow Creek, South
Dakota), less negative values also represent the consumption of a mixture of C3 and C4 diets,
although maize probably contributed significantly to native diets (Bumsted, 1984).
Analysis of carbon isotope ratios in the American Southwest and eastern Great Basin points
to temporally distinctive patterns of dietary variation. Comparison of early (AD 1275–1330)
with late (AD 1330–1400) populations at Grasshopper Pueblo, Arizona, reveals a trend toward
less negative δ13C values, indicating an intensification of maize agriculture (Ezzo, 1993). This
increase coincides with a period of resource stress, decrease in dietary diversity, and environ-
mental disruption associated with climate change during the later occupation of Grasshopper
Pueblo. Ezzo (1993) contends that these disruptions and a decreasing quality of diet may have
contributed to the abandonment of the region in the late fourteenth to early fifteenth
centuries. This pattern of intensification of maize consumption is also present in the Pueblo
period (AD 750–1350) in the Anasazi region of southwestern Colorado (Coltrain et al., 2007).
This gradual increase began when local populations first became reliant on maize at about
400 BC (Coltrain et al., 2007).
At the Pecos Pueblo site, New Mexico, where subsistence practices involved intensive maize
farming (Kellner et al., 2010; Kidder, 1932; Spielmann, 1991; Spielmann et al., 1990),
the comparison of the pre-Plains trade period (AD 1200–1450), Plains trade period
(AD 1450–1600), and historic (mission) period (AD 1600–1838) revealed that there were no
temporal changes in stable carbon isotope ratios (Kellner et al., 2010). Average δ13C values of
−7.7‰, −7.6‰, and −7.7‰, respectively, document a consistent diet for the 400-year period
based on heavy maize dependence and consumption across the population (no differences in
sex or age).
While maize was a dietary staple at Grasshopper and Pecos Pueblo, in the Great Salt Lake
region of the eastern Great Basin, isotopic analysis reveals that Fremont populations experi-
enced a dramatic reduction in C4 consumption and reversion to exploitation of wild resources
in later prehistory. That is, for the period of c. AD 400–1150, there is an increase in
δ13C values, expressing a shift and increased commitment to maize farming (Coltrain &
Leavitt, 2002; Coltrain & Stafford, 1999) (Figure 8.4). As with the Southwestern population,
these eastern Great Basin groups relied on maize as an important component of diet and
subsistence economy. However, unlike the Southwestern populations, there is a sharp drop in
δ13C values (to −18.1‰), revealing the complete shift to a foraging adaptation in later
prehistory. After AD 1150, native populations abruptly and completely eliminated a food
source that heretofore had been a central part of their diet. While various factors were
involved in this rapid shift, analysis of climate data derived from tree rings and pollen profiles
shows a marked reduction in summer rainfall and truncation of the growing season (Coltrain
& Stafford, 1999; Grayson, 2011). This profound climate impact rendered the continuation of
large-scale maize production impossible, and native populations greatly curtailed agricultural
productivity. This climate shift, combined with a reduction in rainfall, contributed to an
adaptive response involving a return to foraging.
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8.2 Isotopic analysis 309
-10
-12
-14
d13 C‰
-16
-18
-20
Figure 8.4 δ C values from bone collagen samples exhibit diverse plant consumption from
13
AD 400 to AD 1150. After AD 1150, Great Salt Lake region populations ate more C3 plants,
shown in more negative δ13C values. (From Coltrain & Leavitt, 2002; reproduced with
permission of authors and the Society for American Archaeology.)
(DeNiro & Epstein, 1981; Farnsworth et al., 1985). In contrast, analysis of stable carbon
isotope ratios in lower Central America reveals that maize use was very minor or altogether
missing in Panama for the same time period (5000–3000 BC) and was adopted considerably
later in Costa Rica (AD 300–1550) (Norr, 1991).
Analysis of prehistoric and contact-era Mayan skeletons from Belize, Honduras, and
Guatemala provides an important picture of dietary variation in Mesoamerica. The Maya
lowlands contain a wide diversity of plants and animals with potential dietary importance,
but poor preservation of food remains – plants and animals – greatly limits understanding of
foodways in this otherwise well-documented region. It has long been recognized that maize
was an important food source in past Mayan populations. Mayan skeletal series show an
abundance of skeletal pathology indicating poor-quality nutrition and elevated disease
burdens. Populations consuming the greatest proportion of maize have an especially elevated
prevalence of skeletal stress indicators, including dental caries, enamel hypoplasia, and cribra
orbitalia/porotic hyperostosis (see Chapters 2 and 3) (Scherer et al., 2007; White, 1997;
Whittington & Reed, 1997; Wright, 2006; Wright & White, 1996). Due to the paucity of
subsistence-related data from archaeological localities, it is difficult to link these indicators of
health to any one particular subsistence strategy.
The picture of dietary ecology in the ancient Maya is informed by a robust record of stable
isotope variation. Temporal and spatial comparisons of δ13C values from environmentally
heterogeneous locations of Lamanai and Pacbitun, Belize, suggest that prehistoric Mayans
placed less emphasis on maize than did some other Mesoamerican groups (e.g., Tehuacan)
(White & Schwarcz, 1989; White et al., 1993, 1994). Nonlinear temporal changes suggest
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310 Isotopic and elemental signatures of diet, nutrition, and life history
variable emphasis on maize in the region. Pre-Classic populations (1250 BC–AD 250) at
Lamanai have relatively high δ13C values (−12.4‰) at both sites, suggesting strong reliance
on maize. Late and Terminal Classic Lamanai, however, have increasingly negative
δ13C values, indicating a decrease in C4 consumption (White, 1997; White et al., 1993). In
contrast to Lamanai, the record from Pacbitun shows an increased reliance on maize begin-
ning in the Preclassic and peaking in the Late Classic periods (White, 1997; White et al., 1993).
These elevated levels of C4 diets in the Late Classic are similarly displayed at Copán (Reed,
1994, 1999; Whittington & Reed, 1997) and at inland sites in Petén, Guatemala, such as
Piedras Negras and Altar de Sacrificios (Scherer et al., 2007; Wright, 1997, 2006).
Following the collapse of the Classic period Maya in the eighth to ninth centuries AD,
δ C values at Lamanai (Pacbitun was abandoned by this time) show a marked reversal in the
13
trend of decreased maize consumption documented for the Late and Terminal Classic periods.
Post-Classic and Historic period Mayan skeletons have substantially less negative values
(−9.3‰ and −9.9‰, respectively), indicating an increased reliance on maize. White and
Schwarcz (1989) interpret this trend as representing a doubling of maize consumption in less
than a century. Thus, in late prehistoric and early contact-era Maya, maize consumption was
high, a finding similar to estimates of 65% to 86% of diet based on maize in Maya Indians in
contemporary society (White et al., 1994).
In addition to the temporal trends, populations with access to marine resources show lower
δ13C values than interior populations having limited or no access to marine foods (Gerry &
Krueger, 1997; Scherer et al., 2007). For example, Lamanai, a coastal population, has a
considerable amount of variation in δ13C ratios, with some individuals consuming relatively
less maize than others. In contrast, in the interior, Copán populations show less variation and
generally high levels of commitment to maize across the community, and consequently higher
δ13C ratios. This pattern is strikingly similar to analyses showing regional variation in coastal
and interior settings and with the localized nature of diet focusing on extraction of resources
that are nearby (and see earlier discussion).
In the entire record of dietary patterns for the Maya region, little evidence for an association
between social collapse and dietary change can be provided (White, 1997; Wright, 2006;
Wright & White, 1996). Based on isotopic analysis, prehistoric diets appear to have been
linked in a general sense with major events in Maya history, but the variation is so great that
no argument can be made tying resource stress – as indicated by dietary change – to the
collapse and abandonment of the southern Maya lowlands (Wright & White, 1996).
A growing record of past diets based on stable isotope variation is available from South
America, especially in the Peruvian Andes. In order to track key aspects of dietary
variation in diverse ecological and cultural settings in this region, Tykot and coworkers
(2006) documented the record at Pacopampa in highland northern Peru and Cardal and
Tablada de Lurin in the Lurin River valley of the central coast of Peru. These sites give an
indication of dietary variation for the Early Horizon period of the first millennium BC, a
time during which maize has traditionally been regarded as a staple crop. Predictably,
analysis of stable carbon and nitrogen isotopes reflect a highly diverse set of diets in this
complex setting. Overall, the record shows that, like coastal populations of Peru and
Ecuador (Ericson et al., 1989; Tomczak, 2003; Tykot & Staller, 2002; van der Merwe
et al., 1993), there is a strong commitment to marine foods with a later increase in maize
in diets.
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8.2 Isotopic analysis 311
-3.0
Material
Bone apatite
Tooth enamel
-5.0
-7.0
d13 C‰
-9.0
-11.0
-13.0
~
Guanape Salinar Pre-structural Structural Post-structural
Gallinazo Gallinazo Gallinazo
Figure 8.5 Bone apatite and tooth enamel δ13C values increase from the Guañape to
Post-Structural Gallinazo periods in Cerro Oreja, Peru. These dietary changes correlate
with transitions in sociopolitical complexity up to and after state formation in the
Gallinazo phase. (From Lambert et al., 2012; reproduced with permission of authors
and the Society for American Archaeology.)
In later coastal settings, maize production and consumption intensified over time and
played a crucial role in the rise of complex societies serving as an important precursor to
state development. In the Moche Valley of north coastal Peru, for example, Lambert and
coworkers (2012) tested the hypothesis that intensified maize production preceded the
appearance of the complex political economy fueling the state system associated with the
Early Intermediate period (c. 400 BC–AD 600). Diachronic analysis of bone apatite and tooth
enamel from the Cerro Oreja site documents dietary changes associated with the rise and
establishment of increasing sociopolitical complexity and possible state-level organization
(Figure 8.5). During this time frame, there appears to have been no change in δ13C from the
Guañape phase (c. 1800–400 BC) to the Salinar phase (c. 400–1 BC). This period of dietary
stasis was followed by progressive and swift enrichment of 13C in the Gallinazo phase
(AD 1–200) by about 2.5‰. The change in isotope ratios coincides with a marked increase
in dental caries in the final period representing the state system (Gagnon & Wiesen, 2013).
The record overall supports the hypothesis that intensified production of this storable crop
was likely an important antecedent to the rise of the Moche state during the AD 200s or
300s. Similar peaks representing maize intensification are also documented in the rise of
later state systems, such as the Inca Empire (Burger et al., 2003; Hastorf, 1990; Hastorf &
Johannssen, 1993).
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312 Isotopic and elemental signatures of diet, nutrition, and life history
15
d15 N‰
PIT
10
URN CRYPT/CIST
5
-15 -10 -5
d13 C‰
Figure 8.6 Carbon isotope ratios from bone collagen vary among individuals from urn,
pit, and crypt/cist burials at Pacbitun, Central America. These distributions indicate that
high-status individuals (crypt/cist burials) consumed more C4 plants in their diet than
lower-class individuals (urn and pit burials). (From White et al., 2006; adapted and
reproduced with permission of authors and Elsevier Ltd.)
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8.2 Isotopic analysis 313
In the La Florida site in the northern highlands of Ecuador, carbon isotope analysis shows
distinctive differences between high- and low-status individuals during the Chaupicruz
phase (AD 100–450) (Ubelaker, 2000; Ubelaker et al., 1995). Elite individuals recovered from
elaborate tombs having emerald beads, marine shells, textiles, and other exotic items have less
negative carbon isotope values than low-status individuals (δ13C ¼ −10.3‰ and −11.6‰,
respectively). Ethnohistoric and archaeological (mortuary) evidence suggests that these dif-
ferences may reflect a greater consumption of maize beer in high-status adults, especially
males. Moreover, if high-status individuals were consuming more tortillas and other maize-
based food than were low-status individuals, the caries prevalence would be similar in the two
social groups (Ubelaker, 2000).
The dietary signatures of carbon isotope values and inferences drawn from dental caries
prevalence provide congruent results in comparisons of high- and low-status individuals. At
Copán and Lamanai, dental caries prevalences are substantially higher in low-status adults
than in high-status adults, and very high-status adults have relatively low δ13C values. Both
data sets indicate greater consumption of maize in low-status individuals (White, 1994). At
Copán, the greater range of values and lower caries prevalence in high-status individuals
suggest a more varied, less carbohydrate-rich diet (Reed, 1994).
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314 Isotopic and elemental signatures of diet, nutrition, and life history
10.0
Female Male
9.0
8.0
d15N‰
7.0
6.0
5.0
4.0
-12.0 -11.0 -10.0 -9.0 -8.0 -7.0 -6.0
d13C‰
Figure 8.7 Stable nitrogen and carbon isotope ratios show dietary distinctions between
males and females in the pre-Puebloan American Southwest. Slight variability in carbon
isotopes suggests differential maize consumption between sexes. Even more profound is the
dichotomization in meat consumption between sexes: males have more enriched nitrogen
ratios than females, which suggests the consumption of deer/sheep by the former and of
small game (e.g., rabbits) by the latter. (From Coltrain & Janetski, 2013; reproduced with
permission of authors and Elsevier Ltd.)
Lamanai females and males show no difference in stable carbon isotope values, suggesting
a similarity in their diets (Coyston et al., 1999; White, 1997; White & Schwarcz, 1989). Reed
(1994, 1999) and White and coworkers (1993) contend that sex differences in diet at Pacbitun
and Copán represent variation in socioeconomic status, with males consuming more maize
than do females. With regard to Copán, Reed observes that this difference “parallels the
observation of higher frequencies of anemia, infection, and a statistically significant higher
rate of caries in females than in males” (1994:216). These findings of reduced consumption of
maize in females relative to males contradict the results presented from analysis of these other
data sets. For example, higher caries prevalence in females than males suggests greater dietary
emphasis on maize-based foods among women than among men in this setting. Thus, while
both isotopic and caries analyses point to possible differences in diet by gender, the contra-
dictory results do not provide a clear picture of what these differences may have been. At
Lamanai, like the isotope evidence, dental caries prevalence data suggest distinctive differ-
ences in diet between females and males (White, 1994).
The variable nature of diet in comparing isotope and dental caries suggests that these data
sets may reflect somewhat different aspects of diet. Dental caries is caused by acids produced
by bacterial metabolism of carbohydrates (sugar). Carbon isotope data derived largely from
collagen – the basis for most of the Mayan isotopic investigations – provides information on
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8.2 Isotopic analysis 315
protein consumption as well as diet generally (Ambrose & Norr, 1993; but see White et al.,
2001 advocating use of carbonate). Thus, changing patterns of diet – especially with regard to
the consumption of maize – will not necessarily be expressed in the same way in the analysis
of dental caries and stable carbon isotope ratios. The general congruence of temporal trends in
caries prevalence and in carbon isotopes in the Maya lowlands with other regions argues for a
generally overlapping picture of these data sources and dietary shifts. For example, caries
prevalence declines dramatically in the Terminal Classic, but then peaks in the Post-Classic,
indicating respective decrease and increase in maize consumption. This pattern is identical to
dietary changes indicated by δ13C values (White, 1988; White et al., 1994). The interpretations
of dietary patterns are enhanced by use of both direct (isotope) and indirect (caries)
approaches.
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316 Isotopic and elemental signatures of diet, nutrition, and life history
maize-dependent native populations living in the region prior to European contact (Katzen-
berg et al., 1995; Schwarcz et al., 1985). The mean δ13C value is −18.7‰, which is similar to
fifteenth- to seventeenth-century European values of −18‰ to −19‰ (compare with
B. Kennedy, 1989). These findings suggest, therefore, that the Harvie family did not eat maize,
but rather, mostly C3 plants, such as wheat, barley, rye, and oats. The Harvie dietary
composition is much more in line with conventional interpretations of nineteenth-century
foodways: farm animals ate maize; farmers did not.
Analysis of nineteenth-century US military buried at the Snake Hill site, Ontario, indicates
a wide range of isotopic variation in contrast with the Cross and Harvie pioneer families
(Katzenberg, 1991b). The mean value from the Snake Hill series is −15.8‰, with a range of
−18.5‰ to −12.5‰. This variation reflects the presence of individuals recruited from all over
the northeastern United States and the high degree of variability in regional cuisines,
especially with regard to use of C3 (e.g., wheat and rice) and C4 (maize) plants. Thus, it is
not surprising that these values range between a mostly C3 diet and a mixed C3/C4 diet
(Katzenberg, 1991b).
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8.2 Isotopic analysis 317
13.0
Male, high status
12.0 Male, low status
Female, high status
11.0
Female, low status
d15N‰
10.0
9.0
8.0
7.0
6.0
-20.5 -19.5 -18.5 -17.5
d13C‰
Figure 8.8 Dietary differences are distinguishable between sexes and statuses at Medieval
Trino Vercellese, Italy. Although high-status males demonstrate less consumption of C3
millet plants than do low-status males, no significant discrimination in diets is apparent
between high- and low-status females. (From Reitsema & Vercellotti, 2012; reproduced
with permission of John Wiley & Sons, Inc.)
Millet became a central part of the diet in northern China by 9000 BC (Barton et al., 2009;
Pechenkina et al., 2005; van der Merwe, 1992; Yang et al., 2012). Analysis of δ13C values
indicates that millet contributed well over 50% of carbon in the diets of some groups. The
record of first domestication is situated in the Yellow River (Huang He) basin, initially with
incipient and short-term use and later involving intensive production and consumption by
humans and as fodder for domesticated pigs and dogs, and perhaps chickens. Analysis of
stable carbon isotope ratios in a number of sites reveals remarkably high δ13C values
(Pechenkina et al., 2005), but also with considerable variation, even within a tightly restricted
geographic area. The implications of these heavily millet-based diets are similar to those
settings where maize became a central component of diet (North and South America). That is,
millet is an incomplete protein, is among the poorest sources of protein, and is especially
deficient in the amino acid lysine. Anemia and other physiological disorders occur in these
kinds of dietary regimens. In this setting, relatively little evidence of porotic hyperostosis is
associated with early farming populations (Pechenkina, Benfer et al., 2007). Indeed, at the
Jiangzhai site (6900–6000 yBP) where δ13C values are relatively high, no individuals dis-
played lesions that are associated with anemia. However, enamel hypoplasias and other
lesions representing growth disruption and compromised health are highly prevalent (Pechen-
kina, Benfer et al., 2007).
From the earliest exploitation in China, millet cultivation spread into the Eurasian steppes.
In the Minusinsk Basin in southern Siberia, comparisons of earlier and later cultures reveals
a rapid uptake in millet at c. 1400 BC or slightly later (Svyatko et al., 2013), a shift that
took place within a century or less. The speed of adoption is remarkable, and is analogous
to the fast adoption of maize in some New World settings (Larsen, Griffin et al., 2001;
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318 Isotopic and elemental signatures of diet, nutrition, and life history
Larsen, Schoeninger et al., 1992; and the next section) or the shift from marine to
terrestrial foods in Neolithic Europe (Tauber, 1986; and following). On the other hand, when
millet is adopted in the Minusinsk Basin, it occurs in a highly variable way, involving
some individuals consuming it and others not. The adoption of millet in this setting occurred
simultaneously with the adoption of the horse and increasing social stratification
in southern Siberia (Svyatko et al., 2013). Therefore, this disparity may reflect variation in
social status in this setting.
Along with millet, sorghum – also a C4 domesticate – dominated the archaeological record
for much of Nubian prehistory (White & Schwarcz, 1994). Analysis of bone and other tissues
shows that although these plants were present in substantial amounts, diets were based
principally on C3 cultigens (e.g., wheat and barley) (White & Schwarcz, 1994), but analysis
along the length of hair shafts from mummies indicates seasonal shifts from C3 to C4 staples
(White, 1993). Isotopic evidence indicates a shift toward consumption of more C4 plants in the
X-group (AD 350–550), a period characterized by political instability, alterations in trade
patterns, and decreased water availability as the Nile River lowered. In the following Christian
period (AD 500–1400), more negative isotope values reveal a decline in C4 plant consumption.
This shift in diet appears to have accompanied an increase in elevation of the Nile and
generally improved economic conditions.
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8.2 Isotopic analysis 319
16
14
12
d15N‰
10
RIA MED
6 VIKA MES
NEO
4
-22 -21 -20 -19 -18 -17 -16 -15 -14 -13 -12 -11
d13C‰
Figure 8.9 Diachronic dietary changes in Denmark: RIA (Roman Iron Age), VIKA (Viking
Age), NEO (Neolithic), MED (Medieval), and MES (Mesolithic). Decreases in δ13C over time,
from the Mesolithic to the Neolithic and subsequent periods, demonstrate an increase in
13
C-depleted marine foods. (From Jørkov et al., 2010; reproduced with permission of
authors and John Wiley & Sons, Inc.)
Prowse et al., 2004). In northern England, a significant return to use of marine foods is
observed only after the rise of deep sea fishing, c. AD 1000 (Müldner & Richards, 2007a,
2007b). These analyses reveal general dietary patterns, geographic and temporal, but they also
show how specific diets can be within restricted areas (Herrscher & Le Bras-Goude, 2010;
Polet & Katzenberg, 2003; and see Hutchinson et al., 1998 for a New World setting).
The overall record shows that the foraging-to-farming (Mesolithic to Neolithic) transition in
Europe was not a gradual process, but was rapidly achieved in many of the settings where
there is a substantial isotopic record of diet. Even coastal populations saw some adaptive
advantage to abandoning – either wholly or mostly – the use of marine foods in diet and
deriving protein and energy from terrestrial foods. This also implies a general reduction in the
quality of diet and a more homogeneous diet relative to Mesolithic forager-fisher-gatherers
(Schulting, 2011). Moreover, in the Baltic region where agriculture is adopted later in the
Neolithic, there is evidence for a greatly reduced marine component as terrestrial domesticated
resources become emphasized (Lillie & Budd, 2011).
The quick shift in diet is not restricted to northern Europe but it is also well documented in the
Mediterranean region, indicating that the earlier record of the agricultural transition taking
place in this setting was also rapid. In particular, terminal Neolithic populations from Alepo-
trypa Cave, coastal southern Greece, have very negative δ13C values, for both collagen (mean ¼
−19.9‰) and apatite (mean ¼ −13.1‰) (Papathanasiou, 2001, 2003, 2011; Papathanasiou
et al., 2000). These findings indicate that the diets of these groups were largely focused on
terrestrial C3 plants and animals with near abandonment of marine resources. The narrow range
of values (collagen: −19.5‰ to −20.2‰) indicates a remarkably high degree of homogeneity in
diets. Moreover, this dietary adaptation is associated with a marked increase in stress indicators
(e.g., porotic hyperostosis) (Papathanasiou, 2001, 2003, 2011; Papathanasiou et al., 2000),
similar to that found in other regions of the globe (e.g., Mesoamerica).
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320 Isotopic and elemental signatures of diet, nutrition, and life history
Correspondingly, an abrupt shift from marine to terrestrial food resources in the compari-
son of Mesolithic and Neolithic populations (c. 8500–4500 yBP) in Portugal is indicated by
isotopic analysis (Lubell et al., 1994). Values for the samples drawn from various coastal and
near-coastal sites range from −15.3‰ to −20.4‰. The more negative values are predomin-
antly Neolithic and the less negative values are Mesolithic. The presence of less variability in
the Neolithic period suggests increasing homogeneity in diet during the period where plant
and animal domesticates were more heavily relied upon (Lubell et al., 1994). Unlike the
pattern of increased prevalence of dental caries in New World settings with the increasing
reliance on plant domesticates, there is a marked decline in dental caries prevalence, number
of carious tooth surfaces, and premortem tooth loss in permanent mandibular molars in late
Mesolithic and Neolithic Portuguese in comparison with earlier populations from the region.
The relatively high prevalence of dental caries in the Mesolithic period is probably related to
the consumption of cariogenic plants such as nondomesticated figs (Lubell et al., 1994; and
see Chapter 3).
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8.2 Isotopic analysis 321
TERRESTRIAL MARINE
Carnivores
(n = 30)
Primary
Carnivores (n = 26)
(n = 6)
Herbivores
(n = 21)
Precision of
isotopic analysis
Plants
(n = 32)
(n = 136)
Figure 8.10 Terrestrial and marine plants and animals are represented in broad trophic levels
according to δ15N values in bone collagen. Each trophic level is associated with substantial
enrichment of δ15N. Nitrogen ratios, in turn, can be compared (site-specifically) to these
trophic models to determine primary dietary protein contributions. (From Schoeninger &
DeNiro, 1984; reproduced with permission of authors and Elsevier Ltd.)
vary considerably, depending on the setting. Omnivores, such as humans, have values that are
mostly between those of herbivores and carnivores (Minagawa & Wada, 1984; Schoeninger &
DeNiro, 1984; Schoeninger & Moore, 1992). Therefore, this increasing enrichment of 15N as
one moves up the food chain provides the opportunity to determine the trophic level and to
infer the dietary source based on analysis of bone (collagen, in particular) (Figure 8.10). Key to
interpreting these diets, however, is understanding the biochemistry of plants and animals
within the food webs of specific regions (Katzenberg, 2008). Within these specific regions, key
patterns of dietary consumption involving access to protein show important tendencies that
document broad patterns. For example, at Neolithic Çatalhöyük, there are increases in
δ15N values over the site’s 1400-year occupation (Hillson et al., 2013; Pearson, 2013; Pearson
et al., 2013), likely reflecting increasing protein derived from animal sources, peaking in the
terminal phase of the site’s occupation (Figure 8.11), an increase which coincides with the
presence of domesticated cattle and a greater likelihood of the consumption of cattle protein
in day-to-day contexts as well as special feasting (Russell et al., 2013). In contrast, consider-
ably lower δ15N values are documented at Neolithic Nevalı Çori (Lösch et al., 2006), empha-
sizing a different set of local factors – dietary and cultural – that play into understanding
regional patterns of food use. These findings show that communities in some settings
employed food to differentiate between men and women, or the sexes had somewhat different
lifestyles and eating patterns relating to subsistence activities.
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322 Isotopic and elemental signatures of diet, nutrition, and life history
14.0
Female
13.5
Male
13.0
12.5
d15N‰
12.0
11.5
11.0
10.5
10.0
Early Middle Late
Period
Figure 8.11 Stable nitrogen ratios from Neolithic Çatalhöyük increase from the Early to Late
periods, indicating increased dietary consumption of animal meat. (From Hillson et al.,
2013; reproduced with permission of authors and Cotsen Institute of Archaeology.)
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8.2 Isotopic analysis 323
Schwarcz et al., 1999). Other contexts produce generally high δ15N soils, as in areas with a
history of evaporation (e.g., saline soils) and those enriched in organic materials (e.g., guano).
The very high values reported from analysis of human bone samples in some desert settings
may be explained by these factors (Ambrose & DeNiro, 1986a, 1986b; Aufderheide, Tieszen
et al., 1988; Heaton et al., 1986; Schoeninger, 1995b). In some settings, terrestrial animals
living in arid environments have higher δ15N values than marine animals. Because of climatic
and/or other ecological variables, Late Stone Age coastal and interior foragers in South-
western Cape Province, South Africa, have nitrogen isotope ratios inverse to those expected,
but carbon isotope values clearly identify differences in the exploitation of marine versus
terrestrial resources (Sealy, 1986; Sealy & van der Merwe, 1985, 1986; Sealy et al., 1987). In
other settings, stable nitrogen isotope ratios are useful for identifying consumers of terrestrial
versus marine foods (Schoeninger & DeNiro, 1984; Schoeninger et al., 1983; and see following
discussion).
The life cycle and physiology: breastfeeding, weaning, and the adoption of adult diet
Behaviors impacting the earliest periods of life, not the least of which are the duration of
weaning and the length of time it takes to adopt a complete or near complete adult diet, have a
profound influence on a number of facets of a population’s demographic dynamics. A large
and expanding body of evidence from demographic and anthropological sources, especially
from study of traditional societies, shows that the shorter the duration of reliance on
breastfeeding as the infant’s sole or dominant food source, the sooner the mother can become
fertile again and produce another infant. That is, nursing suppresses ovulation and female
fecundity (Ellison & O’Rourke, 2000; Vitzthum, 1994), and increases the inter-birth interval.
Hypothetically, populations that, on average, have a long-duration breastfeeding regimen will
have fewer offspring than populations with a short-duration breastfeeding regimen. Thus, in
the present world, as in the past, population stasis and growth is very much influenced by
breastfeeding practices as they relate to duration, weaning, and timing of adoption of adult,
non-breast milk diets.
There are many factors that affect fecundity and fertility in a human society (Ellison &
O’Rourke, 2000; Gage, 2010; Vitzthum, 1994). However, all else being equal, breastfeeding
and an infant’s dependence on the mother as the sole source of energy and nutrition, the
duration of this part of early life history, and the time it takes to adopt a wholly adult diet all
play vital roles in shaping demographic characteristics of a population, especially its size and
growth rate. Demographic analysis provides especially important insight into understanding
population growth, stasis, or decline, including the remarkable demographic transition that
accompanied the shift from foraging to farming beginning in the early Holocene. In this
regard, might the dramatic population growth in the Neolithic be explained in part or in whole
by a change in weaning timing?
Buikstra and coworkers (1986) addressed this question by hypothesizing that agriculture
and the development of more efficient ceramic cooking vessels in later prehistory in the lower
Illinois River valley facilitated the production of maize-based gruel. The increased use of this
improved technology for preparing these soft weaning foods thus would shorten the length of
time to transitioning to adult foods. This factor, therefore, might explain the growth in
population in the later intensive farmers compared to the earlier less-intensive farmers (see
Chapter 10). In order to test the hypothesis regarding reduced duration of breastfeeding,
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324 Isotopic and elemental signatures of diet, nutrition, and life history
weaning, and the complete shift to adult foods, Fogel and coworkers (1989, 1997), in their
classic study, documented the δ15N values from fingernails (composed of the protein keratin)
of nursing infants and their lactating mothers over the course of the first year of life. They
found a distinctive trophic effect involving an average increase in δ15N values when compar-
ing the infants to their mothers. When the infants were fully weaned, they exhibited values
within 1‰ of their mothers; within three to five months following cessation of consumption
of mother’s milk, the infant values were identical to those of their mothers. In the interim–the
weaning process – as the milk provided to the infant via breastfeeding is replaced by other
foods, the δ15N values decline.
Fogel and coworkers (1989, 1997) also compared δ15N values in juveniles and adults from
hunter-gatherers and maize farmers from eastern North American archaeological contexts,
predicting that the hunter-gatherers would express a longer duration in early life of elevated
δ15N values than the farmers. The former are represented by Archaic period remains from Eva,
Cherry, and Ledbetter sites in the Tennessee River valley, and the latter are from the historic-
era Sully site in the northern Missouri River valley. The age changes in δ15N followed exactly
what would be predicted: (1) newborns had values strongly similar to adults, reflecting the
fact that the breast milk is derived from the mother’s diet and has isotope values largely the
same as all tissues in the mother; (2) within several months, infant values increased, reflecting
trophic differences between the consumer (the infant) and the consumed (the mother); and (3)
values increase during the period of breastfeeding, and decrease thereafter, reflecting the
weaning process. Thus, the pattern of δ15N values clearly tracks the duration of breastfeeding
and the subsequent weaning process leading to fully adult diets. Contrary to their expect-
ations, however, Fogel and coworkers found no difference in the timing of weaning between
the hunter-gatherers and the agriculturalists. Nevertheless, their study accomplished an
important goal; namely, establishing the essential link between the weaning process and
how the process is documented via δ15N values in early life. Moreover, the values showed that
weaning is not an abrupt event. Rather, these findings point to a gradual reduction in
δ15N values over young childhood rather than a sudden decline, suggesting that the introduc-
tion of foods – generally having lower δ15N values – and replacement of breast milk is a
gradual process. While many anthropologists and other social scientists tend to regard age-at-
weaning as a specific life history milestone marking the complete termination of breast
feeding, their study also confirmed the growing realization of nutrition scientists that
weaning is a process, sometimes of lengthy duration (Herring et al., 1998). In this way, work
by Fogel and collaborators (1989, 1997; and see Fuller et al., 2006; Nitsch et al., 2011; Schurr,
1998) allows us to model and predict life history patterns relating to early life (Figure 8.12).
Children in the twenty-first century are typically weaned from breast milk completely at
some point between the ages of one and five years. However, it is common in developed
countries for weaning to occur considerably earlier (Dettwyler & Fishman, 1992). Documen-
tation of age profiles of δ15N values in relation to the breast feeding of infants is demonstrated
via stable nitrogen isotopic analysis, revealing the variation and timing from diverse arch-
aeological contexts globally. All of these records reveal a strikingly similar pattern in the spike
in δ15N values over the life course, with an increase in values tracking the period of bone
development, a peak representing full dependence on mother’s milk, the period of weaning,
and the assumption of a fully mature diet. Most of the settings where δ15N ratios have been
documented by age from neonate to adult show very similar patterns overall, but with
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8.2 Isotopic analysis 325
11
-16
Exclusive d15N
breastfeeding
d13C
10 -17
Weaning
9 -18
d15N‰ d13C‰
8 -19
Introduction of
solid foods
-20
7
6
0 2 4 6 8
Age (years)
Figure 8.12 The process of weaning, as modeled with stable carbon and nitrogen ratios,
depicts the incorporation of solid foods into a child’s diet followed by gradual weaning
from maternal milk resources. (From Nitsch et al., 2011; reproduced with permission of
authors and John Wiley & Sons, Inc.)
considerable variation in the timing of the weaning process. For example, at Neolithic
Çatalhöyük and nearby Aşıklı Höyük, in southern Turkey, documentation of δ15N values
reveals no enriched isotope values after the age of one year, suggesting that the weaning
process started before 1.5 years or somewhat later (Hillson et al., 2013; Pearson et al., 2010,
2013; Richards, Price et al., 2003) (Figure 8.13). In contrast, the earlier Neolithic site of Çayönü
Tepesi in southeastern Turkey shows a pattern indicating that the weaning process com-
menced by as much as a year later than at Aşıklı Höyük (Pearson et al., 2010). This represents
significant variation within the larger region, reflecting an entirely different timing of and
approach to weaning. Likely, these differences reflect a combination of factors, including
environmental differences (more arid to the west and more temperate to the east) and dietary
differences (narrower range of foods to the west and wider range to the east) (Pearson et al.,
2010; and see Lösch et al., 2006).
Analysis of δ15N values in a range of settings in the Roman Empire reveals similar patterns
in breastfeeding and weaning, but with significant variation, generally supporting the notion
that weaning is locally determined, primarily by environmental and cultural factors. At Kellis
in the Dakleh Oasis, Egypt, the stable isotope record indicates that weaning foods were likely
introduced by about six months and breast feeding cessation occurred no later than three
years of age, the time at which there is no apparent trophic effect (Dupras, 2010; Dupras &
Tocheri, 2007; Dupras et al., 2001). The record from δ15N in bone collagen reveals the general
pattern, but the use of δ13C in the analysis using the comparison of deciduous and permanent
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326 Isotopic and elemental signatures of diet, nutrition, and life history
(a)
-15
-16
-17
-18
-19
d13C‰
-20
-21
-22
-23
-24
-25
in utero birth 5 10 15 20
Age (years)
(b)
18
16
14
12
10
d15N‰
0
in utero birth 2 4 6 8 10 12 14 16
Age (years)
Figure 8.13 Stable carbon (a) and nitrogen (b) isotope ratios from Neolithic Çatalhöyük show
a decrease in isotope enrichment after about 1.5 years, which indicates that weaning
processes began before this age. (From Hillson et al., 2013; reproduced with permission of
authors and Cotsen Institute of Archaeology.)
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8.2 Isotopic analysis 327
teeth provides an additional measure of trophic level that is similar to δ15N. That is, there is a
trophic effect in δ13C representing about 1‰ (DeNiro & Epstein, 1978; Post, 2002), also
documented in the comparison of mother and infant pairs (Fuller et al., 2006). There are some
differences between δ15N and δ13C in the rate of return to the mother’s baseline values.
Namely, δ13C appears to return to the mother’s baseline value sooner than δ15N. At Isola
Sacra, the largest port located near Rome, the strong shift in and variation of δ15N and
δ13C values occur between one and two years of age, indicating the time of weaning and the
process of full adoption of adult diets, perhaps by 2.5–3 years (Prowse et al., 2008).
Documentation of dental caries and tooth wear indicates the kinds of high-carbohydrate
foods that were used in the weaning process. For example, calculus deposits are first observed
in individuals older than two years of age, and occlusal surface wear, albeit slight, is present
in deciduous teeth in the one- to two-year age range. Carious teeth are not present until the
age of two years, and clearly point to the minimal level to which infants and young children
are first exposed to cariogenic foods.
On the northern periphery of the Roman Empire, patterns of nitrogen and stable carbon
isotope variation reveal a very similar outcome, with a decline in δ15N values after the age of
two years. Relatively low δ15N values in infants aged 1.5–2 years indicate an earlier age of
complementary feeding in a series from Queensford Farm, England (Fuller et al., 2006). In this
setting, nearly all juveniles have low (adult) values by the age of three to four years. Overall,
therefore, stable isotope values from the handful of Roman period sites and estimated ages of
peak reliance on the mother’s milk, the duration of weaning, and the full adoption of adult
diet are similar and consistent with written accounts of weaning practices during the Imperial
period.
More recent populations from historical contexts in Europe and North America show the
same pattern of age change in δ15N values, involving an initial and fast-paced rise when the
isotopic signal of breastmilk is reflected in recently formed bone, indicated by the highest
values at about one year of age during the peak period of complete dependency of the nursing
infant on the mother’s milk, followed by a decline in values. It is the period of decline that
provides the picture on the timing and duration of the weaning process. In contrast to the
Roman-period populations, a later Medieval series from the Wharram Percy site (c. AD
900–1500), England reveals an earlier and faster decline in δ15N and δ13C values such that
by two years of age, the values are strongly similar to adult values, indicating that weaning
had commenced well prior to the age of two years (Mays et al., 2002; Richards et al., 2002)
(Figure 8.14).
Analysis of δ15N and δ13C values in the an eighteenth- to nineteenth-century urban setting
of Spitalfields, London, England provides important insight into weaning practices, especially
involving a time during which women were more heavily involved in formal employment and
had a considerably greater presence in the workforce in general. Under these circumstances,
women who were employed away from the home were almost certainly likely to either wean
their infants earlier than in prior times or not breastfeed at all. As a result, mothers were
increasingly using artificial means of feeding their infants (Fildes, 1995; Lewis, 2007). The
Spitalfields series represents an affluent population, with individuals interred in a church
crypt between AD 1729 and 1859 (Nitsch et al., 2011). Analysis of nitrogen and stable carbon
isotopes reveals clear age changes representing the key phases of early childhood involving
dependence on breastfeeding and weaning (Figure 8.15). Hypothetically, an urban setting like
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15
14
13
12
d15N‰ 11
10
8
Fetal Adult
7
4
0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Age (years)
Figure 8.14 Infants from Wharram Percy exhibit enriched stable isotope nitrogen ratios
during breastfeeding years. By two years old, weaning has occurred, as demonstrated by the
decreased δ15N values, which fall within the average adult δ15N range. (From Mays et al.,
2002; reproduced with permission of authors and Antiquity.)
(a)
17
Female
Male
16 Unknown
15
14
d15N‰
13
12
11
10
0 5 10 15
Age (years)
Figure 8.15 Ratios of stable nitrogen (a) and carbon (b) isotopes from post-Medieval
Spitalfields reveal definitive dietary shifts in early childhood associated with breastfeeding
(enriched isotopic values) and weaning practices. The dashed line and bar represent the
average ( 1 standard deviation) isotope ratios for adult females from Spitalfields. (From
Nitsch et al., 2011; reproduced with permission of authors and John Wiley & Sons, Inc.)
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8.2 Isotopic analysis 329
(b)
-15
Female
Male
Unknown
-16
-17
-18
d13C‰
-19
-20
-21
-22
0 5 10 15
Age (years)
this would show minimal breastfeeding based on the age profile of the δ15N values, but this
series shows a majority of individuals having a period of exclusive breastfeeding. On the other
hand, there is considerable variation in values ranging from no or little departure from adult
values in the birth to two-year-old age range to moderate departure. These values suggest that
some infants were breastfed on an irregular basis, and some were likely never breastfed. The
stable carbon isotope record shows a similar pattern supporting the notion for a high degree of
variability with at least some individuals being breastfed. Because many children have known
ages at death, there is a more precise picture of when weaning was complete based on the
stable isotope record. This record shows that weaning was complete in the population between
the age of one and two years. Similarly, the later population from the nineteenth century
St. Thomas’s cemetery in Belleville, Ontario, displays a pattern of stable nitrogen isotope
values increasing in the first two months (δ15N ¼12.1‰), peaking, and then declining by the
age of one year (Herring et al., 1998). By two years old, all individuals had adopted a fully
adult diet (post-age two years, δ15N ¼10.4‰). Like the Spitalfields series, there is considerable
variation in the period of birth to 1.5 years, with some showing no evidence of breastfeeding.
The documentation of breastfeeding, weaning, and duration of weaning leading to adop-
tion of a fully adult diet suggests that dependence on breastfeeding and shortening of the
weaning period certainly occurred in historical, urban contexts in Europe and North America.
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330 Isotopic and elemental signatures of diet, nutrition, and life history
However, the examples discussed here likely do not represent the range of circumstances of
weaning as documented in archaeological settings. That is, the limited number of historical
contexts (Medieval and post-Medieval) have documented relatively recent times, and cer-
tainly not rural localities. The record of weaning as based on stable isotope analysis shows
considerable variation and not discernible trends. In this regard, in prehistoric archaeological
settings, there is a remarkable range of variation, from relatively short duration of total
breastfeeding (e.g., Native Americans: Schurr, 1998; Watts et al., 2011) to long duration of
total breastfeeding (e.g., Maya: Williams et al., 2005; Wright & Schwarcz, 1998; Gaya
confederacy, Korea: Choy et al., 2010). Moreover, consistent with Fogel and coworkers’
(1997) findings, there is no clear relationship between weaning and subsistence economy or
demographic pattern, at least as they are measured in bioarchaeological contexts (Schurr,
1998; Schurr & Powell, 2005). These findings suggest that while it has become possible to
document dietary signatures in young juveniles, providing an important perspective on
breastfeeding, weaning, and adoption of adult diets, the linkage between fertility, population
growth, and weaning is complex. Much has yet to be learned.
The terminus of the weaning process, as documented by the δ15N and/or δ13C curves,
corresponds closely with the rise in morbidity indicators of physiological stress, such as
hypoplasias and circular caries, and elevated mortality (Dupras & Tocheri, 2007; Fuller
et al., 2006; Herring et al., 1998; Lewis, 2007; Pearson et al., 2010; Reed, 1994). Fuller and
coworkers, for example, clearly document the association between decreasing δ15N values and
low δ13C values and the spike in childhood deaths. This is because once weaning processes
commence, infants simultaneously lose the passive immunity and complete nutrition pro-
vided by the mother that had been provided during the period of breastfeeding, thus com-
promising the infant’s immune system. This double jeopardy – weaning and reduced
immunity – explains elevated infant and child illness and mortality, both in the archaeo-
logical past and in a large number of settings around the world today, especially in developing
countries (Lewis, 2007). In the modern world, weaning and other associated health risks (e.g.,
poor diets and exposure to contaminated food) contribute to debilitation, diarrheal disease,
and early death (Motarjemi et al., 1993). The bioarchaeological record shows that this peak age
of risk has considerable antiquity.
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8.2 Isotopic analysis 331
Comparison of female average and male average isotopic signatures shows distinctive
trends indicating gender-based dietary differences between women and men in some but not
all settings. In a comprehensive study of age-related dietary changes in Isola Sacra, Italy,
combined δ15N and δ13C values reveal a gender distinction: males have higher average
δ15N and δ13C values than females, for the population as a whole, and within age categories
(Prowse et al., 2005). This record reveals that males were likely consuming more protein
(fish, animal products) and fewer carbohydrates (plants) than were females. Interestingly, the
δ13C values appear to converge in older adulthood with females becoming more like males.
This suggests that older females had greater access to protein in later life. These differences
match the record of status and gender in Imperial Rome. That is, men had considerably
higher status and access to resources than women. Likely, the dietary differences are part of
the status distinctions along gender lines. Similarly, written accounts document the advice
given by physicians, namely that it is preferable to limit womens’ food consumption, and for
them to avoid the so-called “cold and wet” foods, such as fish and meats (Prowse et al.,
2005:9). On the other hand, other stable isotope analysis of two other Imperial Rome series
(Casal Bertone, Castellaccio Europarco) reveals that diet varies along class lines (Killgrove &
Tykot, 2013). That is, higher-status individuals living closer to the center of Rome (Casal
Bertone) consumed more marine sources of protein than did lower-status individuals living
further from the city center (Castellaccio Europarco). Moreover, those living further from the
center ate more C4 food sources, likely millet, than did their periurban counterparts. These
findings show that diet was variable in Rome, and was likely influenced by status.
Patterns of variation in δ15N and δ13C values show generally greater access to animal
protein for males in Roman and Medieval-period England (Poundbury: Richards et al., 1998;
Queenford Farm: Fuller et al., 2006; Fishergate: Müldner & Richards, 2007b), Medieval Poland
(Giecz: Reitsema et al., 2010), early historic South Korea (Choy et al., 2010), the ancient Maya
(Whittington & Reed, 1997), and prehistoric preagricultural and agricultural settings in the
Ohio River valley and tributaries (Schurr & Powell, 2005). In the Ohio River valley region, the
differences were statistically significant only in the maize-based late prehistoric settings from
Sun Watch and Wickliffe sites. The patterns in these diverse settings suggest significant
differences in diet along gender lines, which is also apparent in terms of oral health, especially
caries prevalence (see Chapter 3). In addition, historical accounts document the differences in
gender roles in eastern North America where males hunted while women focused their
attention on horticulture (Swanton, 1946).
Schurr and Powell (2005) offer that while the variation in isotope values – especially for
nitrogen – may well reflect dietary differences, these disparities more likely represent physio-
logical distinctions (i.e., related to pregnancy and lactation) between adult females and males.
However, a physiological explanation does not account for a lack of sex differences in a
number of settings in North America and elsewhere, where females and males are similar in
their isotopic signatures or where females have higher δ15N values than males. For example,
in a comprehensive study of isotopic variability in the Georgia Bight and northern Florida, no
significant sex differences were identified in either δ15N or δ13C values, regardless of setting
(coastal, terrestrial) or period (Hutchinson et al., 1998; Larsen, Griffin et al., 2001; Larsen,
Schoeninger et al., 1992; and see Kellner & Schoeninger, 2008; Kjellström et al., 2009;
Pechenkina et al., 2005; Tafuri et al., 2009; Ubelaker, 2000). This variation is consistent with
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332 Isotopic and elemental signatures of diet, nutrition, and life history
the conclusions drawn by Schwarcz and Schoeninger (1991) that there is no or very limited
physiological basis for the differences seen in the comparison of adult females and males. On
the other hand, among the ancient Maya, there are clear sex differences, especially in elite
individuals, throughout the sequence from Pre-Classic through the Historic period (White,
2005). Even in the Historic period (Lamanai, Belize) when considerable social disruption and
other influences considerably altered native foodways, males showed greater consumption of
animal sources of protein than did females, as expressed in carbon and nitrogen stable isotope
ratios. This pattern of greater carnivory in males than females expresses clear gender differ-
ences and differential access to socially valued foods.
11
10
d15N‰
8 Males
Females
7
-20 -19 -18
d13C‰
Figure 8.16 δ13C and δ15N values from the Medieval population of Giecz (Poland) express
differential dietary patterns between the sexes. Males consume diets with more nitrogen
isotopically enriched animal protein. (Data from Reitsema et al. 2010.)
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8.2 Isotopic analysis 333
course of pregnancy in living humans (Fuller et al., 2006). In this regard, owing to morning
sickness and appetite reduction following conception, the loss in body weight occasions an
increase in δ15N values, but following recovery of appetite in later stages of pregnancy, body
weight increased and δ15N values increased sharply. These later changes reflect an increas-
ingly positive nitrogen balance and an attendant shift of nitrogen to tissue growth instead of
being excreted with urea. Similarly, various disease states (e.g., liver disease, diabetes, and
osteoporosis) affect isotope balances.
Stress and compromised health clearly have an influence on isotopic signatures in all of the
body tissues, which is displayed in the variation documented in the bones and teeth studied by
bioarchaeologists. The study of bone collagen from Nubian X-group skeletons reveals that
osteoporotic females have higher δ15N values than non-osteoporotic females, especially in the
third and fifth decades of life (White & Armelagos, 1997). This pattern is consistent with
histomorphometric differences between X-group individuals with osteoporotic and normal
skeletal tissue. In this setting, it appears that the differences in δ15N values reflect differences
in urea nitrogen excretion or altered renal processing and clearance of phosphorus and
calcium (White & Armelagos, 1997). In the hot desert setting of Nubia, water stress may
contribute to elevation in δ15N values. Lactation contributes to urea loss, which has been
linked theoretically to enrichment of 15N in bone collagen (Ambrose & DeNiro, 1986b). These
findings suggest that physiological disruption, more so than diet, may be responsible for the
variation in nitrogen stable isotopes (and bone loss) in this population (White & Armelagos,
1997). More generally, it strongly suggests that stable nitrogen isotopes reflect, at least in part,
non-dietary – especially physiological – factors and may serve as an indicator of osteoporosis
and likely other health conditions in past human populations.
8.2.4 Bivariate use of carbon and nitrogen stable isotope ratios: sorting out diets
in coastal environments
In many coastal areas of the world where marine and terrestrial foodways may be difficult to
identify owing to the complexities of local and regional food webs, the simultaneous analysis
of δ13C and δ15N is very informative. This has been especially so for coastal settings involving
maize production and consumption in New World contexts. Where marine foods and maize
were simultaneously consumed by human populations, the overlapping carbon isotope ratios
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334 Isotopic and elemental signatures of diet, nutrition, and life history
16
Early Prehistoric
14
Late Prehistoric
12
d15N‰
10
Mission
6
-20 -15 -10 -5
13
d C‰
Figure 8.17 Diachronic changes in isotope ratios from Early (black circle) and Late (white
circle) Prehistoric to Mission (hatched circle) periods on St. Catherines Island, Georgia,
reveal dietary shifts away from marine resources and toward terrestrially based plants
and animals. (From Larsen, Griffin et al., 2001; reproduced with permission of authors
and University Press of Florida.)
of marine and C4 foods preclude the discrimination of diets, at least with regard to assessing
the relative contribution of marine foods versus maize (Figure 8.1). That is, what might appear
to be a marine diet may simply be mimicking a C4 diet, and vice versa. In order to circumvent
this complication, the use of bivariate plots of δ15N and δ13C values has served to clarify diet
(Brown & Brown, 2011; Cooke et al., 1996; Katzenberg, 2008; Tomczak, 2003). In the Georgia
Bight, archaeological evidence indicates that marine foods continued to be heavily used
throughout prehistory and into the contact period (Larsen, 1982; Thompson & Turck, 2009,
2010; Thompson and Worth, 2012). Maize consumption is largely based on circumstantial
evidence in this region, including changes in settlement (increased population size and
aggregation), increasing social complexity, and increasing morbidity (e.g., dental caries,
periosteal reactions; see Chapters 2 and 3). Owing to extraordinarily poor preservation of
plant remains in late prehistoric archaeological sites, dietary reconstruction based on the plant
record is inconclusive as to the relative importance of maize as a routine food source.
Analysis of collagen samples from prehistoric foragers and farmers and early and late
contact mission native populations alleviates the incomplete and vague picture of diet in this
setting. Isotopic analysis reveals a distinctive temporal trend showing increasingly less
negative δ13C values and less positive δ15N values, both in local settings (e.g., St. Catherines
Island: Figure 8.17) and in the larger region (Figure 8.18) (Hutchinson et al., 1998; Larsen,
Griffin et al., 2001; Larsen, Hutchinson et al., 2001; Larsen, Schoeninger et al., 1992;
Schoeninger, 2009; Schoeninger et al., 1990). This trend indicates an increasing focus on
terrestrial plants (maize) and animals, coupled with a decreasing reliance on marine resources.
This shift commences during the twelfth century AD with the adoption of maize as a dietary
staple, reaching its peak and extreme degree of focus in Spanish mission native populations
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8.2 Isotopic analysis 335
14
13
12
11
N‰
10
15
d
6
-18 -16 -14 -12 -10 -8
13
d C‰
Georgia early prehistoric, coast Florida mission, inland
Georgia late prehistoric, coast Georgia early prehistoric, inland
Florida late prehistoric, coast Florida early prehistoric, inland
Georgia mission, coast Georgia late prehistoric, inland
Florida mission, coast Florida late prehistoric, inland
Figure 8.18 Mean stable carbon and nitrogen isotope values from coastal and inland Florida
and Georgia skeletal series demonstrate the significant change in diets from prehistoric to
mission periods. The mission-period samples (box) show increased consumption of C4
plants relative to earlier periods, along with a narrower dietary range.
inhabiting St. Catherines Island, Georgia, in the early to mid-seventeenth century, and in the
later descendant groups on Amelia Island, Florida (Hutchinson et al., 1998, 2000; Larsen,
Griffin et al., 2001; Schoeninger, 2009). Overall, this temporal record reveals a near abandon-
ment of marine foods in the mission era (post-AD 1600) in the Georgia Bight. Moreover, at
several of the missions, more negative δ13C values and/or more positive δ15N values in elite
individuals (those interred closer to the church altar) than in the non-elite (those interred
further from the church altar) suggest differential access to high quality foods (Larsen, Huynh
et al., 1996; Stojanowski, 2013a, 2013b).
The prehistoric dietary changes documented in the Georgia Bight are, in some respects, also
present in the western Florida panhandle. In this regard, the late prehistoric Mississippian
center at Lake Jackson and Waddells Pond sites have δ13C values representing a significant
maize diet (mean ¼ −13.7‰; Larsen, Griffin et al., 2001). These are inland localities, and as
predicted, there is a non-marine signature of nitrogen. In sharp contrast to the late prehistoric
Georgia coastal localities, late prehistoric coastal and non-coastal stable carbon isotope ratios
for populations from the northern and central peninsula of Florida are noticeably negative,
expressing results that indicate little C4 (maize) consumption. Maize may have been present in
these diets and there was considerable variation in diet, depending on habitat (Kelly et al.,
2006), but the isotope record suggests that maize or other C4 foods were not predominant in
diet, certainly not to the same extent as in most of the rest of eastern North America. However,
as with the Georgia coastal mission populations, other historic settings in Florida, mission and
non-mission, express considerably higher δ13C values, indicating the spread and adoption of
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336 Isotopic and elemental signatures of diet, nutrition, and life history
maize after European contact (Hutchinson, 2006; Hutchinson & Norr, 1994; Hutchinson et al.,
1998, 2000; Larsen, Griffin et al., 2001). The sudden shift in diet in contact-era Florida and
Georgia suggests that dietary change was a largely cultural, economic, and political endeavor
occasioned by the arrival of Europeans and exploitation of native populations in food
production (Worth, 2001). In contrast, the shift to maize in the prehistoric era likely involved
an adaptive strategy.
Additional analysis of collagen from the late prehistoric Irene Mound site on the north
Georgia coast indicates that although maize generally increased in importance, there is a
marked decrease in δ13C values for the period immediately prior to European contact,
suggesting that use of maize temporarily decreased following its initial introduction in the
region (Larsen, Schoeninger et al., 1992). The decline in maize consumption may be linked
with social and environmental disruption in the Mississippian period in this region during late
prehistory. Similar declines in use of maize during the period preceding contact by Europeans
has been documented in at least one other major Mississippian center in the American
Southeast, Moundville, Alabama (Schoeninger et al., 2000). These findings from Georgia
and Alabama point to the highly variable pattern of maize use in eastern North America.
Analysis of carbon and nitrogen stable isotope ratios helps to clarify the relative consump-
tion of marine foods and maize in other coastal settings in the Americas, including New
England (Bourque & Krueger, 1994; Little & Schoeninger, 1995; Medaglia et al., 1990), Gulf
coast Florida (Hutchinson, 2004; Hutchinson & Norr, 1994), coastal North Carolina (Hutch-
inson, 2002; Hutchinson et al., 2007), Panama and Costa Rica (Norr, 1991), Belize (White &
Schwarcz, 1989), and Peru (Tomczak, 2003). Comparisons of inner and outer coastal popula-
tions from North Carolina show that, in contrast to Georgia groups where agriculture was
adopted regardless of location (inner vs. outer coast), the outer coastal North Carolina groups
continued to rely on oceanic and estuarine resources throughout later prehistory, whereas inner
coastal groups adopted maize farming after AD 1000 (Hutchinson et al., 2007). Similarly,
Chiribaya culture (AD 1000–1450) sites representing coastal (San Geronimo), head of the
coastal valley (Chiribaya Alta), near coastal, but with material culture showing little focus on
marine resource extraction (Chiribaya Baja), and middle valley, located some distance from the
coast with material culture indicating terrestrial exploitation (Yaral), provide an important
context for addressing the degree to which populations with access to marine resources
committed to maize agriculture (Tomczak, 2003). Bivariate comparison of nitrogen and carbon
reveals that diets in this setting show a progressive decrease in both δ13C and δ15N values.
Importantly, this variation indicates that diets are determined on the basis of local extractive
strategies with consumption being highly localized. Predictably, the δ15N values show that
populations living closest to the coast consume the highest amount of seafood (San Geronimo).
The proportion of seafood that was consumed by these populations, at progressive points
further from the coast, reaches its lowest at Yaral, where diets included marine foods, but were
dominated by consumption of C4 foods and animals eating C4 foods (e.g., llama).
Even where maize is not utilized in New World settings, the isotopic compositions involv-
ing carbon and nitrogen are not always clear. Walker and DeNiro (1986) showed that the
simultaneous use of carbon and nitrogen ratios serves to clarify the potentially conflicting
signatures of either element alone in marine environments of the Santa Barbara Channel
region. Archaeological and biocultural evidence indicates a shift in dietary emphasis from
terrestrial to marine resources in later prehistory in this region as well as a generally heavier
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8.2 Isotopic analysis 337
emphasis on marine foods on the islands than on the nearby mainland coast or interior.
Isotopic analysis reveals that the carbon and nitrogen ratios in native populations progres-
sively decrease from the islands to the mainland coast and mainland interior, indicating a
strong correlation of diet with geographical location (Walker & DeNiro, 1986). These geo-
graphic distinctions in isotopic signatures show that terrestrial diets were emphasized in
mainland populations, and marine diets were emphasized in island populations. Additionally,
isotope values from older sites on the mainland coast and interior are lower for both carbon
and nitrogen than those from younger sites in the same region. This finding is consistent with
other archaeological and biocultural evidence suggesting that mainland-oriented diets
became more marine-oriented later in time.
In the Bahamas where terrestrial, reef, and deep-ocean habitats were utilized by prehistoric
populations, Keegan and DeNiro (1988) analyzed numerous potential foods from these
settings. Some reef fish have higher δ13C values and lower δ15N values than other ocean fish.
The analysis of carbon and nitrogen isotope ratios in archaeological human samples from this
setting indicates that prehistoric populations fished primarily in seagrass and coral-reef
ecosystems. The most negative δ13C values are in the initial period of occupation; later
populations have less negative values. This temporal shift in stable carbon isotope ratios
suggests that Caribbean populations became increasingly marine-oriented in later prehistory.
In coastal settings in both the western and eastern Pacific Rim, there has long been
confusion over the role of marine and terrestrial foods consumed. For example, in Japan
and the American Northwest coast, while it has long been understood that both marine and
terrestrial foods were consumed by Jomon foragers, the importance of each has been unclear
(Chisholm & Koike, 1999; Chisholm et al., 1992; Minagawa & Akazawa, 1992). Isotopic
analysis of human remains from a series of sites revealed that hunter-gatherer-fishers from
Hokkaido of northern Japan, from Jomon through Ainu periods, showed few temporal
distinctions and with a pattern of carbon and nitrogen isotope ratios indicative of maritime
mammals. Jomon-period individuals from coastal sites on Honshu, however, consumed a
lesser amount of maritime mammals, and instead relied on combinations of plant food,
terrestrial mammals, and fish. In contrast, populations from the inland site of Kitamura (Late
Jomon, 4000 yBP) on the main Japanese island display very negative δ13C and less positive
δ15N values, signaling a largely terrestrial diet. Interestingly, some complexity in these dietary
patterns is observed for the inland populations: individuals from the Boji site (Late Jomon,
c. 4000 yBP) have isotopic signatures that are consistent with the consumption of freshwater
fish (Yoneda et al., 2004).
Although representing a very different ecosystem, some island populations of the western
Pacific that acquired agriculture show a similar temporal pattern of shifts from heavily marine
to heavily terrestrial diet. Non-isotopic archaeological data from a number of settings show
that populations likely adopted agricultural subsistence within the last 2000 years of prehis-
tory, but marine foods remained important dietary targets (Field et al., 2009). In order to
identify dietary change in the southwest Pacific, Field and coworkers (2009) documented
stable isotope ratios for carbon and nitrogen in Fiji, from the period of 2700–300 yBP from
both coastal and terrestrial contexts. The general pattern is clear: the earliest samples from
Waya Island show an isotopic pattern that is fully marine, similar to other Pacific settings,
such as Rota, Guam, and Saipan in the Marianas (Ambrose et al., 1997) and Tutuila in
American Samoa (Valentin et al., 2011). In contrast, the later samples from the Sigatoka
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338 Isotopic and elemental signatures of diet, nutrition, and life history
Valley display values that characterize a terrestrial adaptation involving domesticated plants.
Overall the record for Fiji reveals a shift from marine to terrestrial (Field et al., 2009; Valentin
et al., 2006, 2011), with the change occurring earlier on larger islands having more arable land
than smaller islands. Interestingly, however, while the general trend toward terrestrialization
of diet is clear, exceptions to the pattern suggest local variation in diet based on access to
marine foods (Jones & Quinn, 2009).
Much of this record documenting coastal environments and resource acquisition shows the
complex nature of dietary interpretation based on the analysis of stable isotope ratios.
However, the use of bivariate plots of stable carbon and nitrogen values alleviated some of
the issues with dietary interpretation. There are many coastal environments worldwide where
use of these plots is quite complex, such as in large freshwater lakes (Katzenberg & Weber,
2009; Katzenberg et al., 2012). For example, in the Lake Baikal region (Siberia, Russian
Federation), Katzenberg and coworkers used stable isotopes to reveal variation in use of
terrestrial mammals versus freshwater marine sources (fish and seals), largely owing to
differences in the stable carbon and nitrogen isotope signatures. Adding to the complexity
of the setting are findings relating to isotopic signatures of freshwater fish. The analysis of
Katzenberg and coworkers (2012) reveals considerable variation. However, a number of
patterns emerge, including important distinctions between specific settings that provide a
means to show where fish are caught. With this record in hand, temporal comparisons of the
human samples show a more intensive focus on freshwater fish in the Early Neolithic
(8000–6800 yBP) compared to a wider dietary breadth in the Late Neolithic and Early Bronze
Age (6200–4000 yBP).
The record of dietary reconstruction via bivariate plots has clearly provided an invaluable
reference point for interpreting past diets. Bivariate plots of δ13C from collagen and carbonate
have also been established, based on controlled feeding experiments (Kellner & Schoeninger,
2007). Regression models derived from these experiments provide a powerful source for
developing considerable precision to dietary reconstructions and serve to explain why a
bivariate characterization of isotopes is important. That is, collagen and carbonate captures
the respective fish and millet in the diet, but the model captures both.
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8.2 Isotopic analysis 339
Strontium isotopes
Strontium has proven especially robust in identifying individuals who may be migrants and
for identifying migration during a person’s lifetime by comparing isotopic variation of earlier
forming tissues with later forming tissues, such as teeth and bone in adults or earlier forming
teeth and later forming teeth (or bone) in juveniles. Bones and teeth are both used, but owing
to its lower potential for diagenesis, tooth enamel is the preferred tissue for analysis.
Strontium is present naturally in the geological environment, including four naturally occur-
ring isotopes – three stable (84Sr, 86Sr, and 88Sr) and one radiogenic, unstable (87Sr) isotope
that is formed by the decay of 87Rb. Strontium enters the ecological system, originating in
bedrock and then moving into soil and groundwater, and subsequently enters the food chain
(Price et al., 2002). However, unlike the other elemental isotopes described earlier, strontium
isotopes have very small relative mass differences – the isotope composition remains rela-
tively unchanged from its origin through the food chain (but see Knudson et al., 2010). Thus,
unlike carbon and nitrogen stable isotope ratios, there is not a diet-to-tissue spacing seen, for
example, in trophic level changes in nitrogen. The strontium isotope ratios determined from
archaeological skeletal and dental tissues directly reflect the local geochemistry, and the
geochemical isotope ratio is passed unaltered from food to consumer (Ericson, 1985; Price
et al., 2002; Sealy et al., 1991).
Geochemistry of the decay process has revealed that the ratio of 87Rb/87Sr is directly
proportional to the ratio of 87Sr/86Sr, and it shows considerable variation in local geology
in some regions, with earlier forming rocks having higher ratios of 87Sr/86Sr than in later
forming rocks (Bentley, 2006). When local and regional geochemistry show distinctive
patterns of values in local settings and regions, the ratio of 87Rb/87Sr provides an invaluable
tool to identify migrants in a population. For example, owing to the common geological
history in Denmark and across southern Scandinavia, northern Germany, and northern
Poland, the 87Sr/86Sr values are generally homogeneous with values ranging from about
0.7090 to 0.7108 (Price et al., 2011).
The values of human tissues derived from archaeological contexts are compared with the
local geological values. However, in practice, values derived from analysis of bones and teeth
are compared to bioavailable values of the burial context, including soil, archaeological
fauna, modern animals, and archaeological bones and teeth (Price et al., 2002, 2011). The
immediate analytical goal is to characterize the local population isotopically and to identify
individuals who are outliers (“foreigners”) from the local isotopic composition. Presumably,
these outliers represent persons originating from another provenience and having matured in
another geographic setting. These individual non-locals are identified as those whose stron-
tium isotope values fall outside the local range of strontium isotope variation (Bentley, 2006),
defined by some as plus or minus two standard deviations from the mean of modern, non-
ranging fauna isotopic ratios (Knudson, 2008) (Figure 8.19).
Initial bioarchaeological applications of these biogeochemical approaches to reconstructing
and interpreting residence in a handful of settings (Ericson, 1985, 1989; Grupe, 1995; Price,
Grupe et al., 1994; Price, Johnson et al., 1994) revealed that variation in strontium isotope
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340 Isotopic and elemental signatures of diet, nutrition, and life history
0.720
Tiwanaku Tilata, Coya Coya-3 Solcor-3 Chen Chen
0.719 Kirawi, Oriental
Iwawe
0.718
0.717
0.716
0.715
0.714
87Sr/ 86Sr
0.713
0.712
0.711
0.710 87
Sr/ 86Sr = 0.7087−0.7106 for
southeastern Lake Titicaca Basin
0.709
0.708
0.707
0.706
0.705
Figure 8.19 Strontium isotope ratios (87Sr/86Sr) from human tooth enamel samples across
Tiwanaku sites, South Central Andes. The horizontal gray bar captures the local range in
strontium isotope ratios for Lake Titicaca Basin, which is determined by sampling strontium
isotope ratios from local archaeological fauna. Isotopic values from human teeth enamel,
which fall within this range (e.g., seven individuals from the Tiwanaku sample), indicate
these individuals lived within the Lake Titicaca Basin during childhood. (From Knudson,
2008.)
ratios can inform our understanding of population dynamics as they relate to identification of
non-locals and locals in archaeological skeletal series. A pilot study of isotope ratios of dental
enamel and bone from late prehistoric Grasshopper and Walnut Creek sites, Arizona shows
that only some individuals share values similar to local geology (Price, Johnson et al., 1994).
These individuals likely spent their lives at their birth residence, whereas others appear to have
been born elsewhere and moved to their place of residence at some later time. Comparison of
strontium ratios in tooth enamel and bone from the Bell Beaker period (2500–2000 BC) of
southern Bavaria also reveals significant variation (Price, Grupe et al., 1994), which appears to
be more pronounced in the earlier than the later part of the period (Grupe, 1995). This finding
suggests that populations in the later period may have been more sedentary, which is
consistent with settlement analysis from conventional archaeological data.
From both New World and Old World settings, there is a rich body of work showing
important elements of variation in strontium isotopes. Analysis of this variation reveals key
interactions ranging from intercommunity relationships to long-distance migration involving
relatively few individuals, such as in trading, or recruitment of individuals into some aspect of
society, such as the formation of military groups. In summary, identification of non-local
individuals via their bone chemistry in general, but especially via strontium stable isotope
ratios, provides a powerful means of illuminating social, political, and economic dynamics
locally and regionally.
In the Americas, there has been a long-standing interest in the influence of dominant,
complex societies on a region. In the South American Andes, during the last two millennia
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8.2 Isotopic analysis 341
prior to European conquest, a number of powerful state systems rose and dominated all or
parts of the modern countries of Peru, Ecuador, Bolivia, and Chile. In most of these settings,
the record of the rising state is well documented archaeologically by ceramic styles, iconog-
raphy, architecture, settlement pattern, and other material evidence. As the state develops, it
first originates in a “heartland” at a particular locality or set of localities representing various
communities. Over time, the state influences surrounding regions, sometimes covering many
thousands of square miles, including part or all of the Andes Mountains and coast. Anthro-
pologists and other social scientists are keenly interested in the mechanics of the rise and
spread of these early South American state systems. Did these states begin to dominate
surrounding societies via migration, the spread of ideology, violence and subjugation (see
Chapter 4), trade and commerce, or by some other means that is simply invisible in the
archaeological record? For example, in the Middle Horizon period of the South Central Andes
(AD 500–1000), a very detailed record shows that societies throughout the region were
influenced by the heartland of the Tiwanaku state, centered at the site of Tiwanaku near Lake
Titicaca, Bolivia. A number of archaeologists argued that this influence resulted from state
expansion and incorporation of a remarkably large region of the Peruvian Andes by the
Tiwanaku polity. By gradually colonizing the region, the state played a role in spreading
material culture and political, cultural, and economic systems (Kolata, 1993). Other authorities
propose that the conquest of the region was accomplished by commercial trade (Dillehay &
Núñez, 1988) and spread of ideology via what Browman (1978:327) referred to as “proselyt-
izing merchant missionaries.”
The general hypothesis that Tiwanaku and other state systems in the Andes in prehistory
involved colonization can be tested by documenting whether “foreign” individuals were
present in these communities via analysis of isotope ratios in the skeletons of the people
who were once members of the communities comprising the Tiwanaku state system. If it can
be demonstrated that non-local isotope ratios are present in a population displaying a
particular pattern of variation of local isotope ratios, then a case can be made for the presence
of foreign persons. However, that alone does not tell the entire story. Rather, it is the context
provided by other data that present the case for mechanisms of state development, rise, and
domination of a larger geography that included once-independent societies.
Analysis of strontium isotope ratios from sites representing communities having Tiwanaku-
style material culture and architecture in the Moquegua Valley of southern Peru – Omo, Río
Muerto, and Chen Chen – provides important documentation of the potential role of actual
movement of people (Knudson & Price, 2007; Knudson et al., 2004). In these communities,
architecture and material culture provide evidence to suggest that individuals living there
were either affiliated with or were originally from the Tiwanaku heartland. Omo, for example,
has the only Tiwanaku-style temple outside of the Tiwanaku heartland (Goldstein, 1992).
While the Tiwanaku signature in material culture – from pots to buildings – is ubiquitous
throughout the region, it remains possible that local people exclusively adopted it as they
came under the influence of the Tiwanaku state in the absence or very marginal presence of
individuals from the heartland. Analysis of strontium isotopes in dental enamel of individuals
from Chen Chen, Moquegua Valley (southern Peru) and Tiwanaku (Bolivia) reveal several
findings that suggest the movement of actual people. The 87Sr/86Sr baseline biogeochemical
ratios are distinctively different on comparing the two regions. That is, the 87Sr/86Sr ratios for
the Moquegua Valley range from 0.7059 to 0.7067, and the 87Sr/86Sr ratios for Tiwanaku,
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342 Isotopic and elemental signatures of diet, nutrition, and life history
Lake Titicaca basin range from 0.7083 to 0.7111. Analysis of strontium ratios from eight
individuals from Chen Chen reveals ratios that fall well outside the range for the Moquegua
Valley (Knudson & Price, 2007; Knudson et al., 2004). Two of the ratios, in fact, match the
isotope ratios that would be expected if they were from Tiwanaku. Both are women, and
the data suggest that when they were three to four years of age they were residents of the
Tiwanaku heartland. At the site of Tiwanaku, based on strontium isotope ratios, three of
the 10 individuals were non-local. Interestingly, all three were interred in an elite quarter of
the monumental district of Tiwanaku. They included a young adult female and a young adult
male, both likely sacrificial offerings for the dedication of important ritual structures. The
third individual, also an adult, was a dedicatory offering at another ritual structure.
Weathering on the skeleton suggests that this body may have been displayed publically prior
to interment (Blom et al., 2003; Knudson & Price, 2007; Knudson et al., 2004).
The isotopic record from Chen Chen and Tiwanaku reveals that non-local individuals are
present at both centers. While the data sets are small, they nevertheless imply the presence of a
larger network involving movement of people from a center (Tiwanaku), taking place after the
formation of teeth. Regardless of scope, these analyses support the hypothesis that non-local
individuals are present at two important communities in the larger sphere of the
Tiwanaku state.
This is not to say that all of Tiwanaku territory – heartland and hinterland – involved actual
movement of people as a means of spreading political, social, and economic influence from
core to periphery. In this regard, communities from the San Pedro de Atacama oases and the
Loa River valley of northern Chile, while within the Tiwanaku sphere of influence, have no
individuals expressing non-local stable isotope ratios (Knudson, 2008; Knudson & Torres-
Rouff, 2009). Thus, in consideration of the Tiwanaku polity as a whole, while the influence of
the polity clearly shows that some communities were partially inhabited by individuals
originating from the core Lake Titicaca region and migrating to the periphery (e.g., the
Moquegua Valley), in other settings, local communities adopted elements of the dominant
society via other means (Knudson, 2008).
Isotopic variation as it relates to strontium documents the presence of non-locals within
communities and evidence of movement, likely explaining, in part, mechanisms of cultural,
social, and economic influence involving a dominant core polity. Similarly, an isotope record
of regional mobility documents the widespread influence of a dominant polity and long-
distance interaction in the Andes, including in states, such as the Wari empire (Conlee et al.,
2009; Knudson & Tung, 2011; Slovak et al., 2009; Tung & Knudson, 2008) and the Inca empire
(Turner et al., 2009). Other settings in pre-Hispanic Latin America also reveal a record of
movement of people, explaining, at least in part, the spread of a dominant system over
remarkable distances. At Teotihuacan (AD 1–650) in the Basin of Mexico near Mexico City,
archaeologists have identified foreign architecture centered in two ethnic compounds. The
skeletal remains are assumed to be foreigners living in these compounds (Price et al., 2000).
A relatively high degree of variation in stable isotope values in the tooth enamel but little
variation in the values in bone suggests that some individuals were born elsewhere and
migrated to the city in later life. This finding is consistent with the notion that the fast growth
of the city, while perhaps due, at least in part, to elevated fertility and birthrate, must also
have been fueled by significant migration (Price et al., 2000; and see Blanton, 1981). In at
least one setting in the city, numerous individuals display evidence of cutmarks and
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8.2 Isotopic analysis 343
mutilation patterns consistent with sacrifice. For example, at the Pyramid of the Moon, only
three of the 40 victims have strontium isotope ratios that suggest local derivation; all others
have values from a range of settings throughout Mesoamerica (Price et al., 2007).
While much of this record emphasizes aspects of empire building and social dynamics
under a variety of circumstances, it is also the case that inter-regional trade between cultures
in prehispanic Latin America is an important element in both of the settings described from
highly complex societies (Price et al., 2000; Wright et al., 2010). Similarly, it is important in
simple societies that are often incorrectly thought of as being isolated, such as island inhabit-
ants. However, stable isotope analysis reveals significant mobility and exchange in these
settings (Hoogland et al., 2010). Moreover, the strontium isotope record provides a means of
tracing the origins of enslaved individuals and patterns of forced migration of persons in the
African diaspora of postcolonial North America (Goodman et al., 2009; Price et al., 2006).
Investigation of stable isotope variation provides a window onto the role of migration in
the formation of new societies in the Old World, such as low mobility Holocene hunter-
gatherers in the southern Sahara Desert (Stojanowski & Knudson, 2011), late prehistoric
southeastern Arabia (Gregoricka, 2013a), Neolithic and Copper Age eastern Europe (Giblin,
2009), and Iron Age northeast Thailand (Cox et al., 2011). In the Oman Peninsula during the
third millennium BC, an abundance of foreign grave goods in local communal tombs led to
speculation that non-locals were frequently included as tomb members, an assumption
corroborated by the region’s growing involvement in inter-regional trade across the Persian
Gulf with major powers, including Mesopotamia and the Indus Valley. However, radiogenic
strontium isotope analysis revealed a different pattern, in which the majority of those interred
in these Bronze Age cairns were indeed local and displayed little isotopic variability indicative
of an increasingly sedentary lifestyle with the adoption of date palm agriculture (Gregoricka,
2013b) (Figure 8.20). Nevertheless, the few non-locals identified were buried according to
local customs, suggesting a form of “fictive kinship” as a means of strengthening political or
economic relationships (Gregoricka, 2013b).
This record contrasts with other settings showing relatively high levels of residential
mobility and population movement, colonization, and trade, such as in northwestern Africa
and the Near East (Buzon et al., 2007; Perry et al., 2008, 2009, 2012), and Imperial Rome
(Killgrove, 2010). Folklore, historical documentation, sagas, and archaeological documenta-
tion record the famously mobile Vikings of Scandinavia, who moved throughout northern
Europe, with travel extending throughout the continent and the northern Atlantic, and
including colonization of the British Isles, Iceland, Greenland, and North America, all by
the turn of the eleventh century AD (Fitzhugh & Ward, 2000; Jones, 1984). While widespread
movement was the case, identification of locals and non-locals in a number of settings reveals
alternative strategies of colonization and domination of regions that came under the influence
of the Vikings. In northwest Scotland in the New Hebrides islands, an area colonized by
Norwegian Vikings, four individuals in a sample of 20 were clearly distinctive in their isotope
ratios, enough to be identified as non-local (Montgomery & Evans, 2006). Although non-
local, their isotope ratios are strongly dissimilar to those from Norway, but may source these
individuals to either further south in England or Denmark. In contrast, a series of remains
from Viking-period Dublin reveal that all but one of the isotope values are local (Knudson,
O’Donnabhain et al., 2012). All values – from both enamel and bone – are remarkably
homogeneous and suggest that none of the individuals are from Viking populations
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344 Isotopic and elemental signatures of diet, nutrition, and life history
0.7210
0.7195
0.7190
0.7185 MW 197
TA 161
0.7180
2700 BC 2500 BC 2400 BC 2300 BC 2200 BC 2000 BC
Figure 8.20 Strontium isotope ratios from human tooth enamel are presented for Bronze Age
tombs in Arabia. The local strontium isotope range, represented by the horizontal line with
standard deviations, was established by local faunal references. As the strontium ratios for
tooth enamel depict, individuals associated with these tombs spent their childhood years
within this local region. (From Gregoricka, 2013b; reproduced with permission of author
and Elsevier Ltd.)
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8.2 Isotopic analysis 345
0.720
0.716
87Sr/ 86Sr
0.712
0.708
Figure 8.21 Strontium isotope ratios from human (black bars) and faunal (gray bars, far left)
remains from Medieval Trelleborg, Denmark. Over two-thirds of the human isotope values
are outside the local geological strontium isotope range (horizontal gray bar), which
suggests that the majority of these individuals grew up on foreign, non-Scandinavian soil.
(From Price et al., 2011; reproduced with permission of authors and Antiquity.)
Harald Bluetooth’s military stronghold. In contrast to the Viking community at Dublin, the
Danish record as it is revealed in a military context is consistent with the high degree of
residential mobility in Viking-period Scandinavia – the military were recruited from a wide
variety of settings and there was considerable movement of individuals, sometimes involving
all ages and both sexes and with different health profiles in comparison with local and non-
local persons (Groves et al., 2013).
Interestingly, this pattern of widespread recruitment contrasts with the Imperial Roman
period pattern in the northern frontier British Isles where isotopic signatures indicate that
military recruiting was done on a local level, certainly mostly within the British Isles (Chenery
et al., 2011; regarding the diversity of the Roman diaspora in Britain, see Eckardt et al., 2009;
Evans et al., 2006; Hakenbeck et al., 2010; Leach et al., 2009; Montgomery et al., 2011;
Pollard et al., 2011).
Much of this record of individual movement and identification of foreign individuals
documents patterns of variation that speak to the movement of persons within the context
of wider population movements. Analysis of sex differences in adults (tooth enamel) provides
insights into marital residence patterns in past societies. In this regard, during the period of
agricultural intensification in Ban Chiang, Thailand, there is a strontium signature showing
considerably more variation in males than females (Bentley et al., 2005). This suggests a
matrilocal residence pattern where women grow up in the community consuming the same
foods, whereas men are drawn from a wider geographic spectrum and consumed different
foods. In contrast, females from Neolithic Central Europe have considerably more variation in
their strontium values than males, suggesting a patrilocal residence pattern (Bentley et al.,
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346 Isotopic and elemental signatures of diet, nutrition, and life history
2002). More precise information on biological patterns relating to marriage patterns and
inferences of residence are provided by biodistance analysis of genetic variation (see
Chapter 9). However, these highly distinctive patterns in strontium are strongly suggestive. In
this regard, at Ban Chiang, the strontium signatures may simply reflect differences in female
and male diets where girls and women restrict their food consumption to local sources and boys
and men have access to foods hunted some distance away. The combined records of strontium
isotopic signatures with biodistance analysis will help test these important hypotheses, espe-
cially with respect to processes behind the introduction and intensification of farming, which
must have been influenced by social structure and marriage residence patterning.
Oxygen isotopes
Climate has a tremendous influence on human adaptation, impacting such factors as resource
productivity and where people live. Thus, climatic patterns are potentially useful for inter-
preting land use patterns. δ18O values (determined from stable isotopes, 18O and 16O) of
terrestrial water vary in relation to climate, especially by temperature and humidity (Kolodny
et al., 1983; Luz et al., 1990). Because mammalian skeletal and dental phosphate and
carbonate are in equilibrium with body water, the phosphate oxygen isotope composition of
these hard tissues directly reflects the temperature and climate at the time of tissue formation
(Fricke et al., 1995). In order to test the hypothesis that hard tissues from archaeological
settings can be used to track the history of regional climates, Fricke and coworkers (1995)
determined temporal change in values from archaeological dental enamel (Eskimos and
Europeans) in coastal western Greenland and Denmark for the periods preceding, during,
and following the so-called “Little Ice Age” of the Medieval period (AD 1400–1700). There is a
3‰ decrease in δ18O values from earlier to later periods, which is consistent with increased
cooling documented in historical records describing climates in the North Atlantic
region. These changes are in accordance with other studies indicating increasing dietary
and climatological stress and eventual abandonment of Greenland by Vikings during the
Medieval period (Buckland et al., 1996; Scott et al., 1992).
There is a growing record to document residential mobility, place of origin (from enamel),
and later migration to a new geographic region (from bone or teeth that develop later in life),
thus making it possible to distinguish locals from non-locals (Katzenberg, 2008; White,
Spence et al., 2004). One of the neighborhoods at Teotihuacan on the western edge of the
city contains archaeological evidence – architecture and material culture – suggesting that it
was occupied by immigrants from Oaxaca. Consistent with the strontium isotope evidence
(Price et al., 2000), analysis of oxygen isotopes reveals values that document the clear
presence of non-locals, both in children (based on enamel) and in later life (based on bone),
including but not limited to immigration from Monte Albán in the Valley of Oaxaca (White
et al., 2007). Overall, the high degree of variation seen in the stable oxygen isotope values
expresses a picture of wide circulation of people in the region. Much of the relocation over the
course of lifetimes of individuals is due to normative behaviors related to trade, labor, and the
wide variety of behaviors responsible for movement in any society, past or present. In this
setting, analysis of oxygen isotope variation coupled with strontium isotope variation is
related to some very specific social behaviors. That is, analysis of one distinctive set of human
remains from Teotihuacan – those military or other guardians sacrificed and interred at key
ritual temples – provides important perspective on social events (White et al., 2007). While the
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8.3 Elemental analysis 347
motivation for the movement of individuals across the region to Teotihuacan is complex, at
least some of the movement of individuals is related to highly specific social practices,
including sacrifice.
As with strontium isotopes, analysis of oxygen isotopes facilitates the testing of long-
standing hypotheses about past social interaction, sometimes involving travel over consider-
able distances. While archaeological evidence based on material culture provides a snapshot
of regional interaction, in the absence of historical records, it does not provide the documen-
tation for mechanisms involving movement of people or the spread of ideas and influence in
the absence of migration. The strontium isotope record reveals important evidence for of
regions having complex social and population histories (see earlier). As with Mesoamerica, the
oxygen isotope record is consistent with the strontium isotope record showing variation in
immigration patterns, such as in Imperial Rome (Prowse et al., 2007) and in Roman colonies
such as the British Isles (Eckardt et al., 2009; Evans et al., 2006; Leach et al., 2009).
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348 Isotopic and elemental signatures of diet, nutrition, and life history
et al., 1990; Pate et al., 1989; Price et al., 1992; Sandford, 1993b; Sillen, 1989). Analyses
generally focus on three areas: (1) alkaline-earth elements (strontium and barium); (2) multi-
element analysis; and (3) single-element analysis.
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8.3 Elemental analysis 349
Piepenbrink, 1988; Lambert et al., 1983, 1984; Nelson & Sauer, 1984; Nelson et al., 1986; Pate
& Hutton, 1988; Pate et al., 1991; Price, 1989; Sillen, 1981, 1992; and others). In addition to
the aforementioned factors influencing strontium in bone, there are a number of behavioral
and dietary influences. A variety of evidence shows that weaning (Sillen & Smith, 1984),
pregnancy and lactation (Blakely, 1989; Radosevich, 1989, 1993), consumption of some
nondomesticated plants (e.g., nuts) and shellfish (Benfer, 1984; Buikstra et al., 1989; Byrne
& Parris, 1987; Kyle, 1986; Schoeninger & Peebles, 1981), and health (Dolphin et al., 2005;
Lane & Peach, 1997) will influence values. In summary, reliable biological signals can only be
acquired through careful consideration of a range of factors influencing strontium in bone
tissue. Much of the literature documenting strontium in archaeological bone has not con-
trolled for diagenesis and other factors. Thus, the results based only on strontium should be
viewed with skepticism (Katzenberg, 1993).
Studies of strontium provide results that are suggestive of dietary strategies (Katzenberg,
1984, 1993; Radosevich, 1989, 1993; Runia, 1987b; Schoeninger, 1981, 1982; Sillen, 1981;
and others). Using strontium values from herbivore and carnivore controls, comparisons of
Aurignacian foragers with later Natufian farmers at Hayonim Cave, Israel, show dietary
change consistent with the shift to agriculture (Sillen, 1981). Analysis of other Middle
Eastern populations (Schoeninger, 1981) and comparisons of archaic and modern humans
(Schoeninger, 1982) also demonstrate marked distinctions in dietary focus between foragers
and farmers. Both the Sillen and Schoeninger investigations demonstrate the necessity of
using herbivore and carnivore control samples when attempting to characterize human diets
in local settings. Through her comparison of human and herbivore strontium values, Schoe-
ninger concluded that “[w]ithin the Levant, the nonagricultural populations. . .were including
a large proportion of plant material in their diets. This represents increased use in plant
material when compared to the earlier human population [in the region]” (1981:87). This
conclusion was independently confirmed using a similar research protocol based on analysis
of strontium/calcium ratios from humans, herbivores, and carnivores at Hayonim Cave
(Sillen, 1981; Smith, Bar-Yosef et al., 1984).
In prehistoric Ontario populations studied by Katzenberg (1993), where diagenesis and
other factors influencing bone strontium are controlled for, there is a decrease in values that
reflects a shift from plants high in strontium (leafy plants and nuts) to low-strontium grasses
(i.e., maize) (Figure 8.22). The results are consistent with findings based on nitrogen and
carbon stable isotopes (see earlier), although the approaches – isotopic and elemental – refer to
different parts of diet, those that contribute to the organic and inorganic fractions of bone
tissue.
Schoeninger (1979) determined strontium values in members of different status groups at
Chalcatzingo, Mexico. Her study suggested lower values in higher-ranked individuals, which
is consistent with the hypothesis that the elite in this society likely had greater access to
animal protein whereas lower-ranked individuals consumed relatively more plants, such as
maize (Blakely & Beck, 1981; Hatch & Geidel, 1985; Jacobs, 1995, for alternative patterns).
Weaning patterns have also been estimated on the basis of Sr/Ca ratios by mapping the
variation in values from earliest to latest forming enamel. Spatial analysis of Sr/Ca ratios from
deciduous teeth in healthy European children of known dietary/nursing histories shows clear
patterns of change (Humphrey et al., 2008). The pattern is highly specific, documenting the
onset of breastfeeding at birth (decrease in Sr/Ca ratios), its duration, and the introduction of
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350 Isotopic and elemental signatures of diet, nutrition, and life history
250
200
Strontium (‰)
150
100
50
0 500 1000 1500 2000
Year (AD)
Figure 8.22 Plot of trace element (strontium) values in prehistoric and historic Ontario
native populations, showing an increase in maize in diets after AD 1000. (Data from
Katzenberg, 1984.)
solid foods (increase in Sr/Ca ratios). While not applied to archaeological remains, this
approach has considerable promise for tracking life history as it relates to diet.
Barium
Barium has received little attention in bioarchaeological chemistry and paleodietary studies.
This underrepresentation is surprising as a significant body of evidence indicates that the
element is a highly sensitive indicator of past foodways (Burton & Price, 1990a, 1990b; Ezzo,
1992, 1993, 1994a; Francalacci & Borgonini Tarli, 1988). Like strontium, barium has no
known biochemical function, it is nontoxic, it is structurally similar to calcium, it is not
tightly regulated metabolically, and it undergoes biopurification with increasing trophic level
(Burton & Price, 1990a, 1990b; Burton & Wright, 1995; Elias et al., 1982). Barium values and
Ba/Ca ratios are lower in herbivores in relation to the plants they consume, and carnivores
have lower values and ratios than herbivores. Most barium in the geological environment
comes in the form of barite (BaSO4), a chemical that is relatively less soluble than the apatites
that are associated with strontium (Ezzo, 1992). This suggests that the soil-to-plant discrimin-
ation should be greater for barium than for strontium. Consistent with this hypothesis,
analysis of hundreds of archaeological bone samples indicates that barium separates organ-
isms into their trophic positions better than does strontium (Burton & Price, 1990a, 1990b).
Thus, barium may be an even more sensitive indicator of diet than strontium.
The importance of barium has been shown by the comparison of the analysis of Sr/Ca and
Ba/Ca ratios in animal and human bone from early prehistoric foragers from Carrier Mills,
Illinois. The ratio Sr/Ca indicates only slight differences between animal species; the overlap-
ping ranges of values in herbivores and carnivores prevent any meaningful dietary interpret-
ation (Burton & Price, 1990a). Barium ratios (Ba/Ca) show clear interspecies differences.
Human values at Carrier Mills are closer to values for carnivores than herbivores, suggesting
that meat is a dominant food in the diets of these prehistoric foragers.
Barium analyses yield an important perspective on temporal shifts in diet in relation to
social and environmental circumstances (Ezzo, 1992, 1993, 1994a, 1994b). Increased
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8.3 Elemental analysis 351
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352 Isotopic and elemental signatures of diet, nutrition, and life history
0.25
0
12
7 11
–0.25 10
8 9
–0.50
Log (Ba/Sr)
-1.0
-1.25 13 14
4
-1.50 6
1 3
5
-1.75 2
-2.0
Figure 8.23 Ba/Sr values in archaeological bone, showing the distinction between marine
and terrestrial consumers: (1) Paloma, Peru; (2) Rolling Bay, Alaska; (3) Kiavak, Alaska;
(4) Three Saints Bay, Alaska; (5) Chaluka, Alaska; (6) Port Möller, Alaska; (7) Rio Viejo,
Mexico (coastal); (8) Cerro de la Cruz, Mexico (coastal); (9) Fabrica San Jose, Mexico;
(10) Monte Albán, Mexico; (11) Poland (multiple sites); (12) Pirincay, Ecuador;
(13) Pueblo Grande, Arizona; (14) Stillwater Marsh, Nevada. (From Burton & Price,
1990b; reproduced with permission of authors and Academic Press Ltd.)
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8.3 Elemental analysis 353
iron status where skeletal lesions have been documented (Edward & Benfer, 1993; Fornaciari
et al., 1981; Sandford et al., 1988; Zaino, 1968). Based on limited samples, the presence of
lower iron in individuals with cribra orbitalia in Punic Carthage (third century BC) compared
with nonpathological individuals suggests a link between iron status and pathological indi-
cators of anemia (Fornaciari et al., 1981; but see Richtsmeier & Sheridan, 1996).
Juveniles from later Carthage (seventh century AD) have significantly higher levels of iron
than adults, which may indicate that iron in archaeological bone represents a biogenic signal
(Sandford et al., 1988). Comparison of iron levels in the third century BC and seventh century
AD Carthaginians shows higher values in the later period. This suggests that either iron status
improved or iron intakes increased in the later period (Sandford et al., 1988).
Ezzo (1994a) has questioned the biological significance of iron measurement in archaeo-
logical human bone. Except with regard to marrow, iron has no apparent function in bone
tissue, nor does it act as a reservoir for the element (Ezzo, 1994a). The lack of physiological
evidence suggests that varying amounts of iron in archaeological bone likely do not correlate
with prevalence of iron deficiency anemia. Additionally, iron content in soil is considerably
higher than in bone, indicating that unless all dirt is removed – a virtual impossibility in
archaeological remains – iron content in the skeletal tissue is highly exaggerated (Carvalho
et al., 2004; Ezzo, 1994a).
Zinc
Following Gilbert’s (1977) lead, there has been a general acceptance of zinc as an important
discriminator of diet (e.g., amount of meat versus plant consumption) and inferences about
nutritional health in past populations. Unlike strontium, calcium, or barium, a theoretical
basis has not been developed for zinc as a paleodietary indicator (Ezzo, 1994a, 1994b). Thus,
although zinc may be sensitive to dietary history, the lack of theoretical models limits its
usefulness in bioarchaeological chemistry and dietary inference (Burton, 2008). Underlying
the use of zinc is the unsubstantiated assumption that the element is present in higher levels in
meat and shellfish than in plants. Zinc acquired in the diet will influence the levels in skeletal
tissues of growing animals, such that diets deficient in zinc will result in low values in bone
and other tissues (Ezzo, 1994b). Zinc appears to play an essential role in growth: severe
deficiencies of the element result in markedly reduced growth and dwarfism. Thus, in contrast
to strontium and barium, the essential nature of zinc and its rather tightly controlled
metabolic regulation suggest that it has little use as a paleodietary indicator.
Lead
Lead is among the most discussed elements in bioarchaeological chemistry, mostly in
attempting to document chronic toxicity and excess intake in past groups (Aufderheide,
1989; Aufderheide et al., 1981, 1985; Aufderheide, Wittmers et al., 1988; Ericson et al.,
1979, 1991; Jaworowski, 1968; Keenleyside et al., 1996; Lalich & Aufderheide, 1991; Mont-
gomery et al., 2010; Nakashima et al., 2011; Perry et al., 2012; Polet et al., 2000; Reinhard &
Ghazi, 1992; Reinhard et al., 1994; Smrčka, 2005; Waldron, 1981, 1982, 1983; Wittmers et al.,
2008). Like barium and strontium, lead shows preferential bone deposition in comparison with
other tissues (more than 95% of absorbed lead is stored in the skeleton), it is not readily
excreted, and it has an unusually low turnover rate in skeletal tissue (Aufderheide, 1989;
Montgomery et al., 2010; Moore, 1986; Sandford, 1992; Wittmers et al., 1988). These factors
indicate that adult skeletal lead values represent a measure of lifetime exposure. Because
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354 Isotopic and elemental signatures of diet, nutrition, and life history
exposure is only rarely due to natural sources, lead found in bones and other tissues originates
from anthropogenic factors (Aufderheide, 1989; Burton, 2008; Montgomery et al., 2010;
Smrčka, 2005), but only if diagenetic sources can be ruled out (Wittmers et al., 2008).
In Colonial-era North America, the use of pewter – a lead-based material used in cooking
and eating implements – resulted in increased exposure to lead, in some instances at toxic
levels (Aufderheide et al., 1981, 1985; Aufderheide, Wittmers et al., 1988). Owing to greater
access to pewter by elite individuals in Colonial society, they had greater risk of exposure to
lead than did non-elite individuals. The relatively higher exposure is well illustrated in the
comparison of the bones of land owners and their slaves. For example, family members of the
owner of the Clifts Plantation, Virginia, had a considerably higher lead content than their
slaves (185 ppm vs. 35 ppm) (Aufderheide et al., 1981). This distinction reflects the use of
pewter by the landowner class and wood or unglazed ceramics by the non-elite class. Analysis
of lead content in military burials from the War of 1812 Snake Hill sample, Ontario, also show
low values suggesting that recruits were drawn from lower socioeconomic levels of the
civilian population (mean ¼ 31.3 ppm; Lalich & Aufderheide, 1991). Similarly, high-status
individuals from Medieval Belgium (c. AD 1100–1500) and Edo-period Japan (AD 1603–1867)
have elevated lead content, reflecting lead toxicity (Nakashima et al., 2011; Polet et al., 2000).
The former may be due to use of tin pewter eating utensils. The latter is exhibited as lead
bands in the metaphyses of femora in young juveniles. These indicate the incorporation of
lead in the growth plates during the developmental years. While the source is unclear, given
the very young age of these individuals (<3 years old), breastfeeding mothers may have
inadvertently poisoned infants and young children.
Slave skeletons from a Colonial-era Barbados sugar plantation have very high skeletal lead
concentrations (mean ¼ 118 ppm; values up to 424 ppm) (Handler et al., 1986, 1988). Skeletal
lead in this setting is apparently derived from various origins (e.g., pewter and lead-glazed
vessels used for storage, preparation, or serving of food and drink). The major source of lead
originated from the processing of sugar into rum via lead stills. In a similar vein, unusually high
intake of lead in Romano-British and later populations in Great Britain is well documented
(Montgomery et al., 2010; Waldron, 1981, 1983) (Figure 8.24). The source of lead is unknown,
but it appears to have been acquired from use of lead-based eating utensils, water pipes, and
simply living in a setting involving mining or metal-using, thereby increasing lead exposure.
During the late eighteenth and early nineteenth centuries, Omaha populations from north-
eastern Nebraska used lead for a variety of purposes – such as for the production of body
ornaments and musket balls (Reinhard & Ghazi, 1992; Reinhard et al., 1994). Analysis of
skeletal remains from this group reveals unusually high lead concentrations in some individ-
uals. Analysis of stable lead isotope ratios (206Pb/204Pb) suggests that lead was traded to the
Omaha from present-day Missouri. Lead was also used to make red facial pigment, which would
have been readily absorbed. Thus, facial paint is likely the primary source of lead for many of
these individuals. Some skeletal lead may have also originated from activities taking place after
death, since red paint was also applied to deceased individuals prior to their interment.
Mercury
Mercury is a highly toxic heavy metal that, when ingested either in its element (Hg) form or in
compound form (e.g., mercury sulphide), causes mercury poisoning (hydrargyria), a condition
resulting in neurological disruption and death. Mercury can be ingested, either through food
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8.4 Methodological issues in bioarchaeological chemistry 355
4
1st–3rd c. AD Rome
0
c e e l
hi Ag Ag
e
AD AD AD va
lit Ag ie
eo ze Iro
n n ie
s
ie
s
ie
s ed
N on Iro ur ur ur M
Br sh nt nt nt te
/Iri ce ce ce La
h h h
ti sh 4t 7t 1t
ot t– h– 1
1s 5t h–
Sc 8t
Period
Figure 8.24 Mean lead concentrations in human tooth enamel showing increased exposure
to anthropogenic lead from Neolithic to Late Medieval periods in Britain. The average
lead concentration in enamel from the first to third century AD Rome is depicted for
comparison. (From Montgomery et al., 2010; reproduced with permission of authors and
Cambridge University Press.)
or directly. Regarding the latter, during the Medieval period, mercury was prescribed for
treatment of venereal syphilis and leprosy. Atomic absorption analysis of a large series of
skeletons from southern Denmark having osteological evidence of syphilis and leprosy (see
Chapter 3) revealed that nearly half of the individuals likely having syphilis showed elevated
mercury levels (Rasmussen et al., 2008). These elevated levels were considerable and suggest
that in life these individuals had been treated with mercury. Similarly, nearly all individuals
having evidence of leprosy had mercury levels strongly suggesting that they were also treated
with mercury. With several exceptions, individuals lacking leprous or syphilitic lesions had
either no or minimal mercury levels. The exceptions pertain to three individuals recovered
from the friars’ cemetery of the Svenborg Friary who showed toxic levels of mercury.
Rasmussen and coworkers (2008) suggest that these individuals absorbed mercury, either
through the skin or via inhalation, while using mercury-laden red ink. Alternatively, these
individuals may have been exposed to mercury during the process of its administration to sick
individuals in the Abbey hospital.
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356 Isotopic and elemental signatures of diet, nutrition, and life history
Stable isotope (i.e., carbon and nitrogen) analysis of collagen has several advantages over
elemental and stable isotope analysis of apatite in dietary documentation and interpretation.
Importantly, because bone collagen is not subject to ionic exchange for carbon and nitrogen,
diagenetic effects are not as influential in determining values as in bone apatite (Ambrose,
1987; Grupe et al., 1989). Within the mineral component of bone, the apatite fraction is
subject to extensive diagenetic change (Schoeninger & DeNiro, 1982; Wright & Schwarcz,
1996). It is a more straightforward process to identify and remove contaminants (fats,
particulate plant matter, and humic matter) in collagen samples that potentially could alter
the biological signal (Ambrose, 1987, 1990; Ambrose & Norr, 1992; Stafford et al., 1988). New
technology and better understanding of the biochemistry of the mineral component (apatite)
of bone for trace element and stable isotope analysis has greatly expanded the range of
possibilities for paleodietary study (Metcalfe et al., 2009). Moreover, tooth enamel apatite is
relatively more resistant to diagenesis than bone apatite, and offers a source of potentially
valuable dietary information (Lee-Thorp, 2008). This is especially advantageous for humid
and/or low-latitude contexts (Krigbaum, 2007). However, there is no generally accepted
means of assessing diagenesis in tooth enamel (Grine et al., 2012; Schoeninger, 2010).
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Biological distance and historical
9 dimensions of skeletal variation
9.1 Introduction
Human remains are a central data source for documenting temporal and spatial patterning
of biological relatedness. Determination of biological relatedness has traditionally been
achieved via analysis of phenotypic skeletal and dental traits and their intra- and inter-
population variation. Biological distance or biodistance analysis is based on the premise
that the variation in morphological traits of the skeleton reflects underlying genotypic
variation. Although biodistance analysis pertains largely to characterizing phenotypic
variation, genotypic variation is becoming increasingly important via the study of ancient
DNA (aDNA).
Regardless of the type of biodistance data, the degree of relatedness presupposes that a
greater frequency of shared attributes indicates closer genetic affinity than a lower frequency
of shared attributes. In this regard, the orientation of the research is linked to the questions
being asked about intra- and inter-population interactions, ranging from continental-scale,
containing many sites, to regional-scale containing multiple sites, to single site analyses
focusing on individual variation within a population (Figure 9.1).
Biodistance analysis is motivated by three key interests (after Buikstra et al., 1990). First,
results are important for investigation of issues relating to evolutionary history, such as
genetic drift and selection, gene flow, and the influence of geography and other isolating
mechanisms on biological relatedness (Conner, 1990; Droessler, 1981; Heathcote, 1994;
Ossenberg, 1986; Rothhammer & Silva, 1990; Sciulli, 1990; Scott & Turner, 1997;
Stojanowski & Schillaci, 2006). By understanding the degree of relatedness, it becomes
possible to characterize temporal relationships in the identification of local in situ develop-
ment, or migration, or some combination thereof (Konigsberg, 2006).
Second, biodistance analysis addresses other fundamental archaeological and biohistorical
issues regarding the degree to which cultural and biological changes are influenced by
extrinsic factors versus local circumstances (Conner, 1990; Droessler, 1981). In such cases,
it is absolutely vital that population history is considered in order to fully assess the potential
contributions of these alternative agents. Biodistance analysis has the potential to identify
population boundaries, postmarital residence patterns, familial and kin groupings, social
groupings, and presence of individuals from other populations, especially in settings involv-
ing contact between biologically distinctive groups (Konigsberg & Buikstra, 1995) or long-
term regional gene flow (McIlvaine et al., 2014). Collectively, the biodistance record provides
important insights into population history, including shifts and stasis over time. Shifts imply
formation of new populations and social groupings (ethnogenesis) and stasis indicates lack of
local biological change with little or no external contribution of variation.
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358 Biological distance and historical dimensions of skeletal variation
(a) (b)
8.0 8
Plains Florida Panhandle
7
6.6 Illinois Florida North Interior
Variable 2
Variable 2
1.0 1
1 2 3 4 5 6 7 8 1 2 3 4 5 6 7 8
Variable 1 Variable 1
(c)
8.0
B. # 1
6.6 B. # 3
B. # 4
B. # 2
5.2
Variable 2
B. # 5
3.8
B. # 6
2.4
1.0
1 2 3 4 5 6 7 8
Variable 1
Figure 9.1 Biodistance analysis, employing the same two variables, on continental (a), regional (b), and
site (c) scales. (From Stojanowski & Schillaci, 2006; reproduced with permission of authors and John
Wiley & Sons, Inc.)
Third, biodistance investigations provide an important context for assessing skeletal and
dental variability where population structure may have an influence, such as disease or
nutrition (e.g., access to key nutrients and other resources by segments of a population; see
Buikstra, 1976a). In this regard, Buikstra noted that “It is well known. . .that the response to
both disease and generalized stress can vary from population to population. Therefore,
the strength of epidemiological statements must be tempered with a consideration of
biological relationships within and between the archaeological series” (1977b:72). The
adoption of new subsistence strategies by a population can have a tremendous influence
on the size of the population and where it lives. Resultant demographic shifts can, in turn,
influence patterns of mating and genetic exchange. In summary, the units of change – the
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9.1 Introduction 359
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360 Biological distance and historical dimensions of skeletal variation
10
–3.
10
–2.
GEG
10
–1.
0
Y
–.1
HES
SAQ
.90
QUR KHA
THE
0
GIZ
1.9
HAW
BAD
TAR
LIS
0
2.9
ABY HRK
2.00 NAQ
1.00
GRM
N
.00
–1.00
–2.00
1.70
.70
–.30
X –1.3
0
–2.3
0
Figure 9.2 Three-dimensional ordination of dental divergence (MMD) values from Neolithic
(GRM, Gebel Ramlah), Predynastic (BAD, Badarian; NAQ, Naqada; HRK, Hierakonpolis),
Early Dynastic (ABY, Abydos), Middle Kingdom (THE, Thebes), New Kingdom (QUR,
Qurneh), and Roman (HES, El Hesa; KHA, Kharga) Upper Egypt; and Early Dynastic
(TAR, Tarkhan), Old Kingdom (SAQ, Saqqara), Middle Kingdom (LIS, Lisht), Late Dynastic
(GIZ, Giza), Ptolemaic (GEG, Greek Egyptian), and Roman (HAW, Hawara) Lower Egypt.
(From Irish, 2006; reproduced with permission of author and John Wiley & Sons, Inc.)
matrices provides an important analytical tool to analyze genetic variation, especially with
regard to genetic distances relating to patterns of relationships between skeletal series within
a regional mating network. The statistic ranges from 0, where the samples are identical
genetically and engaged in considerable migration and genetic exchange, to 1, where all
samples are genetically distinct and have no genetic exchange. That is, if two populations are
exchanging mates frequently, they will express smaller phenotypic or genotypic differences
than two populations exchanging mates infrequently. Over time, changes in these values
represent changes in degree of mate exchange, patterns of migration, and gene flow in
general (Table 9.1).
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9.1 Introduction 361
Table 9.1 Temporal change in genetic diversity (FST) in Spanish Florida (From Stojanowski,
2009)
Period FST value Interpretation
Precontact 0.018 Significantly different from 0, indicating genetic exchange but with
(pre-AD 1550) limited gene flow
Early Mission 0.024 Increased values reflect somewhat increased genetic isolation, limited
(AD 1600–1650) gene flow, and likely little change in mating patterns in comparison
with the precontact populations
Late Mission 0.002 Dramatic decline in genetic variance indicates decrease in biological
(AD 1650–1704) structure and no isolation by distance. The record suggests increased
long-distance migration and gene flow during a time of population
decline and collapse
Biodistance analysis is problematic largely owing to the multifactorial nature of the traits
being studied. As with phenotypic variation in general, skeletal and dental traits are influ-
enced by a combination of intrinsic (genetic) and extrinsic (environmental) factors. Nowhere
is this more apparent than in skeletal tissue, which undergoes considerable modeling and
remodeling over the course of the lifetime of a person. Dental enamel – the hard tissue
containing most variation studied by biological anthropologists – does not remodel, but
nonetheless is complex and influenced during formation by a variety of factors (see Chapter 2).
It is not surprising, therefore, that skeletal and dental heritability – one estimate of the influence
of genetics – is so imprecisely understood (for some estimates of heritability, see Carson, 2006;
Goose, 1971; Hughes et al., 2000; Hughes & Townsend, 2013; Potter et al., 1976; Rizk et al.,
2008; Sjøvold, 1984, 1995; Townsend, 1992; Townsend et al., 2003). Thus, the application of a
biological population model – which assumes at least some degree of genetic control of traits
and that the distances based on these traits are directly proportional to those derived from
gene frequencies – is difficult to verify in human populations drawn from archaeological
settings (Konigsberg, 2006; Saunders, 1989; Scott, 2008; Stojanowski & Schillaci, 2006). On
the other hand, genetic data sets from twin and family studies reveal a highly heritable
component as it pertains to dental morphology (Hughes & Townsend, 2013; Scott, 2008).
Heritability remains an important focus of study, but developmental aspects of variation play
an increasingly important role in understanding the underlying biology of dental morphology
(Hunter et al., 2010).
Unlike DNA, neither metric nor nonmetric traits bear a one-to-one correspondence with the
genome of an individual. For example, the humeral septal aperture, a frequently used post-
cranial nonmetric variant for biodistance analysis, has a high negative association with
robusticity. Similarly, spondylolysis has often been assumed to be a population genetic
marker. However, spondylolysis is strongly influenced by activity load on the lower back
(see Chapter 5). Consistency of findings with other lines of evidence (e.g., archaeological or
historical), especially when care is taken to screen out traits influenced by activity or
representing biomechanical adaptations (Heathcote, 1994), suggests that nonmetric traits
can provide important insight into biological relatedness. Minimally, these traits are import-
ant for testing nonrandomness in skeletal series (Saunders, 1989; Spence, 1994). Familial
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362 Biological distance and historical dimensions of skeletal variation
studies of modern humans (Saunders & Popovich, 1978; Sjøvold, 1995), laboratory mice
(Grüneberg, 1952), and rhesus monkeys (Cheverud & Buikstra, 1981a, 1981b, 1982) support
this assessment. Phenotypic anthropometric analysis seems to provide the same results as for
quantitative genetic analysis (Konigsberg & Ousley, 1995). In assessing relationships, it does
not matter much that heritability of traits may be low. Rather, what matters is that environ-
mental variation is random with respect to the traits being analyzed.
The role of biodistance analysis in addressing archaeological questions and in providing a
context for other bioarchaeological research is central to most of the topics considered in this
book. This chapter discusses the important contributions of biodistance analysis to biohisto-
rical questions, with regard to both diachronic and synchronic perspectives on variation.
Folded into this discussion are the ways in which these data are used to provide contextual
information for interpreting other skeletal and dental variation, some of which are highlighted
in this book (e.g., infectious disease). Although virtually every area of the skeleton and
dentition has been considered to one extent or another, the preponderance of findings are
based on cranial and dental remains (for a discussion of use of postcranial remains, see
Donlon, 2000; Finnegan, 1978; Saunders, 1978).
As reflected in the explosion of investigations of DNA, both ancient and modern, since the
mid-1990s, paleogenetics has become central for understanding biological relatedness and
population history in a wide range of settings (Brandt et al., 2013; Brown & Brown, 2013;
O’Rourke, 2010; Raff et al., 2011; Relethford, 2010; Stone, 2008). Owing to the rapid onset and
degradation of DNA following death, most techniques used for documenting DNA from living
subjects are not appropriate for study of ancient remains, largely because of the very low
concentration and presence of mostly short, damaged fragments of DNA (Brown & Brown,
2011; Kaestle, 2010; O’Rourke, 2000; Pääbo et al., 2004; Stone, 2000; Willerslev & Cooper,
2005). However, polymerase chain reaction (PCR) and the more recent next generation
sequencing (NGS) technologies have opened a new window onto documenting and interpret-
ing genetic variation in archaeological remains. In particular, this technology has made it
possible to amplify single molecules, rendering them available for detailed analysis. Although
interesting and revolutionary when first reported (Doran et al., 1986; Pääbo, 1985), because of
what we now know about contamination and preservation in various environments, much of
the record of early discovery of aDNA has either been disproven or remains unsubstantiated.
However, important advances in improvement of recovery and especially accounting for
contamination are beginning to reveal the full strength of the study of aDNA for documenting
intra- and inter-population relatedness in past populations (Adler et al., 2011; Brandt et al.,
2013; Raff et al., 2011; Rasmussen et al., 2014). On the other hand, this is not to say that the
issues of contamination are solved. Rather, it remains a matter of considerable importance as
researchers determine the authenticity of aDNA (Brown & Brown, 2013).
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9.2 Classes of biodistance data 363
involving digitizing nodes on the cranium and other skeletal elements to produce three-
dimensional images provides increased precision of quantification (McKeown & Jantz, 2005;
Richtsmeier et al., 1992; Ross & Ubelaker, 2010). Nonmetric traits are discrete or quasi-
continuous anatomical entities often expressed as gradations from absence to full expression
(e.g., metopism, incisor shoveling).
A high degree of variation exists in cranial shape in human populations, which is heavily
influenced by the manner in which the teeth and jaws are used in masticatory and nonmasti-
catory functions (see Chapter 7). Nevertheless, heritability studies reflect the considerable
genetic and epigenetic contributions to cranial shape (Carson, 2006; Cheverud et al., 1979;
Sjøvold, 1984; Stefan & Chapman, 2003). As such, craniometric data have been a key resource
in identifying cultural-historical associations and in characterizing relatedness in a wide range
of settings (Droessler, 1981; Hanihara et al., 2008; Heathcote, 1986; Hemphill & Mallory, 2004;
Howells, 1973, 1989; Hubbe et al., 2010; Jantz, 1973; Key, 1983; Matsumura, 2006, 2011;
Pietrusewsky, 1994, 2006, 2008, 2010; Powell & Neves, 1999; Ross & Ubelaker, 2010; Schillaci &
Stojanowski, 2005; Sciulli, 1990; Spence, 1974a, 1974b, 1994; Stefan, 1999; Varela et al., 2008;
and many others). More than 200 cranial nonmetric traits have been identified in humans
(Hauser and De Stefano, 1989) and include four primary types – ossicles (small bones) within
cranial sutures (e.g., lambdoidal ossicles); hyperostotic traits or unusual skeletal proliferations
such as bridges of bone (e.g., atlas bridging); hypostotic traits involving ossification deficiencies
(e.g., metopic suture); and foramen variation (e.g., single or double supraorbital foramen)
(Buikstra & Ubelaker, 1994; Ossenberg, 1970; Saunders, 1989; Tyrrell, 2000) (Figure 9.3).
Similarly, analyses of tooth size, shape, and morphology are also important for understand-
ing population structure and history, and because of the far greater degree of genetic
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364 Biological distance and historical dimensions of skeletal variation
canalization of tooth shape and size, teeth may provide stronger forms of data for many types
of biodistance research. Built on earlier work by Albert Dahlberg (1956, unpublished), there
are some 30 standard nonmetric dental traits presently used in biodistance analysis (Scott,
2008; Scott & Turner, 1997; Turner et al., 1991) (Table 9.3). These traits include morphological
variants on tooth crowns (e.g., incisor shoveling, Carabelli’s trait) as well as root variation
(e.g., lower molar root number) (Figure 9.4). There is a strong genetic signal to these traits
implied by studies of living populations (Biggerstaff, 1970; Harris & Bailit, 1980; Hughes &
Townsend, 2013; Lundström, 1963). Thus, at its most simple level, populations containing a
high frequency of rare traits have a high degree of biological relatedness (Alt et al., 2013).
Coupled with the conservative nature of their evolution, dental traits and other dental
variation have proven important for biodistance analysis (Brace & Hinton, 1981; Calcagno,
1989; Dahlberg, 1951, 1963; Greene, 1982; Griffin et al., 2001; Hanihara et al., 2008; Harris &
Rathbun, 1991; Hemphill, 1991; Irish, 2005; Lukacs, 1983, 1989; Lukacs & Hemphill, 1991,
1993; Matsumura, 2006; Pilloud & Larsen, 2011; Sciulli, 1998; Sciulli et al., 1984; Scott &
Turner, 2006; Stojanowski, 2005b; Turner, 1991, 1993b; Ullinger et al., 2005; and many
others).
The aforementioned PCR and later technology have made it possible to extract, amplify,
and identify DNA from both the nuclear and mitochondrial genomes recovered from ancient
tissues. Controlling for a variety of issues, especially contamination by modern DNA, it has
become possible to identify the unique patterns of DNA in ancient remains associated with
(a)
1. Metopic suture complete
Figure 9.3 Cranial nonmetric traits. (From Buikstra & Ubelaker, 1994; reproduced with
permission of the authors and the Arkansas Archaeological Survey.)
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9.2 Classes of biodistance data 365
both genomes (Willerslev & Cooper, 2005). Nuclear DNA provides the key source of the unique
genetic profile of an individual. This pattern presents itself in a series of tandemly repeated
nucleotides known as short-tandem repeats (STRs), which are a series of two nucleotides
repeated many times (Brown & Brown, 2011). In archaeological contexts, DNA typing based
on STRs has been successful in identification of family groups (Haak et al., 2008; Keyser-
Tracqui et al., 2003; Ricaut et al., 2006). Comparison with biodistance analysis based on
cranial and dental discrete trait analysis in an archaeological series from northern Mongolia
revealed concordance between the genetic evidence based on STRs and discrete traits (Ricaut
et al., 2010). Unfortunately, because there is only one diploid copy of nuclear DNA in each cell
and a high degree of DNA degradation, it is extremely unlikely that the STRs will be preserved
in most circumstances (Brown & Brown, 2011). In contrast, mitochondrial DNA (mtDNA) has
a very high copy number – about 8000 identical copies of the mtDNA genome are present in a
cell – which greatly increases the chance of finding diagnostic DNA, even if degraded. Further,
mtDNA is inherited through the mother only, facilitating the documentation of maternal
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366 Biological distance and historical dimensions of skeletal variation
Table 9.3 Dental traits used in biodistance analysis (See definitions by Turner et al.,
1991; Scott & Turner, 1997)
Dental trait Teeth
Winging Upper first incisor
Shoveling Upper incisors, canine, lower incisors
Labial convexity Upper incisors
Double-shoveling Upper incisors, canine, first premolar, lower incisors
Interruption groove Upper incisors
Tuberculum dentale Upper incisors, canine
Canine mesial ridge Upper canine
Canine distal accessory ridge Upper and lower incisors
Premolar mesial and distal Upper premolars
accessory cusps
Tricuspid premolars Upper premolars
Distosagittal ridge Upper first premolar
Metacone Upper molars
Hypocone Upper molars
Cusp 5 (metaconule) Upper molars
Carabelli’s trait Upper molars
Parastyle Upper molars
Enamel extensions Upper premolars and molars
Premolar root number Upper premolars
Upper molar root number Upper molars
Radical number All teeth
Peg-shaped incisor Upper second incisor
Peg-shaped molar Upper third molar
Odontone Upper and lower premolars
Congenital absence Upper premolars and molars, upper and lower second
premolars, upper and lower third molars
Premolar lingual cusp Lower premolars
variation
Anterior fovea Lower first molar
Groove pattern Lower molars
Cusp number Lower molars
Deflecting wrinkle Lower first molar
Distal trigonid crest Lower molars
Protostylid Lower molars
Cusp 5 Lower molars
Cusp 6 Lower molars
Cusp 7 Lower molars
Canine root number Lower canine
Tome’s root Lower first premolar
Lower molar root number Lower molars
Torsomolar angle Lower third molar
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9.2 Classes of biodistance data 367
(a)
(b)
(c)
Figure 9.4 Three examples of dental nonmetric traits: grades of labial convexity (a),
mandibular first molar anterior fovea size (b), and maxillary second incisor shoveling (c).
(From Turner et al., 1991; photographs by Scott Haddow; reproduced with permission of
authors and Wiley-Blackwell, a division of John Wiley & Sons, Inc.)
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368 Biological distance and historical dimensions of skeletal variation
lineages and showing maternal relationships. Specific patterns of mtDNA are identified on the
basis of haplogroups – groups of mitochondria sharing a specific set of markers (Raff et al.,
2011). Thus, individuals or populations having the same haplogroup are more closely related
biologically than individuals or populations having different haplogroups.
Some mtDNA markers are unique to specific geographical regions as well as to single
populations (Brown & Brown, 2011; Kaestle, 2010; O’Rourke & Raff, 2010). In this regard,
analysis of mtDNA from archaeological contexts has provided important insights into
large regional patterns of population history, such as in the peopling of the Americas
(O’Rourke & Raff, 2010; Raff et al., 2011; Reich et al., 2012), the beginning of human
population expansion across the Pacific (Knapp et al., 2012), the impact of Aztec imperi-
alism on local populations in Mesoamerica (Mata-Miguez et al., 2012), population history
in Neolithic and post-Neolithic Europe (Brandt et al., 2013), and in a wide variety of other
geographical and cultural contexts (Adachi et al., 2009; Alzualde et al., 2007; Bolnick &
Smith, 2007; Carnese et al., 2010; Deguilloux et al., 2011; Fan Zhang et al., 2010; Forgey,
2011; Kaestle, 1995; Kaestle et al., 1999; Kemp et al., 2009; Lertrit et al., 2008; Malhi et al.,
2003; Moraga et al., 2005; Raff et al., 2011; Ricaut et al., 2006, 2010; Shimada et al., 2004;
Stone & Stoneking, 1993).
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9.3 Biohistorical issues: temporal perspectives 369
Nubia
The traditional model of population structure and history in the Nile River valley is based on
the assumption that a series of migrations of “racially” distinct populations replaced one
another at various times. The model posits that in the past there were two racial types living in
the Nile River valley. These include a “Caucasoid” type in the Upper Valley and a “Negroid”
type in the Lower Valley. Strouhal (1971) views biological changes in the region as resulting
from the arrival and greater influence of one type over another at various times (see discus-
sion by Armelagos et al., 1981). Thus, the traditional model invokes population movement
and gene flow to account for biological change in this region. From this perspective, the
reduction in craniofacial robusticity found in X-group populations is interpreted as signaling
the arrival of a “Negroid” population (Batrawi, 1946; and see Chapter 7).
In order to evaluate the traditional model of population replacement in Nile Valley history,
Greene and colleagues (Greene, 1967, 1972, 1982; Greene & Armelagos, 1972; Greene et al.,
1967) analyzed a battery of dental nonmetric traits within samples from Wadi Halfa, Sudan.
Univariate statistical comparisons of nonmetric dental traits revealed no significant differ-
ences between Meroitic and X-group samples. Based on these results, they concluded that
population replacement was unlikely in this setting, “but instead represent(s) a continuum of
(temporally) successive local Mendelian breeding populations” (Greene, 1982:76). Although
suggestive, univariate statistical analyses are problematic, especially because such analyses
incorrectly assume that individual traits are independent. In order to provide a more convin-
cing argument for demonstrating biological continuity, Greene (1982) employed the MMD
distance statistic to dental nonmetric traits to test the hypothesis that inter-population
biological differences are not significant. This analysis revealed extremely low divergence
values, and thus indicates no significant change in biological groupings, or the virtual lack of
infusion of peoples with different biological histories into the region. The implications of this
study are important because they lend strong support to the argument that biological change
in Nubia was due to local circumstances and not gene flow.
Turner and coworkers (Irish & Turner, 1990; Turner & Markowitz, 1990) reassessed Greene
and coworkers’ continuity model of Nubian population history. Although lauding the studies
of Greene and others for using a non-racial, non-typological approach to Nubian population
history, they argued that extra-regional sources of variation have been insufficiently con-
sidered, especially sources that may explain temporal shifts in craniofacial morphology in this
region. Implicit in the work by Greene and others is the assumption that population continuity
in Nubia extends at least as far back as the late Pleistocene (c. 12 000 yBP). Turner and
coworkers contended that continuity can only be claimed if non-Nubian populations are also
considered in statistical analyses of dental trait variation in this region.
Turner and coworkers included in their analysis additional Nubian dentitions from the late
Pleistocene Upper Stone Age, Meroitic, X-group, Christian period, and historic-era European
samples. Computed MMD values, modified for small samples (Green & Suchey, 1976) and
tested for significance (Sjøvold, 1973), reveal few significant differences in comparing the
Meroitic, X-group, and Christian periods, thus confirming Greene’s earlier conclusions
regarding population continuity. However, significant differences between the Pleistocene
and later groups were clearly identified (e.g., frequency of incisor shoveling and three-rooted
mandibular molars). Based on the apparent temporal discontinuity between the Pleistocene
and later populations, Turner and Markowitz (1990) hypothesized that the ancestry of recent
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370 Biological distance and historical dimensions of skeletal variation
Nubians did not derive from local late Pleistocene populations, and that a population
replacement event occurred during the Holocene in Nubia. The origin of these later popula-
tions is unclear, but based solely on dental traits, they argued that populations north of Nubia
containing European and Near Eastern traits are the most likely sources.
In an effort to identify other possible sources of variation, Irish and Turner (1990) compared
their sample of Nubian dentitions (late Pleistocene to Christian period) with historic-period
dentitions from a West African group – the Ashanti. Univariate and multivariate (MMD)
statistical treatment of these samples revealed strong similarities between modern West Africans
and late Pleistocene Nubians. As with previous studies, later Holocene dentitions were found
to be very similar. The late Pleistocene and modern West Africans are strongly divergent from
the Meroitic, X-group, and Christian-period Nubians. Thus, they argued that there is a
population discontinuity between the late Pleistocene and Holocene populations in Nubia,
with the former sharing biological affinities with West Africans. Irish and Turner (1990)
suggested that the discontinuity can be explained by high rates of violence and decline (or
possibly extinction) in late Pleistocene (Mesolithic) Nubian foragers, which may have left
them susceptible to invasion or “genetic swapping” by other groups from West Africa
(Turner & Markowitz, 1990).
In a more detailed analysis containing remains from Nubia not included in previous studies,
Irish (2005) performed additional multivariate analysis to document patterns of discontinuity
and continuity inferred from biodistance analysis. His comprehensive analysis revealed a
clear discontinuity between the late Paleolithic Jebel Sahaba series (14 000–12 000 yBP) and
all later series. That is, the late Paleolithic series displays higher frequency of some specific
dental traits than all other (later) series, such as in UI1 shoveling, six-cusped LM1, and LP1
Tomes’s root. Irish concluded that the Jebel Sahaba population is likely not ancestral to any
later regional population. However, the entire record of Final Neolithic through Christian
periods shows homogeneity in dental variation, suggesting regional population continuity
from at least c. 5600 yBP onward (Irish, 2012). It is during this later time frame that much of
the craniofacial reduction described by Carlson & Van Gerven (1977, 1979; Van Gerven et al.,
1995) took place in Nubia. Thus, although the high degree of robusticity displayed in pre-
Neolithic populations is apparently not the ancestral morphology for the region, there still
remains the presence of significant craniofacial size reduction in the post-Mesolithic popula-
tions, changes taking place within a single population evolving over time in situ (see
Chapter 7). Thus, viewed in the context of changes in diet, food preparation, and technology
in general, these craniofacial modifications were likely due to reduced mechanical loading of
the jaws and teeth in later times. That said, the Nubian populations did not live in isolation,
and the record presented through biodistance analysis by Irish is consistent with mtDNA
studies of modern people showing evidence for gene flow especially from Egypt (Buzon, 2006;
Irish & Konigsberg, 2007) as well as the dynamic nature of population movement in the Nile
Valley corridor in general (Irish, 2000, 2006; Zakrzewski, 2007) and during specific times and
places (Buzon, 2011).
Southern Africa
Early models accounting for the origin of population variation among native peoples of
southern Africa argued for large-scale immigration and complex processes associated with
isolation and development of distinctive populations (Ginter, 2011; Hausman, 1984;
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9.3 Biohistorical issues: temporal perspectives 371
Morris, 1992; Rightmire, 1970, 1976). A range of anthropological and historical evidence
suggests that cultural and biological characteristics of modern native peoples may have arisen
from in situ processes associated with the transition from foraging to nomadic pastoralism
at about 2000 yBP (Ginter, 2011). Morphometric analysis of archaeological crania addresses
the issue of evolution and population history in this very dynamic region (Ginter, 2011;
Hausman, 1984; Irish, 2013; Morris, 1992; and see Rightmire, 1970, 1976).
When first encountered by Europeans, native peoples of southern Africa were described
as consisting of two distinctive groups, including the San, who were relatively small in
body size and who followed an exclusively foraging lifeway. The Khoi, who were described
as taller and more robust than the San, pursued a pastoralist lifeway based on sheep and
cattle husbandry. Khoi pastoralists were widespread, but lived mostly in coastal areas (e.g.,
Eastern Cape Province) and in some major river valleys in the interior of southern Africa.
The origin of pastoralism in recent Khoi is unclear, and revolves around the question: Was
it due to a migration of people into the region or a diffusion of ideas? The consensus
drawn from ethnographic, linguistic, and archaeological data is that people migrated to
southern Africa c. 2000 yBP, bringing with them pastoralism (Deacon & Deacon, 1999; but
see Sadr, 2003).
Multivariate discriminant function analysis of craniometric traits confirms the biological
distinctiveness of Khoi and San peoples (Hausman, 1984; and compare with Morris, 1992;
Rightmire, 1970, 1976). Although separated from other South Africans, crania of these two
groups are morphologically similar, suggesting that they share a common origin. Based on her
analysis of cranial morphology, Hausman proposes that one segment of the population
adopted pastoralism, which contributed to “the biological separation of the groups”
(1984:268). Differences between Khoi and San marriage and kinship systems may have
contributed to this biological variation (Hausman, 1984). Exogamy among the highly mobile
San facilitates gene flow, and thus may explain the high degree of homogeneity across local
San groups within broad regions. Khoi are exogamous primarily along patrilineal clan lines
within individual tribes. These marriage patterns, combined with other practices (e.g., cross-
cousin marriage) would increase interbreeding and homogeneity within small populations.
These social behaviors would discourage interbreeding between Khoi and San, resulting in
greater genetic isolation and morphological differentiation between the two groups. Overall,
social behavior has enormous implications for patterns of gene flow and the way that these
patterns influence biological variation in South Africa (Morris, 1992).
Biodistance analysis of dental variation in sub-Saharan Africa reveals an overall uniform-
ity of variation (Irish, 1997, 2013), so much so that Irish (2013) identified an “Afridont” dental
pattern, including a series of high-frequency traits (e.g., Bushman canine, two-rooted
maxillary first premolar, two-rooted mandibular second molar). Moreover, the craniometric
findings suggest very little evidence for change before and after the shift from foraging to
pastoralism among prehistoric Khoi in the coastal region (Ginter, 2011). In contrast to
traditional models for origins of Khoi coastal pastoralists, this finding argues that population
replacement did not occur in this region. Rather, pastoralism was introduced to and adopted
by local populations (Hausman, 1984). Similarly, analysis of cranial, dental, and postcranial
measurements and cranial discrete traits using multivariate and univariate statistical analysis
of a large data set from the Eastern Cape Province reveals strong evidence for biological
continuity spanning the pre-herding and herding traditions in the region and a general
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372 Biological distance and historical dimensions of skeletal variation
pattern of biological homogeneity (Ginter, 2011). Collectively, these analyses strongly favor
an in situ development of pastoralism in the southern African region.
European Neolithization and the spread of farming: demic or cultural diffusion, or both?
As with much of the rest of the world, the period of the late Pleistocene and early Holocene in
Europe was enormously dynamic, involving movement of people, their technology, and their
ways of acquiring food and other resources. Populations saw a transition from a lifeway
and subsistence economy based on nondomesticated to domesticated plants and animals,
representing a profound alteration in nutrition and lifestyle. Humans first began domesti-
cating animals (goats, sheep, pigs, and cattle) and plants (wheats and barleys) around
13 000–10 000 yBP in the Near East (Pinhasi et al., 2005; Smith, 1998; Zeder, 2008). By
8500 yBP, this Neolithic package had spread to Greece, and by 5000–6000 yBP to Scandinavia
and Great Britain (Burger & Thomas, 2011). Based on modern genetic evidence, a number of
authorities argue for a demic diffusion process whereby the spread of domestication was
occasioned by the migration and dispersion of early farmers from the Near East via Anatolia,
replacing indigenous hunter-gatherers throughout Europe (Barbujani et al., 1998; Cavalli-
Sforza et al., 1996). In contrast, the competing cultural diffusion model argues that while there
may have been population dispersion in some areas, in part originating from the Near East, it
was the movement of the idea of agriculture and domestication that explains the Neolithic
transition (Whittle, 1996; Zvelebil, 1986). The former should be represented by clines of
genetic variation across Europe, whereas the latter would predict similar kinds and frequen-
cies of genetic variants between the earlier foragers and later farmers of Europe (Deguilloux
et al., 2012; Pinhasi & von Cramon-Taubadel, 2009).
A wide variety of non-molecular and molecular data presents a contradictory picture
indicating evidence of both migration and replacement and indigenous adoption of farming
and domestication generally (Deguilloux et al., 2011; Fox, 1996; Harding et al., 1989;
Linderholm, 2011). Analysis of aDNA from a series of Neolithic individuals from Germany,
Austria, and Hungary revealed a substantial presence of sequence-specific genetic markers
particular to haplogroup N1a (25%), found only rarely in modern Europeans (0.2%) (Deguil-
loux et al., 2011; and see Pinhasi et al., 2012). This finding suggests that modern Europeans
may not be the descendants of the first farmers in the continent (Haak et al., 2005; and see
Pinhasi & von Cramon-Taubadel, 2009). Similarly, analysis of aDNA in Sweden provides a
genetic reconstruction showing that Mesolithic hunter-gatherers may have been replaced by
farmers moving into the region (Linderholm, 2011). Moreover, analysis of genetic diversity,
including markers in the Y-chromosome, appears to be best understood in relation to a single
population source in the Near East (Anatolia) during the Neolithic (Deguilloux et al., 2011).
In reality, the process of the adoption of domestication and especially farming was complex
and almost certainly involved different and region-specific circumstances across Europe. The
genetic discontinuities suggest that it is likely that all modern Europeans are not the direct
descendants of either Paleolithic hunter-gatherers or migrating Neolithic farmers. Thus, the
question of demic or cultural diffusion may not be the best one to ask. Rather, it would appear
from the complexity of the record that a much more interesting problem is to explore the wide
variation, such as that observed in modern and ancient DNA. This conclusion about popula-
tion history in Europe is indicated by the likelihood of periods of continuity and discontinuity
based on mtDNA (Brandt et al., 2013). Until complete diachronic data sets of autosomal aDNA
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9.3 Biohistorical issues: temporal perspectives 373
become available, especially those that present evidence for continuity (or discontinuity), the
answers to questions about population origins and evolution – with or without domestication
in pre-Neolithic and Neolithic contexts – remain unclear.
Skeletal biodistance analyses undertaken by Schwidetzky and Rösing for the Neolithic/
Bronze Ages (Schwidetzky, 1967), Iron Age (Schwidetzky, 1972), Roman period
(Schwidetzky & Rösing, 1975), and Medieval and recent periods (Rösing & Schwidetzky,
1977, 1979, 1984; Schwidetzky & Rösing, 1982) also bear out the complex nature of European
population history. Their cluster analysis indicates two primary morphometric complexes –
Western and Eastern – with size gradients between the two. In the Western complex, cranial
length is greater than that in the Eastern complex; in the Eastern complex, cranial width and
bizygomatic breadth, are greater (Rösing & Schwidetzky, 1979). During the Early Middle Ages,
distinctions between the groups become less straightforward. Schwidetzky and Rösing sug-
gested that such a reduction in distinctiveness is due to increased population movement and
increased gene flow. In the following Late Medieval period, the distinctions between Eastern
and Western complexes become even less clear (Rösing & Schwidetzky, 1984).
Other cluster analyses of these European samples reveal little evidence of sharp differenti-
ation between groups during the Medieval period (Sokal et al., 1987), which contrasts with a
record of abrupt gene frequency boundaries in modern Europe (Barbujani & Sokal, 1990).
Sokal and coworkers (1987) contended that the apparent lack of clear definition between
groups in later European history may be related to many factors, including variability in
measurement accuracy among observers and definitions of characters, small sample size for
some sites, the nature of variables used in the study, and incorrect sex identification. Cranial
lengths, breadths, and facial size are developmentally plastic (see Chapter 7). Thus, some of
the variability documented in European populations may also be influenced by shifts in
technology or dietary practices and the craniofacial responses to these shifts. Long-term
contacts (e.g., Gothonic-Slavic groups), admixture (e.g., Turkic-Mongolian groups), wide-
spread migration (e.g., Italian groups), and random genetic drift may have influenced the
patterns of cranial variation seen in these samples. In summary, no clear groupings are
evident when looking at this very broad region.
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374 Biological distance and historical dimensions of skeletal variation
Hemphill and coworkers (1991, 1996) employed multivariate statistical analyses of cranio-
metric, cranial nonmetric, and dental nonmetric traits for Indus Valley samples in an
effort to resolve problems in understanding biocultural change and population origins in
this exceedingly complex region. Cluster and principal components analyses of Harappan-
phase dentitions and skulls from Cemetery R37 at the site of Harappa provide two principal
findings. First, the differences between Neolithic and the later Chalcolithic dentitions from
the Indus Valley suggest a lack of population continuity between about 6000 and 4500 BC.
Second, a general similarity in traits between sites in the region after 4500 BC indicates the
presence of biological continuity during the rise, apogee, and decline of the Harappan
culture. Iron Age populations (c. 200 BC) are markedly different in pattern of dental
nonmetric variation, suggesting a discontinuity during later prehistory after the demise of
Harappan culture. Analysis of cranial nonmetric traits indicates that the source of this
discontinuity may be gene flow into the region from the Iranian Plateau in the Near East.
In summary, the results of this research document the lack of biological continuity from the
Neolithic to the period of Christianity. Rather, two discontinuities existed, the first between
6000 and 4500 BC and the second between 800 and 200 BC. These findings suggest that
although the Harappan culture was an indigenous development within the Indus Valley, the
region nevertheless interacted on a widespread level at various times, especially with
populations to the west.
Unlike the Indus Valley, the biological and archaeological history of central Asia is poorly
documented, in part because the region has long been envisioned as culturally barren, serving
primarily as a buffer between the Harappan culture to the east and the Mesopotamian
societies to the west (Wheeler, 1968). Archaeological research beginning in the 1970s and
1980s revealed that the central Asian borderlands were inhabited by various complex soci-
eties, including the Oxus culture, a culture that was associated with oases in Bactria and
Morgiana, located in Uzbekistan, Turkmenistan, and Afghanistan (Hemphill, 1998, 1999a,
1999b; Hemphill et al., 1996, 1998). This region may have played an important role in the
diversification and spread of Indo-Iranian languages into the Iranian plateau and Indian
subcontinent (Hiebert and Lamberg-Karlovsky, 1992). Based on archaeological evidence, and
especially material culture, some argue that the Bactrian culture is an eastward expansion of
the central Asian oasis society, originating in western Turkmenistan and northwestern Iran.
The presence of Harappan artifacts in Bactrian sites indicates to some authorities that the
origins of Bactrian culture are tied to the Indus Valley to the east.
Univariate, cluster, and MMD statistical analyses based on dental and craniometric traits
from the northern Bactrian oasis sites in Uzbekistan (2300–1650 BC), localities in Turkmeni-
stan, Iran, the Indus Valley, and other South Asian localities, reveal strong separation between
central and south Asians (Hemphill et al., 1996). Unlike the South Asian groups, Bactrian
samples contain low frequencies of reduced buccal-distal cusp (metacone) in the first maxil-
lary molar. This and other distinctions indicate no evidence of a biological link between
central Asian and Harappan populations. Thus, there appears to be no support for a model
positing that the sedentary Bronze Age settlements in Bactria owe their origin to Harappan
settlers from the east; neither is there evidence to suggest that Bactrian populations moved
eastward into the Indus Valley following the decline of Harappan culture. Overall, the rise of
the Oxus culture appears to have occurred locally without significant external influence,
either from the east or from the west.
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9.3 Biohistorical issues: temporal perspectives 375
One of the most sensational archaeological discoveries in Central Asia involved the recov-
ery of a series of some 300 mummies – naturally desiccated bodies – that displayed “Europoid”
or “Caucasoid” facial features, found in the Tarim Basin of the western Chinese Xinjiang
Uyghur Autonomous Region in sites dating to the Bronze Age (c. 1800 BC). In order to
document and explain the origins of these individuals, Hemphill and Mallory (2004) under-
took a comprehensive craniometric biodistance analysis of Tarim Basin crania and numerous
other crania from a range of localities in Central Asia, South Asia, and China. The analysis
revealed that the population, while distinctive, shows no biological relationship to suggest
that it owes its origins to “Europoid” populations of the Russian steppes. Instead, its origins
appear to have been multiple, and the closest affinities may be found to the west among the
inhabitants of the North Bactrian oasis of Uzbekistan and the Iranian Plateau, as well as to the
south among populations of the Indus Valley.
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376 Biological distance and historical dimensions of skeletal variation
A second hypothesis posits that biological changes in the Jomon–Yayoi transition at about
300 BC were due to immigration and gene flow from the Asian mainland during the late
Jomon period (Hanihara, 1994; Hanihara et al., 2008; Nakahashi, 1993). This dual structure
hypothesis argues that biological and cultural influence from the mainland spread quickly
throughout Japan, eventually involving all populations except those inhabiting Hokkaido and
the Nansei Island chain (Aikens & Higuchi, 1982; for archaeological context, see Chard, 1974).
Because of such profound and widespread gene flow, proponents of this model maintain that
the ancestry of living Japanese is mostly linked with mainland Asians, whereas the modern
Ainu populations of northern Japan are descendants of the founding Jomon populations
(Hanihara & Ishida, 2009).
The study of dental traits and size is consistent with craniometric and cranial discrete trait
analyses completed by various workers (Dodo, 1986; Hanihara, 1979, 1985; Howells, 1966,
1986; Nakahashi, 1993; Ossenberg, 1986; Pietrusewsky, 2000; Yamaguchi, 1982). Multivari-
ate analyses indicate closer biological links between Jomon and living Ainu than between
Jomon and living Japanese. Collectively, these analyses support the alternative point of view
that the biocultural discontinuity during the late Jomon period resulted from migration and
gene flow. These analyses emphasize that this discontinuity is not a precipitous population
replacement. The gradient of biological distances in nonmetric cranial traits from Siberians–
Kinki–Kanto–Ainu–Jomon suggests the presence of gene flow and hybridization between
mainland immigrants and indigenous Japanese during the Yayoi and following periods
(Ossenberg, 1986).
The dual structure hypothesis for the peopling of the Japanese Islands is supported by a
growing body of data. However, while Turner’s Sundadonty versus Sinodonty hypothesis
helped to clarify population differences between the Jomon and Yayoi, the record is providing
increasing evidence that Jomon did not derive from affinities in Southeast Asia. Rather,
skeletal evidence for East Asia (Hanihara and Ishida, 2009; Pietrusewsky, 1999) suggests that
the migration responsible for Jomon people moved first from Northeast Asia into Hokkaido,
then through the southerly islands of Japan. These results are supported by aDNA analysis of
Jomon people from Hokkaido, suggesting that they have population affinities in southwestern
Siberia and not Southeast Asia (Adachi et al., 2009).
Population history in Southeast Asia has long been known to be complex and to involve
considerable migration and movement of people and ideas. One model argues for a later
immigration of populations from southern China to Southeast Asia, bringing with them
agriculture and other elements of complex society (Bellwood, 2001; Howells, 1973). This
model indicates hybridization between Neolithic northern East Asians and local resident
Australo-Melanesians. Turner’s (1987) analysis of the dental record and association of simpler
(Sundadont) dental morphology in the region revealed a different population history and
independent trajectory of biological history, the results of which were said to argue for local
evolution of Southeast Asian populations, independent of northern Asian groups, especially
from China. His model contends that groups with the Sundadont pattern spread north and to
Japan (see earlier), Siberia and northeastern Asia, the Americas, and the Pacific.
An alternative multivariate analysis of tooth size and morphology reveals a very different
pattern of biological history, consistent with the traditional immigration model involving
migration from south China and hybridization (Matsumura, 2006, 2011; Matsumura &
Hudson, 2005) versus Turner’s regional continuity model (Pietrusewsky, 1994, 1999, 2006).
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9.3 Biohistorical issues: temporal perspectives 377
As did earlier workers, Matsumura and others observed that the preceramic hunter-gatherers
of Southeast Asia have morphology similar to Australian Aborigines and Melanesians,
namely relatively long, narrow cranial vaults and robust faces and jaws. In contrast, later
ceramic-equipped, farming populations display rounder crania, reduced facial-masticatory
architecture, and smaller teeth, similar in some respects to Neolithic and post-Neolithic
populations from northern Asia. In light of these differences between earlier and later South-
east Asian populations, Matsumura and Hudson’s (2005) reconstruction of population history,
based on analysis of metric and nonmetric variation in a series of 4000 dentitions from Asia,
Australia, and Melanesia, revealed similarity between preceramic Southeast Asia and
Australo-Melanesians and biological dissimilarity between preceramic and later Southeast
Asians (Figure 9.5). In this “two-layer” model, the later dissimilarity is due to gene flow and
admixture derived from immigration of groups from North Asia or East Asia, or both
(Matsumura, 2011).
The question then emerges: Which of the two hypotheses is correct, local development and
dispersal, or population displacement originating from elsewhere in Asia? In consideration of
other biodistance analyses, the answer to this question is likely more complex than either
model indicates. Large-scale craniometric biodistance analysis shows two broad groupings of
prehistoric populations, including (1) Australia, Tasmania, New Guinea, and Melanesia, and
(2) East/North Asia, Southeast Asia, Polynesia, and other areas of Oceania (Pietrusewsky,
2005) (Figure 9.6). Given the distinctiveness of the clusters, they likely relate to two broad
origins of peopling of this vast region of the world. Within the second cluster, East/North Asia
and Southeast Asia are separated. This distinction may provide evidence for long-term in situ
evolution in each of these two regions and not replacement of ancient Southeastern Asian
populations via immigration from East/North Asia (Pietrusewsky, 2005, 2006).
However, genetic (PCR) analysis of mtDNA from teeth of 30 individuals from the Noen
U-Loke (400 BC–AD 500) and Ban Lum Khao (1200–400 BC) sites provides preliminary
evidence linking early agriculture Bronze–Iron Age populations from northeastern Thailand
with modern Thai and other East Asian populations (Lertrit et al., 2008). In the regional
context, inhabitants of these two settings present strong genetic similarity indicating they are
from the same gene pool of this region of Southeast Asia. Moreover, the close genetic affinity
with modern Thai indicates the strong possibility for a genetic continuity having time depth
extending back millennia. On the other hand, the presence of sequences shared with popula-
tions from southern China provides a complex picture of gene flow, and certainly does not
rule out demic diffusion from southern China into Southeast Asia (Lertrit et al., 2008). While
not a replacement process, this record suggests the important role of gene flow in the
development of modern populations in the region.
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378 Biological distance and historical dimensions of skeletal variation
Buriats
Urga Mongolians
East Asians
Northern Chinese
Chifeng Chinese
Koreans
Kofun Japanese
Tokyo Japanese
Kamakura Japanese
Edo Japanese
Neolithic Southern Chinese
Dayak
Laotians
Lesser Sunda & Java Islanders
Early Holocene Laotians
Ban Kao
Malay
Philippines
SE-Asians Sumatra Islanders
Chinese Thailanders
Thailanders
Yayoi
Anyang Chinese
Dong Son Vietnamese
Vietnamese
Prehistoric SE-Asians Negritos
Australians Melanesians
Guar Kepah
Jomon
Amami-Okinawa Islanders
Non Nok Tha
New Britain Islanders
Gua Cha
Loyalty Islanders
Andaman Islanders
Bac Son Vietnamese
Hokkaido Ainu
Da But Vietnamese
Australian Aborigines
Sakhali Ainu
Figure 9.5 Dendrogram of cluster analysis for prehistoric and historic Southeast Asian
populations, depicting relative genetic affinities between various populations. (From
Matsumura & Hudson, 2005; reproduced with permission of authors and John Wiley &
Sons, Inc.)
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9.3 Biohistorical issues: temporal perspectives 379
Tonga-Samoa
Hawaii
Guam
Rapa Nui
Gambier Is.
Marquesas Is.
Society Is.
Tuamotu Arch.
Chatham Is.
Sumatra
Bomeo
L. Sundas
Java
Sulawesi
Sulu
Cambodia-Laos
Philippines
Vietnam
Bachuc
Thailand
Burma
Kanto
Tohoku
Kyushu
Taiwan
Hainan Is.
Korea
Ryukyu Is.
Atayal
Shanghai
Hangzhou
Nanjing
Chengdu
Hong Kong
Manchuria
Ainu
Mongolia
New Zealand
S. Moluccas
Fiji
Biak Is.
Solomon Is.
New Ireland
Caroline Is.
Marshall-Kiribati
Sepik R.
D’Entrecasteaux Is.
Dawson St.
Vanuatu
New Britain
Santa Cruz
Loyalty Is.
New Caledonia
Fly R.
Purari Delta
Admiralty Is.
Murray R.
New South Wales
Queensland
N. Territory
W. Australia
Swanport
Tasmania
Figure 9.6 Dendrogram of cluster analysis for prehistoric Asian and Pacific populations, based on craniometric
variables. (Pietrusewsky, 2005; reproduced with permission of author and RoutledgeCurzon.)
Harper & Laughlin, 1982; Hrdlička, 1912; Turner & Bird, 1981). A large body of archaeological
and genetic evidence indicates that the immigration of Old World peoples to the New World
took place by 15 000 years ago and likely earlier (Meltzer, 2009; Raff et al., 2011). Moreover,
although the routes and number of colonization events are subject to considerable debate,
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380 Biological distance and historical dimensions of skeletal variation
there is a consensus that the original founders migrating to North America derived from a
sampling of populations from Asia prior to the last glacial maximum (Raff et al., 2011).
With the development of molecular genetic methods for accessing and characterizing aDNA
in archaeological skeletal samples, this finding is now well confirmed and has facilitated more
direct ways of testing hypotheses regarding ancestral – descendant relationships for the
Americas generally (Adachi et al., 2009; Chatters et al., 2014; Kemp et al., 2007; O’Rourke
& Raff, 2010; Raff et al., 2010, 2011; Rasmussen et al., 2014; Schurr, 2004) and regionally. For
example, the peopling of the Aleutian archipelago was characterized as a population discon-
tinuity involving replacement of the original founding population (Paleo-Aleuts) by later
immigrants (Neo-Aleuts) by AD 1000 (Hrdlička, 1945). Based on study of aDNA and other
lines of evidence (Coltrain et al., 2006; Smith et al., 2009), the peopling of the region involved
at least two migrations with later groups coexisting with the earlier founding population. On
the other hand, the close genetic relationship between Paleoamerican pre-Holocene skeletons
and native North Americans indicates biological continuity between these early groups and
indigenous populations (Chatters et al., 2014; Rasmussen et al., 2014).
Analysis of dental traits by Turner (1985, 1986a, 1986b) indicates that Native Americans
are mostly derived from the Sinodont group of Northeast Asia. His extensive study – based on
some 9000 individuals – made the case that Native American populations can be further
subdivided into three groups, the Eskaleut (Eskimos and Aleut of the Arctic), Nadene
(Athapaskans, Tlingits, Haida, and Eyaks), and Macro-Indian (all other Native North, Central,
and South Americans) (Greenberg et al., 1986; Turner, 1986a, 1986b, 1987). Based on the
assumption that dental traits evolve at a constant rate, Turner used MMD distance statistics to
calculate a “dentochronology” for the three groups migrating separately from the Old World:
an initial Paleoamerican population divergence taking place 14 000 yBP between the
founding Sinodonts and the subsequent migration of the first migrant group (Macro-Indians);
a second split and migration along the southern coast of the Bering land bridge with
populations (Eskaleut) migrating at about the same time; and a slightly later split representing
a third population migration (Nadene). Turner’s tripartite model suggests that these migrations
were separate and were completed by 9000 yBP. Greenberg and coworkers (1986:485) argued
that there is “a remarkably good fit” between Turner’s tripartite dental groupings, linguistic
divisions of Native American languages, and genetic evidence of living populations for
inferred divergence and migration dates.
The concordance of the three lines of evidence – dental, linguistic, and genetic – has met with
strong criticism. For example, Turner (1985) assigned prehistoric dentitions in the Aleutian
region to the Aleut category, which assumes that the past and present population distributions
are the same. This assumption is unlikely, especially in consideration of the remarkable
time depth and morphological and genetic diversity of the populations in this setting
(Meltzer, 1993, 1995; Raff et al., 2010; Smith et al., 2009; Steele & Powell, 1992; Stojanowski
et al., 2013).
Whatever the shortcomings of Turner’s hypothesis of the peopling of the Americas based
on his analysis of the dental record, it nonetheless reveals the challenges for developing a
broad assessment of an exceedingly complex record involving genetic drift, gene flow and
subsequent admixture, and other factors that result in the variation seen in the recent past
and living populations. Authorities agree that Northeast Asia is the originating point of
departure for populations that are ancestral to all native peoples living in the Americas
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9.3 Biohistorical issues: temporal perspectives 381
(Tamm et al., 2007; Raff et al., 2011; Rasmussen et al., 2014). Moreover, when viewed in the
broader perspective of peopling of the planet following the spread of early modern Homo
sapiens from Africa some 150 000 years ago, the migration to and settling of the Americas
happened rapidly, possibly within several hundred years of the appearance of the founding
population(s) on the eastern side of the Bering land bridge. Analysis of this complex record of
the peopling and history of native populations in the Americas is revealed through the
molecular record. That is, mtDNA sequence data in modern Native Americans indicates that
more than 90% of mtDNA variation is limited to one of four primary lineages (haplogroups
A, B, C, and D) and a fifth lineage (haplogroup X), found throughout the American
continents and intermittently in Asia (Raff et al., 2010; Schurr et al., 1990; Torroni et al.
1993). In Mongolia, Tibet, and Siberia, these haplogroups occur together (Kolman et al.,
1996), confirming the Northeast Asian origin for the native populations of the Americas.
Torroni and coworkers (1993) regarded the main four haplogroups as representing four
founding lineages. One lineage identifies populations containing an Asian-specific nine-base-
pair (9-bp) deletion between the tRNALys and Cytochrome Oxidase II genes, which has also
been identified in skeletal remains from North American archaeological samples (Kaestle,
1995, 2010; Merriwether et al., 1994, 1995; Parr et al., 1996; Stone & Stoneking, 1993).
However, the distribution in Asia of the four haplogroups suggests just one migration from
Asia (Bonatto & Salzano, 1997; Merriwhether et al., 1995). Similarly, there is a relatively
limited range of Y-chromosome variation and the ubiquitous presence of just two lineages–
C and Q – is well documented in Native American men. Furthermore, like mtDNA variation,
the commonness of the Y-chromosome lineages coupled with the high frequencies in northern
Asia reveal the certainty of an ancestral-descendant relationship between Asia and the
Americas and the likelihood of very few migrations from the former to the latter, quite
possibly just one in the range of 18 000–15 000 yBP (Mulligen et al., 2004; Zegura et al.,
2004). Moreover, this record is confirmed in analysis of autosomal DNA (e.g., STRs), suggest-
ing a single migration in the late Pleistocene (Mulligan et al., 2004).
Wallace and Torroni (1992) traced the origin of the 9-bp deletion to southeastern China,
suggesting a common ancestral group for Macro-Indians and populations from this region of
Asia. The distinctiveness of Nadene mtDNA suggests the presence of a separate and more
recent founding population (Wallace and Torroni, 1992). Much of this genetic record and
reconstructed population history of the peopling of the Western Hemisphere derives from
lineages and genetic variation in living people. The ancient DNA record, while certainly much
smaller, is providing confirmation and clarification of some important points, especially
regarding intercontinental colonizations and population movement in the later Pleistocene
(O’Rourke & Raff, 2010; Raff et al., 2011) and regional population histories (see later).
Although the record of aDNA is drawn nearly exclusively from mtDNA and only several
areas of the Americas are sampled to any extent – especially western North America and
western South America – the record is informative about population history following the last
glacial maximum. Overall, the narrow variation in mtDNA is consistent with a single major
migration event from Asia. Analysis of single nucleotide polymorphisms (SNPs; n¼364 470)
from numerous living Native American and Siberian groups, however, suggests that following
the earliest wave, there were two later migrations, including one serving as the ancestry for
speakers of Eskimo-Aleut languages and the other for Na-Dene-speaking Chipewyan from
Canada (Reich et al., 2012) (Figure 9.7). This record suggests that the vast majority of
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382 Biological distance and historical dimensions of skeletal variation
11 168
Yoruba
56 8
3 8 138 3
Algonquin Han
6 65 16 21 32 8%
16 92%
8 2 13
Figure 9.7 Gene flow, reconstructed through modern day single nucleotide polymorphisms,
from Asia to America suggests two migrations occurring after the initial migration into
the Americas: first migration (blue), second migration (green), and third migration (red).
Dotted lines represent mixture events, and solid circles denote hypothesized ancestral
populations. (From Reich et al., 2012; reproduced with permission of authors and
Nature Publishing Group.) A black and white version of this figure will appear in some
formats. For the color version, please refer to the plate section.
New World populations are descended from the first migration. Moreover, the growing
consensus based on genetic evidence for one early migration and dispersal throughout the
Americas is ultimately consistent with the tripartite model proposed by Turner.
A number of authorities identify two broad clusters of craniofacial morphology in the
Holocene of South America (González-José, Neves et al., 2005; Hubbe et al., 2010, 2011;
Neves & Hubbe, 2005; Neves et al., 2003, 2004, 2005, 2007) and North America (Jantz &
Owsley, 2001; Powell & Neves, 1999). In both continents, the authors argue for the presence of
an early group and a late group, with the former having distinctive craniofacial morphology
characterized by greater robusticity, longer, narrower crania, and greater midfacial prognath-
ism than the latter group. In South America, Neves and collaborators see the presence of two
separate founding populations, with the later representing the origin of “Mongoloid” variation
(Neves et al., 2003, 2004, 2005, 2007). In North America, Jantz & Owsley (2001) viewed a
similar pattern of craniofacial differences in earlier and later groups as a discontinuity, with
the earlier not serving as an ancestor for the later.
Thus, while the molecular evidence points to a single early founding population in the
Americas forming the ancestry for most native North and South American populations, the
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9.3 Biohistorical issues: temporal perspectives 383
craniofacial morphological evidence suggests at least two. It may be the case that the
morphological discontinuity is due to genetic drift, natural selection, or alternatively, devel-
opmental plasticity and response to changing masticatory environments and mechanical
loading patterns due to dietary shifts, or some combination of these processes (Eggars et al.,
2011; Perez et al., 2009; and see Chapter 7). In order to address these alternative explanations,
it will be necessary to document and interpret craniofacial variation in populations that are
intermediate between early (Paleoamericans) and late (modern/living native populations).
Thus far, that work has been limited in scope.
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384 Biological distance and historical dimensions of skeletal variation
American Southwest
The American Southwest is among the most dynamic regions of North America. Populations
living there engaged in considerable movement, demographic disruption, and subsistence
change. The remarkable adaptations to extreme environments and successes in subsistence
productivity involving maize agriculture in later prehistory provided the context for the
development of complex Puebloan societies in the region. As with many other areas of
the world, a central issue has been explaining the origins of farming. Did early farmers in
the region migrate from elsewhere, did the idea of farming spread through cultural borrowing,
or was farming the product of local in situ innovation? Biodistance analysis provides an
important perspective that is helping to reveal the histories of the populations of this region
and to increase our understanding of this important issue. A common theme in the explan-
ation of population history in the region is the debate over the origins of agricultural
Puebloan societies. Did these groups originate in Mexico, replacing earlier hunter-gatherer
Basketmaker populations? Beginning with a comparative biodistance analysis of crania from
the Zuni site of Hawikku, Seltzer (1944) questioned the long-held notion that Basketmaker
societies had been replaced by later-arriving groups from Mexico, who carried with them their
agricultural practices. His study, along with later biodistance analyses focusing on cranial and
dental variation, provides an important context for understanding population history in the
region (Bennett, 1973; Corruccini, 1972; Schillaci & Stojanowski, 2003). LeBlanc and co-
workers (2008) used dental discrete traits to test the hypothesis that farming derived from
migrating populations from Mexico. Their analysis of dental series from the North American
Desert West and Basketmaker populations in the northern Southwest reveals that the early
Mimbres farmers share little biological affinity with populations from northern Mexico,
thereby providing little evidence of continuity between the populations of northern Mexico
and the American Southwest. Such findings do not support a model involving migration of
farming groups from Mesoamerica to the American Southwest. Instead, these findings are
consistent with a growing body of molecular data, especially from living societies, that offer
little evidence that populations from Mesoamerica immigrated to the Southwest. In particular,
the distribution of mtDNA haplogroups among modern Native American populations across
the Southwest and northern and central Mexico is strongly inconsistent with a population
expansion from south to north (Malhi et al., 2002, 2003; Watson, 2010). As such, the
bioarchaeological record not only provides additional evidence for the in situ adoption and
intensification of maize farming by local indigenous foragers, but it also provides support to
the growing notion that common ancestry of modern tribes may have considerable antiquity
in the Southwest (Lorenz & Smith, 1996; Torroni et al., 1993; Watson et al., 2010).
Great Plains
The prehistoric and early historic North American Great Plains region was the scene of a
number of population movements that have implications for biological history. One of the
most important of these developments involved the movement of Caddoan speakers (Pawnee
and Arikara) northward up the Missouri Valley into the present-day states of Nebraska and
South Dakota. This movement is well documented on the basis of linguistic and archaeo-
logical evidence (Jantz, 1973; Parks, 1979), providing an important context for biodistance
analysis and identification of indigenous and immigrant groups. Building on preliminary
research by Bass (1964), questions regarding these relationships have been addressed by
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9.3 Biohistorical issues: temporal perspectives 385
various researchers (Jantz, 1972, 1973, 1974, 1977, 1994; Jantz et al., 1981; Key, 1983, 1994;
Key & Jantz, 1990; Owsley & Jantz, 1978; Owsley, Morey et al., 1981; Owsley, Slutzky et al.,
1981; Owsley & Symes, 1981; Ubelaker & Jantz, 1979; and see later).
Multivariate principal components analysis of more than 800 crania from Kansas,
Nebraska, South Dakota, and North Dakota reveals a general continuity of the native
populations from the initial Paleoamerican (c. 10 000 yBP) occupation of the region to the
Woodland period (to c. AD 900) (Key, 1983). In late prehistory and into the late contact
period, local population continuities are suggested, including one linking prehistoric
Woodland groups in the northern Plains with the historic Mandan of North Dakota.
A biological discontinuity is indicated between populations representing the Central Plains
tradition (c. AD 900–1400) and the period immediately preceding it – the Woodland
period. Beginning with the Central Plains tradition and extending to the historic Arikara
period, some craniofacial changes are documented, including an increase in facial height
and a decrease in vault height. This change may represent admixture with the Mandan
(who have distinctively low cranial vaults), the indigenous population encountered by the
immigrating Arikara in the Upper Missouri Valley (Jantz, 1972, 1973, 1977; Jantz et al.,
1981). Thus, unlike craniofacial changes documented in some other North American
settings, the pattern of biological change in the northern Great Plains appears to indicate
the influx of new populations around AD 900.
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386 Biological distance and historical dimensions of skeletal variation
Biodistance analysis based on nonmetric cranial traits from an expanded sample encom-
passing the Middle Woodland, Late Woodland, and Mississippian (post-AD 800) periods is in
general agreement with Buikstra’s and Droessler’s assessments of population history and
biological continuity (Konigsberg, 1987). The striking consistency of temporal ordering of
samples based on biodistance analysis, along with archaeological evidence for continuity,
provides compelling evidence that the rise of complex societies in west-central Illinois was an
in situ process rather than an event involving significant external forces or replacement.
This is not to say, however, that there were neither population movements nor changes in
the patterns of gene flow within the region. Analysis of cranial morphology in the region
reveals a record of changing patterns of gene flow, especially an expansion in the range of
gene flow in the earlier Late Woodland populations relative to that seen among the later
Mississippian populations (Steadman, 2001). Gene flow is so dramatic during the Late
Woodland that biological distance is not correlated with geographic distance. By contrast,
localization of gene flow during the subsequent Mississippian period results in a classic
pattern of isolation-by-distance, such that biological distances are strongly correlated with
geographic distance. This conclusion suggests the later Mississippian populations participated
in patterns of migration and marital exchange that result in more localized gene flow than
those of their predecessors.
This finding is consistent with other analyses of degree of gene flow in the American
Midwest. In the Ohio River valley, Tatarek & Sciulli (2000) suggested that, owing to greater
mobility of foragers than farmers, inter-population biological differentiation should be less
across forager populations than in farming populations due to their greater level of gene flow
between local populations. They tested this hypothesis through a biodistance analysis of a
large series of Late Archaic (pre-1000 BC) hunter-gatherers and Late Prehistoric (AD
1000–1600) agriculturalists. Tatarek and Sciulli used R-matrix models to assess population
structure. Such models provide a robust means of identifying genetic distance, population
divergence, and the overall effects of gene flow into a population (Relethford, 1994, 2012).
Consistent with their hypothesis, the analysis yielded evidence of greater inter-population
differentiation among Late Prehistoric farmers relative to Late Archaic hunter-gatherers, who
show remarkably little differentiation on the whole. Thus, it appears that reduced mobility
among members of these later farming societies contributed to a general reduction in gene
flow between local populations.
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9.3 Biohistorical issues: temporal perspectives 387
0.2
Apalachee Guale
Eigenvector 2 (39.4%)
0.1 Coastal
Irene Mound
0
Timucua
–0.1 Irene Mortuary
–0.2
–0.2 –0.1 0 0.1 0.2
Eigenvector 1 (54.8%)
Figure 9.8 Eigenvector plot depicts the genetic differences, represented by tooth dimensions,
between precontact populations in Georgia and Florida. (From Stojanowski, 2005a;
reproduced with permission of author and the American Anthropological Association.)
analysis of dental biodistance for La Florida indicates a dynamic period of redefinition and
emergence of groups. These ethnogenetic processes resulting in new social ties between
communities and emergence of new groups in northern Florida and coastal Georgia reveal
a complex period of biological history and social interactions (Figure 9.8).
Large-scale regionally based biodistance analysis provides essential perspective on social
and cultural formation, identity, and interaction (Stojanowski, 2004, 2005a, 2005b, 2005c,
2009, 2013a, 2013b; Stojanowski et al., 2007). In this regard, La Florida reveals population
interactions within and among communities in the mission setting on this remarkably
dynamic social landscape. Overall, the process of missionization involved aggregation of
communities resulting in broad patterns of admixture, largely in contrast to the later prehis-
toric period when population boundaries were defined by local chiefdoms that controlled
regional resources. That is, the patterns of differentiation viewed elsewhere in the later
prehistoric Eastern Woodlands (Steadman, 2001; Tatarek & Sciulli, 2000), and likely for this
setting prior to European contact, underwent a rapid change as separate communities were
gathered through various colonial-driven reducción policies into mission centers during the
historic period (Blair, 2013; Hann, 1988, 1996; Larsen, 1990a; Stojanowski, 2005c). While the
strategy of population agglomeration was viewed as facilitation of proselytizing and conver-
sion to Christianity, the underlying practice was viewed as necessary for labor exploitation,
control of native societies, and adjustment to population losses taking place during the
colonial period.
Independent biodistance analyses of dental and cranial variation (discrete traits and
metrics) documented patterns of genetic distance (Griffin et al., 2001; Stojanowski, 2005a,
2005b, 2005c, 2005d, 2009) that show striking similarity in results. In particular, there is an
increase in phenotypic variability in the early mission period followed by a decline in
variability in the late mission period. The earlier trend likely reflects the aggregation of
diverse groups in a single community and thus increases in biological diversity, largely
reflecting the presence of what were originally distinct populations now gathered into a
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388 Biological distance and historical dimensions of skeletal variation
1600–1650 1650–1704
0.2 0.2
San Martin
0.1 0.1
Eigenvector 2
Eigenvector 2
Ossuary Yamassee
0.0 0.0
SCDG-Amelia
Patale San Luis
–0.1 –0.1
SCDG-SGI
–0.2 –0.2
–0.2 –0.1 0.0 0.1 0.2 –0.2 –0.1 0.0 0.1 0.2
Eigenvector 1 Eigenvector 1
Figure 9.9 Eigenvector plots for early mission (left) and late mission (right) populations in La
Florida indicate a reduction in genetic diversity among local groups over time. (From
Stojanowski, 2009; reproduced with permission of authors and University Press of Florida.)
single population. However, in the later mission period of the later seventeenth century, a
population bottleneck and increased broad-scale gene flow among once-distinct populations
resulted in reduced genetic diversity in the larger, aggregated mission community (Figure 9.9).
Although the patterns of genetic diversity presented in these analyses are not a surprise in
consideration of the context of population aggregation and movement during the mission period,
they nonetheless reveal the powerful analytical depth of biodistance analysis for developing
a more informed understanding of biological variation and social change in this setting.
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9.4 Biohistorical issues: spatial perspectives 389
pre-Koniag (1500 BC–AD 1100) and Koniag (AD 1100–1763) periods. Univariate statistical
treatment of crown trait frequencies reveals that only two (of 14) traits exhibit significant
differences between the pre-Koniag and Koniag archaeological populations. Analysis of living
Koniag populations reveals no significant intervillage variation. Trait comparisons indicate
only three statistically significant differences between pre-Koniag and modern Koniag and
two differences between prehistoric and modern Koniag. This finding suggests long-term
population continuity.
In order to frame the biodistance analysis of native Kodiak Islanders in a wider biogeo-
graphical context, Scott (1994) compared crown trait frequencies between Kodiak Island
samples (three prehistoric, three modern) and prehistoric and protohistoric northern popula-
tions (Eskimo, Aleut, Northern Maritime, Central Maritime, Gulf of Georgia, Athapaskan,
St. Lawrence Island). Biodistance analysis based on a symmetrical relationship matrix reveals
important links between prehistoric and historic groups in the larger Arctic region, suggesting
broad patterns of continuity. Importantly, this analysis indicates that questions of local
continuity need to be addressed within larger geographical settings. With regard to questions
of affiliation and cultural patrimony, the dental record provides evidence from skeletal
remains that the prehistoric Kodiak Islanders are the ancestors – in at least some measure –
to native individuals living in the area today.
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390 Biological distance and historical dimensions of skeletal variation
iver
ois R
Illin
Missis
sippi R
RY
iver
EZ
YO
LD KL
GI
SH
KO
HN
HR
Figure 9.10 Map generated from Womble analysis of cranial discrete traits in west-central
Illinois populations. The analysis identifies a series of tiles collectively representing
population boundaries. The map indicates that the Ray site (RY) to the north is separate
from the other sites, and the Yokem (YO) and Elizabeth (EZ) sites lie in a population
boundary separating a northern from a southern group (LD, Ledders; KL, Pete Klunk;
GI, Gibson; SH, Schild; KO, Koster Mounds; HN, Helton; HR, Hacker South Mound 2).
(From Konigsberg & Buikstra, 1995; reproduced with kind permission from Springer
Science and Business Media.)
lower from the central Illinois Valley, and contains a mix of cultural and settlement attributes
from both regions. Overall, this site appears most like lower valley archaeological sites. Thus,
biodistance analysis locates the study of population groupings within a behavioral realm that
has been based previously on conventional archaeological data.
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9.4 Biohistorical issues: spatial perspectives 391
two Egyptian cemeteries, Qubbet el Hawa and Elephantine. Qubbet el Hawa is the location for
burial of nobility from the first upper Egyptian capital on the island of Elephantine. The site of
Elephantine contains the remains of mostly middle- to low-status individuals. Groups of
individuals from specific graves exhibit unusual frequencies of cranial nonmetric traits. In
grave 89, for example, 47% of individuals have occipital precondylar tubercles, normally a
very rare occurrence. This frequency is considerably higher than found in other populations
worldwide (compare with Hauser & De Stefano, 1989:135). The grave owner and lineage
founder – Sebek-Hotep – shares six of seven variants with the other individuals interred in the
family tomb. These findings provide corroboratory evidence for familial relationships and
likely a strong record of endogamy.
The study of skeletons from historic-era rural family Euroamerican cemeteries in North
America facilitates inferences about familial relationships. Analysis of remains from Wise
Cemetery in Ontario (Spence, unpublished data) and Cross Cemetery in Illinois (Larsen,
Craig et al., 1995) reveals an unusual prevalence of traits. In the Wise cemetery, the
association between grave markers and individual skeletons permitted genealogical recon-
struction of specific family relationships, thus providing important context for biodistance
analysis. Father and son (Peter Wise Sr. and Peter Wise Jr.) show evidence of trait
correspondence for some unusual variants, which include divided supraorbital foramen,
absence of zygomaticofacial foramen, open foramen spinosum, temporal squamous fora-
men, and patent mendosal suture, among other traits. The general pattern of consistency in
trait distribution in these two individuals as well as the homogeneity of traits in the sample
is well outside of what would be expected in a random sample of individuals drawn from a
larger population.
The Cross series contains an unusually high frequency of individuals with partial or full
persistent metopic sutures: nine of 14 crania (64.4%) have either partial or full metopic
sutures. This frequency contrasts sharply with what is found in other populations, most of
which are well under 10% (Hauser & De Stefano, 1989; Sullivan, 1922), with few exceptions
(Alt & Vach, 1998). Metopism heritability is unknown, but most workers are confident that it
has a high genetic component (Hauser & De Stefano, 1989). Unlike the Wise series, precise
familial relationships among skeletal individuals could not be determined owing to the lack of
association between grave markers and human remains. Historical records indicate that
individuals interred in the cemetery included primarily members of a nuclear family and their
close kin (Larsen, Craig et al., 1995). Thus, an extraordinarily high frequency of metopism is
consistent with archaeological and historical documentation that the cemetery is dominated
by closely related individuals.
There is a striking degree of homogeneity of traits in skeletal samples from family
cemeteries. When observed in large skeletal samples, such homogeneity suggests the
presence of endogamous mating patterns. Biodistance analysis of dental nonmetric traits
of some 300 individuals from the Early Bronze Age site, Bab edh-Dhra’, in southern Jordan
reveals an unusually high degree of homogeneity – about 80% of the sample shares specific
traits (Bentley, 1991). There is also a high degree of similarity between adult females and
males. Analysis of trait distribution reveals a clustering of rare traits within particular
tombs. For example, only 8% (13/158) of the Bab edh-Dhra’ population possess mandibular
molars with six cusps; all individuals with the trait are clustered in six of 25 tombs. Third
molar agenesis is present in only five tombs. In summary, the high overall degree of
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392 Biological distance and historical dimensions of skeletal variation
homogeneity suggests group endogamy, but with the clustering of traits indicating burial of
related individuals in the same tomb.
The conclusion that trait homogeneity represents group endogamy is based on the assump-
tion that traits become homogeneous through time in biological lineages (Kennedy, 1981;
Konigsberg, 1987; Konigsberg & Buikstra, 1995). Hypothetically, adult males and females in
endogamous societies should exhibit the same degree of intra-sex variability. In contrast,
exogamous populations should exhibit a relatively high degree of variability as well as
significant sex differences. This suggests, therefore, that the natal group or non-migratory
component of the population should be relatively more homogeneous in trait expression than
the migratory component.
The remarkable transformation of mortuary behavior of native populations in regions of the
New World controlled by Spain, especially in those areas having strong missionization efforts,
is revealed not only in burial posture but also in the organization in family units. In this regard,
at Tipu, Belize, and other Spanish colonial cemeteries in the Maya region, family-owned
cemeteries were used (Jacobi, 1997, 2000). In this setting, there are clear tendencies for the
presence of clustering of dental discrete traits in specific regions of the Tipu cemetery,
identifying family or kin groups. For example, there are clusters of rare traits that identify
kin relationships, such as the presence of a labial groove on the maxillary incisors and canines
and winged central incisors. At San Luis de Apalache, a group of three elite burials located
adjacent to the altar, also the only individuals interred in coffins, had similar dental size–shape
profiles suggesting familial relationships (Stojanowski et al., 2007). No other biodistance
patterns are present in the cemetery series, suggesting that only the elite were interred in a
distinctive cluster based on kinship. At the San Pedro y San Pablo de Patale mission, Jones and
coworkers (Jones, Hann et al., 1991; Jones, Storey et al., 1991) hypothesized that interment in
the cemetery was structured according to kin identity. To test the hypothesis, Stojanowski
(2005d) undertook a multivariate biodistance analysis of buccolingual and mesiodistal dimen-
sions of permanent teeth in order to assess size and shape. In contrast to San Luis, a number of
statistically significant intra-cemetery patterns are consistent with the hypothesis of kin
patterning. For example, individuals buried on the right side of the cemetery had statistically
larger teeth than those on the left side of the cemetery. Overall, the statistical analysis supports
the hypothesis that rows were occupied by distinctive kin/family groups, a pattern that is
especially clear in relation to the location of juvenile interments (Figure 9.11).
Postmarital residence is a central area of interest in anthropology, largely due to the
importance of understanding the general organization of human societies and the establish-
ment of kin-based networks, within and between communities and regions. Moreover, post-
marital residence is a fundamental part of social, political, and economic organization (Hubbe
et al., 2009). Therefore, to understand postmarital residence is to understand some very basic
premises of how past societies were organized and how they interacted with surrounding
communities and regions (Hubbe et al., 2009; Stojanowski & Schillaci, 2006). Moreover, the
case can be made that biological data that generate sex-specific patterning in phenotypic
variation are likely more appropriate for the documentation of postmarital residence in
prehistoric contexts because other lines of evidence – such as lineal patterns of material
culture transmission (e.g., ceramic styles) – are not determined by factors that are necessarily
predictable. In contrast, biological variation is governed by principles of genetic inheritance
(Schillaci & Stojanowski, 2003; Stojanowski & Schillaci, 2006).
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9.4 Biohistorical issues: spatial perspectives 393
0.5
63
52
54 7
7
8
0.0 44 8 5
8 8
7
56A
7
PC 2
6 42 8
–0.5
5
–1.0
–3 –2 –1 0 1
PC 1
Figure 9.11 At the mission cemetery of San Pedro y San Pablo de Patale, Florida,
principal components – which explain tooth size and morphological variation between
individuals – indicate definitive similarities between subadult individuals in the same burial
rows and sides. Open circles represent individuals buried on the left side of the cemetery
and filled circles subadults buried on the right side. Data points correspond to burial
rows (e.g., rows 7 and 8). (From Stojanowski 2005d. Reproduced with permission of the
author. © 2005 by the Southeastern Archaeological Conference.)
In his analysis of cranial variation in the late prehistoric Irene Mound site, Georgia skeletal
series, Hulse (1941) observed less craniometric variation in adult females than males, leading
him to speculate that this Mississippian society had a matrilocal postmarital residence pattern.
While Hulse’s analysis was insightful, the idea of postmarital residence and the ability to
document it via bioarchaeological research was undeveloped until the 1970s, beginning with
the influential studies of Corruccini (1972), Lane and Sublett (1972), and Spence (1974a,
1974b). Further inferences have been drawn about postmarital residence patterns by various
researchers, especially in relation to inter-group and intra-group variance in nonmetric traits
(Birkby, 1982; Buikstra, 1980; Cook & Aubry, 2014; Corruccini, 1972; Droessler, 1981;
Kennedy, 1981; Konigsberg, 1988; Nystrom & Malcom, 2010; Schillaci & Stojanowski,
2002, 2003; Tomkzak & Powell, 2003; see review by Stojanowski & Schillaci, 2006).
Spence (1974a, 1974b) emphasized within-group variation based on his modification of a
simple matching coefficient. In his model, a high degree of similarity in traits within a group
represents low variability. Thus, a higher mean similarity coefficient (i.e., decreased variabil-
ity) in adult males of a population in comparison with adult females indicates that males are
more closely related to one another than are females. This pattern suggests co-residence of
related men – patrilocal or virilocal residence. In this marriage system, males remain in the
natal group, whereas female partners immigrate from elsewhere. Because females are drawn
from a greater number of population groupings, they display greater trait variability
than males.
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394 Biological distance and historical dimensions of skeletal variation
Spence’s biodistance analysis of cranial, dental, and postcranial traits in skeletal remains
from the Classic period urban center of Teotihuacan in central Mexico reveals a pattern of less
variability between males than between females (see Chapter 8). Moreover, some trait fre-
quencies are considerably higher in males than in females for specific apartment complexes.
At the La Ventilla B apartment compound, for example, two supraorbital traits – supraorbital
foramen presence and multiple supraorbital foramina – are far more common in males than in
females (see discussion in Spence, 1994). The greater trait similarity among males than among
females suggests that the corporate social group may have been organized around male kin
who maintained their residence over the period of the occupation of the apartment compound.
In this patrilocal residence pattern, adult females would have originated elsewhere, either
from other apartment compounds or outside of Teotihuacan altogether.
Owing to the availability of their rich ethnographic, ethnohistorical, and archaeological
records, a number of regions have developed important and compelling analyses of post-
marital residence-based skeletal and dental parameters. In the American Southwest, for
example, at the ancestral Zuni settlement of Hawikku (New Mexico), dental biodistance (cluster)
analysis reveals individual cemeteries surrounding the community display distinctive pat-
terns, especially in Cemetery 3 where two-thirds of the sample are related (Howell & Kintigh,
1996; and see Birkby, 1982; Corruccini, 1972). Building on this biodistance analysis, Schillaci
and Stojanowski (2002, 2003, 2005; Stojanowski & Schillaci, 2006) have undertaken a
comprehensive craniometric biodistance analysis for a range of late prehistoric communities
in northern New Mexico, addressing the general hypothesis that the matrilocal residence
pattern that is ubiquitous among the modern Puebloan communities today has its roots in
their prehistoric Anasazi ancestors. An R-matrix biodistance analysis reveals a complex
pattern of residence in later prehistory. Consistent with their hypothesis, at least one prehis-
toric Tewa Pueblo – Puye – shows greater male than female variability, indicating the strong
likelihood that the community postmarital residence was matrilocal (Schillaci & Stojanowski,
2005; and see Corruccini, 1972). On the other hand, biodistance analysis of Chacoan culture
Pueblo Bonito reveals two contemporaneous but distinct populations (Figure 9.12).
Biodistance analysis reveals greater female than male variability, reflecting bilocal or non-
subscribed postmarital residence (Schillaci & Stojanowski, 2003). The latter finding contra-
dicts inferences about matrilocal postmarital residence for the Anasazi (Eggan, 1950) and
especially for Pueblo Bonito (Peregrine, 2001). Regardless of the conclusions drawn from the
biodistance analyses, the skeletal record reveals that postmarital residence in the prehistoric
American Southwest was likely considerably more varied than in the native societies in the
region today.
The analysis of relatedness, including postmarital residence, at the household or neighbor-
hood level is only rarely possible, largely because of the predilection for cemetery burial,
where it is far from clear how the cemetery is structured in relation to families or kin-based
groups. In rare instances, analysis at the household level is possible. At the large Neolithic
community of Çatalhöyük, Turkey (7400–6000 BC), the deceased were interred under the
floors of houses, presumably the place of habitation at the time of death (Hodder, 2006).
Pilloud & Larsen (2011) tested the hypotheses that (1) individuals interred within the same
house were biologically related, and (2) neighborhoods, demarcated by clusters of contem-
porary houses, were also defined by biological affinity. Dental biodistance analysis (univariate
and multivariate) revealed remarkably minimal phenotypic patterning in dental size or
morphology for deciduous or permanent teeth either at the household or neighborhood levels.
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9.4 Biohistorical issues: spatial perspectives 395
West Cemetery
North Cemetery
Stress < 1%
Figure 9.12 Approximate location of adult burials in the West and North Cemeteries at
Pueblo Bonito, New Mexico. Linear discriminant functions, using nine craniometric traits,
classified individuals into West (open shapes) or North (shaded shapes) cemeteries. Two
adult females, represented by stars, were misclassified into cemeteries based on
craniometric traits. This evidence suggests that exogamy was practiced between these two
kin-groups. In this figure, triangles represent adult males, and circles represent adult
females. (From Stojanowski & Schillaci, 2006; reproduced with permission of authors and
John Wiley & Sons, Inc.)
There are some relatively rare traits in the Çatalhöyük series that are above the mean for
individual houses (e.g., 50% parastyle in one house). These findings suggest that biological
affinity played only a limited role in who is or is not associated with specific houses and
neighborhoods. Thus, habitation at Çatalhöyük may have been constituted by “practical kin”
rather than “official (biological) kin” (sensu Bourdieu, 1977). That is, affiliation may have
been determined not on biological terms but rather as groups were called together for various
social or cultural reasons. At this early farming community, this arrangement was likely
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396 Biological distance and historical dimensions of skeletal variation
motivated by the adoption of domesticated plants and animals. Individuals may have aligned
themselves with practical kin for economic-related activities such as herding animals,
planting crops, harvesting, and other functions that required groups of individuals. This
interpretation of a very early farming society still highly dependent on nondomesticated
plants and animals is consistent with an analysis of co-residence patterns for a large sample
of present-day foraging societies (Hill et al., 2011). Most individuals in the sample of
residential groups are not genetically related. This finding is in sharp contrast to the long-
held assumption that hunter-gatherer residence patterns are entirely kin based (Service, 1962).
Importantly, it underscores the significance of residence for cooperation and social learning,
which are not necessarily functions that have to be carried out within biological kin groups.
As with foraging societies, biological kinship in at least this community of early farmers was
likely not the sole defining principle of social organization. This lack of phenotypic patterning
as it pertains to identification of kin in Çatalhöyük households and neighborhoods is not to
say that no other phenotypic patterning was present in this community. That is, dental metrics
of the males are considerably less variable than those of the females in this setting (Hillson
et al., 2013; Pilloud & Larsen, 2011), suggesting that for the community as a whole, men had
greater biological affinity to one another than did women. These results are consistent with a
community that practiced patrilocal residence, an important finding that counters earlier
interpretations of a matrilocal residence pattern and a matriarchal society as inferred from the
symbolic assemblages at the site, such as the so-called mother-goddess figures (Gimbutas,
1982). Thus, females may have been moving into the community on a greater scale than males
and perhaps from some distance.
While dental and skeletal variation provides an important window onto reconstruction
of postmarital residence pattern, new insights are becoming available via analysis of
aDNA, especially with the focus on variation in mtDNA. With DNA recovered from
archaeological human remains, it has become possible to evaluate genetic relationships
more directly and to draw conclusions about postmarital residence. Because mtDNA is
inherited from the mother only, its analysis provides an important perspective on the
degree of matrilineal relatedness within populations and between individuals. The recovery
of mtDNA from a number of archaeological settings allows bioarchaeologists to test
hypotheses about postmarital residence in general and matrilineal relatedness in particu-
lar. Earlier research on Middle Woodland (Hopewell) populations in Illinois, based on
skeletal biodistance analysis, revealed greater intra-site variation in females than males,
making the case that postmarital residence was primarily patrilocal (Buikstra, 1980;
Konigsberg & Buikstra, 1995). In order to test this hypothesis about Hopewell postmarital
residence, Bolnick & Smith (2007) analyzed mtDNA collected from 39 individuals from the
Pete Klunk mound group (Illinois). This analysis revealed the presence of all five Native
American haplogroups, and a population dominated by haplogroup C (n¼19). Comparison
of female and male diversity reveals considerably greater haplogroup variation in males.
While some settings suggest patterns of placement of related individuals in cemetery
contexts (see earlier), there were no spatial patterns of mtDNA lineages (Figure 9.13),
indicating that although the postmarital residence patterns were likely dominated by
matrilocal residence, matrilocal relationships were not expressed in mortuary practices.
This work underscores the importance of mtDNA in identifying social relationships in
past societies.
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9.4 Biohistorical issues: spatial perspectives 397
Haplogroup A
Haplogroup B
Haplogroup C
Haplogroup D
Feature A
Feature B
Figure 9.13 Spatial distribution of haplogroups in Mound 6 from the Pete Klunk mound
group, Illinois River valley. Of the 19 haplogroup C individuals in the total sample, eight are
represented here in this mound. There is no clear spatial delineation between individuals of
different haplogroups. (Adapted from Bolnick & Smith, 2007; reproduced with permission
of authors and the Society for American Archaeology.)
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398 Biological distance and historical dimensions of skeletal variation
marriage (Santley’s model). If social classes were mostly endogamous, as Santley contends,
then biodistance analysis should reveal clear distinctions in trait pattern and frequency
between higher- and lower-rank groups. These differences would reflect restrictions on
inter-group migration and mating.
Univariate statistical comparison of dental and craniofacial nonmetric traits from tomb, or
high-status, and burial, or lower-status, contexts at Monte Albán reveals no significant
differences between the two subgroups (Wilkinson & Norelli, 1981). This finding is consistent
with the model that the two subgroups are drawn from the same larger population. Most
importantly, consistent with Blanton’s hypothesis regarding relatively open interclass com-
munication, these results suggest that Monte Albán class structure may not have been
strongly endogamous.
Other social distinctions aside from rank have also been explored via biodistance analysis.
Multivariate craniometric analysis of historic-era Arikara from the Leavenworth site, South
Dakota, reveals a very high degree of biological variability, far exceeding what would be
expected in a population practicing village endogamy (Byrd & Jantz, 1994; Key & Jantz,
1990). Comparisons of this series with populations that were known to have been distinctive
in earlier Arikara history (e.g., Larson site) reveal an unusually high degree of heterogeneity in
craniometric variation. Consistent with archaeological evidence for use of specific burial areas
by groups attempting to maintain corporate identity (O’Shea, 1984) and linguistic evidence
for the presence of two dialects spoken by Leavenworth inhabitants (Byrd & Jantz, 1994),
there is a significant burial area effect in the analysis, specifically identifying two distinctive
groups from two respective areas of the site. This bipartite grouping pattern likely reflects
social subdivisions such as bands or attempts at maintaining earlier village identities. During
the late eighteenth century, different bands with their own social, historical, and linguistic
identities amalgamated into larger groupings, mostly in response to disease, depopulation,
and warfare. This biodistance analysis accounts for the large degree of heterogeneity in the
sample as a whole by identifying those patterns that contribute to variation. This
approach serves as an important means of revealing patterns of intra-population variability
(Raemsch, 1995).
The northern Peruvian coast provides an ideal natural laboratory for social reconstruction
in general. Among the best known cultures in the region is the Sicán (or Classic Lambayeque)
culture, lasting from c. AD 900 to 1100. During the fluorescence of the culture around
AD 1000, a considerable region of northern Peru was controlled or influenced by Sicán. At
Sicán, Huaca Loro, one of six monumental mounds, was constructed covering dozens of
massive shaft tombs, two of which have been systematically studied. They include the East
Tomb containing five individuals and the West Tomb containing 24 individuals (Corruccini &
Shimada 2002; Shimada et al., 2004, 2005). In addition, other burial features at the site –
including a “North Trench” containing five individuals with very few grave goods – may
represent a non-elite sector of Sicán society. The organization of burials and their treatment
involve careful planning and nonrandom placement of individuals within socially relevant
contexts, including a central chamber containing an elite adult male with an abundance of
material culture reflecting his high status in Sicán society, and adjacent niches with females
who may have been sacrificed. To the north and south of the central tomb are two opposing
groups, each containing nine females. The remains of these females were disarticulated and
hence they may represent individuals who were moved from an adjacent region of the West
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9.4 Biohistorical issues: spatial perspectives 399
tomb. A number of questions emerge from this setting where the mortuary architecture and
arrangement of interments was planned: Was this an elite family mausoleum? Are the females
more closely related than the males, suggesting a matrilocal, matrilineal society that formed
an essential framework of this early state society? Is there a pattern of variation suggesting
that the two groups of females derive from a different level of society?
In order to address these questions, Shimada and coworkers undertook a comprehensive
multivariate biodistance analysis based on large dental series. Several distinctive biologically
informed, socially relevant patterns emerge from the analysis, including (1) a high degree of
biological similarity within the south females and northern trench individuals; (2) strong
dissimilarity with all other interments in the site, a pattern based on the sharing of canine
tubercles, maxillary distal premolar paracone expansion, strong development of maxillary
incisor shoveling, and large mesial-to-central fovea maxillary central molar chords; and (3)
dissimilarity among the north females. The basic pattern shows the division of social grouping
of some manner within a carefully planned elite mortuary context.
Although it is not possible to identify the nature of the relationships in this society, it is
clear that the inclusion of specific individuals, both elite and non-elite, was an important
characteristic of the West tomb, carefully planned in all respects. As the dental biodistance
analysis reveals, the two groups of women may reflect different lineages altogether. In order
to test this hypothesis, Shimada and coworkers (2005) undertook a potentially more informa-
tive mtDNA analysis in order to identify degree of relatedness and population origin. Consist-
ent with much of the dental biodistance analysis, the mtDNA analysis revealed a high degree
of affinity in haplogroups, especially among South females. This finding underscores kin
affiliation within each group but not across groups. That is, unlike the Hopewell populations
from North America, the Sicán mtDNA analysis revealed spatial patterning of mtDNA,
especially involving the placement of possibly two maternal kin relationships (Figure 9.14).
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400 Biological distance and historical dimensions of skeletal variation
SOUTH GROUP
ANTECHAMBER
B15 B5
B8
B7
B6
NORTH NICHE
SOUTH NICHE
B21 CENTRAL
B16 B24 CHAMBER B20
B17
B18
B23 B10 B11
B4 PRINCIPAL B12
B9
B22 PERSONAGE
B19
B1
NORTH GROUP
B13 B14
JUVENILE
B3
B2
0 1 2
B = BURIAL N
m
Figure 9.14 Ancient mitochondrial DNA analyses of skeletons from the West Tomb in Sicán,
Peru, show the close spatial distribution of maternally related individuals in both north and
south group burials. Black shapes represent mtDNA-related individuals within the tomb,
and open circles represent individuals who are not related mitochondrially to other
individuals within the tomb. (From Shimada et al., 2004; reproduced with permission of
authors and University of Chicago Press.)
wide cranial base – that contrast with native Easter Islanders. Additionally, this and two other
individuals possess anatomical features – such as forward projection of the midfacial region –
suggesting European admixture. Owsley and coworkers contend that admixture may have
involved a European father and native mother, a conclusion well supported from the historic
accounts of native and non-native interactions.
In the Great Plains, European exploration and settlement increased the opportunity for
admixture between Europeans and native groups. There are a number of historical accounts of
Euroamerican resident traders who married native women and raised offspring (Jantz &
Owsley, 1994b). Multivariate analysis of human remains from Swan Creek, South Dakota,
indicates the presence of a morphologically unique cranium – an old adult male from a
multiple grave including five individuals. The cranium possesses a number of European or
Euroamerican traits, including reduced facial prognathism, triangular parabolic palate, prom-
inent nasal bones, and narrow nasal aperture. Three craniofacial indices that discriminate
between population groups suggest that the outlier male has Euroamerican ancestry (Gill &
Gilbert, 1990). Based on the archaeological and mortuary context, the individual was likely a
resident trader or a captive. The “foreign” origin of the individual is confirmed by his unique
element composition of dental enamel (Schneider & Blakeslee, 1990), suggesting that the
individual’s enamel had formed while he was living in another geographical setting. The
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9.5 Summary and conclusions 401
similarity of tooth wear pattern with other individuals in the Swan Creek series suggests that
the individual lived a predominantly native lifestyle, at least with respect to the use of the
dentition in masticatory functions. The teeth are heavily worn, a pattern typical of native but
not Euroamerican dentitions in this setting.
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Bioarchaeological paleodemography:
10 interpreting age-at-death structures
10.1 Introduction
There has been a long and enduring interest in the demographic structure of human popula-
tions, past and present. Thomas Malthus (1798), founder of the science of demography, linked
mortality, survival, and success of a population to the food supply – populations with adequate
nutrition and lower disease loads will grow, but malnutrition and disease will limit their size.
The dynamic interplay between fertility, mortality, and available food resources formed a key
element of Charles Darwin’s (1859) theory of natural selection. Darwin concluded that
individuals having characteristics that promote their ability to survive, such as the successful
acquisition of food and resources generally, will live to adulthood and reproduce. The various
demographic measures of the success (or failure) of human populations – mortality, fertility,
survivorship, longevity, and life expectancy – have endured as important foci of inquiry by
social, behavioral, biological, and medical scientists into the twenty-first century and will
continue to do so for the foreseeable future.
Although fertility, mortality, and other key parameters of demographic analysis are import-
ant measures of success in today’s world, life expectancy remains the gold standard for
evaluating health and well-being around the globe, especially in comparative perspective.
Life expectancy of 79 years in the United States versus just 58 years in Rwanda (United
Nations, 2011) speaks to the chasm that exists in health and well-being between these two
nations, implying a considerably healthier living environment in the former than in the latter.
Similarly, increased life expectancy is an indicator of improving living conditions. In the
United States, for example, life expectancy increased from 50 years in 1910 to nearly 80 years
in 2010, clearly due, in part, to expansion in the availability of healthcare and improved
nutrition. If life expectancy (and other demographic parameters) can be measured in ancient
populations, then their age structures can provide a powerful perspective on health and
conditions of life.
Age-at-death of groups of skeletons representing past populations provides important
information regarding the demographic structure of ancient populations. As such, age-at-
death of skeletal samples is a powerful structuring phenomenon. That is, the composite age
structures of collections of skeletons are important for understanding many of the patterns of
variation discussed in the previous chapters of this book. For example, on a very simple level,
it stands to reason that a skeletal series of older individuals should be expected to have more
osteoarthritis and more dental disease than a series of younger individuals; the pathological
conditions are cumulative and are tightly linked to age. More broadly, formal analysis of age
structure of a series of skeletons derived from a once-living community or collection of
communities provides a fundamental record of well-being, as in living populations in the
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10.1 Introduction 403
80
60
40
20
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an ull
Sa ibs
et
U ii
ae
co s
Fi ae
i
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C rum
Pa ora
Ti e
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Figure 10.1 Variable frequencies of well-preserved bone elements among St. Estève le Pont
(Berre l’Etang, Bouche-du-Rhône, France), Observance (Marseilles, France), and Spitalfields
(London, United Kingdom) assemblages. This pattern reveals the disproportionate
preservation of smaller, more fragile elements. (From Bello et al., 2006; reproduced with
permission of authors and John Wiley & Sons, Inc.)
world today. For purposes of this chapter, I use Robert Hoppa’s definition of paleodemogra-
phy, “. . .the field of inquiry that attempts to identify demographic parameters from past
populations derived from archaeological contexts” (2002:9). While hypotheses and questions
in this area are dominated by bioarchaeological concerns and its primary focus is on human
remains, paleodemography explores population distributions and densities on the landscape
at any one time as well as over time from a variety of non-skeletal archaeological data sets
and documentary sources (Chamberlain, 2006; Meindl & Russell, 1998; Milner et al., 2008).
This chapter focuses on the interpretation of age-at-death profiles constructed from collec-
tions of skeletons and in relation to their archaeological contexts.
For a variety of reasons, bioarchaeological paleodemography has struggled, in both its
theory and its method, thus partly reflecting the considerable amount of debate among
paleodemographers (Buikstra & Konigsberg, 1985; Frankenberg & Konigsberg, 2006; Hoppa,
2002; Milner et al., 2008; and following). One of the most obvious issues is the relative
preservation of skeletal elements, especially smaller, more fragile elements (Bello et al., 2006;
Stojanowski et al., 2002; Walker and Johnson, 1988) (Figure 10.1). However, as discussed in
this chapter, even under the best of preservational circumstances, other more fundamental
issues abound in how bioarchaeologists interpret age structures. Despite the hand-wringing
and sometimes raucous disagreements that are alluded to in this chapter, when key assump-
tions are addressed and limitations are understood, this area of inquiry gains considerable
importance for understanding past population dynamics. Paleodemography is here to stay, for
whenever estimates of age or sex are made for skeletons from archaeological settings, a
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404 Bioarchaeological paleodemography: interpreting age-at-death structures
demographic statement has been made. As mentioned earlier, however, this chapter makes the
case that paleodemography as it pertains to the study of human remains from archaeological
contexts is far more than estimating age and sex. Rather, the case is made that to understand
age structure is to have a more informed understanding of health, disease, and a multitude of
other factors considered in this book.
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10.2 Reconstructing and interpreting age-at-death profiles: it has been mostly about mortality 405
Fueling much of this interest in paleodemography in the 1960s and 1970s was the notion
that age-at-death composition and structure of living populations are a record of mortality
and life expectancy. That is, the mean age-at-death for an archaeological skeletal assemblage
provides a record of when, on average, individuals of a population die and the age to
which a person in the population can expect to live from a given point in their life history
(e.g., birth). One of the significant attempts to characterize age structure was Johnston and
Snow’s (1961) analysis of the Archaic period Indian Knoll series, Kentucky. At the time of
their study, the Indian Knoll series was one of the best documented and largest series of
archaeological skeletons in North America, ideally suited for bioarchaeological and paleo-
pathological study (Larsen, 2006, 2012). Using various indicators of age estimation to derive
composite individual ages for this extraordinarily large series (n¼873), Johnston and Snow
found that the average age-at-death (a proxy for “life expectancy”) in the Indian Knoll series
was just 18.6 years. Moreover, 20% of the Indian Knoll individuals were less than one year of
age, which is a considerably higher frequency than any age group in the Pecos Pueblo series.
Based on Johnston and Snow’s demographic assessment and in comparison with other
archaeological series from Austria and Egypt, they concluded that the population from Indian
Knoll had remarkably high infant mortality, “. . .similar to the more primitive, nonagricultural
groups. . .(and) little over-all similarity. . .to more economically and technologically advanced
groups” (Johnston & Snow, 1961:243).
Robert Blakely’s (1971) “mortality” analysis of Indian Knoll a decade later involving the
comparison with other Archaic period foragers and post-Archaic agriculturalists (Hopewell,
Mississippian) from Illinois revealed that, like Johnston and Snow’s analysis, the earlier
Archaic series from both Kentucky and Illinois had a higher percentage of children aged zero
to nine years old than in the later series. These earlier researchers concluded that the samples
they studied represented populations having an elevated level of infant mortality and low life
expectancy, a point to which this chapter later returns.
In ancient Greece, Angel (1947) documented a pattern of temporal increase in the percent-
age of older adults from c. 3500 BC to AD 1300, indicating to him an increase in life
expectancy based on these “mortality” profiles. In other words, like Johnston & Snow
(1961) and Blakely (1971), to Angel, higher average age-at-death or presence of a relatively
greater number of older adults translated directly into a picture of greater survivorship, length
of life, and reduced mortality overall.
Similarly, in Neolithic Çatalhöyük, Turkey (7400–6000 BC), based on his study of
remains recovered by James Mellaart, Angel (1971b) found an average age-at-death for
adult males and females of 34.4 and 29.8 years, respectively, higher than estimates he had
derived for the Upper Paleolithic, thus meeting his expectation of greater “longevity” in
the Neolithic. This increased presence of older, longer-lived adults, he argued, allowed the
society “. . .time to develop and stabilize a culture considerably more complex and richer
than that of the Upper Palaeolithic” (1971b:80). Moreover, for women, the increase in life
expectancy in the Neolithic was important in “. . .allowing more time for childbearing as
well as for training of children” (1971b:80). Thus, his idea of increasing longevity – a
conclusion driven by the presence of more older adults and higher mean age-at-death –
had, in his view, important implications for cultural and social developments in the
earliest farmers of western Asia in comparison with earlier foraging societies from this
region and elsewhere.
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406 Bioarchaeological paleodemography: interpreting age-at-death structures
10.3 Paleodemographers adopt the life table for age structure analysis
Age structure analysis built on the use of life tables was developed by demographers and
actuarial scientists in order to calculate probability of survival to any age, typically starting at
birth (Table 10.1). Because life tables provide the most complete characterization of mortality
in living populations, it made considerable sense for bioarchaeologists deriving their age data
from archaeological skeletal series to apply life table analysis. Bioarchaeologists do this by
Table 10.1 Life table for Northern Ache females. The table is calculated from
survivorship data in Hill & Hurtado (1995) (Adapted from Chamberlain, 2006;
Table 2.2)
x lx dx qx Lx Tx ex
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10.3 Paleodemographers adopt the life table for age structure analysis 407
assuming the age-at-death distribution in the cemetery is equivalent to the dx values in the
life table. Indeed, life table analyses of archaeological skeletal series were rapidly adopted in
the late 1960s and 1970s (Bennett, 1973; Blakely, 1977; Buikstra, 1976a; Kobayashi, 1967;
Lovejoy et al., 1977; Owsley & Bass, 1979; Owsley et al., 1977;Ubelaker, 1974). Acsádi and
Nemeskéri’s (1970) monumental work presented their comprehensive series of life tables,
ranging in time from the late Paleolithic to Medieval Hungary. Importantly, their monograph
was among the first to discuss, in great detail, age and sex estimation methods, quality and
composition of the archaeological skeletal series, and other issues relevant to the study of
ancient populations from skeletal remains.
In the application to North American skeletal series, several key studies revealed the
potential of life table analysis, especially with regard to comparison of multiple populations
within geographical regions. In his classic study, Ubelaker (1974) compared two late prehis-
toric ossuary skeletal series from the Nanjemoy site in the Middle Atlantic region of North
America with a Huron series from Canada (Anderson, 1964). His life table analysis of the
Nanjemoy series revealed relatively few infants and numerous mature adults, suggesting
lower infant mortality and greater adult longevity than in the Huron series. The study was
among the first to apply new methods in age estimation based on histological structures (see
Chapter 2), providing new insights into bone remodeling and age estimation techniques
central to modern paleodemographic analysis.
In lieu of comparisons of demographic profiles derived from archaeological skeletal series,
Weiss (1972, 1973) advocated comparison of archaeological series with model United Nations
life tables (United Nations, 2011). Model United Nations life tables are based on numerous
censuses from around the world and provide mortality data from a wide range of economic,
political, and social circumstances, including those that approximate small scale societies that
are more typically studied by anthropologists (see Weiss’ [1973] model life tables developed
from anthropological field studies). Along this line, Buikstra (1976a) compared estimates of
mortality and survivorship derived from her life table analysis of skeletal remains from the
Middle Woodland Gibson-Klunk mound series (c. 50 BC–AD 400) in the lower Illinois River
valley to model United Nations tables. The general shape of the mortality curve for Gibson-
Klunk suggested a pattern of mortality that approximated recent, modern populations.
What is especially striking about the life table analysis of the Nanjemoy, Gibson-Klunk, and
many other archaeological samples is the paucity of the very young (<5 years) and the very
old (>50 years). This pattern largely reflects poor preservation of the former and inaccuracies
in age estimation methods in the latter (Milner et al., 2008; Walker et al., 1988; and
following). With respect to life table analysis, the apparent underrepresentation of young
juveniles results in considerably skewed (and inaccurate) calculations of the proportion of
deaths (“mortality”) occurring in successive age intervals (dx) as well as the probability of
survival (“survivorship”) of an individual from a cohort that is alive at the beginning of the
age category (lx) (Chamberlain, 2006; Moore et al., 1975). Even skeletal series that were
remarkably large and contained numerous very young juveniles did not seem to represent
realistic population profiles. For example, Howell’s (1982) commentary on Lovejoy and
coworker’s (1977) analysis of the Libben series indicated that “survivorship” of middle- and
older-aged adults as displayed in the Libben life table would have been unrealistically low,
certainly too low to maintain the necessary work force and social context in general for
supporting the community with its dietary and economic needs (Howell, 1982).
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408 Bioarchaeological paleodemography: interpreting age-at-death structures
Despite the newfound use of life tables, a range of problems continued to surface. Angel
(1969) declined analyzing and interpreting population structure based on life table analysis,
largely owing to the vagaries of representation – especially younger juveniles – in archaeo-
logical skeletal assemblages and the inability to account for population growth. The former
has profound influence on mortality calculations, and the latter runs counter to the primary
underlying assumptions of life table analysis, namely population stability. In his mind, the
assumptions about collections of human remains from archaeological contexts did not meet
the key tenets of life table analysis (Frankenberg & Konigsberg, 2006).
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10.4 Addressing the assumptions of paleodemography 409
The choice of archaeological recovery methods and site sampling contributes to variable
sample representation. The excavation at Neolithic Çatalhöyük reveals the importance of
the presence of a clearly defined excavation protocol in determining what is or is not
recovered. Angel (1971b), for example, reported a very low presence of young juvenile
remains excavated by Mellaart. Later excavations produced an unusually high number of
juvenile remains, well over half of the collection, including numerous neonates (Hillson et al.,
2013). The remains examined by Angel were recovered using an excavation protocol involv-
ing rapid shoveling of considerable amounts of fill, whereas the more recent excavations
involved highly detailed recovery techniques, such as screening. Thus, the demographic
analysis based on the latter will likely be more accurate and representative than that based
on the former.
Juvenile remains can also be underrepresented under circumstances where the mortuary
treatment is different from that of adults. For example, shallower burial of juveniles than
adults in some instances in Hungary resulted in poorer preservation of the former relative to
the latter (Acsádi & Nemeskéri, 1970; and see Bello et al., 2006). Moreover, entire age groups –
such as young juveniles – may be missing from cemeteries because they are interred separ-
ately from the main population (Pinhasi & Bourbou, 2008; Schwartz et al., 2010, 2012).
In addition, the completeness of excavation of a mortuary context can play a critical role
in which segments of a population are represented in a study series. Buikstra (1976a) notes
that late nineteenth-century excavations at the Pete Klunk mound group in the lower Illinois
River valley produced only 16 burials, with an average of three skeletons per mound. In
contrast, later excavations involving complete recovery and excavation produced an average
of 38 burials per mound.
Third, life table analysis assumes that the ages of the individuals comprising the
collection of remains are accurate. Indeed, it is this assumption that is so very crucial
to the meaning of ancient age structures. The clear breaking point in discussions of
paleodemography is linked to the controversial article by Bocquet-Appel and Masset
(1982) who concluded from their analysis of the record that, “It is not the information
provided by the age indicators which is dangerous; it is the fruitless need to make them
say more than they can” (1982:332). While concluding that ancient skeletons are
certainly a fund of information for understanding past population biology, they single
out the record of demography as a dead end, bidding “farewell to paleodemography”
(1982:332). Simply, age estimation methods as applied to archaeological skeletons are
too imprecise and fraught with problems to provide anything meaningful about age
structures except in the very crudest way. Moreover, Bocquet-Appel and Masset (1982)
concluded that age structures provided by paleodemographers simply represent artifacts
of the age-at-death composition of the original reference sample used to estimate age-
at-death in the target series. For example, if the age estimations for an archaeological
series were made using the Todd (1920) pubic symphysis method, then the age-at-death
profile simply approximates the series Todd used in the development of his method.
Thus, in the view of Bocquet-Appel and Masset, the average age-at-death of a skeletal
series represents a combination of real biological processes of aging and the reference
sample age structure. If this critique held true, then paleodemography has fundamental
(and fatal) flaws.
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410 Bioarchaeological paleodemography: interpreting age-at-death structures
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10.5 New solutions to interpreting age-at-death profiles in archaeological skeletal series 411
Not surprisingly, almost immediately following Bocquet-Appel and Masset’s (1982) publi-
cation, bioarchaeology saw the slowdown of the publication of paleodemographic reports and
life tables, reflecting the growing recognition that considerably more thought had to be put
into identifying the strengths and weaknesses of deriving mortality profiles from archaeo-
logical skeletal series. As is true of any maturing science, Bocquet-Appel and Masset’s
negative assessment had a positive, constructive outcome. Ultimately, it caused researchers
to undertake thoughtful considerations of what age-at-death and age structure reveal about
population dynamics when these population dynamics rely on collections of dead people
versus populations of living individuals.
Tackling this problem directly, Sattenspiel and Harpending (1983) addressed the question:
Is it appropriate to use mean age-at-death in a skeletal series from a single site or collection of
sites as an appropriate estimate of life expectancy at birth? Put another way, does hypothet-
ical population x with a mean age-at-death of 19.6 have a lower life expectancy at birth than
hypothetical population y with a mean age-at-death of 22.3 years? A reading of much of the
bioarchaeological paleodemographic literature would certainly seem to affirm this assump-
tion, and life table analysis appeared to support the notion as well. Sattenspiel and Harpend-
ing (1983) reminded anthropologists that the procedure is a valid one only if populations are
stationary – that birth and death rates remain stable and population size remains unchanged,
neither increasing nor decreasing. They made the compelling case that this assumption is
rarely met in living populations. Thus, by inference, could that be the case for past popula-
tions? Does mean age-at-death reflect life expectancy in past populations? The answer is no.
Using simple algebraic analysis, they analyzed a hypothetical stable skeletal population
having a uniform age-specific death rate. Specifically, the analysis showed that even a small
change in the birth rate results in a proportional change in mean age-at-death, similar to what
Coale (1972) found in his examination of the influence of fertility and mortality changes on
age distribution in living populations. Mean age-at-death in a series of skeletons is strongly
influenced by fertility and not mortality – mean age-at-death is the inverse of birth rate.
Therefore, if mean age-at-death is declining, then birth rate is likely increasing. This break-
through in paleodemography, therefore, shows that fertility has a considerably stronger
influence on the distribution of deaths in a skeletal series than does mortality. Certainly, this
notion of a death assemblage being more informative about fertility than mortality is counter-
intuitive, so much so that it is now far more common in the literature to see bioarchaeologists
interpreting skeletal samples as records of mortality (Coppa et al., 1995; Nagaoka & Hirata,
2007; Nagaoka et al., 2006; Sullivan, 2004).
When one considers that fertility-focused paleodemography is really more biological than
demographic, the concept is far more intuitive. One need only consider the classic population
pyramid with the oldest at the peak and the youngest at the base (Figure 10.2). In virtually all
mammals, the rate of dying is highest at the pyramid’s base and peak, with the youngest and
the oldest dying in higher proportions than the other age groups. However, fertility happens
only at the base of the pyramid, while mortality is spread throughout the rest of the pyramid.
This means that when there is high fertility, the base of the pyramid is wide and the rest of the
pyramid becomes progressively narrower. When there is low fertility, the base is narrow and
the ages are distributed higher on the pyramid. High fertility, therefore, makes for a young
population. Thus, no matter how counterintuitive fertility-based paleodemography may be,
death assemblages tell far more about fertility and birth rate than mortality and death rate.
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412 Bioarchaeological paleodemography: interpreting age-at-death structures
(a) Melanesia
75+
70−74
65−69
60−64
55−59
50−54
Age groups
45−49
40−44
35−39 Male
30−34
Female
25−29
20−24
15−19
10−14
5−9
0−4
10 5 0 5 10
Percent population
(b) Polynesia
75+
70−74
65−69
60−64
55−59
50−54
Age groups
45−49
40−44
35−39 Male
30−34
Female
25−29
20−24
15−19
10−14
5−9
0−4
10 5 0 5 10
Percent population
(c) Australia
75+
70−74
65−69
60−64
55−59
50−54
Age groups
45−49
40−44
35−39 Male
30−34
Female
25−29
20−24
15−19
10−14
5−9
0−4
10 5 0 5 10
Percent population
Figure 10.2 Population pyramids for Melanesia (a), Polynesia (b), and Australia (c) in 2010.
Note the demographic discrepancies in age groups between populations. Higher fertility
populations have a relatively wide base. (Graph produced from data published by United
Nations, Department of Economic and Social Affairs, Population Division, 2012.)
Simply, as noted by Robert McCaa, “Common sense tells us that the age structure of deaths is
completely determined by mortality, but common sense is wrong” (2002:98).
Sattenspiel and Harpending (1983) examined several published studies and derived new
interpretations that make considerably more sense than what had been reported by Acsádi
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10.5 New solutions to interpreting age-at-death profiles in archaeological skeletal series 413
and Nemeskéri (1970) for skeletal series from Maghreb (Paleolithic), Alsónémedi (Copper Age),
Intercisa and Brigetio (Roman period), Sopronköhida (early Medieval), and Hungary (late
Medieval), by Green and coworkers (1974) for Nubia, and by Angel (1971a) for Greece. Their
birth rate revealed a temporal pattern indicating population dynamics and growth (or decline)
that was very different from the conclusions drawn by earlier investigators. Similarly,
reanalysis of the Dickson Mounds age structure reveals the likelihood that the decline in
mean age-at-death is best interpreted as an increase in birth rate and fertility, rather than
reflecting a decline in life expectancy and increased mortality (Johansson & Horowitz, 1986).
The result of the Sattenspiel and Harpending (1983) study was to make the strong case that
mean age-at-death is a simple inverse of birthrate: the lower the mean age-at-death, the
greater the birthrate (and see Acsádi & Nemeskéri, 1970; Bocquet-Appel & Masset, 1982;
Johansson & Horowitz, 1986; McCaa, 2002; Milner et al., 1989; Wood et al., 1992). However,
this fundamental development in bioarchaeological demography still does not handle the
problem facing most bioarchaeologists in assessing growth (or decline) of a population – the
near-universal underrepresentation of young juveniles in archaeological death assemblages.
The link between age structure and birth rate is clear, but how to accommodate the very
thorny issue of incompleteness of skeletal samples, especially regarding the frequent under-
representation of young juveniles, is less straightforward. This is no small matter if one
expects to apply fertility models to explain key population events, such as (and perhaps
among the most important) the dramatic and global rise in population beginning in the
Neolithic with the shift from foraging to farming. The archaeological community has argued
the shift from foraging to farming was accompanied by an increase in population size owing
to increased growth rates, mostly due to the rise in fertility. Moreover, ethnographic analogies
indicate that horticulturalists and incipient farmers have higher fertility and birthrates than
foragers (Bentley et al., 1993; Gage & DeWitte, 2009; Hewlett, 1991; Leslie et al., 1999). How,
then, do we tackle the problem of missing juveniles and still estimate patterns of fertility and
birthrate?
Acknowledging the issue of underrepresentation and still being able to explain change in
population structure, especially with regard to understanding the dynamics of population
increase in recent human evolution, Buikstra and coworkers (1986) offered a case study from
the Illinois River valley, where population increase was clearly a phenomenon of later
prehistory and was almost certainly linked to the introduction of and increased dependence
on maize agriculture. Following Bocquet-Appel’s lead (Bocquet-Appel & Masset, 1982), in
order to circumvent the clear underrepresentation of young juveniles, they used a fertility
estimator – a juvenility index calculated by the proportion of deaths equal to and exceeding the
age of 30 years (D30þ) to deaths equal to and exceeding the age of five years (D5þ). Consistent
with expectations articulated by Sattenspiel & Harpending (1983), there is a decrease in the
D30þ/D5þ ratio from the Middle Woodland through the Late Mississippian periods, reflecting
an increasingly younger population, a factor driven largely by higher birthrates and increased
fertility overall in later populations compared with earlier populations (Figure 10.3). There-
after, a considerable record of use of the juvenility index, either the one proposed by Buikstra
and coworkers or modifications thereof, reveals a consistent pattern in virtually every setting
examined, especially following upon the expected outcomes associated with either the
adoption or intensification of agriculture. In specific regional settings, this general pattern
of elevated fertility and birthrate has now been well documented in the Near East (Guerrero
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414 Bioarchaeological paleodemography: interpreting age-at-death structures
0.90
0.85
0.80
0.75
0.70
D30+/D5+
0.65
0.60
0.55
0.50
0.45
0.40
0.35
LW
LW
s
LW
LW
W
is
rL
M
M
o.
ld
n
s
te
o.
er
to
ld
C
hi
un
C
s
el
dd
hi
Sc
Ko
ke
Kl
ke
Sc
Le
Pi
n/
Pi
so
ib
G
Skeletal Series
Figure 10.3 Juvenility index or the proportion of deaths, D30þ/D5þ, in prehistoric
west-central Illinois skeletons. Middle Woodland sites: Pike County MW and Gibson/Klunk
MW; Late Woodland sites: Pike County LW, Helton LW, Koster LW, Ledders LW, Schild
LW; and Mississippian sites: Schild Knolls Miss. (Adapted from Buikstra et al., 1986;
reproduced with permission of author and the Society for American Antiquity.)
et al., 2008; Hershkovitz & Gopher, 2008; Littleton, 2011), Europe (Bocquet-Appel, 2002;
Bocquet-Appel & Naji, 2006; Jackes & Meiklejohn, 2008) (Figure 10.4), South and Southeast
Asia (Domett & Oxenham, 2010; Pietrusewsky & Douglas, 2002a, 2002b; Robbins Schug,
2011a, 2011b), various settings in North America (Buikstra et al., 1986; Kohler et al., 2008;
Larsen et al., 2007), and North America generally (Bocquet-Appel & Naji, 2006; Bocquet-
Appel et al., 2008) (Figure 10.5). This remarkably consistent pattern, showing an abrupt
increase in the proportion of juveniles in archaeological death assemblages, provides an
essential element for understanding one of the most important adaptive transitions in human
evolution, namely the foraging-to-farming transition, or what has also been called the
Neolithic demographic transition (Bocquet-Appel, 2002, 2008, 2011) (Figure 10.6). This abrupt
spike cannot be related simply to differences in preservation of skeletons or mortuary
behavior. Rather, it signals a worldwide and rapid transition to agriculture, at least in those
settings studied. Importantly, this development in modern bioarchaeological paleodemogra-
phy adds important context for understanding the various indicators of morbidity, dietary,
and lifestyle issues discussed elsewhere in this book and indeed the entire package of the
foraging-to-farming transition.
All of the aforementioned research speaks to the development and growth of food produc-
tion and the expected link to increased fertility. Likely, the underlying theme is the increased
availability of stored foods made possible by storage of grains and the shift to a practice of
producing weaning foods from cooked grains and consumption of soft mushes by young
juveniles. However, it is important to point out that the shift to agricultural production and
intensification is not always linked to an increase in birthrate or population growth more
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10.5 New solutions to interpreting age-at-death profiles in archaeological skeletal series 415
0.7
La_
0.6 Cal
Hei
Bad
0.5 Par
Nor
Ded
Ajd
Haz
Cau
0.4 Len Viln Sch
D19+/D5+
Les
Ais Mor
Nie
Bru Der
Bre Wan Gro
Pon Fon Cas Ca Pie
0.3 Bau Cer
Ved∑ Son Cov Bel
Col∑ Cha Villan Gou
Yas Mont
Mal Jun
Stu Rea Vik
Ta∑ Nit Bro Loi Eyb
0.2 Sam Mon Pec
Rut
Die∑ Cab Lar
Moi∑ ∑
Ska Ait Tre Vill
Vedr Oct Ave Cen
Mai
0.1
zen Dic
0.0
0
00
00
00
00
0
0
00
00
00
00
10
20
30
40
–4
–3
–2
–1
dt (years)
Figure 10.4 Proportion of deaths, D19þ/D5þ, among 68 prehistoric European cemeteries,
plotted to chronological distance from the Neolithic diffusion front (dt, the time elapsed
from the beginning of farming at each locality). The dashed line represents the expected
proportion of subadults if the growth rate is zero. Subadult prevalence data indicate an
increase in growth rate associated with the Neolithic transition. (From Bocquet-Appel,
2002; reproduced with permission of author and University of Chicago Press.)
generally. There certainly is a link between farming and population increase for most settings
studied by bioarchaeologists. Nevertheless, in some settings where agricultural intensification
has taken place, there have been reductions in fertility owing to generally deteriorating
circumstances and life conditions. The increase in morbidity associated with these deterior-
ating circumstances is well outlined elsewhere (see Chapter 3). Suffice it to say that increased
prevalence of infection and infectious disease owing to living in closer, more crowded
conditions of semi-permanent or permanent settlements, decreased nutritional quality, and
other factors must have contributed to increased mortality, thus partially or entirely offsetting
the elevated fertility rates now documented in the bioarchaeological record. This underscores
the important point that population dynamics as they relate to size is a balance between
fertility and mortality.
Colonial Florida is one such setting where agriculture intensified, but depressed living
circumstances and elevated mortality contributed to a decline and eventual extinction of a
population (Larsen, Griffin et al., 2001; Larsen et al., 2007; Russell et al., 1990). In this setting,
key dietary and behavioral changes contributed to an increase in infectious disease, increased
focus on maize, loss of marine resources, and extraordinary labor demands placed upon
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416 Bioarchaeological paleodemography: interpreting age-at-death structures
0.6
0.5
0.4
D19+/D5+
0.3
0.2
0.1
0 0 00 00 0 0 0 00 00 00
00 00 0 0 00 00 10 20 30
-6 -5 -4 -3 -2 -1
dt (years)
Figure 10.5 Proportion of deaths, D15þ/D5þ, among 62 North American samples, relative
to chronological distance from the Neolithic diffusion front (dt). The dashed line represents
the expected proportion of subadults if the growth rate is zero. (From Bocquet-Appel & Naji,
2006; reproduced with permission of authors and University of Chicago Press.)
0.5
0.4
0.3
15p5
0.2
0.1
0.0
0 0 0 00 0 00 00 00 00
00 00 00 0 10 20 30 40
-4 -3 -2 -1
dt (years)
Figure 10.6 Proportion of deaths, D15þ/D5þ, among 133 Northern Hemisphere cemeteries,
relative to chronological distance from the Neolithic diffusion front (dt). A significant
increase in population is observed during the Neolithic period. The dashed line represents
the expected proportion of subadults if the growth rate is zero. (From Bocquet-Appel, 2011;
reproduced with permission from American Association for the Advancement of Science &
from Springer Science and Business Media.)
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10.5 New solutions to interpreting age-at-death profiles in archaeological skeletal series 417
0.8
0.7
0.6
0.5
D30+ / D5+
0.4
0.3
0.2
0.1
0
PP PA EM LM
Guale Skeletal Series
Figure 10.7 Proportion of deaths, D30þ/D5þ, among Prehistoric preagricultural (PP),
Prehistoric agricultural (PA), Early Mission (EM), and Late Mission (LM) Guale populations,
Spanish Florida. These juvenility indices (D30þ/D5þ) reveal a significant increase in the
proportion of older adults to subadults from Prehistoric and Early Mission to Late Mission
periods, which is indicative of reduced population fertility.
native societies by European powers. During the later seventeenth century, combined pres-
sures from slaving forays into Spanish Florida by native enemies and encroachment south-
ward by British interests led to considerable declines in the population. We know, therefore,
that increased mortality, morbidity, and decaying conditions likely contributed to increased
deaths and declining population size. Occurring over a period of decades, this would have
depressed the size of the population, which, indeed, has been well documented in historic
sources (Worth, 2001, 2007). However, every demographer knows that mortality represents
only one side of the demographic coin – fertility rates also play an important role. In order to
test the hypothesis that fertility and birthrates declined in native populations of Spanish
Florida, application of the juvenility index (D30þ/D5þ) was calculated for a skeletal series
representing the Guale tribal group, including prehistoric foragers, prehistoric farmers, early
mission, and late mission populations. The ratios for the prehistoric foragers, prehistoric
farmers, and early mission Native Americans are statistically indistinguishable, but show
some fluctuation in the juvenility index (0.3790, 0.3500, 0.2823, respectively; Figure 10.7). In
particular, there is a suggestion of increased fertility in the late prehistoric period when maize
farming was present, followed by a further increase in fertility in the earliest mission
occupation. However, in the late mission sample, there are very few children and a consider-
able number of older adults, resulting in a juvenility index of 0.7623 – a significant depression
in fertility. It is also during this period that there is a highly elevated morbidity profile (e.g.,
dental caries, cribra orbitalia, porotic hyperostosis) (Larsen, Griffin et al., 2001; Larsen et al.,
2007). Moreover, historic records on population size, living conditions, increased population
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418 Bioarchaeological paleodemography: interpreting age-at-death structures
nucleation, and peak disruption are consistent with the notion that fertility declined. Coupled
with increased mortality and morbidity, it is little wonder that the population in Spanish-
controlled Florida had reduced from hundreds reported to just scores of individuals, all within
a period of a few decades. This outcome of declining fertility per the juvenility index is similar
to the pattern identified in early contact-era Huron ossuary samples (Kleinburg, Uxbridge,
Ossossané) from early seventeenth-century Ontario (Jackes, 1994) and the colonial north
coast of Peru (Klaus & Tam, 2009). While the pattern of colonialism is different in the Spanish
Florida, Spanish Peru, and Ontario settings, the outcome of contact and population collapse is
clearly revealed in estimates showing declining fertility.
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10.7 Summary and conclusions 419
Pubic Symphysis
Actual Dashes and Estimated (Lines) Ages
100
80
60
40
20
Figure 10.8 Age range estimates for modern day skeletons (vertical lines) based on pubic
symphysis standards. The black dashes represent the actual age-at-death of the individual.
(From Milner & Boldsen, 2012c; reproduced with permission of authors and John Wiley &
Sons, Inc.)
A newly emerging method advocates using the large suite of skeletal characteristics that
undergo progressive age changes, and establishing ages of transition over the course of these
changes for each trait, such as for the pubic symphysis (Boldsen et al., 2002; Godde & Hens,
2012; Milner & Boldsen, 2012a, 2012b, 2012c). The distinctions in age for individual skeletal
traits provide minimal information on age (e.g., “young” versus “old”), but as a collective
approach, transition analysis provides excellent results, especially when numerous traits are
employed simultaneously to assess the transitions (Milner & Boldsen, 2012c; Weise et al.,
2009). Validation of the method suggests that this approach extends the limit of more precise
age estimates from 40 years to more than 60 years old, thus providing a more meaningful
demographic reconstruction of age-at-death profiles in archaeological skeletal series (Milner &
Boldsen, 2012c) (Figure 10.8). Use of this method, combined with probabilistic (Bayesian)
approaches to age distribution of the target population independent of the reference popula-
tion, presents new opportunities for improving the accuracy of age estimates in order to
reconstruct and interpret past demographic patterns in a more informed manner (Boldsen
et al., 2002; Godde & Hens, 2012; Hoppa & Vaupel, 2002; Milner & Boldsen, 2012c).
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420 Bioarchaeological paleodemography: interpreting age-at-death structures
population are consistent with the record. Similarly, the growth in population in the American
Midwestern Illinois River valley and elevated birthrates identified in this well-studied region
are consistent with expectations of birthrates in early farming populations that exhibit
increasing settlement size and distribution. There is some evidence to suggest specific patterns
of age compositions in the comparison of catastrophic and attritional death assemblages
(Gowland & Chamberlain, 2005; Margerison & Knüsel, 2002; but see Paine, 2000; Roberts and
Grauer, 2001). However, it falls on the shoulders of the investigator to fully understand the
social, economic, political, behavioral, and taphonomic contexts when interpreting age
structures.
The study of historical skeletal and demographic profiles involving catastrophic mortality
events clearly shows the importance of understanding the circumstances surrounding death
and who is or is not at risk of dying (Grayson, 1990, 1993; Williams, 2009). An example of
this can be seen in the infamous disaster of starvation and death in the Donner Party, a group
of pioneers who in 1846 were stranded in the Sierra Nevada Mountains when snow made it
impossible for them to reach their destination in California. The party of men, women, and
children ran out of food, resulting in the deaths of 35 of 83 individuals due to starvation
(Grayson, 1990, 1993). The demographic record provides important perspective on frailty and
those most vulnerable to death. Modern demographic data would predict that male mortality
would be higher than female mortality. Indeed, that was the case in this setting where the
number of starvation deaths was greater for males than females, especially younger males. In
addition, males died earlier than females in the disaster episode. Especially striking about the
age profiles were the apparent risks associated with not having kin – those without kin died
earlier during the course of the disaster than those with kin (Grayson, 1990, 1993). Thus, the
survival in these difficult circumstances was related to sex, age, and kin group size (and see
Grayson, 1996 for a similar catastrophic context in western North America).
Analysis of the demographic profile of victims of catastrophic flooding following the
Johnstown Flood of 1889 provides a completely different demographic profile than the
starvation deaths in western North America (Williams, 2009). In this setting, deaths occurred
mostly within hours following the bursting of a dam located upriver from Johnstown,
Pennsylvania. Flood waters reached the town virtually without warning, resulting in the
deaths of about 2200 men, women, and children, roughly 10% of the town’s inhabitants.
Analysis of demographic profiles derived from archival records revealed important perspec-
tive on vulnerability in relation to both natural and culturally mediated hazards. In particular,
the rapidly occurring deaths in Johnstown were more heavily weighted toward females than
males, especially younger females (Figure 10.9). Williams (2009) suggests that circumstances
relating to geographical, social, or biological vulnerabilities were likely important in influ-
encing the significant differences between men and women as pertaining to death and injury
in this setting. In particular, the higher mortality rates in women were likely related to more
women being at home than men at the time the floodwaters struck the town and their
considerably bulkier clothing, rendering women less able to maneuver in the flooded setting.
Other factors may also explain the sex differences in deaths. That is, an analysis of a variety
of circumstances involving rapid catastrophic deaths in a range of settings globally reveals
that women (and children) have higher death rates, perhaps due to less endurance and
strength. In addition to these sex differences, there was considerably greater mortality in
children than adults, and especially in individuals less than five years of age. Like most
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10.7 Summary and conclusions 421
35
Victims of the Johnstown Flood
Females
30
Males
All
25
Percentage
20
15
10
0
0−9 10−19 20−29 30−39 40−49 50−59 60−69 70−79 80−89 90+
Age (in years)
Figure 10.9 The age distribution for Johnstown, Pennsylvania flood victims demonstrates
that adult females (20–40 years old) were more vulnerable to the effects of the flood than
their adult male counterparts. (Data from Williams, 2009; reproduced with permission of
author.)
disaster settings, children are more vulnerable owing to their smaller size, and minimal
stamina and strength (Glass et al., 1977; Seaman et al., 1984). These studies of historical
populations underscore the importance of examining the natural, social, cultural, and eco-
nomic contexts in the analysis of demographic profiles. As per Richard Paine’s (2000:182)
advice, “. . .to make robust conclusion from paleodemographic data, we must develop explicit
models of how cultural and biological processes may manifest themselves in the skeletal
record.”
Many caveats surround paleodemographic study. Contrary to earlier assumptions about
manner and rate of aging and senescence, these processes vary both within and across human
populations. Therefore, it is reasonable to state that the ages of reference populations have an
influence on age estimation and thus age structure of target (archaeological) populations.
Moreover, the age-at-death profiles generated – no matter how accurate – are not the same in
composition as those generated for living populations. The developments in what can and
cannot be said about aging and age structures of archaeological skeletal samples and the
impressive advances in theory and method behind their interpretation are cause for welcom-
ing a new and invigorated paleodemography – earlier reports of its death were greatly
exaggerated.
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11.1 Framing the contextual record 423
and sickness in comparison with the larger population from which they are drawn. Moreover,
this does not even take into account the varied cultural and social circumstances that
determined why a person’s remains were interred in the cemetery, be it their age, gender, or
level of wealth or status in complex or otherwise ranked societies. Death assemblages,
therefore, represent a complex composite of individual circumstances that require careful
consideration.
Wood and coworkers evaluated some potential issues of biological selectivity in their
landmark 1992 study (Wood et al., 1992). They argued that documentation and interpretation
of health levels derived from archaeological human remains is neither a straightforward
exercise nor an intuitive process. That is, temporal increases in skeletal lesion prevalence
(e.g., periosteal reactions, enamel defects) in a series may not be a simple measure of declining
community health. The opposite may be the case instead: increasing numbers of infectious
lesions may indicate an improvement in health. Wood and coworkers contend that “the
frequency of active lesions in a skeletal sample is greater than the fraction of affected
individuals in the living population from which the sample was drawn” (1992:349), as
individuals suffering from such afflictions would be at an increased risk of death compared
to the population as a whole. They also contend that individuals within a population having
lesions may be drawn from the biologically advantaged group rather than the disadvantaged
group, as they were able to survive one or more episodes of chronic stress (e.g., disease,
malnutrition). On the one hand, the implications of this “osteological paradox” are important
in that skeletal evidence for declining health in populations undergoing transitions – such as
from foraging to farming – might just as easily be interpreted as representing improvement in
health. On the other hand, Wood and collaborators (1992) did not intend for future gener-
ations of bioarchaeologists to uncritically throw their hands up in the air, and cry “it’s an
osteological paradox – more pathology, better health!” every time they encountered a
population with a high prevalence of lesions. Quite the opposite: the intention is to highlight
the point that bias, selectivity, and scrutiny of the contextual factors underlying an archaeo-
logical skeletal series must be taken into account. Contemporary bioarchaeology has greatly
benefited from their caution.
Therefore, how might such issues of selectivity and bias be best addressed in the study of
archaeological human remains? Bioarchaeologists are not merely counting the lesions on
skeletons and the shapes of skeletal elements in order to characterize variation. Rather, once-
living people resided within a complex web of circumstances involving social relations,
cultural behaviors, settlement systems, environmental conditions, and a range of other
interlinked contexts that we, as anthropologists, routinely take into account. In interpreting
health patterns, Goodman (1993) advocated use of multiple health indicators, rather than
single skeletal indicators. It is the record of these multiple indicators viewed within these
larger contextual domains that provides for a more informed understanding of the bioarch-
aeological record.
Bioarchaeology today is committed to the understanding of the social context and its role
in framing skeletal biology. Social contexts provide an important opportunity for interpreting
the range of variation in the behavioral record discussed throughout this book. The strength
of “social” bioarchaeology is especially well illustrated in the range of investigations
discussed, but one of the theoretical breakthroughs regarding the relationship between
old approaches and new ways of looking at variation as it pertains to biodistance
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424 Bioarchaeology: skeletons in context
research. Anthropologists have had a centuries-old interest in historical issues – Where did a
specific population or society come from? Within a population, who is related to whom? These
are baseline questions, fueling not an inconsiderable interest in what we do by both the public
and scientists alike. Historically, these questions have been asked in the context of classifica-
tion and racial typology. Certainly, new methods of answering questions relating to identity
have been developed and may have their historical origins in the eighteenth and nineteenth
centuries, and result simply in another rendition of old and debunked notions about human
variation (Armelagos & Van Gerven, 2003; Armelagos et al., 1982). A new generation of
bioarchaeologists advocates the view that biodistance analysis offers an important opportun-
ity to address, in compelling ways, socially relevant problems having meaning for anthropol-
ogy in particular and social science in general (Stojanowski, 2013a). There are a number of
reasons why we do biodistance analysis in the twenty-first century (see Chapter 9), but among
the most important is the study of group identity, whether it is based on family, community,
ethnicity, or some other affiliative context. This ethnogenetic approach focuses on diachronic
processes of group emergence or reaffiliation. These processes are expressed synchronically
between individuals and social groups, such as between communities or tribes at any
particular time. These ethnogenetic events occur in many forms, including via intermarriage
between communities, large-scale migration and gene flow, and sometimes, by replacement
of entire populations. As is so well articulated by Stojanowski (2013a:72), “Bioarchaeology,
by virtue of the temporal resolution afforded archaeological assemblages, has the ability to
elucidate both synchronic and diachronic elements of ethnogenesis in past populations – the
former by considering patterns of genetic/phenetic variation against socially defined criteria
of group distinction; the latter by reconstructing patterns of gene flow among populations
through time.”
11.2 Framing the problems and questions: it is all about the hypothesis
Understanding data sets derived from the study of ancient skeletons and their broader
context is only one part of how best to approach the bioarchaeological record. Of equal
importance is the development of hypotheses that serve to frame the research. By construct-
ing research around a hypothesis, bioarchaeologists are able to develop a set of clearly
articulated questions and to generate expectations, to select skeletal series appropriate in
size and composition for the questions asked, to identify or develop the methods –
sometimes appropriately involving very simple methods, but sometimes involving complex
methods and supporting technology – to collect the data, analyze the data, and interpret the
results. Whether we arrive at the responses to our hypotheses through inductive or deductive
approaches, we must ask of our own research or of the research of others: Are the results
consistent with the original expectations that were established at the beginning of the
project, especially taking into account demographic composition, taphonomic history and
biases, and cultural, social, and behavioral factors that enter into who is or is not included in
mortuary assemblages?
There are a number of well-known examples of hypotheses tested in bioarchaeology. One
that has dominated much of my own research agenda pertaining to recent human evolution is
the foraging-to-farming transition, a global event beginning about 10–12 thousand years
ago. Working in a wide range of areas of the world in different geographic and temporal
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11.2 Framing the problems and questions: it is all about the hypothesis 425
scales over the last half century, bioarchaeologists have undertaken independent regionally
based research programs in North and South America, Europe, Asia, and Africa that have
investigated health and behavioral outcomes in populations that underwent the transition
from foraging to farming in earlier prehistory or from less intensified to more intensified
farming in later prehistory (Cohen, 1989; Cohen & Armelagos, 1984; Cohen & Crane-Kramer,
2007; Eshed et al., 2010; Pechenkina & Oxenham, 2013; Pechenkina et al., 2013; Pinhasi &
Stock, 2011; Roberts & Cox, 2003; Steckel & Rose, 2002; Steckel et al., 2002; and see
summaries in Larsen, 1995, 2006; but see Douglas & Pietrusewsky, 2007; Oxenham, 2006;
Tayles et al., 2009 for an alternative perspective). All of this work addresses the hypothesis
that the foraging-to-farming transition led to an elevation in morbidity in particular and a
decline in the quality of life in general. These investigations showed a consistent pattern of
increasing morbidity, namely, higher prevalence of infectious lesions, enamel growth arrest,
reduced growth velocity, and other outcomes that all point to a decline in health. In a highly
consistent manner, these findings meet the expectation of negative health impacts involving a
shift from foraging and collecting to one where diet either shifted to or increased reliance on
domesticated cereals. This expectation is derived from the fact that domesticated cereals have
the following attributes: (1) they are deficient in one or more essential amino acids (e.g., lysine
for maize, millet, and wheat); (2) they lack of sufficient bioavailable iron; (3) they are deficient
in one or more vitamins; for example, B1 (thiamine), B2 (riboflavin), C (ascorbic acid); (4) they
have links with malnutrition, immunosuppression, and susceptibility to a variety of patho-
gens, rendering the individual prone to infection; and (5) they are carbohydrates, creating
circumstances that promote a cariogenic oral environment and dental caries, oral infection,
and susceptibility to degenerative conditions in later life (e.g., cardiovascular disease and
diabetes).
Within this larger question, one area of study that has demonstrated the consistency of
outcomes is skeletal growth. It is well understood that human populations experiencing
elevated stress are generally growth-retarded, with respect to both attainment per age in
juveniles and terminal height in adults (see Chapter 2). Temporal comparisons within arch-
aeological series reveal reduction in growth velocity and ultimately, adult height, and reflect
negative responses to elevated disease, physiological stress, and nutritional insult. Based on
comparison of attainment of growth in archaeological skeletons and in living populations, it
is difficult to tease apart the precise roles of nutrition and genetics, which are the primary
determinants of body size (Bogin, 1999). Thus, based on growth attainment per age, we do not
know precisely what factors are involved in its determination, or, for the purposes of this
discussion, whether growth attainment observed in skeletal series simply reflects a sample of
nonsurvivors who represent mortality bias rather than being an actual indication of stress
(Lovejoy et al., 1990; Saunders & Hoppa, 1993).
Saunders and Hoppa (1993) addressed this conundrum via their comparison of survivors
and nonsurvivors in living populations and their discussion of the implications for past
populations. They observed that nonsurvivors in living populations have higher morbidity
per age than survivors; nonsurvivors may have had high levels of stress, indeed resulting in
shorter height-for-age than survivors. Modeling the magnitude of mortality bias in growth by
projecting various survivor versus nonsurvivor distributions of height-for-age in living
populations shows that survivors are taller for age than nonsurvivors. This is consistent
with expectations, as it is important to realize that nonsurvivors are not selected against
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426 Bioarchaeology: skeletons in context
simply because they are short. Rather, they are selected against because of the suite of
biological stressors (e.g., disease, undernutrition) that contributed to their reduced height-
for-age in the first place. However, simple comparisons of long bone length at various ages
reveal minimal differences, below 3 mm for femur length at 12 years of age (Saunders &
Hoppa, 1993: Table 4). These findings strongly suggest that the biological mortality bias
observed in long bone length of juvenile skeletons from archaeological settings is minimal.
Other factors far outweigh growth differences that might be present between survivors and
nonsurvivors, including bias in age estimation techniques, sample size, and preservation
status (Saunders & Hoppa, 1993).
There are fundamental differences in growth between human populations that are reflected
in variation in developmental patterns of long bones. Comparisons of the prehistoric Libben
series with modern populations from Denver, Colorado, reveal potentially large growth
differences (attainment per age) between homologous skeletal elements (Lovejoy et al.,
1990). In view of the fact that the Libben and Denver populations are from very different
environmental and sociocultural settings, these differences are to be expected. The pattern of
growth during the postnatal period between the two groups is remarkably similar, despite high
stress levels due to elevated systemic infection in the Libben series (Saunders et al., 1995).
These findings concur with Saunders and Hoppa’s (1993) conclusion that mortality bias is not
a significant deterrent to the study of growth in past human groups.
Other comparative analyses of body mass from juvenile remains based on articular dimen-
sions of lower limb bones (Ruff et al., 2013; Sciulli & Blatt, 2008) and midshaft femoral polar
second moments of area (Robbins et al., 2010) also match expectations in comparison with
modern populations. In addition, estimates of body mass and stature in juveniles from
historic-era Arikara from the Sully site, South Dakota, Neolithic Çatalhöyük, Turkey, and
twentieth-century Americans (Denver, Colorado) show similarities and differences that are
important for interpreting variation in health and well-being in these samples. In particular,
Arikara children had slower growth than Çatalhöyük and modern children in earlier age
groups (Ruff et al., 2013). These findings match the record from evidence of relatively poor
health in historic-era Arikara (Jantz & Owsley, 1984a) and relatively better health in Denver
and Çatalhöyük children (Hillson et al., 2013).
My point here is not that the growth record (and health generally) in recent Homo sapiens
either deteriorated or improved with agriculture or that historic-era Arikara were relatively
more worse off than Neolithic Çatalhöyük inhabitants. Rather, the consistency of patterns
observed by various researchers suggests that selectivity has not appreciably hampered
bioarchaeologists’ efforts to evaluate and characterize health in the past. Nevertheless,
researchers must be diligent in their evaluations of skeletal series by drawing on a range
of contextual observations, including cultural and social setting, multiple skeletal and
dental stress indicators, and health status in contemporary settings that might inform our
understanding of past groups (Wood et al., 1992; and see Goodman, 1993, and Wood &
Milner, 1994).
In summary, this book promotes a science-based approach to bioarchaeology. In the
absence of testable hypotheses, studies of skeletons or application of technology, simple or
complex, are mere description. Rather, the construction of bioarchaeological research should
be informed by hypotheses and questions that address fundamental issues about human
behavior viewed broadly.
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11.2 Framing the problems and questions: it is all about the hypothesis 427
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428 Bioarchaeology: skeletons in context
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11.4 Bioarchaeology looking forward 429
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430 Bioarchaeology: skeletons in context
much greater prevalence of S. mutans and lower microbial diversity in modern Europeans
than in early foragers and the first farmers (Adler et al., 2013). The reduced diversity of
microbiota, but including significant S. mutans, complements genomic analysis showing
elevation in particular strains of pathogenic oral microbiota. These important breakthroughs
underscore the relationship between carbohydrate consumption – a cultural event – and the
ecology of the human mouth. Importantly, the discovery reveals the challenges of oral health
in the world today, all within the context of the co-evolution of the human host and it
pathogens.
Other significant directions pertain to the use of experimental data in interpreting skeletal
morphology and its relationship with activity. Pioneered by Sherwood Washburn (1953) as an
element of the “New Physical Anthropology,” experimental approaches provide important
perspectives on skeletal morphology in functional context. As discussed in Chapter 6, there is
a growing understanding that populations pursuing a lifestyle with high levels of mobility
have distinctive signatures of diaphyseal morphology, which involved a general anterio-
posterior elongation of the femoral midshaft diaphysis, which is also revealed in distinctive
ratios of Ix/Iy or Imax/Imin (second moments of area about the x- and y-axes). Ruff and others
have shown that archaeological series comprising individuals having active, mobile lifestyles
have higher Ix/Iy or Imax/Imin ratios than do less active, sedentary populations (see Chapter 6).
Assessment of loading histories via computed tomographic analysis of midshaft tibiae of male
university cross-country runners, field hockey players, and non-athletes revealed that the
runners and hockey players have significantly higher values of areas and second moments of
area (J, %CA, Imax, and Imin) than the non-athletes (Shaw & Stock, 2009b; and see Macdonald
et al., 2009). These and other experimentally based analyses of behavioral outcomes are a
direction from which bioarchaeology will benefit (Schmitt & Churchill, 2003; Teaford & Lytle,
1996).
Similarly, the importance of collaborative study involving ethnographers and bioarch-
aeologists can be especially informative from the perspective of behavior. For example,
ethnobioarchaeological research on modern pygmy groups (Aka, Mbuti, and Efe) in the
Central African Republic revealed remarkable differences in access to food and nutritional
quality in regard to both gender and status, a finding that runs counter to the long-held
assumption that pygmies are emblematic of egalitarian societies (Walker & Hewlett,
1990). Moreover, the ethnobioarchaeological study of dental caries and carbohydrate
consumption showed a clear link between the amount of carbohydrates consumed and
caries rates, but oral hygiene and differences in food preparation practices (e.g., cleaning
of eating utensils) also play an important part in oral health outcomes. Walker and
collaborators’ (1998) ethnobioarchaeological study of diet and dental caries among trad-
itional foragers/agriculturalists – Yanomami (Venezuela), Yora (Peru), and Shiwiar
(Ecuador) – revealed that the relationship between oral health (especially dental caries)
and food consumption practices is complex. For example, the Yanomami have lower
caries rates than the Yora, even though the Yanomami consume less meat than the Yora.
Unlike the Central African pygmies, there are quite low differences between men and
women, despite the more frequent presence of manioc in womens’ mouths as they chew it
for beer production. However, the constant movement of manioc in the mouth and
frequent spitting appear to reduce the cariogenic effects of the food. These investigations
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11.4 Bioarchaeology looking forward 431
are illuminating because they show the complex nature of food consumption and tech-
nology, and by doing so, they provide key insights into behavioral reconstructions. What
is especially important about this work is that it controls for variables observable in the
living that may be applied through carefully constructed analogies to the study of skeletal
remains.
The aforementioned research also speaks to the profound importance of understanding
the inextricable links between biology, culture, and society. A large and impressive record
involving biocultural perspectives on the study of human remains from archaeological
contexts has revolutionized the field, promoting the transition from what was once
largely a descriptive enterprise to one that is informed by social, economic, cultural,
and political circumstances of once-living societies. Importantly, this work on social and
behavioral circumstances in past populations can be used to document the circumstances
of health disparities. Similarly, circumstances resulting in health disparities in the living
form an experimental platform for interpreting the record of past health and well-being
(Goodman et al., 1992; Harrod et al., 2012; May et al., 1993; Walker & Hewlett, 1990;
Walker et al., 1998).
Among the most promising areas of bioarchaeology moving forward is the study of
changing social relations, and especially the essential role that violence has played in past
societies and at all levels, from interpersonal relations to developments as complex as state
formation (various in Martin et al., 2012; Tung, 2012) and colonization of New World
native societies (Klaus, 2012; Klaus et al., 2009; Stojanowski, 2009, 2013b). In addition to
these developing and emerging frontiers, the field is making tremendous strides in moving
away from one that is focused solely on understanding the past to one that is engaged with
and socially relevant to the present. It is doing this by providing a large and robust record
of the history of the human condition, including especially key aspects of health and well-
being.
This is important because it provides a picture of the human experience laying the
foundation for understanding today’s world. That is, the study of the evolution of pathogens
provides answers to questions about key diseases today, including those now re-emerging
(e.g., tuberculosis). It also provides a means for testing hypotheses about health in early life
and its impact on later life. There is a growing record that early life experiences (e.g., low birth
weight and various forms of fetal stress) predict elevated prevalence of degenerative diseases,
such as cardiovascular disease and diabetes in adulthood. In this regard, the bioarchaeological
record provides evidence of morbidity during the period of the growing dentition. Analysis of
enamel defects as a record of growth arrest in response to environmental perturbations
demonstrates a link between early life stresses and increased mortality (Armelagos et al.
2009).
Bioarchaeology holds considerable promise for addressing other issues relating to current
and ongoing concerns about climate change and its role in health outcomes. Clearly,
climate has played an essential role in explaining at least some of the variation in distribution
of specific diseases in the past and today (Drake & Oxenham, 2012; McMichael, 2010;
Patz et al., 2002; Stewart, 1960). Although climate change may not affect health
patterns directly, the alteration in foods available and loss of resources are important elements
for interpreting dietary and health change. For example, bioarchaeological research in the
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432 Bioarchaeology: skeletons in context
American Great Basin and west-central India reveal changing climate that resulted in a
pattern of increasing aridity and abandonment of agriculture. Both settings show clear
patterns of health and lifestyle change associated with major climatic events, to include
altered growth patterns, mobility and workload, and living circumstances overall (Coltrain &
Leavitt, 2002; Coltrain & Stafford, 1999; Robbins Schug, 2011b; Ruff, 1999; Temple, 2008).
There is a clear opportunity in hand to apply bioarchaeological inquiry and understanding of
the fundamental dynamics of the past in a way that powerfully informs the present and
future.
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INDEX
abrasion (tooth) 277 and fertility/birth rate 411–418 alignment of fractures 169
Abu Hureyra (Syria) 205, 280, 295 decline with agricultural Amelia Island (Santa Catalina de
accidental injuries 115–117, intensification 414–418, 417 Guale de Santa Maria,
127–130, 157, 172 juvenility index 413–414, 414, Florida) 70, 235–237,
forearm (may be due to violence) 415, 416 236, 303
122–124, 123, 124, 157 population pyramids 411, 412 ameloblasts 44
humerus 128 infant mortality 330, 405, America see Mesoamerica; North
pre-existing, in soldiers 161–162, 407–408 America; South America
169 life table analysis 406–409 amino acids 16, 317
rib 117–118, 118 mortality profiles 404–406 Anasazi people (USA) 146–148,
treatment 168–171 problems in age estimation 407, 308, 394
vertebral 118–122, 119 409–410, 418–419, 419 anemia 30–31
Admiralty Island (Alaska) 145 stress indicators 54, 65 cranial porotic lesions 30–41, 32,
adolescent growth rates 9–10 underrepresentation of juveniles 34, 317
Africa 407–408, 413 age distribution 34, 39–41, 41
central African tribes 74–75, variability in preservation of elemental analysis of iron
78, 430 skeletal elements 403 352–353
South Africa 264, 287, 323, aggression see violence/violent genetic 31, 33
370–372, 428 injuries Angel, J. Lawrence 404
see also Nubia agriculturalists Anglo-American War (1812) 167
African–Americans accidental injuries 121, 129–130 Anglo-Saxons 12–13, 162
craniofacial morphology 258 bone infections 88–94 animal skins, chewing 84, 188, 287
during enslavement bone remodeling due to ankylosis 181
lead levels 354 mechanical forces 225, 226, apatite 302–304, 338, 355–356
tooth enamel defects 51 228–229 Arabia 343, 344
osteoarthritis 191–192, 196 dentition archaeological context 3–4, 422–424
periosteal reactions 95 caries 69–73, 70 archers 193, 211, 248
age enamel defects 53–54 Arctic peoples
bone mass/loss 57, 58, 63–64, periodontal disease/tooth loss bone remodeling due to
125–126 81–83 mechanical forces 225–227
bone remodeling 241–246, 244, tooth size 26 craniofacial morphology
245 tooth wear 279–282, 284–287, 260–261, 265–268
C and N isotope ratios 330–332 285, 286, 292–297 dentition 266, 287, 290
cranial porosities 34, 39–41, 41 fertility decline 414–418, 417 damaged teeth 297–298, 299
craniofacial morphology 261, nutritional stress 10–11, 16–18, 37 entheseal changes in the arm
268–270 osteoarthritis 180, 192, 194–196 209–210
dentition pastoralists 54, 253, 370–372 Harris lines 43
caries 327 see also cereal diets; foraging-to- osteoarthritis 188, 191, 193, 196
enamel defects 49, 50, 50–51 farming transition; maize osteon structure 63
tooth loss 82 Ainu (Japan) 376 spondylolysis 119
entheseal changes 210 Aka tribe (Africa) 74–75, 78, 430 violent injuries 143–146
Harris lines 44 Alabama (USA) see Koger’s Island; Argentina 263, 266
osteoarthritis 180–181, 196–197 Moundville; Pickwick Basin; Arikara people (USA)
violent injuries 135–136, 139–140, Tombigbee River valley biodistance analysis 385, 398
155 Alaska (USA) 145–146, 209–210, bone remodeling due to
of weaning 323–330, 325, 326, 225–227, 260–261, 380, mechanical forces 237–238
328, 349–351 388–389 nutritional status 11, 11, 59–60,
age-at-death 402–404, 421 Aleutian islands (Alaska) 209–210, 174–175, 426
disaster events 420–421, 421 225–227, 380 violent injuries 136–138
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Arizona (USA) 194, 211, 308, maize consumption 309–310, loading forces 215, 216
350–351 312, 312, 314 mobile lifestyle 250–253, 251,
arm see upper limb Belleville (Ontario) 12–13, 118, 192, 252, 254
arrow injuries 134, 142–143 329 osteoarthritis 232
arthritis see osteoarthritis bending rigidity (I ) 220 sex differences
articular joints see joints bending stresses 215, 217, 218, foraging-to-farming transition
ascorbic acid (vitamin C) 31 219–220 228–229, 232–233, 238,
scurvy 39, 40 Beringa (Peru) 151 240–241, 241
Asia biodistance analysis mobility 223–224, 251, 252, 253
biodistance analysis 373–377, data types and socioeconomic status in the
378, 379 genetic 359–360, 362, Mesolithic 230
dentition 367–368 socioeconomic status 230, 248
caries 71–73 metric 359, 362–363 soldiers 230, 248
tooth loss 82 nonmetric 359, 360, 361–364, and terrain 225, 253
tooth wear 281, 284, 288, 291, 364, 367 birth rate in paleodemography
296 overview 357–362, 358, 423–424 411–418
rice consumption 71, 281 spatial relationships decline with agricultural
violent injuries 159 ethnic boundaries 389–390, 390 intensification 414–418, 417
see also China; Indian interbreeding between juvenility index 413–414, 414,
subcontinent; Japan; Siberia Europeans and native 415, 416
athletes 214, 243, 250 peoples 399–401 population pyramids 411, 412
atlatls 193–194, 210–211 kinship and marriage 390–396, birth weight, low 7–8
attrition (tooth) 277 393, 395, 397 blunt force trauma 143, 168
Australian Aborigines social distinctions 397–399, 400 see also cranial injuries
cranial porosities 36–37 temporal changes: population Boaz, Franz 257
injuries 124, 157–158 movement and cultural bog corpses 160
treatment 169 transition 368 bone density 216
osteoarthritis 203 into America 377–383, 382 bone mass
periosteal reactions 94 within American regions and cortical area 219, 249, 250
repatriation of remains 428 383–386 decreased
tooth wear 287–288 Central Asia 374–375 in adults 57–60, 58, 63–64, 223
Australopithecus spp. 186–187 East and Southeast Asia and fracture risk 125–126
Averbuch (Tennessee) 94, 195–196 375–377, 378, 379 inactivity-related 215
axial loading 219–220 Europe 372–373 in juveniles 13–14
Ayalan (Ecuador) 205 Indus River valley 373–374 bone remodeling see biomechanics
linking living groups to and skeletal morphology
Bab edh-Dhra’ (Jordan) 205, ancestors 388–389 bows and arrows
391–392 Nubia 369–370 injuries inflicted by 134, 142–143
back see vertebrae South Africa 370–372 upper limb changes associated
Bactrian culture 374 Spanish Florida 361, 386–388, with archery 193, 211, 248
Bahamas 337 387, 388 brachycephalic head shape
Ban Chiang (Thailand) 345–346 biomechanics and skeletal morphology 258–259
Bantu farmers 74–75 214–215, 255, 430 Brazil 119, 149, 290–291
Barbados 354 age changes 241–246, 244, 245 breastfeeding 38, 323
barium (Ba) 350–351, 352 agriculturalists 225, 226, 228–229 see also weaning
Barker hypothesis bilateral asymmetry 214, Bronze Age 130, 343, 368, 375
see also developmental origins of 227–231, 231, 248 see also Indus Valley civilization
disease hypothesis cross-sectional geometry brow ridge (supraorbital torus)
Barlowe, Arthur 279 215–222, 218, 219 264–265
basketmaking 289–290 as applied to specific groups bubonic plague 112
battered-child syndrome 174, 177 222–246 burial practices
beer (chicha) 76, 313 external measurements 247–255 prone position 162–163
beheading 133, 149, 160, 162–163, femoral neck-shaft angle 253–255 and status 95, 197
167, 175 foragers 222–227, 223, 225, 251 treatment of juveniles 409
trophy heads 152 foraging-to-farming transition butchery marks 145–148, 147, 149
bejel (endemic syphilis) 96–97 228–229, 231–241, 234,
Belize 236, 239, 241, 247–249 C3 plants, δ13C values 302–303, 303
environmental stress 54–55 histomorphometry 60–64, 61, 62, C4 plants, δ13C values 302–303, 303
family burials 392 246–247 see also maize; millet
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Index 595
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596 Index
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Index 597
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598 Index
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Index 599
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600 Index
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Index 601
Jebel Sahaba (Sudan) 370 leprosy 108–111, 110, 125 Lucy (australopithecine) 186–187
Johnstown Flood (Pennsylvania, mercury treatment 354–355 lumbar spine 119, 195
1889) 420–421, 421 rhinomaxillary syndrome 109,
joints 109 Mac Bac (Vietnam) 72–73
anatomy 179 Levant Magdalenska Gora (Slovenia) 316
kneeling 205, 206 Bronze Age–Iron Age transition Mahalanobis D2 statistic 359
morphology and mechanical 368 Maitas-Chiribaya (Chile) 19, 96
loading 240 craniofacial morphology 264 maize
osteoarthritis see osteoarthritis dentition caries 70–71, 77–78
squatting 204 caries 71 consumption as charted by carbon
Jomon period (Japan) 51, 53, 272, tooth size reduction 273–274 isotope ratios
337, 375–376 tooth wear 280, 281, 295–296 bivariate δ13C/δ15N analysis
Jordan 205, 391–392 endogamy 391–392 333–338, 334, 335
juveniles see children entheseal changes 210–211 comparison with marine diet
foraging-to-farming transition 333–336, 334, 335
kayaking 188, 205, 209, 225–227 210–211, 264, 273–274, 280, in Euroamericans 315–316
Kentucky (USA) 28, 195–196, 269, 349 in Mesoamerica 308–310, 312,
405 metatarso-phalangeal 312–314
Kerma (Sudan) 156 modifications 205 in North America 304,
Khoi pastoralists 370–372 thalassemia 31, 33 304–308, 305, 309, 313
kinship patterns, biodistance see also Çatalhöyük (Turkey) sex differences 313–315, 314
analysis 390–396, 393, 395, Libben site (Ohio) 12, 56, 127, 169 socioeconomic status 306, 312,
397 life expectancy 402 313–315
kneeling 205, 206 see also age-at-death in South America 310–311,
Kodiak Island (Alaska) 145–146, life tables 406–409 311, 313
388–389 lingual surface attrition of the nutritional stress 16–18
Koger’s Island (Alabama) 138–139, maxillary anterior teeth and anemia 35, 37
272 290–292, 291 and bone remodelling 60–63
Kulubnarti (Sudan) 34–35, 156–157, linguistics, Great Basin area 383 and lower bone mass 59–60
243 Little Bighorn, battle (1876) 167–168 processing
Little Ice Age 346 degenerative joint conditions
La Chapelle-aux-Saints (France) loading forces on bone 215, 216 194, 205
187–188 London (UK) effect on nutritional value 16,
La Florida (Ecuador) 313 industrial era 35
La Florida (North America) osteoarthritis 180 and tooth wear 279
see Florida (USA), Spanish era rickets 21 upper limb bone remodeling
La Plata Valley (New Mexico) 95, spondylolysis 120 228, 232–233, 238
148 weaning 327–329, 328 protein/cereal balance 74–75,
La Real (Peru) 151 Medieval 24–25, 85–86, 230 313–316, 314, 330–332,
Lake Baikal region (Russia) 203, 210, lower limb 332, 350–351
282, 338 bone remodeling due to malaria 33, 112
Lamanai (Belize) 54–55, 309–310, mechanical forces 223–224, males
312, 314 225 bone remodeling due to
Lambayeque Valley (Peru) age-related changes 241–246, mechanical forces
colonial period 13, 37, 176–177, 244, 245 foraging-to-farming transition
201 asymmetry 230–231 228–229, 232–233, 240–241,
Sicán culture 96, 152, 153, 197, inactivity 215 241
398–399, 400 mobility 250–253, 251, 252, mobility 223–224, 251, 252,
Lapa do Santo (Brazil) 149 254 253
Larson Village (South Dakota) temporal trends 231–241, 234, and socioeconomic status 230
137–138 236, 239, 247–249 death due to starvation 420
latte (megaliths) 121 diaphyseal bowing (rickets) 21–23 dentition
Lawson, John 101–102 diaphyseal flattening 23–24, 249 caries 73–77
lead (Pb) 353–354, 355 femoral neck-shaft angle 253–255 periodontal disease 83–85, 84
Leavenworth (South Dakota) 398 kneeling 205, 206 tooth wear 287–290
leg see lower limb periosteal reactions 88–89, 97, 98 diet
leguminous plants 320 squatting 204 caries 74–76
Lengua people (Paraguay) 29 sword cuts 161 entheseal changes 209–211
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602 Index
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Index 603
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604 Index
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Index 605
Oxus culture (central Asia) 374 diet 310–311, 311, 336 postmarital residence 394,
oxygen (O) isotopes 346–347 kinship in a Sicán burial 398–399, 395
400 bone histomorphometry
Pacatnamú (Peru) 150, 150 osteoarthritis 193, 197, 201 246–247
Pacbitun (Belize) 310, 312, 312 periosteal reactions 95–96 cranial porosities 35
Pacific islands see Easter Island; Fiji; socioeconomic status 47–48, 78, 96, diet 63, 308, 350–351
Hawaii; Marianas Islands 197 entheseal changes in the arm
paleodemography 402–404, 421 Tiwanaku state 341–342 211
disaster events 420–421, 421 trephination 169–170, 170 osteoarthritis 191
and fertility/birth rate 411–418 tuberculosis 104–105, 107–108 paleodemography 404
decline with agricultural violent injuries 123, 150, violent injuries 146–148, 166
intensification 414–418, 417 150–152, 151, 153, 154–155 pygmy forager tribes (Aka, Mbuti,
juvenility index in the Pete Klunk mound (Illinois) 197, Efe) 74–75, 78, 430
foraging-to-farming 396, 397, 407
transition 413–414, 414, pewter 354 racial groups, craniofacial
415, 416, 416 photosynthetic pathways morphology not a good
population pyramids 411, 412 (δ13C values) 302–303, 303 guide 257–259
infant mortality 405, 407–408 Phum Snay (Cambodia) 159 radius
life table analysis 406–409 physical activity Colles’ fractures 122, 124
mortality profiles 404–406 and bone mass 59 entheseal changes 207
problems entheseal changes 206–212, 207 Rapa Nui (Easter Island) 155–156,
age estimation 407, 409–410, and female fertility 178 399–400
418–419, 419 foragers 178 reduction of fractures 169
excavation techniques 408–409 osteoarthritis see osteoarthritis relatedness, analysis see biodistance
underrepresentation of and remodeling of bone analysis
juveniles 407–408, 413 see biomechanics and remodeling of bone
variability in preservation of skeletal morphology due to mechanical forces
skeletal elements 403 rib fractures 117–118 see biomechanics and
timing of weaning 323, 330, 408 spondylolysis 118–122, 119 skeletal morphology
Panama 309 phytates 31, 34–35 histomorphometry 60–64, 61, 62,
Paraguay 29 Pickwick Basin (Alabama) 193, 207, 246–247
parasitic infestations 34, 37 229, 231–233, 272 repatriation of human remains
parry fractures of the forearm pinta 96–97 388–389, 428–429
122–124, 123 plague 112 residential change 338–347, 340,
pastoralism 54, 253, 370–372 Point Hope (Alaska) 260–261 344, 345
Patagonia (South America) 266 Poland 332, 332 rhinomaxillary syndrome (leprosy)
Pecos Pueblo (New Mexico) 166, poor house inhabitants 248 109, 109
191, 246–247, 308, 404 population density rib lesions
pelvic shape/deformity 21 and spread of disease 89–90, 93–94 fractures 117–118, 118
percent cortical area index (PCCA) and violence 172–173 tuberculosis 103, 105, 107
219, 249 see also urbanization rice 71, 281
perikymata 45, 47, 52 population pyramids 411, 412 rickets 21–23, 23
periodontal disease and lost teeth porotic hyperostosis 30–41, 32 riding 194, 238
78–81, 79, 82 Portugal 72, 280, 284–287, 320 rigidity of bone 218–220, 219
in pregnancy 76 posture ritualized violence 148–154, 150,
sex differences 83–85, 84 kneeling 205, 206 153, 177
socioeconomic status 84–85 squatting 204 Riviere aux Vase (Michigan)
and systemic health 85–86 Pott’s disease 103, 105 135–136
temporal trends in prevalence pregnancy Rodeo athletes 128
81–83 dentition 76–77 Roman Empire
periostitis/periosteal reactions 12, nitrogen balance 333 decapitation 162
86, 86–96 prevalence, definition 66 diet 330–331
Peru principal component analysis 359 fractures 60, 169
colonial period projectile injuries lead levels 354, 355
environmental stress 13, 37 arrows (or spears) 134, 142–143, 160 migration 345
osteoarthritis 201 firearms 155, 164, 167–168 weaning 51, 325–327
periosteal reactions 95 Pueblo Bonito (Florida) 394, 395 Ruff’s mobility index 250–253, 251,
structural violence 176–177 Puebloan societies 252, 254
dentition 28, 47–48, 78, 82 biodistance analysis 384 Russia see Siberia
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606 Index
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Index 607
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608 Index
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