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Ortner’s Identification of Pathological Conditions
in Human Skeletal Remains
Ortner’s Identification of
Pathological Conditions in
Human Skeletal Remains

Third Edition

Edited by

Jane E. Buikstra
Arizona State University, Tempe, AZ, United States
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List of Contributors
Amanda M. Agnew, School of Health and
Rehabilitation Sciences, The Ohio State University,
Columbus, OH, United States
Megan B. Brickley, Department of Anthropology,
McMaster University, Hamilton, ON, Canada
Jane E. Buikstra, Arizona State University, Tempe, AZ,
United States; Arizona State University, Phoenix, AZ,
United States
Morgana Camacho, Pathoecology Laboratory, School of
Natural Resources, University of Nebraska - Lincoln,
United States
Mary E. Cole, Department of Anthropology, The Ohio
State University, Columbus, OH, United States
Sharon DeWitte, University of South Carolina, SC,
United States
Bruno Frohlich, Department of Anthropology,
Smithsonian Institution, Washington, DC, United
States; Department of Anthropology, Dartmouth
College, Hanover, NH, United States
Anne L. Grauer, Loyola University Chicago, Chicago,
IL, United States
Rebecca Kinaston, Department of Anatomy, School of
Biomedical Sciences, University of Otago, Dunedin,
New Zealand
Haagen D. Klaus, Department of Sociology and
Anthropology, George Mason University, Fairfax, VA,
United States
Mary Lewis, University of Reading, Reading, United
Kingdom
Niels Lynnerup, Department of Forensic Medicine,
University of Copenhagen, Copenhagen, Denmark
Carina Marques, Research Centre for Anthropology and
Health (CIAS), Department of Life Sciences,
University of Coimbra, Coimbra, Portugal; Department
of Anthropology, William Paterson University, Wayne,
NJ, United States
Simon Mays, Historic England, Portsmouth, United
Kingdom
Justyna J. Miszkiewicz, School of Archaeology &
Anthropology, Australian National University,
Canberra, ACT, Australia
Marc F. Oxenham, School of Archaeology &
Anthropology, Australian National University,
Canberra, ACT, Australia
Andrew T. Ozga, Center for Evolution and Medicine,
Tempe, AZ, United States; Institute for Human
Origins, Tempe, AZ, United States
Rebecca Redfern, Centre for Human Bioarchaeology,
Museum of London, London, United Kingdom
Karl Reinhard, Pathoecology Laboratory, School of
Natural Resources, University of Nebraska - Lincoln,
United States
Charlotte A. Roberts, Department of Archaeology,
Durham University, Durham, United Kingdom
Anne C. Stone, School of Human Evolution and
Social Change, Tempe, AZ, United States; Center
for Evolution and Medicine, Tempe, AZ, United
States; Institute for Human Origins, Tempe, AZ,
United States
Samuel D. Stout, Department of Anthropology,
The Ohio State University, Columbus, OH, United
States
Richard Thomas, School of Archaeology and Ancient
History, University of Leicester, Leicester, United
Kingdom
Monica Tromp, Department of Anatomy, School of
Biomedical Sciences, University of Otago, Dunedin,
New Zealand; Department of Archaeology, Max
Planck Institute for the Science of Human History,
Jena, Germany
Chiara Villa, Department of Forensic Medicine,
University of Copenhagen, Copenhagen, Denmark
Tony Waldron, University College London, London,
United Kingdom
Anna Willis, College of Arts, Society & Education,
James Cook University, Townsville, QLD, Australia
xiii
Preface
One of the last times I saw Don Ortner in his office at the
Department of Anthropology of the Natural History
Museum, he gestured to the shelves and filing cabinets
where he had been beginning to accumulate sources for
the third edition of Identification of Pathological
Conditions in Human Skeletal Remains. As we who
mourn him know only too well, he died unexpectedly on
April 29, 2012, and this task remained undone. Hoping
that at least a partial manuscript existed, I asked Bruno
Frohlich, a close colleague of Don’s at the National
Museum of Natural History, about evidence of the
volume’s progress. As the person who assumed the chal-
lenging task of sorting Don’s office, Bruno indicated that
there was nothing of substance, no outline, no negotia-
tions with a press.
Thus, it was obvious that organizing a new edition
would require starting with the Ortner (2003) volume and
revising. The alternative, letting the fine second edition
become increasingly out of date, a piece of history but
not a useful teaching and research aid, seemed an
unhappy choice. New volumes by other authors would no
doubt appear, but in my opinion that energy and expertise
could be better directed toward advancing knowledge in
other ways rather than “reinventing the wheel.”
Following discussions with colleagues in paleopathol-
ogy and the Ortner family, primarily Don’s widow Joyce
and son, Don, Jr., I agreed to explore publication options
and consider how the volume might be revised to reflect
new knowledge and the further integration of the study of
health into perspectives on the past. After discussions
with several presses, it seemed prudent to choose
Elsevier, as they could readily provide the text and image
files from the second edition. I wish to thank them most
sincerely for their support and patience throughout this
protracted process.
There have been many decisions along the way.
Initially, and with sage advice from many colleagues,
such as Anne Grauer and Charlotte Roberts, I generated a
proposal for Elsevier, which included an outline of the
volume, as it appears here. Recognizing that a collabora-
tive effort would be needed to update the core chapters on
pathological conditions, specialists in the paleopathology
of specific conditions were invited to take Don’s (and
Walter Putschar’s) chapters and rework them to reflect
new knowledge. Each invited author accepted, which is a
measure of their professionalism and their respect for
Don.
In creating the current organization, I have deleted
information about basic osteological methods, such as esti-
mating age-at-death and biological sex. These are now
covered in much greater detail in a variety of basic and
advanced osteology texts. We have introduced distinctive
chapters on normal and abnormal bone development,
imaging, radiology, and ancient pathogen DNA and micro-
biomes. The chapter on dental disease now also includes
biochemical methods for estimating diet (paleodiet). In
some cases, conditions have been shuffled between chap-
ters, their realignment reflecting contemporary thought.
Faced with deciding whether to follow my vision of
paleopathology in the 21st century or to attempt to guess
what Don might have wanted 15 years after the previous
edition and 6 years after his death, I have chosen the for-
mer. In reflecting upon the many stimulating and open
discussions that Don and I have had about the field, I am
convinced that he would approve. I have therefore de-
emphasized “classification” in the diagnostic process, and
I have added a chapter that recognizes social theoretical
approaches to interpreting pathological conditions. In
addition, attempting to recognize related specialties, chap-
ters on mummy science and animal paleopathology have
also been added. It is my personal view that the 21st cen-
tury will witness remarkable new knowledge of disease
histories and disease transmission that unites the study of
zoonotic and human infections, facilitated by molecular
studies. The biomolecular “revolution,” however, will
continue to complement and augment our studies of
human remains, which will continue to be as fundamen-
tally important to the study of ancient disease as Don and
Walter recognized in their 1981 volume.
A final word should be added about authorship.
Several chapter authors asked that Don be included as a
co-author, as I also felt appropriate for the volume as a
whole. As there are prohibitions against attributing post-
humous authorship, I decided to follow the biomedical
xv
xvi Preface

model and entitle this volume Ortner’s Identification of
Pathological Conditions in Human Skeletal Remains, 3rd
edition.
So here it is! It wouldn’t have been possible without
Don’s (and Walter’s) exemplary prototype, as well as the
many colleagues who so willingly contributed revisions
and original chapters. I sincerely hope that you find it
useful in your research and teaching, as you advance the
vibrant field of paleopathology during the 21st century.
Jane E. Buikstra
Arizona State University
A Tribute to Don Ortner
It is a great honor to be asked to write this tribute for the
third edition of Don’s Identification of Pathological
Conditions in Human Skeletal Remains. Charlotte had
attended Don’s 1985 Short Course in Paleopathology at
the Smithsonian Institution, Washington, DC (the fifth
and final one there), and we both met Don at the
Paleopathology Association European Meeting in Madrid
in 1986. He indicated that he was “looking for a hook on
which to hang his hat” in Europe and do research and
teaching. We proposed the University of Bradford, and he
accepted the University’s invitation to be an Honorary
Visiting Professor. Thus began a long and enduring rela-
tionship and collaboration with the Smithsonian
Institution, and a long and close friendship between the
Ortners and the Manchesters, and Charlotte and family.
This friendship has endured to the present, long past
Don’s untimely tragic death, and is exemplified by the
endearing label applied to Don, with typical Yorkshire
bluntness, by Keith’s wife’s aunt: “the Big Bug from
America.”
Research collaborations at the University developed,
especially in tuberculosis and leprosy, and in 1988 the
first (Bradford) Short Course in Paleopathology was run.
It ran seven times, with the final one in 2008. Don’s
involvement at Bradford had continued for several years
by then, and during that time he and his wife Joyce had
explored much of the county of Yorkshire, and become
the owners of “Yorkshire passports”!
Don started his career with an undergraduate degree
(BA) in Zoology with a minor in chemistry. This is interest-
ing when compared to the late Don Brothwell, whose first
degree was a BSc in Anthropology and Archaeology
(including zoology and geology), and the fact that Don
Ortner was inspired by a primatologist to move into looking
at past disease. A Masters degree in Anthropology followed
at Syracuse University, where he also did the physical
anthropology course taught by Gordon Bowles, who had
studied under EA Hooton. He then completed a PhD in
1969 at the University of Kansas (the effects of ageing and
disease on the micromorphology of human compact bone).
He worked for some time as a Museum Technician in the
Department of Anthropology at the National Museum of
Natural History at the Smithsonian Institution, Washington,
DC (with JL Angel and TD Stewart), and then was
recruited as an Assistant Curator in 1969, becoming
Curator of Physical Anthropology in 1976.
Following a strongly influential meeting with Adolph
Schultz at the University of Zurich in Switzerland who
studied wild shot primate pathology, he was convinced
that “paleopathology could make a valuable contribution
to science if the research was founded on a thorough
knowledge of anatomy, physiology and the mechanisms
of disease processes” (Powell, 2012: 91). The rest is his-
tory. This set the stage for the rest of his career. His meet-
ing with pathologist Walter Putschar led to Ortner and
Putschar (1981) and Don’s considerable work for that first
edition benefited from his experience working with
pathology reference collections in European museums.
Highlighting these collections as beneficial to understand-
ing how disease processes affect bones has led to much
more work on documented skeletal collections in paleo-
pathological research. Don was deeply involved with
paleopathology at many levels, including service to the
field, and he headed up the Paleopathology Association
(PPA) as President from 1999 to 2001. In terms of
research, Don has contributed much to the literature
beyond his books. He was particularly proud of his
achievements in developing diagnostic criteria for scurvy
and rickets, and documenting the effect of the early stages
of leprosy on the facial bones. He was open to debates in
paleopathology, and welcomed interactions with younger
scholars where he could help. He was always willing to
talk to anybody about paleopathology, young, old, ama-
teur or highly experienced.
In particular, we would like to emphasize Don’s com-
mitment to research-led education in paleopathology, epit-
omized by many activities. Three are prominent. Firstly,
the hugely successful short courses in paleopathology
with a worldwide participation helped many “graduates”
along the road to successful careers, including one of the
authors (Ortner et al., 2012). Secondly, these courses ran
alongside the many workshops in paleopathology Don led
at the annual meetings of the PPA, starting in 1985, and
gave people the opportunity to engage with different path-
ological conditions at theoretical and practical levels.
There is no underestimating the time Don (and his compa-
triot Bruce Ragsdale, a pathologist) spent putting the
workshops together. They remain a legacy for PPA
xvii
xviii A Tribute to Don Ortner
meetings today. Thirdly, this volume has become the
mainstay for scholars working in paleopathology.
The first edition of this book had been published in
1981, well before his link with Bradford began (“Don’s
Bible”). This marked a turning point in the global evolution
and development of paleopathology. Previous publications,
whilst important in establishing paleopathology as a disci-
pline and documenting global evidence for disease in antiq-
uity, lacked the scientific and clinical rigor of Don’s book
in elucidating diagnostic and differential diagnostic paleo-
pathological criteria for different diseases. The second edi-
tion was produced in the prime years of his involvement at
Bradford (Ortner, 2003). In that edition Don wrote 20 of the
23 chapters; authoring virtually all the chapters was no
mean achievement. These two editions had focused on a
classificatory system of disease, whilst incorporating and
integrating clinical and epidemiological aspects.
His writings on the basic biology of bone, on patho-
logical processes, and on clinical and scientific methodol-
ogy create a baseline for this third edition which, whilst
maintaining a classificatory base, has diversified and
expanded into broader aspects and concepts of paleopa-
thology. This appropriately includes methodological
developments. We believe that this edition is a just and
fitting tribute to Don’s immense and unequaled contribu-
tion to the totality of paleopathology, making it an
accepted and important component of anthropology,
archeology, and clinical medicine. The chapters of the
current edition, by necessity, have been reworked by a
range of authors from both the Old and New Worlds, but
the work Don put into the chapters of the previous
volumes provided a very strong base with which the new
chapter authors could work. We are sure that Don would
have been incredibly pleased to see this new edition and
the developments the volume has taken, and happy to see
Jane head it up.
This new edition of Don’s seminal work in paleopa-
thology will clearly take us well into the 21st century and
set the stage for research and teaching in this field. In so
doing, it takes into account developments in the field over
the last 15 years, showing particularly how nonhuman
paleopathology, paleoparasitology, and biomolecular anal-
yses have an increasing part to play in the reconstruction
of the origin, evolution, and history of disease. It also
illustrates that paleopathology is rapidly progressing as a
multimethod-driven discipline fit for the future, and one
that embraces other disciplines across the arts, humanities,
social sciences, and sciences.
Charlotte Roberts and Keith Manchester
REFERENCES
Ortner, D.J., 2003. Identification of Pathological Conditions in Human
Skeletal Remains, second ed. Smithsonian Institution Press,
Washington, DC.
Ortner, D.J., Putschar, W.G.J., 1981. Identification of Pathological
Conditions in Human Skeletal Remains. Smithsonian Institution
Press, Washington, DC.
Ortner, D.J., Knüsel, C., Roberts, C.A., 2012. Special courses in human
skeletal paleopathology. In: Buikstra, J.E., Roberts, C.A. (Eds.), The
Global History of Paleopathology. Pioneers and Prospects.
University Press, Oxford, pp. 684 693.
Powell, M.L., 2012. Donald J. Ortner. In: Buikstra, J.E., Roberts, C.A.
(Eds.), The Global History of Paleopathology. Pioneers and
Prospects. University Press, Oxford, pp. 89 96.
Chapter 1
Introduction
Jane E. Buikstra
Arizona State University, Tempe, AZ, United States
This third edition of the Identification of Pathological
Conditions in Human Skeletal Remains updates and
expands upon the topical coverage of earlier works pub-
lished by Ortner and Putschar (1981) and Ortner (2003).
In this chapter, we develop a “roadmap” for the structure
and organization of this volume. First, we present the his-
tory of this landmark volume from the perspectives of
Donald J. Ortner (first and second editions) and Jane E.
Buikstra (third edition). In these sections, and elsewhere,
our goals have included retaining Don’s voice, so there
are many portions of the second edition that are retained
throughout the volume. We also acknowledge those indi-
viduals and institutions who have contributed to its devel-
opment over the past 30 1 years. We then introduce the
objectives for this third edition, outlining those chapters
that have been reorganized as well as those chapters that
have been added to this edition, which cover a new range
of related fields integral to the development of 21st cen-
tury paleopathology. Finally, we will introduce and
review the format of the volume and its organization.
HISTORY OF THE FIRST EDITION FROM
DONALD J. ORTNER
The first edition of this book was the result of a joint col-
laboration between Dr. Walter G. J. Putschar and me. Dr.
Putschar was an internationally known, consultant pathol-
ogist at Massachusetts General Hospital in Boston, MA,
who had a special interest in diseases of the human skele-
ton. We began our professional relationship in 1970 when
he accepted my invitation to be the principal lecturer in a
seminar series on human skeletal paleopathology that I
was organizing at the Smithsonian Institution. The first
Paleopathology Seminar Series was held in 1971 and
brought several leading authorities on skeletal disease,
paleopathology, and related subjects to the Smithsonian
Institution to present a series of lectures to a select group
of scholars interested in skeletal paleopathology.
Ortner’s Identification of Pathological Conditions in Human Skeletal Remains. DOI: https://doi.org/10.1016/B978-0-12-809738-0.00001-6
© 2019 Elsevier Inc. All rights reserved.
The seminar series was held yearly through 1974. By
that time the logistics of obtaining funds to offer the
series, arranging for students to come from many univer-
sities, including those outside the United States, and
assembling an outstanding faculty for the 10-week series
of lectures and laboratory sessions raised serious ques-
tions about whether this was the most cost-effective
method for enhancing the quality and direction of
research in skeletal paleopathology. It also highlighted the
need for a comprehensive reference work on diseases of
the skeleton that might be encountered in archeological
skeletal remains. I discussed this issue with Dr. Putschar
and we decided that many more scholars interested in
skeletal paleopathology would have access to the sub-
stance of the seminar series if the information in the lec-
tures and laboratory sessions was incorporated into a
well-illustrated and comprehensive reference work on
pathological conditions that affect the human skeleton.
In the summer of 1974, with the support of a grant
from the Smithsonian Research Foundation (now the
Smithsonian Scholarly Studies Program), Dr. Putschar
and I, accompanied by our wives, Florence Putschar and
Joyce E. Ortner, and my three children, traveled exten-
sively in Great Britain and several European countries for
more than three months visiting educational and research
centers that had significant collections of documented
human skeletal pathology. In selecting these centers, we
leaned heavily on the advice of the late Dr. Cecil J.
Hackett, a physician who had worked for several years in
Uganda where he had treated hundreds of patients suffer-
ing from yaws. This experience led to a research interest
in treponematosis, and Dr. Hackett wrote his doctoral dis-
sertation on the clinical, radiological, and anatomical
manifestations of yaws (Hackett, 1947). Following his
career in Uganda, Dr. Hackett settled in England where
he continued his research on treponematosis, its history
and skeletal manifestations. As part of this research he
visited many of the major European collections of ana-
tomical pathology that contained documented cases of
1
2 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
syphilis. Hackett’s research on these cases resulted in the
publication of his classic monograph (Hackett, 1976) on
the skeletal manifestations of syphilis, yaws, and trepo-
narid (bejel). His knowledge of these collections and
which ones were likely to serve the objectives Dr.
Putschar and I had set out to achieve was an invaluable
asset.
During our visit to these institutions, Dr. Putschar and
I studied and photographed hundreds of cases of skeletal
disease. In addition to the photographic record we made
of these cases, we often were able to obtain autopsy or
museum records that provided descriptive details and a
diagnosis for the cases. Radiographic films were acquired
for some of the cases. Dr. Putschar dictated his observa-
tions about each case and these observations were subse-
quently transcribed and organized by Mrs. Putschar. In
some cases, Dr. Putschar’s diagnostic opinions were at
variance with the diagnosis given in the catalog and this
difference was duly noted in his observations. Most often,
however, the diagnosis given in the catalogs was plausible
if not reasonably certain.
We began the task of writing the book shortly after
completing our European research in 1974. In 1979, we
submitted the completed manuscript to the Smithsonian
Institution Press for publication as part of the Smithsonian
Contributions to Anthropology series. The manuscript
was reviewed by the Department of Anthropology, exter-
nal reviewers, the Director’s office of the National
Museum of Natural History, and the Press. After approval
on all levels, editing and production took an additional
several months and the book was published in December
of 1981 as Smithsonian Contributions to Anthropology,
Number 28. A hard-cover edition was published in 1985
that was identical to the first edition except for the addi-
tion of an index.
Acknowledgments for the First Edition
The initial research conducted for the first edition of this
book was an extensive survey in 1974 by Dr. Putschar
and me of documented skeletal pathology in 16 European
pathology and anthropology collections in six countries.
This survey was supported by the Smithsonian Research
Foundation and Hrdlička Fund. The following list of these
institutions and the staff members who assisted our survey
of their collections is inadequate recognition of the many
courtesies extended during our work. Sadly, many collea-
gues who provided this assistance have since retired or
died. Furthermore, some of the collections have been
moved from the site where we studied them and some
probably no longer exist. However, it remains appropriate
to acknowledge the contribution they have made to both
editions of this book. Austria: Federal Pathologic-
Anatomy Museum, Vienna (Dr. Karl von Portele and Dr.
Alexander Müller); Pathology Museum of the University
of Graz (Prof. Dr. Max Ratzenhofer); Pathology Museum
of the University of Innsbruck (Prof. Dr. Albert Probst
and Prof. Dr. Josef Thurner, Salzburg, Austria).
Czechoslovakia: National Museum, Department of
Anthropology, Prague (Dr. Emanuel Vičk, Dr. Milan
Sfloukal and Dr. H. Hanākovā). England: The Natural
History Museum, London (Dr. Theya Molleson and
Rosemary Powers); Guy’s Hospital Medical School,
Gordon Pathology Museum, London; The Royal College
of Surgeons of England, Wellcome Museum, London (Dr.
Martin S. Israel); The Royal College of Surgeons of
England, Hunterian Museum, London (Elizabeth Allen);
St. George’s Hospital Medical School, Pathology
Museum, London; Westminster Hospital School of
Medicine, Pathology Museum, London. France: (Prof. Y.
Le Gal and Prof. Andrè Batzenchlager). Scotland: The
Royal College of Surgeons of Edinburgh (Prof. Eric C.
Mekie, Dr. Andrew A. Shivas, Violette Tansy, Turner,
McKenzy). Switzerland: Anthropological Institute of the
University of Zurich (Dr. Wolfgang Scheffrahn);
Historical Museum, Chur (Dr. H. Erb); Institute of
Pathological Anatomy of the University of Zurich (Prof.
Dr. Erwin Uehlinger, Prof. Dr. Christoph E. Hedinger,
and Aschwanden); Natural History Museum, Bern (Prof.
Dr. Walter Huber). Dr. Cecil J. Hackett, an associate of
the Royal Orthopaedic Hospital, did much to expedite our
work in London, England, and offered several helpful
suggestions regarding collections in other countries that
proved valuable to our study.
The product of this 1974 survey was more than 1200
photographs, both black and white and color (taken by
me) of approximately 500 pathological specimens jointly
studied. For some cases, we were able to obtain x-ray
films as well. Dr. Putschar described the specimens in
detail on tape, and included original autopsy and clinical
data where available. This collection of photographs,
radiographs, and the transcripts of case descriptions is
available for study at the Department of Anthropology,
National Museum of Natural History, Smithsonian
Institution, Washington, DC. Many of them are used as
illustrations in this book.
A number of people made significant contributions
during the preparation of the manuscript. Paula Cardwell,
Elenor Haley, and particularly Katharine Holland typed
initial drafts. Marguerite (Monihan) Guthrie and Elizabeth
Beard typed the final draft. Marcia Bakry prepared some
of the drawings. A special note of appreciation goes to
Jacqui Schulz for the many unpaid hours spent preparing
the remaining drawings and getting the photographic
illustrations ready for publication. Photographic enlarge-
ments were prepared by H.E. Daugherty and Agnes I.
Stix. Stix also assisted in editing and typing the manu-
script. David Yong, Edward Garner, and Dwight Schmidt

provided valuable technical assistance. The staff of the
library of the Smithsonian Institution, particularly Janette
Saquet, was most helpful. Dr. J. Lawrence Angel, Dr. T.
Dale Stewart, and Dr. Douglas H. Ubelaker, members of
the Department of Anthropology, Smithsonian Institution,
have made valuable suggestions, as have Dr. Saul Jarcho
(New York City) and Dr. George Armelagos (University
of Massachusetts, Amherst, MA). The staff of the
Smithsonian Institution Press, particularly Albert L.
Ruffin, Jr., managing editor, series publications, and, Joan
B. Horn, senior editor, deserve special recognition for
their assistance from the conceptualization through publi-
cation of the book. Finally, the wives of both authors
have been intimately involved with the preparation of the
book. Florence Putschar spent hundreds of volunteer
hours organizing photographs, typing, preparing the bibli-
ography, editing, and otherwise making her remarkable
abilities available to the project. Joyce Ortner has also
assisted in obtaining illustrative material and skeletal
specimens.
HISTORY OF THE SECOND EDITION
FROM DONALD J. ORTNER
Since Dr. Putschar and I completed the manuscript for the
first edition, much has changed in the study of ancient
skeletal diseases. The Paleopathology Association, estab-
lished in 1973 with fewer than two dozen members, is
now a thriving international scientific association with
more than 600 members worldwide that holds annual
meetings in the United States and biennial meetings in
Europe. There is now a scientific journal devoted to
paleopathology1 and another new journal in which this
subject is an important emphasis. A bibliography of
paleopathology (both the published edition and the sup-
plements) contains more than 26,000 citations, many of
which were published in the last 20 years (Tyson, 1997).
My own research interest and experience has devel-
oped as well. In 1984 I received a 3-year grant from the
National Institutes of Health (NIH; grant AR 34250) to
conduct a survey of pathological cases in the human skel-
etal collections at the National Museum of Natural
History (NMNH). This survey was superimposed on a
major effort by the Museum to create an electronic data
base of our catalog that required that the anthropological
collections be inventoried. Several people were involved
in this inventory, but three members of the technical staff
deserve particular mention: Marguerite (Monihan)
Guthrie, who typed much of the manuscript of the first
edition of this book, was responsible for creating, editing,
1. Refers to the Journal of Paleopathology, founded by Luigi Capasso,
which has been published by the Abruzzo Anthropological Association
since 1987.
Introduction Chapter | 1 3
and maintaining the data base. Dwight Schmidt and
Stephen Hunter were responsible for doing the actual
inventory of the human skeletal collection. This inventory
required that all human remains in the collection be com-
pared with the catalog record to ensure that the skeleton had
been cataloged and that the catalog record was accurate.
This meant opening thousands of drawers and handling
more than 36,000 partial to complete human skeletons.
While they were engaged in this task, Schmidt and
Hunter were encouraged to identify any cases of skeletal
pathology and bring them to my attention. Both Schmidt
and Hunter were enthusiastic and highly motivated. They
became skilled at identifying pathological cases and this
added immeasurably to the quality and quantity of archeo-
logical and anatomical cases of skeletal disease in the
human skeletal collection of the NMNH. One of the frus-
trating aspects of the research Dr. Putschar and I had con-
ducted on the NMNH pathological materials was the lack
of accessible and reliable information on the archeologi-
cal dating of the human remains. The grant from NIH pro-
vided funding to hire an archeologist, Dr. James Krakker,
to review the archeological field records and publications
to determine as accurately as possible the archeological
dates for much of the human skeletal collection.
After a cluster of pathological cases had been identi-
fied, Dr. Putschar would come to the Museum for several
days and the two of us would review each one, and he
would dictate his observations on the pathogenesis and
differential diagnosis. During these visits, Mrs. Putschar
would transcribe the dictation and organize the notes. The
result was the identification and documentation of many
additional cases of skeletal paleopathology that added
greatly to our knowledge of disease in antiquity and our
ability to diagnose diseases encountered in archeological
remains.
One of the interesting dimensions of this exercise was
the enthusiasm with which Dr. Putschar reviewed these
cases. Virtually every pathological specimen brought new
knowledge and insight about pathogenesis to both of us.
Because of Dr. Putschar’s vast previous experience with
skeletal disease in many countries, it surprised me that he
was still finding new insights as he studied these cases.
The lesson he repeatedly emphasized was that archeologi-
cal remains offer the potential to see the expression of
disease in an entire skeleton and usually in the untreated
state. This is rarely possible in a modern clinical context.
He also stressed that careful observation of the type and
distribution pattern of lesions within the skeletal specimen
provided insight regarding pathogenesis that complemen-
ted other sources of information about the disease process.
Since 1979, research methodology has also benefitted
from some major breakthroughs in technology. Computed
tomography has brought new understanding to our knowl-
edge of skeletal radiology and pathology. Archeological
4 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
skeletal tissue has been found to be a remarkably good
substrate for the preservation of ancient biomolecules,
including DNA. Recovery of identifiable genetic material
from pathogens has been reported (e.g., Kolman et al.,
1999) and this is only the early stage of this research. The
remarkable power of the personal computer has provided
new ways to manage data and visualize the patterns of
pathology that we encounter in archeological skeletal
remains. The first edition of this book was prepared using
an electric typewriter. I am using a computer word pro-
cessing system for this edition and I often listen to the
music of Mozart being played through my computer while
I work. I doubt that Dr. Putschar would have approved of
listening to Mozart while writing. Among many other
interests, he had a passionate enthusiasm for classical
music and especially the music of Mozart, a fellow
Austrian by birth. Mozart, in his view, must be listened to
and appreciated without distractions.
We also know much more about the skeletal manifes-
tations of disease in archeological human remains and
this has led to greater diagnostic certainty for many patho-
logical conditions. Medical knowledge has continued to
grow, with new insight about the causes of and relation-
ships between skeletal diseases. Not surprisingly the ter-
minology in medicine and paleopathology has continued
to change to reflect the new knowledge acquired about
skeletal diseases.
All of these changes argue for a revision of the first
edition that will address the new knowledge about both
skeletal pathology and paleopathology that has developed
in the last 20 years. Regrettably, Dr. Putschar did not live
to see the development of many of these innovations or to
participate in this revision. While attending professional
meetings in Scotland in early October 1985 he and Mrs.
Putschar visited a medieval castle site near Edinburgh.
During the visit he fell and hit his head on the stone ruins.
He developed a hematoma on the brain that subsequently
required surgery. On their return to the United States he
and Mrs. Putschar received more bad news when she was
diagnosed with terminal cancer. Despite these health pro-
blems they both insisted that before Dr. Putschar’s sur-
gery he go ahead with the lectures he had promised to
deliver on skeletal disease for the last seminar series on
skeletal paleopathology held at the Smithsonian
Institution from October 21 through November 8, 1985.
Although his balance was affected by his injury, and he
was deeply troubled by Mrs. Putschar’s illness, his lec-
tures were models of clarity and provided a remarkable
learning experience for all who heard him. Mrs. Putschar
died on December 31, 1985. The Putschars had a wonder-
ful marriage and her death was a devastating loss for him.
Dr. Putschar’s health declined following two surgeries to
control the bleeding in his brain and he died on April 5,
1987 at the age of 83.
Inevitably the progress made in both medical knowl-
edge and paleopathology during the past 20 years means
that the revisions for this edition are substantial. However,
much of the insight and understanding of pathology that
Dr. Putschar brought to the first edition remains relevant
and wherever possible I have retained his language and
perspectives on skeletal disease. This second edition owes
much to his knowledge and experience.
Acknowledgments for the Second Edition
In the first edition of this book, I acknowledged the assis-
tance of those who contributed so substantially to its prep-
aration. Some of these people have since died, but the
kindness of all who gave of their time and expertise
remains a wonderful memory. Since the publication of the
first edition many additional people have shared their
knowledge and made collections and many additional
cases of pathology available for my research. These
include the following institutions and people. Australia:
The Shellshear Museum, Sydney (Prof. Jonathan Stone
and Kenneth Parsons); The Australian Museum, Sydney
(Phillip Gordon and Dr. Ronald Lampert); The South
Australian Museum, Adelaide (Dr. Graeme Pretty).
Denmark: The Danish National Museum, Cophenhagen
(Prof. Vilhelm Møller-Christensen). England: The
Department of Archaeological Sciences, The University
of Bradford, Bradford (Arnold Aspinall, Dr. Keith
Manchester, Dr. Charlotte Roberts, Anthea Boylston,
Jason Maher, Prof. Mark Pollard, and Dr. Carl Heron);
The Rheumatology Unit, Bristol University, Bristol (Dr.
Juliet Rogers and Prof. Paul Dieppe); The Canterbury
Archaeological Trust, Canterbury (Paul Bennett and
Trevor Anderson); English Heritage, Ancient Monuments
Laboratory, London (Dr. Simon Mays). Norway: The
Department of Anatomy, University of Oslo (Prof. Dr.
Per Holck and Inger Saelebakke); The Leprosy Museum
of Bergen (Prof. Lorentz M. Irgens). Scotland: The Royal
College of Surgeons of Edinburgh (Dr. I. S. Kirkland).
Switzerland: The Institute of Pathological Anatomy,
University of Zurich (Prof. Dr. Ph. U. Heitz and Prof. A.
R. von Hochstetter). United States: The Bishop Museum,
Honolulu, Hawaii (Dr. Donald Duckworth, Dr. Yosiniko
H. Sinoto, and Toni Han); The Peabody Museum,
Harvard University (Dr. David Pilbeam and Dr. Lane
Beck); The San Diego Museum of Man (Rose Tyson);
The Lowie Museum (now the Phoebe Apperson Hearst
Museum of Anthropology), University of California,
Berkeley, California.
In 1987 I was appointed Visiting Professor of
Paleopathology at the University of Bradford, Bradford,
England. Since 1988, I have been in residence in the
Department of Archaeological Sciences at the University
for varying lengths of time almost every year. This has

been a remarkably valuable experience and I am very
grateful for the wonderful collegial relationships that have
developed over the years and the generous hospitality
extended to me and my family. These colleagues include
Arnold Aspinall, the Chairman of the Department when I
was first appointed, Dr. Keith Manchester, Dr. Charlotte
Roberts (now at the University of Durham), Prof. Mark
Pollard, who followed Mr. Aspinall as Department
Chairman, and Dr. Carl Heron, the current Department
Chairman. The skeletal collection in the department, par-
ticularly the remarkable collection of human remains
from the medieval cemetery in Chichester, England, asso-
ciated with the Hospital of St. James and St. Mary
Magdalene have been of great help in furthering my
knowledge of human skeletal paleopathology. Many of
the people buried in this cemetery were lepers and their
skeletons provide crucial insight regarding the skeletal
manifestations of this dreaded disease.
In 1992 I had a casual conversation about my research
with a friend of many years, David Malin, a sales repre-
sentative for Siemens Medical Systems, Inc. He offered to
try and arrange access to CT equipment at a Siemens
facility. His efforts put me in contact with Matthew
Riemann (now retired), the director of the Training and
Development Center for Siemens Medical Systems, Inc.
in Iselin, NJ. Riemann was supportive and asked two
members of his staff, Valere Choumitsky and Blaise
Falkowski, to do what they could to assist my research.
At that time Mr. Falkowski was the senior instructor for
technical training of engineers and service technicians
who service Siemens CT scanners in North America.
When the facility was not being used for training we were
able to use the equipment to scan paleopathological cases.
Eventually the Training and Development Center moved
to Cary, North Carolina, and I and my Smithsonian col-
league, Dr. Bruno Frohlich, continued to use the equip-
ment at no cost during windows in the training schedule.
Access to this equipment proved to be a powerful
research tool and most of the CT images included in this
edition were generated on Siemens equipment.
CT scanning equipment at the Siemens training facility
is upgraded periodically to the newest models manufac-
tured by Siemens. On one occasion Dr. Frohlich learned
that a Siemens Somatom AR-T scanner was to be replaced
with a new model. He suggested that Siemens donate the
older model to the Smithsonian. After approval on all rele-
vant levels the equipment was given to the Museum and is
now used in support of the research endeavors of the
museum staff. The expertise and the many hours of assis-
tance provided by Mr. Falkowski and his colleagues at
Siemens continues to be of major value to my research.
Agnes Stix, Museum Specialist, and Janet Beck,
Volunteer Research Assistant, Department of
Anthropology, National Museum of Natural History,
Introduction Chapter | 1 5
Smithsonian Institution, have invested countless hours in
organizing bibliographic source materials and illustrations
for this book. They have created computer data bases for
the references and photographs that greatly facilitated my
work. Stix in particular has had the responsibility of orga-
nizing the various electronic files of figures, tables, text,
figure legends and references and keeping changes in one
file congruent with the other. Their contributions to this
edition are substantial and I am in their debt. Marcia
Bakry, Scientific Illustrator, Department of Anthropology,
National Museum of Natural History, Smithsonian
Institution, is responsible for preparing the digitized
figures for the book. Using the powerful software avail-
able today for manipulating digitized photographic
images, she has been able to improve significantly the
quality of the figures used in this edition and deserves my
deepest thanks and that of the reader who will benefit
from her skilled work. Dr. Margaret R. Dittemore, Branch
Librarian, Anthropology Branch Library, Smithsonian
Institution Libraries, and her colleagues in the library
were crucial in identifying and obtaining source materials
used in the book. I am also indebted to Roxie Walker and
the Institute of Bioarchaeology (formerly the
Bioanthropology Foundation) for grants that partially sup-
ported the preparation of this edition.
OBJECTIVES OF THE FIRST AND SECOND
EDITIONS
There are many sources of information on the history of
disease, including ancient medical documents, historical
records, art, and the physical remains of ancient people
including both soft tissues and skeletons. Undoubtedly,
human skeletons represent the most ubiquitous source of
information on ancient diseases. This fact must be tem-
pered with the knowledge that relatively few morbid con-
ditions affect the skeleton in a way that leaves visible
changes in dry bones. In spite of this limitation, the study
of skeletal pathology in archeological materials can pro-
vide time depth to our understanding of disease and con-
tribute to our knowledge regarding the role of disease in
human adaptation. In addition, skeletal paleopathology
may also broaden our understanding of disease as it
affects bone tissue. The paleopathologist often has access
to all portions of the skeleton, a situation rarely realized
in modern pathology or radiology. This means that the
gross pattern and distribution of the morbid condition in
all areas of the skeleton can be studied in detail.
To provide reliable standard specimens for dry bone
diagnosis, the reference cases used as a basis for the first
two editions of this book were primarily from the period
between AD 1750 and 1930. Ortner felt that earlier than
this range the medical data were too ambiguous and later,
6 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
the pathologic manifestations were too altered by surgery,
chemotherapy, radiation therapy, and, above all, by the
use of antibiotics. For this reason, the first two editions
turned to the great medical and anatomical collections of
Great Britain and of continental Europe. The British col-
lections proved in many ways to be the most useful,
because they were made by physicians and surgeons, who
were at all times interested in documenting clinical and
historical data. Yet even this material is not necessarily
identical to manifestations seen in archeological specimens.
As Ortner noted, his compiling these editions highlighted
the fact that even the great pathological anatomists of ear-
lier times made mistakes in differential diagnosis.
This book was intended mainly to serve as a text and
atlas of dry bone pathology, regardless of whether or not
each entity had been identified in paleopathology. For
that reason, as many aspects as possible of documented,
dry bone pathology were illustrated, especially because
the original skeletal collections can never be duplicated
and may ultimately disappear. In the paleopathological
discussions in these earlier editions, emphasis was laid on
careful and critical study of published reports and of
actual specimens, bringing a variety of types of evidence
to bear on arriving at a reasonable diagnostic assumption.
Even so, multiple possibilities and uncertainties often
remained. Not the least of these problems was the ambig-
uous and confusing terminology about the nature of path-
ological conditions and the chronology of archeological
specimens in published reports.
This book was written primarily with the needs of the
biological anthropologist and archeologist in mind, with
the hope that they would be able to recognize the abnor-
malities seen in archeological human skeletal material
they excavate or study. This book was also meant to high-
light the importance of recovering all mineralized tissues,
including the small bones of the hands and feet, during
excavation of a burial. Ortner also was interested in gen-
erating a broader readership with different backgrounds,
though, and emphasized the importance of including his-
torians of medicine and disease, orthopedic surgeons,
radiologists, pathologists, and physicians, who may be
called upon to interpret skeletal lesions in dry specimens
or who are interested in extending their understanding to
the more detailed gross expressions of skeletal disease.
HISTORY OF THE THIRD EDITION FROM
JANE E. BUIKSTRA
Don Ortner was just embarking upon the third edition of
this important volume at the time of his unanticipated
death on April 29, 2012, following a brief illness. For
those of us who had been close to Don professionally
and/or personally, our grief was profound. For example,
I—who had enjoyed so many pleasant lunches with Don
when I could sneak away from meetings of the National
Museum of Natural History’s Repatriation Committee or
other Washington responsibilities—could not bear to walk
by his office door for nearly a year, finding other circui-
tous routes to reach the Rose Seminar room of the
NMNH’s Anthropology Department. Don’s achievements
were celebrated both at the Smithsonian, during an event
held during the autumn of 2012, and at the annual meet-
ing of the Paleopathology Association, held during the
2013 annual meeting, April 9 and 10. Fortunately, Powell
(2012) had been able to convince Don to be interviewed
for a chapter in the Global History of Paleopathology
(Buikstra and Roberts, 2012), wherein details of his life
and scholarly contributions may be found. I can add only
that he was an enthusiastic supporter of the fledgling
International Journal of Paleopathology, ably contribut-
ing one of the Inaugural Essays and serving as an
Associate Editor. He rolled up his sleeves upon many
occasions to review articles and offer sage advice to
junior colleagues.
In discussions with Don’s family, especially his part-
ner Joyce and Don Jr., who sounds remarkably like his
father, it became clear that they would be supportive of a
third edition of Identification of Pathological Conditions
in Human Skeletal Remains, under my editorship.
Discussions with Bruno Frohlich, who was helping the
Department of Anthropology in archiving the materials
from Don’s office, made it clear that Don had only just
embarked on the project. No publisher had been identi-
fied, nor was there a proposal. Given this situation, I
began plans for the project. In creating the proposal, first
discussed with the Smithsonian Press, who were not
enthusiastic about the project due to concerns with copy-
right issues, I reflected upon the many productive discus-
sions in which I had engaged with Don. These convinced
me that he would have wanted the volume revision not to
second guess “what Don might have wanted,” but rather
to reflect the status of paleopathology at the time the revi-
sion appeared. This meant continuing to emphasize the
basic empirical evidence upon which paleopathological
identifications are based, but also to reflect the dynamic
nature of paleopathology today. Given the mentorship and
encouragement that Don had so freely provided to so
many of us, I also believe that he would have wanted our
generation(s) to leave our imprint upon the work—giving
it our best effort. It is with this spirit that we have
approached the volume.
When I approached Elizabeth Brown, Senior
Acquisitions Editor at Elsevier, about the project, she was
enthusiastic in support. We have tried to maintain the many
strengths of the earlier editions, while also adding new
methodological advances (molecular and parasitology),
mentioning closely related and increasingly convergent

research topics (animal paleopathology; mummy science)
and emphasizing the interdisciplinarity of paleopathology
in exploring themes based in the social sciences and human-
ities. When approached, colleagues in paleopathology and
related disciplines signed on enthusiastically, bringing their
special expertise to this important initiative.
Don and I agreed about most aspects of paleopathol-
ogy, especially the need for detailed descriptions of path-
ological changes, for standard terminology, to appreciate
limitations of early clinical accounts as well as those of
the antibiotic era, and for rigorous applications of differ-
ential diagnostic methods. I am less concerned than he
about classification, and therefore this topic will be less
visible in this third edition. I sincerely hope that we have
done justice to Don’s fundamental contributions to the
discipline of paleopathology, while recognizing key
developments since his seminal 2003 publications.
Acknowledgments for the Third Edition
First and foremost, I would like to acknowledge the
Ortner family in their support of this initiative. The
Department of Anthropology, especially its Chair during
the period of project development, Torben Rick, along
with Don’s long-term collaborator, Bruno Frohlich have
been immensely reassuring. The editor is extremely
appreciative of the enthusiasm and expertise of the colla-
borators, whose wisdom is represented here. The editorial
and content editorial assistance of Katelyn Bolhofner has
improved clarity and accuracy throughout the develop-
ment of the volume. Additional polish has been added by
the skills of Sylvia Cheever in final stages of the process.
Anne Grauer’s careful proof-reading and apt suggestions
have improved the final production, which is deeply
appreciated. Many of the authors wish to express their
gratitude to Don Brothwell for his scholarship and per-
sonal encouragement of our research, both in human and
in animal paleopathology. Finally, the assistance and
encouragement from Elsevier, including Elizabeth Brown,
Pat Gonzalez, and the production team have been essen-
tial to the success of the project.
OBJECTIVES OF THE THIRD EDITION
More than 30 years have passed since the landmark
Identification of Pathological Conditions in Human
Skeletal Remains (Ortner and Putschar, 1981) was pub-
lished, followed by the second edition (Ortner, 2003) over
a decade ago. The field and the profession of paleopathol-
ogy have changed markedly over this period, in no small
part due to the influence of these volumes. Ortner had
planned but not begun writing a third edition at the time
of his sudden death, and this volume represents the com-
pletion of this project, reflecting his (and Walter
Introduction Chapter | 1 7
Putschar’s) core contributions in bone disease through
revised and new chapters that manifest the contemporary
breadth and depth of the discipline of paleopathology.
This third edition updates the previous volumes
through the addition of recent medical information on
skeletal disorders and the latest relevant literature on
human skeletal paleopathology. This work also adds chap-
ters on current methods being used in research on skeletal
paleopathology. These include increased reliance on
imaging, including CT methods, histology, and analysis
of ancient DNA. In addition, chapters covering closely
related subjects, such as diet (including isotopes, micro-
wear, colon contents, (macro/micro fossils; pollen), dental
calculus, dental caries), mummy science, animal paleopa-
thology, and paleoparasitology have been added. Given
the contemporary availability of numerous texts covering
basic osteology, in this edition chapters on biological pro-
filing and osteobiographical methods have been deleted.
These topics are now introduced briefly in Chapter 3, and
Chapter 2 now offers an extended history of paleopathol-
ogy, current issues in the field, and the importance of rig-
orous differential diagnosis. The volume is further framed
by an expanded discussion of important themes for con-
sideration in this paleopathological research (Chapter 3).
As was the case for the first two editions of this vol-
ume, the most fundamental objective of this third edition
is to provide an integrated, detailed discussion of the
gross pathology of the human skeleton to facilitate rigor-
ous differential diagnosis of these pathologies in human
skeletal remains from archaeological contexts. In addition
to this foundation, the objectives of this third edition
include: emphasizing careful consideration of contempo-
rary clinical literature in diagnosis, encouraging knowl-
edge in epidemiology, animal paleopathology,
parasitology, and molecular and chemical advances in
contextualizing skeletal analyses, and presenting advances
in imaging, data collection, and diagnostic approaches
arising from such related fields as forensic science, dental
anthropology, biogeochemistry, and molecular science.
FORMAT OF THE VOLUME
While texts in paleopathology all agree that classification
is an important aspect of disease diagnosis, there is no gen-
eral agreement upon the number of classes of disease. As
Ortner (2012) notes, Reznick’s orthopedic radiology text
recognizes 17 categories. Aufderheide and Rodrı́guez-
Martı́n (1998)’s paleopathology text recognizes 13, while
both editions of the Ortner volumes focus upon 12.
Influenced by Lent Johnson, Ragsdale and various cowor-
kers (Ragsdale and Miller, 1996; Ragsdale and Lehmer,
2012) have asserted the utility of seven basic disease cate-
gories, readily recalled through the use of the acronym
VITAMIN (see Table 1.1, adapted from Ragsdale and
8 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
TABLE 1.1 Ragsdale’s Seven Basic Disease Categories
1 V Vascular
2 I Innervation/mechanical
3 T Trauma/repair
4 A Anomaly
5 M Metabolic
6 I Inflammatory/immune
7 N Neoplasms
Lehmer, 2012: 230). Roberts and Manchester (third
edition, 2010) also organize their discussion in The
Archaeology of Disease into seven categories.
After emphasizing the need for histology and recogni-
tion of disease processes, Ragsdale and Lehmer (2012:
247) close with the assertion, “that only through detailed
descriptions and diagnoses to general disease categories,
will a stronger methodological basis for comparative
research in paleopathology be reached.” They argue, based
upon evidence from four workshops held at the paleopa-
thology meetings (Miller et al., 1996) that assignments to
disease categories are more accurate than specific diagno-
ses. While these conclusions do reflect the empirical data
from the Workshops, questions about the relative experi-
ence of the participants remains. In addition, the degree to
which comparisons across 7, 12, 13, or 1 categories are
meaningful in interpreting the past must, of course, depend
upon the research question addressed or the hypothesis
posed. Further, the issue of contexts—environmental, tem-
poral, cultural—must be considered.
As Ortner (2012) emphasizes, disease classifications
emphasize cause or pathogenesis of a disease. In that,
e.g., bacterial pathogenesis can be a cause, pathogenesis
would seem to be the overarching category. Many dis-
eases have multiple causes, and classifications become
complex. Metabolic diseases, due to disturbances in oste-
oid formation and mineralization, are often associated
with nutritional deficiencies. Similarly, erosive arthropa-
thies are typically classified as joint disorders, even
though an infection may trigger the response.
This volume will follow the previous editions in its
classification of disease conditions: trauma, infectious dis-
eases, circulatory disorders, reticuloendothelial and
hematopoietic disorders, metabolic disorders, endocrine
disorders, congenital and neuromechanical disorders, dys-
plasias, tumor and tumor-like disorders, joint disorders,
dental and jaw disorders, and miscellaneous disorders. A
few specific disorders have been moved to more
completely reflect contemporary knowledge of pathogene-
sis. In reference to the process of classification, Ortner
(2012: 263) emphasized that the important point “is the
need to understand the pathogenesis and, where possible,
the cause of the disorder and not let the assignment to a
specific category of disease obscure our understanding of
the basic bone biology of disease.”
ABBREVIATIONS
The illustrations in this book are of specimens from many
institutions. The following abbreviations are used in the
legends to avoid repetition of lengthy institutional names
and locations. This list includes institutions that had path-
ological cases used in both the first and second editions.
AFIP Armed Forces Institute of Pathology, Washington,
DC, United States
AIUZ Anthropological Institute, University of Zurich,
Zurich, Switzerland
ANM National Museum of Anthropology, Prague, Czech
Republic
BMNH British Museum, The Natural History Museum,
London, England
CGH Department of Pathology, Charleston General
Hospital, Charleston, WV, United States
CISC Coimbra Identified Skeletal Collection,
Departamento de Ciências da Vida, Universidade de
Coimbra, Portugal
DPUS Department of Pathology, University of Strasbourg,
Strasbourg, France
FM Field Museum of Natural History, Chicago, IL,
United States
FPAM Federal Pathologic-Anatomy Museum, Vienna, Austria
HM Hunterian Museum, The Royal College of Surgeons
of England, London, England
IEC International Exchange Collection, Departamento
de Ciências da Vida, Universidade de Coimbra,
Coimbra, Portugal
IPAZ Institute of Pathological Anatomy, University of
Zurich, Zurich, Switzerland
LLAC- Luı́s Lopes Anthropological Collection, Museu Bocage,
MUHNAC Museu Nacional de História Natural e da Ciência,
Lisbon, Portugal
MGH Department of Pathology, Massachusetts General
Hospital, Boston, MA, United States
NHMB Natural History Museum, Bern, Switzerland
NMNH National Museum of Natural History, Smithsonian
Institution, Washington, DC, United States
PMES Pathology Museum, The Royal College of Surgeons
of Edinburgh, Edinburgh, Scotland
WM Wellcome Museum, The Royal College of Surgeons
of England, London, England
REFERENCES
Aufderheide, A.C., Rodrı́guez-Martı́n, C., 1998. The Cambridge
Encyclopedia of Human Paleopathology. Cambridge University
Press, Cambridge.

Buikstra, J.E., Roberts, C.A. (Eds.), 2012. The Global History of
Paleopathology: Pioneers and Prospects. Oxford University Press,
New York.
Hackett, C., 1947. The Bone Lesions of Yaws in Uganda. Thesis.
University of London, London.
Hackett, C., 1976. Diagnostic criteria of syphilis, yaws and treponarid
(treponematoses) and of some other diseases in dry bones.
Sitzungsberichte der Heidelberger Akademie der Wissenschaften
Mathematisch-naturwissenschaftliche Klasse, Abhandlung 4.
Springer-Verlag, Berlin.
Kolman, C., Centurion-Lara, A., Lukehart, S., Owsley, D., Tuross, N.,
1999. Identification of Treponema pallidum subspecies pallidum in a
100-year-old skeletal specimen. J. Infect. Diseases 180, 2060 2063.
Miller, E., Ragsdale, B.D., Ortner, D.J., 1996. Accuracy in dry bone
diagnosis: a comment on palaeopathological methods. Int. J.
Osteoarchaeol. 6 (3), 221 229.
Ortner, D.J., 2003. Identification of Pathological Conditions in Human
Skeletal Remains. Academic Press, New York.
Ortner, D.J., 2012. Differential diagnosis and issues in disease classifica-
tion. In: Grauer, A. (Ed.), A Companion to Paleopathology. Wiley-
Blackwell, New York, pp. 250 267.
Introduction Chapter | 1 9
Ortner, D.J., Putschar, W.J.P., 1981. Identification of Pathological
Conditions in Human Skeletal Remains. Smithsonian Institution
Press, Washington, DC.
Powell, M.L., 2012. Donald J. Ortner (1938 ). In: Buikstra, J.E.,
Roberts, C.A. (Eds.), The Global History of Paleopathology:
Pioneers and Prospects. Oxford University Press, New York,
pp. 89 96.
Ragsdale, B.D., Lehmer, L.M., 2012. A knowledge of bone at the cellu-
lar (histological) level is essential to paleopathology. In: Grauer, A.
(Ed.), A Companion to Paleopathology. Wiley-Blackwell, New
York, pp. 227 259.
Ragsdale, B.D., Miller, E., 1996. Workshop A. Skeletal Disease
Workshop VIII: several of the seven basic categories of disease.
In: Cockburn, E. (Ed.), Papers on Paleopathology Presented at the
23rd Annual Meeting of the Paleopathology Association, Durham,
North Carolina. Paleopathology Association, Detroit, p. 1.
Roberts, C.A., Manchester, K., 2010. The Archaeology of Disease.
Cornell University Press, New York.
Tyson, R. (Ed.), 1997. Human Paleopathology and Related Subjects.
An International Bibliography. San Diego Museum of Man,
San Diego.
Chapter 2
A Brief History and 21st Century
Challenges
Jane E. Buikstra1 and Sharon DeWitte2
1
Arizona State University, Tempe, AZ, United States, 2University of South Carolina, SC, United States
In this chapter, we consider the history of paleopathology
and a few of the fundamental issues faced by practitioners
in the development of this field. We then turn to a discus-
sion of the current state of paleopathology, reviewing
methodological and theoretical issues encountered in 21st
century paleopathology. In this regard, we discuss the dif-
ferential diagnosis of pathological conditions in archeolo-
gical skeletal remains, suggesting avenues by which
paleopathologists may pursue more rigorous diagnosis.
Finally, we discuss the important contribution of paleoe-
pidemiology in the advancement of this field, as well as
considering the ramifications of the osteological paradox
in such work.
A BRIEF HISTORY OF PALEOPATHOLOGY
Paleopathology has been defined in recent decades as the
study of disease, both human and nonhuman, in antiquity
using a variety of different sources, including human
mummified and skeletal remains, ancient documents,
illustrations from early books, painting and sculpture
from the past, and analysis of coprolites (Ortner, 2003: 8)
More recently, this definition has been reevaluated and
expanded to reflect the crucial interplay of biomedical
and social sciences and the humanities in the development
and future of the field (Buikstra et al., 2017). A compre-
hensive history of paleopathology has recently been writ-
ten (Buikstra and Roberts, 2012), and there are several
other older summaries of this history that readers who
have a specific interest in the subject may wish to consult
(e.g., Jarcho, 1966; Angel, 1981; Ubelaker, 1982;
Armelagos, 1997; Aufderheide and Rodriguez-Martin,
1998). Thus, a detailed history of paleopathology that
includes research using all the varied sources of potential
information is beyond the scope of this book. Here, we
offer a brief summary of the history of paleopathology,
Ortner’s Identification of Pathological Conditions in Human Skeletal Remains. DOI: https://doi.org/10.1016/B978-0-12-809738-0.00002-8
© 2019 Elsevier Inc. All rights reserved.
highlighting some of the issues and major developments
in the field over the past 200 years.
The history of paleopathology in many ways parallels
the development of most other scientific disciplines. The
early publications consist of a body of descriptive litera-
ture in which abnormalities encountered by an observer
are described against the background of what is normal.
Much of this early research was no more than an anatomi-
cal account of these abnormal conditions with little if any
attempt to explore the biological or pathological signifi-
cance of what was being described. The earliest work
focused on nonhuman paleontological specimens (e.g.,
Esper, 1774; Cuvier, 1820). Warren (1822) included a dis-
cussion of artificial cranial deformation in human skulls
of indigenous North Americans in his book titled, A
Comparative View of the Sensorial and Nervous Systems
in Man and Animals. In 1861 in Paris, Gosse published
another study of artificial cranial deformation. In the fol-
lowing decades, the question of the origin of syphilis
began to be debated with intensity (e.g., Jones, 1876;
Virchow, 1898). This debate marks one of the earliest
attempts to use archeological human remains to resolve
an important biomedical problem. And toward the end of
the 19th century, R.W. Shufeldt proposed that the term
“paleopathology” be used to describe “all diseased or
pathological conditions found fossilized in the remains of
extinct or fossil animals” (Shufeldt, 1892: 679).
As the term “paleopathology” began to be used in the
early 20th century, this period witnessed a marked expan-
sion of published reports on ancient disease. Particularly
notable is the work of Sir Marc Armand Ruffer (1910) on
Egyptian mummies, and the studies on Nubian skeletal
material by Wood-Jones (1908, 1910) and Elliot-Smith
and Wood-Jones (1910). In the United States, Aleš
Hrdlička (1914) published some observations on the
pathology of ancient Peruvian skulls. In 1923, Moodie’s
11
12 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
introduction to the study of ancient disease, which empha-
sized nonhuman paleontological specimens, appeared. A
brief, general review of human paleopathology was pub-
lished by Williams in 1929. This review included obser-
vations on bones and teeth as well as on mummy tissue
and ancient art. Pales (1930) followed with his book on
paleopathology and comparative pathology. Most of his
cases and discussions concerned European human speci-
mens. In the same year, Hooton (1930) published his clas-
sic study of North American Indian skeletal material from
Pecos in which he included an extensive description of
pathological specimens. Hooton’s study is notable in its
descriptive detail, in the statistical treatment of different
types of disease in the skeletal population, and in his
efforts to show trends in disease frequency through the
time period of human occupation at the site.
In the 1960s, Wells (1964) published a review of evi-
dence of human paleopathology from skeletal material,
mummies, and art that brought paleopathology to the
attention of a more general audience. But in the preced-
ing decades, paleopathological studies had fallen into a
pattern of inclusion in archeological research as descrip-
tive addenda or appendices. Thus, calls for further
advances in the field of paleopathology were made
(Jarcho, 1966; Brothwell and Sandison, 1967; see also
Grmek, 1983/1989), resulting in the establishment of
professional organization, international journals, and pro-
fessional meetings and training seminars (Buikstra and
Roberts, 2012).
Throughout the development of human skeletal paleo-
pathology as a scholarly discipline there have been recur-
ring problems in both theory and methodology. In the
early stages of paleopathology, most of the research was
conducted by physicians who had little knowledge of
archeology, thus context often was overlooked. As studies
of pathological skeletal specimens began to be conducted
primarily by biological anthropologists, whose formal
training and experience in skeletal pathology and radiol-
ogy may be deficient, pathological conditions were at risk
of being attributed incorrectly to the wrong time period
by those unfamiliar with the complexities of archeological
dating. Further, bone lesions were incorrectly diagnosed
through ignorance of anatomy and the total range of dis-
eases that affect bone (see Stewart’s comments on this
problem in Jarcho, 1966: 43). These problems were com-
plicated further due to the slow formulation of a theoreti-
cal context for interpreting the meaning of
paleopathological data. [See, e.g., the debate (Wood et al.,
1992; Goodman, 1993) about what can and cannot be said
about prevalence data and the inferences made about the
health of past human populations.]
In 1988, Ortner and Aufderheide (1991) organized a
symposium held as part of the International Congress of
Anthropological and Ethnological Sciences in Zagreb,
Yugoslavia (now Croatia) that attempted to assess (1)
how far paleopathology had developed as a scientific dis-
cipline, (2) some of the theoretical and methodological
problems that needed to be resolved, and (3) directions
that research might take in the future. Methodological
issues included an inconsistent descriptive terminology
that precluded comparison between published reports, and
the lack of diagnostic criteria that fully utilized the infor-
mation available in archeological human skeletons
(Ortner, 1991). Theoretical issues included the need for
greater understanding of what skeletal disease meant in
terms of the general morbidity that existed within the liv-
ing population in which the person with skeletal disease
lived (Ortner, 1991).
Much of the emphasis in paleopathology until fairly
recently has been on descriptions of pathological speci-
mens, and there had been little effort to relate the evi-
dence of disease to the broader problems of human
adaptation. Early hints of such an emphasis exist in
Hooton’s Pecos Pueblo monograph (1930), in the consid-
eration of epidemiological factors in evaluating the data
on pre-Columbian tuberculosis in the New World (Morse,
1969), and in discussions on the origin of treponemal dis-
eases (Hackett, 1963; Hudson, 1965). But not until
recently has the trend toward population studies of ancient
disease become a significant part of the literature on
paleopathology as these methodological and theoretical
problems are resolved (e.g., Larsen, 1997).
Much of the descriptive literature in skeletal paleopa-
thology depended upon the scholar’s knowledge of gross
bone pathology. Unfortunately, where this knowledge was
inadequate there were few reference sources that could be
of assistance. Jarcho (1966) organized a symposium on
human paleopathology that addressed this problem,
among others. The participants called for the establish-
ment of a paleopathology registry and improved diagnos-
tic methodology to partially correct these problems.
Steinbock’s reference book (1976) on diagnosis of ancient
bone disease represented the first integrated attempt to
establish diagnostic criteria for the paleopathologist that
addressed the broad range of diseases that affect the
human skeleton. The first two editions of this volume
(Ortner and Putschar, 1981 and slightly revised in 1985;
Ortner, 2003) provided a complimentary treatment of
skeletal disease. Both these reference works represented
important steps in improving the knowledge regarding the
types of diseases that affect bone and the morphological
features associated with the disease.
Since the publication of the first two editions of this
book, there has been a substantial increase in research on
broader scientific problems, particularly those related to
paleoepidemiology, as we will review later in this chapter.
There has also been significant progress made on several
crucial methodological problems. One of the most
 A Brief History and 21st Century Challenges Chapter | 2
important of these has been the improvement in our appli-
cation of differential diagnosis, which we will discuss in
detail in the following section. As we face a new suite of
issues and advances in the 21st century, we argue that
paleopathology should be an interdisciplinary endeavor,
incorporating expertise from the humanities, the social
sciences, and the biomedical sciences (Buikstra et al.,
2017).
21ST CENTURY PALEOPATHOLOGY
Our vision of 21st century paleopathology is of a pro-
foundly interdisciplinary endeavor, drawing knowledge
and professionals from the biomedical and social
sciences, as well as the humanities. We use knowledge
about past health to address the coevolution of humans
and pathogens, and we anticipate much more knowledge
about both human and animal disease will soon be
reviewed through molecular study. This volume therefore
is meant to be an entry point for knowledge that necessar-
ily extends well beyond these pages.
First of all, we must recognize that paleopathology
proceeds primarily through scientific methods. Our obser-
vations of ancient remains should be drawn carefully, fol-
low standard descriptive terminology, and be designed to
minimize both intra- and interobserver error. A general
overview of terminology appears on the Paleopathology
Association’s website (https://paleopathology-association.
wildapricot.org/Nomenclature-in-Paleopathology). While
this overview generally follows medical terms, methodo-
logical and application issues arise due to the fact that
most of our observations are made upon materials that
emerged from a burial environment. Taphonomic changes
are frequently described in terms also used for vital pro-
cesses, “abraded” and “eroded” being two apt examples.
Therefore, when using such terms, the observer should be
careful to indicate whether the process occurred ante- or
postmortem.
We continue to follow Ragsdale and colleague’s
(1981) descriptions of periosteal bone reactions (see also
Weston, 2012), familiar to those of us who have been
humbled during Ortner/Ragsdale and Ragsdale workshops
at the annual meetings of the Paleopathology Association.
It is crucial not only to describe, but also to understand,
the processes that have led to the observed change. As we
consider our observations, we should report whether or
not the process was active at the time of death, or
quiescent.
There are published standards (Buikstra and Ubelaker,
1994) and freely available databases (Osteoware) for
recording pathological changes in human bones.
Whatever system is used, an explicit key that explains the
coding system is crucial. While this issue may not seem
so important to those starting their research careers, its
13
significance will become apparent as the need for consis-
tency across years of data collection and comparative
approaches emerge.
Observing pathological changes and distinguishing
these from postmortem alterations is one crucial step in
assessing ancient disease (see Chapter 5: Abnormal Bone:
Considerations for Documentation, Disease Process
Identification, and Differential Diagnosis). Once these
have been coded by individual, and then across a skeletal
sample, the identification of a condition assumes signifi-
cance. Remembering that observations may be compli-
cated by comorbidities, i.e., that two or more diseases
may affect a given individual, the survey of possible con-
ditions should begin. In most cases, this assessment can
begin with this volume, but it should not necessarily end
here. To fully appreciate the manner in which bones (and
other tissues) may react to a given insult requires an
appreciation of the variable manner in which a person
may be affected and the fact that the person may have
died prior to the most extreme manifestation of the dis-
ease, as recorded here or in the clinical literature.
Certainly medical interventions, especially antibiotics and
chemotherapy, have changed the course of disease over
the past century profoundly. Earlier medical procedures,
such as treating venereal syphilis with mercury or malaria
with high-temperature baths, may or may not have altered
the course of disease. Such treatments, however, may
have introduced their own diagnostic sequelae.
Earlier editions of this volume have recommended
clinical diagnoses found in books and medical museums
between 1750 and 1930. We are inclined to a more con-
servative perspective, particularly in reference to infec-
tious diseases. The most reliable sources, in our
experience, have been clinical reports from the period fol-
lowing the identification of the pathogen causing the con-
dition and prior to the development of effective
interventions. In the absence of documented collections,
of course, autopsies and radiographic records are seldom
sufficiently complete to provide the desirable, complete
skeletal record. Even those practitioners using documen-
ted collections should be careful to read all the supporting
documentation to discern the degree to which the “diag-
nosis” was based upon clinical observations rather than
posthoc skeletal observations.
Again, we emphasize that, in most cases, this book
should be considered a secondary source. Anyone wishing
to develop a definite differential diagnosis should consult
the primary literature, which engages the clinical litera-
ture. Web-based searches are important, especially in dis-
covering primary source documents from an earlier era.
Identifying a disease process in archeologically recov-
ered human remains is only part of the process of inter-
preting past lives. A practitioner of paleopathology needs
to appreciate concepts drawn from the social sciences
14 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
(Buikstra et al., 2017; Chapter 3: Themes in
Paleopathology) and the humanities (Mitchell, 2011,
2012, 2017). Given the myriad branches of knowledge
required for most studies of ancient disease, collabora-
tions are crucial and training in professional cooperation
and respect are important for paleopathologists. Among
other essential considerations are ethical issues relating to
patients and descendent groups (Lambert, 2012). These
are an essential part of any training program and any
development of a project involving peoples from the past.
In closing, we will more closely examine the potential
contribution of paleoepidemiology to 21st century paleo-
pathology research.
PALEOEPIDEMIOLOGY
Epidemiology and Paleoepidemiology
Epidemiology is the study of the distribution of health-
related states or events (including, but not limited to, dis-
ease) within populations and of the factors that affect
them, and the application of this information in efforts to
control diseases and other health problems (WHO, 2018).
Paleoepidemiology is the study of population-wide pat-
terns of human health and disease in the past, typically
done using data derived from skeletal or mummified
remains excavated from archeological sites or from docu-
mented skeletal collections. For many populations, skele-
tal data provide the only remaining evidence of health in
the past, and paleoepidemiology thus provides invaluable
insights into how human health has varied within and
between populations or subpopulations throughout human
prehistory and history. Hooton’s (1930) examination of
pathology in Pecos Pueblo is often credited as the first
paleoepidemiological study, providing a model for the
application of quantitative analyses of paleopathological
data that became more widely used in bioarcheological
research beginning in the 1960s (Armelagos, 2003;
Mendonça de Souza et al., 2003). Since then, paleoepide-
miologists have addressed such topics as the Neolithic
and the second epidemiological transitions (e.g.,
Armelagos and Cohen, 1984; Wilson, 2014; Zuckerman,
2014), the effects of European contact on indigenous
populations (e.g., Klaus and Tam, 2009; Larsen et al.,
2001), and mortality patterns during and health patterns
following infectious disease epidemics (e.g., DeWitte,
2018; DeWitte and Wood, 2008).
Though the ultimate goal of paleoepidemiology—
understanding how and why health-related states vary
within a population—is shared with epidemiology, and
though both fields focus on groups rather than individuals
as the fundamental units of analysis, the data and analyti-
cal methods available to scholars in these fields are quite
different. Epidemiologists use experimental or
observational data derived from longitudinal or cross-
sectional studies of living populations; however, only
cross-sectional data are available to paleoepidemiologists.
Because paleoepidemiologists work with samples of the
dead, they cannot follow individuals over time to deter-
mine how their health-related states change in response to
exposure to a particular variable. It is possible for paleoe-
pidemiologists to examine the within-individual effects of
variables over time (i.e., the life course) in a typical
archeological skeletal sample by assessing later-life out-
comes associated with developmental stress markers or
isotopic signature of diet or mobility that form relatively
early in life and can be assigned ages-at-formation.
Alternatively, paleoepidemiologists can study documented
skeletal collections for which they have information both
about exposures early in life and later health or mortality
outcomes. With respect to the former approach, there are
unfortunately a limited number of developmental skeletal
stress markers (e.g., enamel hypoplasia, neural canal
dimensions, tooth size, cribra orbitalia, stature), and they
generally suffer from low specificity. Further complicat-
ing paleoepidemiological studies is the fact that skeletal
samples are typically accumulated over multiple genera-
tions, and often it is difficult or impossible to determine
more precisely, within the general period of use of a cem-
etery, the date of death of each individual in the sample
(Mendonça de Souza et al., 2003). This is even further
complicated by the lack of accuracy and precision associ-
ated with adult skeletal age-estimation methods (Bocquet-
Appel and Masset, 1982; Milner and Boldsen, 2012). As a
consequence of these issues, paleoepidemiologists rarely
examine true cohorts of individuals (a cohort is a group
of individuals who all experience a particular event at the
same time; e.g., a birth cohort is a group of people who
were all born at the same time) as is possible for epide-
miologists. Thus, paleodemographic studies might be con-
founded by temporal changes in exposure variables that
cannot be detected and thus which cannot be controlled
for.
Epidemiologists are interested in and generally capa-
ble of measuring health-related states and disease out-
comes relative to a particular population at risk. That is,
epidemiologists can identify not only those individuals
who have the specific conditions of interest, but also
those alive at the same time (and at the same age) who do
not have those conditions or who do not develop them
over the course of a study. Combined with good temporal
control, information about the population at risk allows
epidemiologists to better contextualize, among other
things, the incidence and prevalence of conditions.
Incidence is the number of new or newly diagnosed cases
of a condition within a specified period of time, and prev-
alence is the actual number of individuals with the condi-
tion alive at a particular point or during a particular
 A Brief History and 21st Century Challenges Chapter | 2
period of time. In theory, paleoepidemiologists share an
interest in these phenomena with respect to past popula-
tions. However, paleoepidemiologists have much more
limited information about past populations at risk, as they
observe only those individuals who died and ultimately
became part of excavated skeletal samples (see more
about selective mortality below), not the actual once-
living populations to which they originally belonged.
Paleoepidemiological reconstructions of past populations
at risk, incidence, prevalence, and other measures are
thus, at best, biased.
Epidemiologists are also better able to identify health-
related states or diseases of interest because they have at
their disposal data collected from living people using a
variety of diagnostic tools (the nature of which depends
on the condition of interest), including physical examina-
tions of living patients or decedents, health history and
behavior questionnaires, immunoassays, histological anal-
yses, and cell cultures. Diagnostic criteria or tests for
identifying diseases or conditions are typically described
in terms of their sensitivity and specificity. Sensitivity,
which is also referred to as the true positive rate, is the
proportion of people with a condition who are correctly
identified by a test as having the condition, i.e., the extent
to which true positives are not overlooked by the test
(Boldsen, 2001; Waldron, 2007). Diagnostic tests with
high sensitivity produce few false negatives, so if people
test negative for the disease of interest, it is likely that
they do not, in fact, have that disease. Specificity (also
called the true negative rate) is the proportion of people
without a condition who are correctly identified by the
test as not having it (Boldsen, 2001; Waldron, 2007); i.e.,
specificity is the extent to which people who test positive
really represent the condition of interest. Diagnostic tests
with high specificity produce few false positives, so if
people test positive for a condition, it is likely that they
actually have the condition. Because of controlled labora-
tory and field experiments, it is possible to accurately
assess the sensitivity and specificity of diagnostic criteria
used in living populations, so epidemiologists know how
confident they can be in their diagnoses and research find-
ings based thereon. Though epidemiologists often work
with data derived from tests having relatively low sensi-
tivity and specificity, they are at an advantage in knowing
something about the level of uncertainty they face in their
research.
Paleoepidemiologists, on the other hand, most often
rely solely upon skeletal lesions or stress markers to
assess health-related states, which provide relatively lim-
ited information about health and disease (compared to
the data available to epidemiologists) and for which there
is often limited, if any, information about sensitivity and
specificity. The specificity of skeletal lesions for diagnos-
tic purposes is limited in large part by the fact that bone
15
can respond to disease and trauma in just a few general
ways: bone is deposited, removed, or deformed in
response to these deviations from normal physiology or
structural integrity. Paleoepidemiologists are faced with
the dilemma that many diseases can lead to the produc-
tion of skeletal lesions that look similar, if not identical,
across those etiologies. For example, Weston (2008)
assessed periosteal new bone formation macroscopically
and using radiographs from individuals with known meta-
bolic, infectious, and other conditions, such as chronic
osteomyelitis, fracture, syphilis, and rickets. She did not
identify any location, size, shape, or form characteristics
of the lesions that were specific to those conditions,
which indicates that responses to diseases are determined
by the nature of the affected bone and the periosteum
rather than by the diseases themselves (Weston 2008: 56).
Even in cases of diseases that produce pathognomonic
lesions (many examples of which are provided in this vol-
ume), differential diagnosis can be severely hampered if
the preservation of the relevant elements is poor. The sen-
sitivity of skeletal lesions is limited because not everyone
with conditions that have the potential to affect the skele-
ton will, in fact, develop skeletal lesions in response
(Milner and Boldsen, 2017). For example, tuberculosis
can cause the production of diagnostic bony lesions, but
only approximately 3% 5% of people with untreated
tuberculosis develop such lesions (Resnick and
Niwayama, 1995). This low proportion means that many
people with tuberculosis in skeletal samples will not be
diagnosed based on skeletal pathology alone.
As has been discussed elsewhere (see, e.g., Mays,
2018; Zuckerman et al., 2016), few paleoepidemiological
studies have estimated the sensitivity and specificity of
skeletal lesions. For example, Smith-Guzmán (2015)
assessed the sensitivity and specificity of a suite of skele-
tal lesions with respect to malaria-associated anemia using
clinical samples of individuals with known cause of death
or malaria exposure. Boldsen (2001) estimated the sensi-
tivity and specificity of skeletal indicators of leprosy
based, in part, on samples drawn from medieval cemeter-
ies associated with lepers’ hospitals. Konigsberg and
Frankenberg (2013) illustrate the general approach to esti-
mation using a hypothetical example. Often, however,
paleoepidemiologists face an unquantified level of uncer-
tainty regarding how many false negatives and false posi-
tives with respect to a particular condition exist within
their samples.
As emphasized in Chapter 8, it is increasingly possible
to use ancient biomolecular approaches to identify dis-
eases such as bubonic plague, tuberculosis, leprosy,
malaria, hepatitis, and enteric fever (Salmonella) in skele-
tal or mummified tissue samples (Bos et al., 2011, 2014,
2016; Donoghue et al., 2015; Marciniak et al., 2016;
Patterson Ross et al., 2018; Vågene et al., 2018).
16 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
However, these approaches are not without their own pro-
blems; e.g., they are expensive (prohibitively so for many
scholars) and destructive, and it can be difficult to inter-
pret negative findings given the myriad factors that inter-
fere with DNA and other biomolecule preservation,
extraction, or amplification. As a result, ancient biomole-
cule studies tend to yield small sample sizes of indivi-
duals who test positive for the pathogens of interest, and
thus, to date, few paleoepidemiological studies have been
based solely on ancient bimolecular data.
Despite the issues associated with identifying specific
conditions in skeletal samples, paleoepidemiological stud-
ies have examined a variety of specific infectious, meta-
bolic, and degenerative conditions in past populations,
including leprosy (Boldsen, 2001), syphilis (Harper et al.,
2011), tuberculosis (Buikstra, 1999), vitamin D deficiency
(Snoddy et al., 2016), developmental dysplasia (Blatt,
2015), and degenerative joint disease (Klaus et al., 2009).
The existence of documented historical plague burials has
also facilitated paleoepidemiological studies of bubonic
plague in the absence of diagnostic skeletal pathology
(DeWitte and Wood, 2008; Kacki, 2017). By choice or
necessity, however, rather than attempt to diagnose spe-
cific etiologies, many paleoepidemiologists use skeletal
lesions as general (i.e., nonspecific) indicators of exposure
to physiological stress or developmental disturbance.
Thus, many paleoepidemiological studies focus on the
general health of populations using nonspecific indicators
rather than attempt to assess health in the context of spe-
cific diseases. This approach skirts some of the issues
associated with low sensitivity and specificity, but still
must contend with the fundamental issue that skeletal
samples are inherently biased and that the presence or
absence of lesions can be difficult to interpret (as framed
by the osteological paradox, described below).
The Relationship Between
Paleoepidemiology and Paleopathology
With its focus on skeletal pathology, paleoepidemiology
is clearly aligned with its sister discipline, paleopathol-
ogy. Both fields focus on health, disease, or well-being in
the past, and both make use of the same skeletal patholo-
gies and stress markers (and thus both ultimately grapple
with the same limitations associated with these data).
However, paleopathology is primarily concerned with the
differential diagnosis of pathologies in individual skele-
tons, establishing the antiquity of specific diseases, or
documenting the presence of particular conditions in past
populations via case studies of one or a few individuals
(Boldsen and Milner, 2012). Paleoepidemiology, as
detailed above, focuses on populations as the unit of anal-
ysis, and though some practitioners might not entirely
agree with Goodman’s (1993: 282) claim that “paleoepi-
demiologists are rarely interested in individuals,” it is cer-
tainly true that from an analytical and interpretive
perspective, individuals are of interest because they con-
tribute to the observed aggregate patterns (Milner and
Boldsen, 2017).
Because of these different scales of focus, paleopa-
thology and paleoepidemiology typically use different
analytical approaches. Paleopathology tends to be more
descriptive, whereas paleoepidemiology applies quantita-
tive analyses to a greater extent (indeed, quantitative anal-
yses are impossible to apply to paleopathological case
studies involving isolated individuals). Like epidemiol-
ogy, paleoepidemiology is inherently comparative; in
order to interpret the broader implications of the presence
of pathologies, rates of pathological lesions are compared
in paleoepidemiological studies between groups, such as
male versus female, urban versus rural, or high status ver-
sus low status. Paleopathology, however, can be success-
fully done without the application of a comparative
framework. Milner and Boldsen (2017) emphasize the
unique paleoepidemiological focus on estimating the risks
of death associated with skeletal pathologies. Their defini-
tion of paleoepidemiology is inherently demographic.
Informative paleopathological research does not necessar-
ily require information beyond the presence (or absence)
of pathology, and thus paleopathology can be done inde-
pendently of demographic data.
Paleoepidemiology and the Osteological
Paradox
The focus on population-level health and disease dynam-
ics in paleoepidemiology provides scholars in the field
the opportunity to actively engage with and attempt to
resolve some of the issues associated with the osteological
paradox, which was described over 25 years ago by
Wood et al. (1992). The osteological paradox centers
around two important phenomena: heterogeneous frailty
and selective mortality. Frailty, in this context, refers to
the age-standardized relative risk of death (Vaupel et al.,
1979). Variation in frailty (i.e., heterogeneous frailty)
exists in populations because of a variety of factors, such
as differences in immune competence (associated with
nutritional status, genetic variation in regions of the
genome associated with immunity or disease susceptibil-
ity, the effect of sex hormones, etc.), differences in risk-
taking behavior (e.g., smoking or heavy drinking) or
exposure to occupational hazards, or variation in exposure
to disease vectors or environmental pollution. Wood et al.
emphasized the potential for “hidden” heterogeneity in
frailty to complicate reconstructions of health from skele-
tal samples. Epidemiologists, because they have access to
 A Brief History and 21st Century Challenges Chapter | 2
observational, interview, and clinical data from living
people, can potentially identify and control for numerous
factors known or suspected to influence frailty. However,
even in living populations, not all sources of variation in
frailty are known and thus controlled for in studies of
population health. This problem of hidden heterogeneity
is exacerbated when we rely on biased samples of the
dead for whom behavioral and clinical information is
nearly or (more often) totally nonexistent. Without the
ability to control for many potential sources of variation
in frailty in skeletal samples, we cannot be certain that
the aggregate patterns we observe in these samples accu-
rately represent the health or disease experiences of all of
the subgroups that comprise the larger sample.
Heterogeneous frailty strongly influences the composi-
tion of the skeletal samples. With respect to many causes
of death, mortality does not behave indiscriminately, kill-
ing all individuals at each particular age at the same rate.
Instead, mortality is often selective: disproportionately
affecting individuals with the highest frailty at each par-
ticular age. It is these individuals, with the highest frailty,
who are most likely to become part of the skeletal sam-
ples that are eventually available to paleoepidemiologists.
This phenomenon makes it difficult, if not impossible, to
estimate the prevalence of conditions in once-living popu-
lations based on the observed frequencies of associated
pathologies in a skeletal sample, particularly if those con-
ditions are associated with elevated risks of mortality.
Using this approach would tend to result in the overesti-
mation of the prevalence of the causative conditions.
Because of the potential effects of heterogeneous frailty
and selective mortality, Wood et al. urge caution in the
interpretation of health from observations of skeletal
pathologies or stress markers, particularly avoiding the
conventional assumption that the presence of skeletal
pathologies is an indicator of poor health and a lack
thereof reflects good health.
As had previously been addressed by Angel (1975),
Ortner (1991,1992), and Harpending (1990), Wood et al.
discussed the relationship between skeletal lesion forma-
tion and survivorship. Specifically, they raise the possibil-
ity that because skeletal pathologies take time to form,
they might, at least in some cases, indicate relatively good
health rather than high frailty. That is, the presence of a
skeletal pathology reflects survival, at least temporarily,
with or beyond the causative condition. The absence of
pathology might indicate relatively poor health if indivi-
duals without skeletal lesions succumbed to illness,
trauma, or malnutrition and died before the lesions could
form. Wood et al. (1992) do not argue that stress markers
are necessarily or even typically reflective of good health;
instead they urge scholars to consider multiple, equally
plausible, interpretations rather than uncritically hewing to
conventional interpretations that might not be appropriate.
17
One way to address these fundamental difficulties is to
leverage aggregate demographic data to assess the effects
of skeletal pathologies (Milner and Boldsen, 2017). Rather
than making assumptions about how pathologies reflect
health, paleoepidemiologists can establish whether (in a
particular context) a positive association exists at the pop-
ulation level between a skeletal pathology and risk of
death (or a negative association exists between the pathol-
ogy and survival). Such an association would support
interpretations of the pathology as an indicator of poor
health. This approach reduces uncertainty about what skel-
etal pathologies indicate about health at the population
level, but we must still be cautious about the inferences we
make for individuals in the sample. Estimation of survivor-
ship or the risk of death associated with pathologies is
only possible using paleoepidemiological data; it cannot
be done using isolated individuals. This approach requires
a comparative approach and access to information about
the demographic outcomes for people with and without
pathologies. With structured aggregate data, paleoepide-
miologists are also in a position to directly assess hetero-
geneous frailty, as least with respect to those factors that
are detectable in the skeleton or burial context, such as
age, sex, social status, or nutritional status. Being able to
compare mortality outcomes across these and similar cate-
gories does not entirely alleviate the problem of hidden
heterogeneity in frailty, but at the very least, paleoepide-
miologists can, with large enough samples, control for
some sources of heterogeneity that might otherwise con-
found reconstructions of population health. Examples of
paleoepidemiological research that have addressed the
osteological paradox include Boldsen’s (2005) study of the
association of skeletal indicators of leprosy and risk of
mortality in medieval Denmark; Wilson’s (2010, 2014)
assessment of the health and demographic effects of the
intensification of maize agriculture, the adoption of
Mississippian lifeways, and increased interpersonal vio-
lence and warfare in Illinois; and DeWitte and colleagues’
evaluation of selective morality during the medieval Black
Death in London (DeWitte and Hughes-Morey, 2012;
DeWitte and Wood, 2008).
REFERENCES
Angel, J., 1975. Paleoecology, paleodemography and health. In: Polgar,
S. (Ed.), Population, Ecology, and Social Evolution. The Hague,
Molton, pp. 167 190.
Angel, J.L., 1981. History and development of paleopathology. Am. J.
Phys. Anthropol. 56 (4), 509 515.
Armelagos, G., 1997. Paleopathology. In: Spencer, F. (Ed.), History of
Physical Anthropology, 2, M-Z. Garland, New York, pp. 790 796.
Armelagos, G., 2003. Bioarchaeology as anthropology, Archaeological
Papers of the American Anthropological Association, vol. 13.
pp. 27 40.
18 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
Armelagos, G.J., Cohen, M.N., 1984. Paleopathology at the Origins of
Agriculture. Academic Press, Orlando, FL.
Aufderheide, A.C., Rodrı́guez-Martı́n, C., 1998. The Cambridge
Encyclopedia of Human Paleopathology. Cambridge University
Press, Cambridge, UK.
Blatt, S.H., 2015. To swaddle, or not to swaddle? Paleoepidemiology of
developmental dysplasia of the hip and the swaddling dilemma
among the indigenous populations of North America. Am. J. Hum.
Biol. 27, 116 128.
Bocquet-Appel, J.P., Masset, C., 1982. Farewell to paleodemography. J.
Hum. Evol. 11, 321 333.
Boldsen, J.L., 2001. Epidemiological approach to the paleopathological
diagnosis of leprosy. Am. J. Phys. Anthropol. 115, 380 387.
Boldsen, J.L., 2005. Leprosy and mortality in the medieval Danish vil-
lage of Tirup. Am. J. Phys. Anthropol. 126, 159 168.
Boldsen, J.L., Milner, G.R., 2012. An epidemiological approach to
paleopathology. In: Grauer, A.L. (Ed.), A Companion to
Paleopathology. Wiley-Blackwell, Oxford, pp. 114 132.
Bos, K., Schuenemann, V., Golding, G., Burbano, H., Waglechner, N.,
Coombes, B., et al., 2011. A draft genome of Yersinia pestis from
victims of the black death. Nature 478, 506 510.
Bos, K.I., Harkins, K.M., Herbig, A., Coscolla, M., Weber, N., Comas,
I., et al., 2014. Pre-Columbian mycobacterial genomes reveal seals
as a source of new world human tuberculosis. Nature 514, 494 497.
Bos, K.I., Herbig, A., Sahl, J., Waglechner, N., Fourment, M., Forrest, S.
A., et al., 2016. Eighteenth century Yersinia pestis genomes reveal
the long-term persistence of an historical plague focus. eLife 5,
e12994.
Brothwell, D.R., Sandison, A.T. (Eds.), 1967. Diseases in Antiquity: A
Survey of the Diseases, Injuries and Surgery of Early Populations.
Springfield, Ill.: C. C. Thomas.
Buikstra, J.E., 1999. Paleoepidemiology of tuberculosis in the Americas.
In: Pálfi, G., Dutourk, O., Déak, J., Hútas, I. (Eds.), Tuberculosis
Past and Present. Golden Book Publishers and Tuberculosis
Foundation, Budapest/Szeged, pp. 479 494.
Buikstra, J.E., Ubelaker, D. (Eds.), 1994. Standards for Data Collection
from Human Skeletal Remains: Proceedings of a Seminar at the
Field Museum of Natural History. Arkansas Archaeological Survey
Press, Fayetteville.
Buikstra, J.E., Roberts, C.A. (Eds.), 2012. The Global History of
Paleopathology: Pioneers and Prospects. Oxford University Press.
Buikstra, J.E., Cook, D.C., Bolhofner, K.L., 2017. Scientific rigor in
Paleopathology. In: Buikstra, J.E. (Ed.) Rigor in Paleopathology. pp.
80-87. Special Issue of the International Journal of Paleopathology,
19, 80 141.
Cuvier, G., 1820. Recherches sur les Ossemens Fossiles, vol. 4. Dufour,
Paris.
DeWitte, S.N., 2018. Stress, sex, and plague: patterns of developmental
stress and survival in pre- and post-black death London. Am. J.
Hum. Biol. 30. Available from: http://dx.doi.org/10.1002/
ajhb.23073.
DeWitte, S.N., Hughes-Morey, G., 2012. Stature and frailty during the
black death: the effect of stature on risks of epidemic mortality in
London, A.D. 1348 1350. J. Archaeol. Sci. 39, 1412 1419.
DeWitte, S.N., Wood, J.W., 2008. Selectivity of black death mortality
with respect to preexisting health. Proc. Natl. Acad. Sci. U.S.A. 105,
1436 1441.
Donoghue, H.D., Spigelman, M., O’Grady, J., Szikossy, I., Pap, I., Lee,
O.Y.C., et al., 2015. Ancient DNA analysis—an established
technique in charting the evolution of tuberculosis and leprosy.
Tuberculosis 95, S140 S144.
Elliot-Smith, G., Wood-Jones, F. (Eds.), 1910. The Archaeological
Survey of Nubia Report for 1907-1908, Vol II: Report on the
Human Remains. National Printing Department, Cairo.
Esper, J. 1774. Ausführliche Nachricht von Neuentdeckten Zoolithen,
Unbekannter Vierfüssiger Thiere, und denen sie Enthaltenden, so
wie Verschiedenen Andern Denkwürdigen Grüften der
Obergebürgischen Lande des Marggrafthunis Bayreuth. Nürnberg.
Goodman, A., 1993. On the interpretation of health from skeletal
remains. Curr. Anthropol. 34, 281 288.
Gosse, L.A., 1861. Présentation d’un Crane défermé de Nahoa. Bulletins
de la Société d’Anthropologie de Paris 2, 567 577.
Grmek, M.D., 1983/1989. Diseases in the Ancient Greek World. Johns
Hopkins University Press, Baltimore, Maryland.
Hackett, C., 1963. On the origin of the human treponematoses (pinta,
yaws, endemic syphilis and venereal syphilis). Bulletin of the World
Health Organization 29, 7 41.
Harpending, H., 1990. Review of “health and the rise of civilization” by
MN Cohen. Am. Ethnol. 14, 799 800.
Harper, K.N., Zuckerman, M.K., Harper, M.L., Kingston, J.D.,
Armelagos, G.J., 2011. The origin and antiquity of syphilis revisited:
an appraisal of old world pre-Columbian evidence for treponemal
infection. Am. J. Phys. Anthropol. 146, 99 133.
Hooton, E.A., 1930. The Indians of Pecos Pueblo: A Study of Their
Skeletal Remains. Yale University Press, New Haven, CT.
Hrdlička, A., 1914. Special notes on some of the pathological conditions
shown by the skeletal material of the ancient Peruvians. Smithsonian
Miscellaneous Collections 61, 57 69.
Hudson, E., 1965. Treponematosis and man’s social evolution. American
Anthropologist 67, 885 901.
Jarcho, S., 1966. Human Paleopathology. Yale University Press.
Jones, J. 1876. Explorations of the aboriginal remains of Tennessee. In
Smithsonian Contributions to Knowledge, 259.
Kacki, S., 2017. Influence de l’état sanitaire des populations du passé sur
la mortalité en temps de peste: Contribution à la paléoépidémiologie.
BMSAP 29, 202 212.
Klaus, H.D., Tam, M.E., 2009. Contact in the Andes: bioarchaeology of
systemic stress in Colonial Mórrope, Peru. Am. J. Phys. Anthropol.
138, 356 368.
Klaus, H.D., Spencer Larsen, C., Tam, M.E., 2009. Economic intensifi-
cation and degenerative joint disease: life and labor on the postcon-
tact north coast of Peru. Am. J. Phys. Anthropol. 139, 204 221.
Konigsberg, L.W., Frankenberg, S.R., 2013. Bayes in biological anthro-
pology. Am. J. Phys. Anthropol. 152, 153 184.
Lambert, P.M., 2012. Ethics and issues in the use of human skeletal
remains in paleopathology. In: Grauer, A.L. (Ed.), A Companion to
Paleopathology. Wiley-Blackwell, Chichester, West Sussex, UK,
pp. 17 33.
Larsen., C.S., 1997. Bioarchaeology: Interpreting Behavior from the
Human Skeleton. Cambridge University Press, Cambridge, UK.
Larsen, C.S., Griffin, M.C., Hutchinson, D.L., Noble, V.E., Norr, L.,
Pastor, R.F., et al., 2001. Frontiers of contact: bioarchaeology of
Spanish Florida. J. World Prehistory 15, 69 123.
Marciniak, S., Prowse, T.L., Herring, D.A., Klunk, J., Kuch, M.,
Duggan, A.T., et al., 2016. Plasmodium falciparum malaria in
1st 2nd century CE Southern Italy. Curr. Biol. 26, R1220 R1222.
Mays, S., 2018. How should we diagnose disease in palaeopathology?
Some epistemological considerations. Int. J. Paleopathol. 20, 12 19.
 A Brief History and 21st Century Challenges Chapter | 2
Mendonça de Souza, S.M.F., Carvalho, D.Md, Lessa, A., 2003.
Paleoepidemiology: is there a case to answer? Mem. Inst. Oswaldo
Cruz 98, 21 27.
Milner, G.R., Boldsen, J.L., 2012. Transition analysis: a validation study
with known-age modern American skeletons. Am. J. Phys.
Anthropol. 148, 98 110.
Milner, G.R., Boldsen, J.L., 2017. Life not death: epidemiology from
skeletons. Int. J. Paleopathol. 17, 26 39.
Mitchell, P.D., 2011. Retrospective diagnosis and the use of historical texts
for investigating disease in the past. Int. J. Paleopathol. 1, 81 88.
Mitchell, P.D., 2012. Integrating historical sources with paleopathology.
In: Grauer, A.L. (Ed.), A Companion to Paleopathology. Wiley-
Blackwell, Chichester, West Sussex, UK, pp. 310 323.
Mitchell, P.D. 2017. Improving the use of historical written sources in
paleopathology. In Rigor in Paleopathology: Perspectives from
across the Discipline, J. E. Buikstra, ed., International Journal of
Paleopathology 19, 88 95.
Moodie, R., 1923. Paleopathology: An Introduction to the Study of
Ancient Evidences of Disease. University of Illinois Press, Urbana.
Morse, D., 1969. Ancient disease in the Midwest. Springfield, Illinois
State Museum Reports of Investigations, No. 15.
Ortner, D.J., 1991. Theoretical and methodological issues in paleopathol-
ogy. In: Ortner, D.J., Aufderheide, A.C. (Eds.), Human
Paleopathology: Current Syntheses and Future Options. Smithsonian
Institution Press, Washington, DC, pp. 5 11.
Ortner, D.J., 1992. Skeletal paleopathology: probabilities, possibilities,
and impossibilities. In: Verano, J.W., Ubelaker, D.H. (Eds.), Disease
and Demography in the Americas. Smithsonian Institution Press,
Washington, DC, pp. 5 13.
Ortner, D.J., 2003. Identification of pathological conditions in human
skeletal remains. Academic Press, San Diego, CA.
Ortner, D.J., Aufderheide, A. (Eds.), 1991. Human Paleopathology:
Current Syntheses and Future Options. Smithsonian Institution
Press, Washington, D. C.
Ortner, D.J., Putschar, W.G.J., 1981. 1985 Identification of pathological
conditions in human skeletal remains. Smithsonian Institution Press.
Pales, L., 1930. Paleopathologie et Pathologie Comparative. Masson et
Cie, Paris.
Patterson Ross, Z., Klunk, J., Fornaciari, G., Giuffra, V., Duchêne, S.,
Duggan, A.T., et al., 2018. The paradox of HBV evolution as
revealed from a 16th century mummy. PLoS Pathog. 14, e1006750.
Ragsdale, B.D., Madewell, J.E., Sweet, D.E., 1981. Radiologic and
Pathologic Analysis of Solitary Bone Lesions, Part II: Periosteal
Reactions. Radiologic Clinics of North America 19, 749 783.
Resnick, D., Niwayama, G., 1995. Osteomyelitis, septic arthritis, and
soft tissue infection: organisms. In: Resnick, D. (Ed.), Diagnosis of
Bone and Joint Disorders, third ed. W. B. Saunders, Edinburgh,
pp. 2467 2474.
Ruffer, M., 1910. Remarks on the histology and pathological anatomy of
Egyptian mummies. Cairo Sci. J. 1 5.
Shufeldt, R.W., 1892. Notes on paleopathology. Popular Science
Monthly 42, 679 684.
Smith-Guzmán, N.E., 2015. The skeletal manifestation of malaria: an
epidemiological approach using documented skeletal collections.
Am. J. Phys. Anthropol. 158, 624 635.
Snoddy, A.M.E., Buckley, H.R., Halcrow, S.E., 2016. More than meta-
bolic: considering the broader paleoepidemiological impact of vita-
min d deficiency in bioarchaeology. Am. J. Phys. Anthropol. 160,
183 196.
19
Steinbock, R., 1976. Paleopathological Diagnosis and Interpretation.
Charles C. Thomas, Springfield, IL.
Ubelaker, D.H., 1982. The Development of Human Paleopathology.
In: Spencer, F. (Ed.), A History of American Physical Anthropology
1930-1980. Academic Press, New York, pp. 337 356.
Vågene, Å.J., Herbig, A., Campana, M.G., Robles Garcı́a, N.M.,
Warinner, C., Sabin, S., et al., 2018. Salmonella enterica genomes
from victims of a major sixteenth-century epidemic in Mexico. Nat.
Ecol. Evol. 2, 520 528.
Vaupel, J.W., Manton, K.G., Stallard, E., 1979. The impact of heteroge-
neity in individual frailty on the dynamics of mortality. Demography
16, 439 454.
Virchow, R., 1898. Knochen aus Alten Gräbern von Tennessee.
Verhandlungen der Berliner Gesellschaft für Anthropologie 30,
342 344.
Waldron, T., 2007. Paleoepidemiology: The Measure of Disease in the
Human Past. Left Coast Press Inc, Walnut Creek, CA.
Warren, J., 1822. A Comparative View of the Sensorial and Nervous
Systems in Man and Animals. Ingraham, Boston.
Wells, C., 1964. Bones, Bodies and Disease. Thames and Hudson,
London.
Weston, D.A., 2008. Investigating the specificity of periosteal reactions
in pathology museum specimens. Am. J. Phys. Anthropol. 137,
48 59.
Weston, D.A., 2012. Nonspecific Infection in Paleopathology:
Interpreting Periosteal Reactions. In: Grauer, A.L. (Ed.), A
Companion to Paleopathology. Wiley-Blackwell, Chichester, West
Sussex, UK, pp. 492 512.
Williams, H.U., 1929. Human paleopathology, with some original obser-
vations on symmetrical osteoporosis of the skull. Arch. Pathol. 7,
839 902.
Wilson, J.J., 2010. Modeling Life Through Death in Late Prehistoric
West-Central Illinois: An Assessment of Paleodemographic and
Paleoepidemiological Variability [PhD]. Binghamton University,
SUNY, Binghamton, NY.
Wilson, J.J., 2014. Paradox and promise: Research on the role of recent
advances in paleodemography and paleoepidemiology to the study
of “health” in Precolumbian societies. Am. J. Phys. Anthropol. 155,
268 280.
Wood-Jones, F., 1908. The pathological report. Archaeological Survey
of Nubia Bulletin 2, 55 69.
Wood-Jones, F., 1910. Anatomical variations, and the determination of
the age and sex of skeletons. In: Elliot-Smith, G., Wood-Jones, F.
(Eds.), The Archaeological Survey of Nubia Report for 1907 1908,
Vol II: Report on the Human Remains. National Printing
Department, Cairo, pp. 221 262.
Wood, J.W., Milner, G.R., Harpending, H.C., Weiss, K.M., 1992. The
osteological paradox: Problems of inferring prehistoric health from
skeletal samples. Curr. Anthropol. 33, 343 370.
World Health Organization, 2018. Health Topics: Epidemiology.
,http://www.who.int/topics/epidemiology/en/. (accessed
15.01.18.).
Zuckerman, M.K., 2014. Modern Environments and Human Health:
Revisiting the Second Epidemiological Transition. Wiley-Blackwell,
Hoboken, NJ.
Zuckerman, M.K., Harper, K.N., Armelagos, G.J., 2016. Adapt or die:
three case studies in which the failure to adopt advances from other
fields has compromised paleopathology. Int. J. Osteoarchaeol. 26,
375 383.
Chapter 3
Themes in Paleopathology
Anne L. Grauer1 and Jane E. Buikstra2
1
Loyola University Chicago, Chicago, IL, United States, 2Arizona State University, Phoenix, AZ, United States
The identification and diagnosis of pathological conditions
in human skeletal remains is a key component of paleopa-
thology. As discussed in Chapters 1 and 5, detailed
descriptions of lesions and the application of differential
diagnoses have allowed researchers to more closely align
skeletal lesions with the clinical manifestations of disease.
A number of themes emanate from interpretations of
pathological lesions, which extend beyond diagnosis. In
Chapter 1, we discussed the need for paleopathologists to
appreciate perspectives and theories drawn from the social
sciences and humanities. Especially important are cautions
about attributing social status based exclusively on counts
of grave wealth without consideration of broader contextual
issues. This approach became popular, especially in
American archeology, during the 1960s with the work of
Louis Binford, Arthur Saxe, and James Brown. Binford
(1971), working within a cross-cultural processual paradigm
(see also Carr, 1995) and using subsistence as a proxy for
social complexity, argued that dimensions of mortuary
behavior correlated predictably with social status. Arthur
Saxe’s (1970) unpublished dissertation also interrogated the
funereal ethnographic record cross-culturally. His
“Hypothesis 8,” which explores the relationship between
the presence of formal cemeteries and resource ownership
has been expanded and applied by researchers, including
Charles and Buikstra (1983), Goldstein (1980), and Morris
(1991). During the 1980s, a “postprocessual” rebuttal of
processual archeology, including mortuary studies (Hodder,
1982, 1984), led to diminished visibility for funerary arche-
ology (Rakita and Buikstra, 2005), in spite of its important
role in paleopathological analyses.
Despite cautionary tales, some researchers continue to
equate the quality and quantity of grave goods with levels
of social status, and tie these closely to the presence of
pathological conditions, often without considering other
contextual factors (Grauer, 2019). However, research such
as that conducted on the Andean Chiribaya peoples of
Peru (AD 800 1350), serves as a great example of the
importance of nuanced interpretations of funerary
Ortner’s Identification of Pathological Conditions in Human Skeletal Remains. DOI: https://doi.org/10.1016/B978-0-12-809738-0.00003-X
© 2019 Elsevier Inc. All rights reserved.
behavior. Here, we find elaborate subsurface tombs with
metals, feathers, and other material culture in sites such
as Chiribaya Alta (Buikstra, 1995). Contemporary inland
Chiribaya sites at higher elevations, such as Yaral
(1000 m above sea level), present far less elaborate tombs
with small numbers of ceramics and other vessels. Before
beginning to assume greater economic wealth (and thus,
status) among coastal Chiribaya, however, we need to
appreciate the presence of elaborate public buildings at
Yaral, where public rituals likely took place. Rather than
Yaral graves serving as symbols of wealth or status, per-
haps the grand public spaces are where social differences
were displayed. Hence, we must avoid embracing counts
of material items within tombs as reflecting the status of
the dead. As with the dedicatory offerings to the Maya
woman in the “Margarita” tomb at Copán, grave accom-
paniments may reflect pilgrimages of mourners, who—in
this case—elevated and probably changed the status of
the decedent from a biological to a primordial progenitor
of the Copán royal dynasty during the Classic period
(BAD 400 800). Clearly, the interpretation of social sta-
tus requires careful and nuanced interpretations of archeo-
logical and historical contexts.
Social status is only one aspect of human identity that
impacts and informs paleopathological analysis. Gender,
age, religion, ethnicity, and disability, inferred from
bioarcheological data, also play important roles in paleo-
pathological research (Grauer, 2018). In the following
sections we discuss social attributes increasingly synthe-
sized with paleopathological research: sex, gender, and
age; and discuss research into structural violence, disabil-
ity, and care, and the role of individual and populational
studies within paleopathology.
SOCIAL AND IDENTITY THEORY
For paleopathologists and bioarcheologists, bones serve as
the nexus of interpretation, as pathological, biomechani-
cal, and physiological factors acting upon and within the
21
22 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
individual are evaluated alongside psychological and
social responses. Whether viewed within the fields of
sociology, psychology, or anthropology, identity studies
share a common goal: to explore and understand intersec-
tionality between an individual and larger social spheres.
Feminist and Gender Theory
A key dimension in the integration of social and identity
theory in paleopathological and bioarcheological research
is the influence of feminist and gender theory on skeletal
analysis (Grauer, 2018). Similar to the effects of proces-
sual and postprocessual paradigms (discussed earlier),
which profoundly influenced the interpretation of skeletal
remains, feminist and gender theories have evoked
changes to the definitions and the operationalization of
our concepts of sex and gender. Feminist theory is argued
to have developed in three waves. The first wave, stem-
ming from entrenched political and economic inequities
of the 19th and early 20th centuries that denied women
the right to vote, sought to fight for basic rights (Sanders,
2006; Sharlach, 2009). Its influence on skeletal analyses
would be felt decades later as women such as Mildred
Trotter, Lucille St. Hoyme, and Alice Brues, became pio-
neers in the growing field of physical anthropology
(Buikstra and Roberts, 2012). The second wave of the
feminist movement focused on social empowerment
(Baxandall and Gordon, 2005). Concepts of sex (based on
biological determinants) and gender (argued to be the
social role adopted by an individual based on sex) were
disentangled as a means to draw attention to and end
repression. This perspective deeply influenced skeletal
analyses, as it established binary definitions of sex and
gender: i.e., that there are two discrete biological sexes
that can be identified skeletally, and based on ascribed
sex, there are two polemical social roles (one masculine,
one feminine) that become socially enacted.
Investigation into skeletal manifestations of sexual
dimorphism bolstered this perspective. As osteologists
isolated key anatomical features, such as the sciatic notch
and subpubic angle, which qualitatively differed in adults,
and developed quantifiable measures of variance to test
predictability, the perceived chasm between female and
male widened. Paleopathological studies increasingly
mapped the presence of lesions onto female and male ske-
letons. As a careful control, skeletons in too poor condi-
tion for adequate evaluation, or morphologically falling
into the “undeterminate,” category, were often excluded
from analyses as a means to clearly discern patterns of
disease by sex.
This approach proved productive. Cribra orbitalia and
porotic hyperostosis, for instance, often associated with
the presence of iron-deficiency anemia (Stuart-Macadam,
1987), were predicted and found to be common in female
skeletons. The cause, it was asserted, was females’ fluctu-
ating physiological need for iron due to pregnancy, lacta-
tion, and menstruation (e.g., Cybulski, 1977; Webb,
1984). Social roles of females were also argued to con-
tribute to sex-demarcated patterns of skeletal lesions.
Biological differences between females and males were
expected to be exacerbated by an assumed ubiquitous
gender hierarchy, whereby males held power and main-
tained access to resources (Cohen and Bennett, 1993),
leaving females vulnerable to fluctuating access to nutri-
tion. Blood loss, alongside disruption to dietary iron
acquisition and absorption, would therefore contribute to
higher rates of iron-deficiency anemia in females. The
entangled effects of sex and gender have also been
explored using nonspecific indicators of stress, such as
enamel hypoplasias. The presence of childhood growth
disruption, mapped onto sex in adult skeletons, might
infer differential treatment of children based on sex, but
varying rates of enamel hypoplasias have alternatively
been used to explore the axiom of male biological vulner-
ability (Stinson, 1985) The argument here is that females,
due to demands of pregnancy and lactation, have evolu-
tionarily been selected to physiologically buffer the
effects of fluctuating environments, rendering males, in
comparison, more vulnerable. Tackling this precept head-
on, Guatelli-Steinberg and Lukacs (1999), in their meta-
analysis of human and nonhuman primates conclude that
“in most studies, sex differences in EH prevalence are sta-
tistically nonsignificant. However, when sex differences
are significant, there is a slight trend for them to be
greater in males than in females, suggesting a weak influ-
ence of greater male vulnerability. Cultural practices of
sex-biased investment in children appear to have greater
impact on EH expression than does male vulnerability/
female buffering” (Guatelli-Steinberg and Lukacs, 1999:
73).
Innate differences between females and males, and
their consequences for the paleopathological record,
played a prominent role in Ortner’s second edition of this
volume (2003: 114 118). Focus was placed on skeletal
indicators of infectious disease and human immune reac-
tivity. Ortner asserted that bearing children posed an
undeniable biological risk for women that was exacer-
bated by the effects of agriculture: i.e., sedentism and its
concomitant increase in infection. Culturally determined
differential access to food, especially during times of fam-
ine, placed women in a particularly precarious position.
What were the skeletal outcomes of these conditions?
Ortner’s model predicting paleopathological ramifications
of differing male and female immune reactivity tackled

these issues of sex, gender, and the skeletal record. Ortner
(2003: 115 116) contended that:
1. “If all factors were equal,” meaning that social and/or
environment variables influencing health were the
same for women and men, then “women might be
expected to survive to the chronic stage of infectious
disease more often than men.”
2. “Given the known sex difference in immune reactiv-
ity, the male and female subsamples of the population
might be arranged as two partially superimposed, nor-
mal distributions (Fig. 3.1). The mean (X1) of the
male subsample would be positioned more toward the
poor end of the scale than the female mean (X2).”
3. “An additional variable is the hypothetical range on
the immune response scale where skeletal involvement
occurs. At the poor end of the scale we may designate
as R1 the point below which skeletal involvement does
not occur and death is the typical event with R2 as the
point on the other end of the scale beyond which skel-
etal manifestations do not occur because of complete
recovery from infectious disease.”
4. “If, for the moment, we assume that all other variables
(most particularly, exposure to infectious agents) that
affect the expression of infectious disease in the
human skeleton are constant and the distribution
approximates normality, it is apparent that the position
and range of X1 and X2 and R2 on the scale will affect
the sex ratios for the prevalence of infectious disease
in the skeletal sample.”
5. When modern clinical male/female ratios in various
infectious diseases are evaluated, “they do tend to
cluster and consistently show greater male morbidity
for infection.” When the prevalence of periostitis is
Frequency
R1 X1 R2X2
Poor Good
Immune response
FIGURE 3.1 Graph showing hypothetical male and female distribu-
tions on an immune response scale from poor to good. X 1 designates the
mean of the male distribution and X 2 the mean of the female distribu-
tion. R1 is a theoretical point below which skeletal manifestations of dis-
ease do not occur and death is a typical event. R2 is the point on the
other end of the scale where no skeletal disease occurs because of com-
plete recovery from infectious disease. R1 and R2 are positioned to create
a male/female ratio of approximately 3:1, similar to what one finds in
the clinical ratio of infectious disease.
Themes in Paleopathology Chapter | 3 23
examined within archeological populations, males fre-
quently display higher rates of morbidity compared to
women (p. 116).
Ortner’s model crystallized the perceived dichotomy
between biological sex and socially established notions of
gender. His conclusions, however, extend beyond sex- or
gender-based determinism. For instance, he asserted that,
“males have higher morbidity than females. One question
is whether this is simply the result of greater exposure to
infectious agents among males or a more effective
immune response among females. In some infectious dis-
eases, such as mycotic infection, a case can be made for
greater exposure to infectious agents by males in at least
some agricultural societies. However, in at least some cul-
tural contexts and in some age ranges, males and females
seem to have equal exposure to infectious agents but
males seem more vulnerable to disease” (p. 116). Hence,
while Ortner implies that sex and gender can be indepen-
dently isolated, he does not depend on codified gender
roles based on sex to explain differential morbidity. He
recognizes social complexity and its impact on disease.
So perhaps the perceived complexity of social roles,
along with definitions and applications of our concepts of
“sex” and “gender” need revision. Enter third-wave femi-
nist, gender, and queer theory. These paradigms focus on
dispelling heteronormative assumptions, and often refocus
attention onto experiences of the individual. The para-
digms assert that biological sex is not inextricably linked
to an immutable gender role, and gender roles are neither
binary nor absolute. For instance, Sofaer argues that “col-
lapsing sex and gender renders associations between bod-
ies and objects unproblematic. . .” (Sofaer, 2006: 101) and
that “in practice the classification of gender often tends to
assume that gender is stable thereby precluding the fluid-
ity that is a particularly useful element of the concept and
that is inherent in understanding it as culturally
dependent. . .” (Sofaer, 2006: 100). Gender, it is asserted,
is an element of identity, and is malleable and continually
negotiated throughout the life course of the individual.
Adding to an even greater complexity, gender identity is
simultaneously personal and social. An individual’s emic
gender identity (an internalized identity) is shaped and
may differ from their etic identity (attitudes, views, or
interpretations of the individual made by others).
Paleopathological research is impacted by these para-
digmatic shifts, with studies of skeletal trauma leading the
change due to the perceived direct role that behavior
plays in its etiology. Although interpersonal aggression,
violence, and warfare have long been viewed as direct
manifestations of gendered behavior (Holliman, 2011),
the promise of the new theoretical direction lies in the
reinterpretation of the causes and cultural meaning of
traumatic lesions. Notable contributions include Knüsel
24 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
(2011), whose rigorous archeological contextual analysis
of British medieval warfare and differential diagnosis of
humeral medial epicondylar avulsion fractures is accom-
panied by careful exploration of the construction and
effects of masculinity from childhood to adulthood. Work
by Bengtson and O’Gorman (2017) warns us against
adopting the deterministic view of sex and gender roles,
as warfare need not be limited to men and inextricably
tied to masculinity. Their analysis of mortuary data, sub-
sistence, and skeletal data from prehistoric sites in the
Central Illinois River Valley suggests that “Morton
Village women may have regularly and actively partici-
pated in violent encounters as part of their engagement
with the broader socio-politics of the region without being
formally celebrated as warriors in their mortuary disposi-
tion” (Bengtson and O’Gorman, 2017).
The constructed dichotomy between victim and assail-
ant is also being reevaluated in light of feminist and gender
theory. Tung (2014), in her analysis of gender-based vio-
lence against women during Wari imperial rule (AD
600 1100) and post-Wari rule (AD 1100 1400) of south-
ern Peru, argues that “a temporal view of the frequency
and patterning of violence among males and females may
illuminate how social norms related to violence may have
changed from one cultural era to the next, if they changed
at all. In particular, a bioarchaeological study that consid-
ers the role of gender in when and how violence is enacted
can further clarify how one’s sex structured the likelihood
that one might become a victim of violence and whether or
not the violence would be deadly” (Tung, 2014: 335).
Martin et al. (2010) and Harrod and Martin (2014) opera-
tionalize this approach by closely examining the types and
patterns of traumatic injury in ancestral Pueblo skeletal
remains from La Plata (AD 850 1150) and integrating
their data with a carefully constructed model that reflects
proposed social, economic, environmental, demographic,
and archeological manifestations of human subordination
and captivity. They, like a growing number of paleopathol-
ogists and bioarcheologists, are leading the way toward the
development of nuanced and highly contextual interpreta-
tions of engendered violence and aggression (see discus-
sion of structural violence below).
New feminist and gender paradigms also impact
research into the paleopathology of infectious disease. As
emphasized by Zuckerman and Crandall (unpublished
manuscript), “stigmatized and morally-loaded diseases,
including syphilis and leprosy (Hansen’s Disease), present
profound opportunities for detecting the play of sex, gen-
der, and sexuality in the spaces between biological, histor-
ical, and archaeological data,” and “in some instances,
using a biocultural approach and embedding skeletal data
into a highly specific archaeological and historical inter-
pretive framework can reveal inconsistencies that generate
novel, otherwise inaccessible insights into the effects of
past ideologies into the lived experiences of disease, sex,
gender, and sexuality.” In DeWitte and Stojanowski’s
(2015) evaluation of the impact and reaction within paleo-
pathology to the osteological paradox (Wood et al.,
1992), they argue that “frailty—and its causes and conse-
quences—is actually one of the more intriguing topics in
the health sciences today, one to which researchers across
a number of domains contribute. Furthermore, under-
standing the nature of human frailty and how it relates to
social inequality and social complexity is a highly rele-
vant topic that crosscuts disciplinary boundaries”
(DeWitte and Stojanowski, 2015: 428). Applying this dic-
tate, Yaussy et al. (2016) examine the effects of famine
and its influence on frailty using attritional and famine
burials denoted in the medieval cemetery of St. Mary
Spital in London. They conclude that “the significant
association between sex and periosteal lesions suggests
that some aspects of life were different for the two sexes,
resulting in different exposures to traumas, infections, or
other stressors” (Yaussy et al., 2016: 279). Moving
beyond the common binary construction of sex and gen-
der, they warn that confounding factors must be taken
into account. These include socioeconomic and gender-
based decisions of medieval men and women to migrate
to urban centers, which influences the sex ratios of skele-
tal populations and statistical decisions made by research-
ers which render the detection of sex differences in
mortality across the human life span impossible.
The Intersectionality of Sex, Gender, and
Age
When the biological and social ramifications of sex and
gender are examined, the inextricable variable of age
becomes evident. Most recently, this recognition has been
manifested in the development of a life course approach
in paleopathology (see Gilchrist, 2000; Gowland and
Knusel, 2006). Since, as Sofaer (2011) asserts, humans
embody both a chronological and socially defined age,
the life course approach seeks to address the intersection-
ality of sex, gender, multidimensionally defined age, and
health and disease. Focusing on childhood, for instance,
has allowed researchers interested in human disease to
appreciate the complex interactions between childhood
biological development and children’s dynamic social
spaces and roles (see Halcrow and Tayles, 2008, 2011;
Perry, 2008; Lewis, 2007; Mays et al., 2017; Gowland
and Penny-Mason, 2018). Penny-Mason and Gowland
(2014: 185), for instance, in their assessment of over 4600
British medieval skeletons determined to be less than 16
years old at time of death, find that “those aged 6 11
years exhibited similar levels of disease and trauma to
0 5 year olds, suggesting that although children were

developing into adult roles, the majority were likely to
have experienced an extended period of childhood roles
into puberty.” Conversely, Barrett and Blakey (2011) find
that childhood mortality rates were high for enslaved
Africans buried in the 18th century New York African
Burial Ground and that in 1731, an 11-year-old enslaved
African was categorized as an “adult,” thereby profoundly
affecting children’s exposure to trauma and pathogens.
Adopting a life course approach has also been influen-
tial in the exploration of the relationships between age, sex,
gender, and disease in adults. Grauer et al. (1999) for
instance, noted in their inquiry into the interplay between
sex, gender, and the detectable presence of disease in skele-
tons from the 19th-century Dunning Poorhouse Cemetery,
that no statistical differences appeared between adult
females and males in the frequencies of lesions such as
porotic hyperostosis, periosteal reaction, enamel hypopla-
sias, and fractures. However, when lesions were mapped
onto age at death, and social history was carefully interwo-
ven, different results emerged. “Young women, it appears,
were not entering and dying in the poorhouse with a legacy
of childhood anemia, bouts of infections, evidence of endur-
ing severe nonspecific stress, and poor dental health. More
likely, they were dying of acute conditions contracted
shortly before their deaths. Their presence in the cemetery
sample suggests that poor health and difficult childhoods
did not bring them to the facility, and that regardless of rea-
sonable health upon entering, their prolonged (if not eternal)
residency put their lives in jeopardy” (Grauer et al., 1999:
161). As Agarwal asserts, “it must be understood that socio-
cultural influences on the body are not layered on top of the
primary influences of sex and age; rather, they mold and
determine the sex- and age-related trajectory of bone health.
Although this makes the analysis of skeletal variation in
bone maintenance and loss harder to complete, it widens
the potential to visualize skeletons and bodies that are the
result of developmental processes that have acted at the
level of the individual, generations, or entire communities.
This has great relevance for how bioarchaeologists observe
variation in not only bone maintenance but also all aspects
of bone morphology, as well as how we reconstruct age and
gendered identity in the past” (Agarwal, 2012: 331).
Skeletal studies evaluating disease frequency or lesion
susceptibility based on sex appear to rely on two compet-
ing premises: human females are more susceptible to
some diseases such as hematopoietic disorders and gener-
alized bone loss due to reproductive demands (pregnancy,
lactation, menstruation) and fluctuating hormone levels,
while simultaneously being more “naturally” resistant to
infectious diseases due to greater immune responsivity.
Key variables, however, are often overlooked: changing
social and biological effects of the life course and negoti-
ated gender identity. Recent work seeks to tackle these
inextricable associations. Agarwal and Beauchesne (2011)
Themes in Paleopathology Chapter | 3 25
and Agarwal (2016), for instance, examine the plasticity
of bone development and maintenance using variables
such as body build, diet and activity, production of sex
hormones, and hereditary factors, and overlay them onto
the human life course: fetal life, childhood and adoles-
cence, young adulthood, and middle-older adulthood.
Qualitative and quantitative aspects of vertebral trabecular
bone microstructure were assessed from a British rural
medieval skeletal population and from two urban medie-
val sites dated from approximately the 11th 16th centu-
ries. Young women in rural environments, where higher
parity appears to occur, display greater bone loss during
reproductive years. Nevertheless, this does not appear to
impact long-term bone maintenance, perhaps due to the
physical demands of rural life, as witnessed by similar
bone density in older males and females. Hence, the com-
plexities of age, biology, sex, social and gender roles, and
environment, force us to recognize that skeletally deter-
mined sex cannot alone predict health outcomes.
To further confound us, current gender theories sug-
gest that inadvertent effects of second-wave feminist the-
ory, which emphasized differences between women and
men and led skeletal analysts to hone their ability to accu-
rately differentiate female and male skeletons, actually
limited our understanding of health and disease in the
past. Agarwal argues that “if gender is dynamic over the
life course, the reading of gendered patterns of bone loss
in past populations will inevitably be obscured by a static
mapping of gender to biological sex in skeletal analysis”
(Agarwal, 2012: 323). Can skeletons that fall between our
discrete sex categories provide us with information about
the past? The Developmental Origins of Health and
Disease Hypothesis (DOHaD), which focuses on pheno-
typic plasticity would have us saying, “Yes!” Built upon
Barker’s work (2002), which examined the influences of
fetal development and prenatal conditions on postnatal
health and disease, the DOHaD forces us to reevaluate the
biological and social precursors to sexual dimorphism.
Morphological differences between female and male ske-
letons may be influenced by many developmental and epi-
genetic factors. Removing from analysis adult skeletons
that emerge in-between morphological expectations of
female/male differences denies us the opportunity to
explore roles that fetal, childhood, and adolescent life
experiences may play upon the later life course
(Armelagos et al., 2009; Watts, 2013). Kirkpatrick (2000)
and Hollimon (2006, 2011) show us that there are numer-
ous ethnographic examples of social groups that include
labile third or fourth genders which are navigated, negoti-
ated, and performed throughout the life course. Hence,
adult skeletons whose sex cannot be morphologically
determined, and individuals whose social roles are not
heteronormative, are essential contributors to our under-
standing of health and disease in the past.
26 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
Indeed, Ortner’s (2003) ground-breaking model pre-
dicting paleopathological ramifications of differing male
and female immune reactivity can serve as a foundation
for new research informed by current feminist and gender
paradigms. Predicted disparities between female and male
immune reactivity can be tested. Do, for instance, skele-
tons with well-delineated sexual dimorphic features com-
ply with the expected immune responses? If so, does this
mean that ascribed biological sex is the cause of the cor-
relation? Might social, environmental, genetic, and devel-
opmental variables play key roles or even obscure
variation from the prediction? Do individuals falling
toward the center of the morphological spectrum between
female and male display similar patterns of periosteal
reaction? Might social, environmental, genetic, and devel-
opmental variables be key in interpreting these results?
How might interpretations of immune responses be influ-
enced by life course analyses? How might detailed and
highly contextualized understandings of human interac-
tions and experiences influence our interpretation of the
skeletal record? Clinical data indicate that immune
responses vary greatly over the human life span and in
reaction to the life course (Boraschi et al., 2013; Giefing-
Kröll et al., 2015), rendering Ortner’s ubiquitous snapshot
of sex differences a model from which new hypotheses
can be drawn.
STRUCTURAL VIOLENCE
Structural violence is a term that originated in peace stud-
ies (Galtung, 1969; see also Farmer et al., 2006; Klaus,
2012). It refers to social circumstances, frequently aspects
of social structures or institutions that keep individuals
from meeting basic needs—from a healthy existence. The
intimate relationship between structural and behavioral
violence is underscored by Gilligan (1996: 196), who
argues that the “question as to which of the two forms of
violence—structural or behavioral—is more important,
dangerous, or lethal is moot, for they are inextricably
related to each other, as cause to effect.”
While much has been published about behavioral vio-
lence detected by the presence of fractures and trauma in
past populations, only recently has the concept of struc-
tural violence been integrated into paleopathological
research. Its incorporation into skeletal analysis is an
important one, as it allows us to move beyond the recog-
nition of interpersonal aggression and begin to witness the
life-long and postmortem effects of social inequity
(Klaus, 2012). One growing body of research explores the
ramifications of human exploitation and marginalization
(Tegtmeyer and Martin, 2017). For instance, while the
term “structural violence” was not expressly used,
enslavement, low socioeconomic status, and other struc-
tural issues have figured heavily in de la Cova’s (2011,
2017) studies of documented collections. She began using
the term formally in 2012. Examining the skeletal remains
of individuals retained in the Hamann-Todd Human
Osteological Collection, the Terry Collection, and the
William Montague Cobb Collection, she argues that skel-
etal health disparities are evident between 19th-century-
born African Americans and Euro-Americans due to
“environmental conditions related to enslavement, post-
liberation migration to the industrialized North, crowded
urban living conditions, and poor sanitation” (de la Cova,
2011: 536). Such richly embedded studies hold excellent
promise for nuanced perspectives on the complex nature
of human health in situations wherein individuals are dis-
advantaged in circumstances beyond their control (de la
Cova, 2017).
The effects of structural violence are also revealed in
the postmortem treatment and disposition of human
remains. Blakely and Harrington (1997), Mitchell (2012),
and Nystrom (2017a: 16), are just a few of the researchers
who have explored the “systemic political, economic, and
social inequalities” that clearly influenced the bodies cho-
sen for autopsy or dissection, and the means by which
medical colleges, individuals, and organizations obtained
human remains. Autopsy was performed to understand the
cause of death or conditions impacting an individual.
Dissection was performed on individuals stripped of their
identity, rendering their bodies material objects, subjected
to experimentation and display. Adding to the complex
effects of structural violence is the fact that the racially
and socially biased use and collection of skeletal remains
in the past inherently affects our analyses today. These
individuals, who suffered the effects of structural violence
during and after their lives, problematically serve as the
baseline skeletal series long assumed to be representative
of human variability, and thus used for the development
of standards used today for estimating age-at-death and
sex (Nystrom 2014, 2017b).
ANCIENT HUMANS AND IMPAIRMENT,
DISABILITY, AND CARE
The presence of bone change associated with trauma or
disease tells us much about human life in the past by pro-
viding insight into the actions or force needed to fracture
bone, or the presence of pathogens responsible for lesions.
More recently, the presence of bone change has become
the foundation upon which evidence for impairment, dis-
ability, and care is extrapolated. Solecki, for instance, in
1971, concluded that Shanidar I, classified as Homo sapi-
ens neandertalensis, with “sustained injuries to the right
frontal squama, the left lateral orbit, the right humerus
and right fifth metatarsal. . . hypoplasia or atrophy of the
right clavicle, scapula, and humerus, osteomyelitis of the

right clavicle, degenerative joint disease at the right knee,
ankle, and first tarsometatarsal joint, and remodeling of
the left tibia” (Trinkhaus and Zimmerman, 1982: 61), was
“crippled” and at a “distinct disadvantage” in the harsh
Middle Paleolithic environment of modern-day Iraq
(Solecki, 1971). His ability to thrive was attributed to
group compassion and cooperation. As another example,
Stirland (1997: 588) argued that the presence of two
“remarkable examples of disabled individuals,” one dis-
playing juvenile polyepiphyseal disease and the other a
neuromuscular disorder with paraplegia, buried in a poor
medieval parish churchyard in England, implicitly sug-
gested that “care in the community” was not a modern
precept, as neither of the individuals would have survived
into adulthood unaided. Emphasis in these and many
ensuing studies is placed on situating the individual
within a larger social context. Hence, these rising themes
succeed in shifting emphasis away from case study
descriptions of lesions toward complex social interactions
and the surmised implications of trauma and disease on
the individual and community.
Important issues arise from paleopathological and
bioarcheological research into impairment, disability, and
care. The first involves definitions. What exactly constitu-
tes “impairment” or “disability”? In many published
works, the terms are used interchangeably. However, the
amalgamation assumes that there are universal cultural
norms which predicate our understanding of the terms.
Do both terms imply lack of mobility or the presence of
pain? Are they quantifiable? Tilley (1999: 3) asserts that
“‘Disability’ refers to a state (temporary or longer-term)
arising from an impairment in body function or structure
that is associated with activity limitations and/or partici-
pation restrictions. This state is given meaning by both
the individual and the community in relation to the life-
ways in which it is experienced.” Hence, the term
“impairment” may be used to explain the types or extent
of change in the body of an individual, such as the discor-
dant endochondral and intramembranous bone formation
of achondroplasia leading to alterations in body propor-
tions and subsequent morphological and biomechanical
complications. For the paleopathologist, types of move-
ment, ramifications of bone changes, and visual appear-
ance of the individual might be used to qualify and
quantify the term “impairment.” The term “disability,”
however, can then be used to posit ways in which skeletal
changes impacted the individual’s social interactions, or
identity. Cross (1999) offers further points of distinction
between the terms in her construction of two models: the
medical model and the social model of disability.
“According to the medical model, disability is viewed as
a personal, individual medical tragedy amenable to either
a medical intervention, cure, or control. . . the medical
condition, illness, or disease is seen as being ‘the
Themes in Paleopathology Chapter | 3 27
disability’. . .” (Cross, 1999: 181). In contrast, the social
model emphasizes the social response to impairment,
which both creates and frames the notion of disability.
The concept of “disability,” therefore, is highly contextual
and relational. It is formed and manipulated by the social
actors: the individual and the community being researched
by the paleopathologist, and the paleopathologists her/
himself. As Roberts (2000: 48) astutely points out, “defor-
mity does not always lead to disability.” “Health, disease
and disability are perceived very differently in different
cultures, and in many situations caregiving can only be
inferred with reference to what is known about the con-
temporary social, cultural, economic and physical envir-
onments, and only when indicators of a serious challenge
to functioning ability are present” (Tilley, 1993: 3).
The constructed meanings of the terms “impairment”
and “disability” might be operationalized best by appre-
ciating their inextricable connections within careful arche-
ological contexts (Byrnes and Muller, 2017; Tilley and
Schrenk, 2017). Kieffer’s recent work (2015) with an
assemblage of apparently sacrificial skeletal remains
pointedly seeks to “connect the physical condition of two
ancient Maya individuals who suffered from Klippel-Feil
syndrome with how they may have been treated differ-
ently, excluded from society and ultimately documenting
a condition that may have led to them being chosen for
ritual sacrifice.” Similarly, Boutin (2016), offering a care-
ful assessment of pathological conditions in a skeleton of
a young woman from the Early Dilmun period
(c.2050 1800 BCE) from modern Bahrain, interprets the
complexity of her life through careful integration of
archeological context and the development of the
“Bioarchaeology of Personhood” model. Tenets of this
model include “(bio)archaeologists should not expect
fixed conceptions of self across history and prehistory,”
“identity cannot be parsed finely into gender or religion
or class (or disability, for that matter),” and that an “open-
ness to alternative modes of interpretation,” is essential
(Boutin, 2016: 18). “Consequently,” assert Buikstra and
Scott (2009: 42), “through the study of human remains
and their archaeological contexts, we may be able to
address societal definitions of disability for those indivi-
duals who register infirmities skeletally.”
Defining the term “care” has been equally challenging
to paleopathologists. Tilley (2015: 1) argues that “care
provision is a conscious and purposive practice that
involves caregiver(s) and care recipient(s), and it does not
take place in a void. In any community, at any point in
time, the perception of what constitutes ‘health’ and ‘dis-
ease’ and the related response . . . are shaped by a combi-
nation of cultural norms, values and belief systems;
traditions; collective skills and experience; political,
social and economic organization; environmental vari-
ables; and access to recourses.” In the paleopathological
28 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
literature, the recognition of skeletal conditions deemed to
have been incapacitating to an individual are often
assumed to denote the presence of “care.” The argument is
framed something like this: if an individual with a serious
impairment lives longer than might be expected, then she/
he must have been cared for by others. Lebel et al. (2001),
for instance, argue that the recovery of a partial Middle
Pleistocene mandible from France exhibiting substantial
antemortem tooth loss thrived, in part, due to provisioning
and actions by others. Similarly, Oxenham et al. (2009)
document the presence of ankylosed vertebrae, signifi-
cantly reduced diaphysial diameters of postcranial long
bones, and morphological superior inferior compression
of hand phalanges in a Southeast Asian Neolithic skeleton
(known as M9) from modern Vietnam. The proposed dif-
ferential diagnosis of “congenital segmentation disorder as
a child, leading to fusion of his cervical spine and concom-
itant or subsequent severe neurological impairment (likely
quadriparesis or quadriplegia)” led the authors to unequiv-
ocally assert that “this would have left M9 completely
dependent on others for every aspect of daily living”
(Oxenham et al., 2009: 111). The fact that an individual
might have required assistance during their life is not nec-
essarily the contested issue in these and other studies.
Rather, the issue is how “care” is interpreted.
In most instances, the term “care” implies that the
recipient is a “less functioning” individual. The subtext
being that healthy, “normal” individuals are self-reliant
and independent. But humans, like most social animals,
require care and cooperation throughout their lives in
order to survive. Is an infant abnormal if she/he requires
constant care? Is a woman or man abnormal if intense
focus on the production of material goods renders them
reliant on others for food acquisition? Is it unusual for a
relative to assist a family member who has a fractured
limb? No. What, then, does survival of a physically inca-
pacitated individual really tell us? Ironically, it may sim-
ply tell us that humans in the past were remarkably
similar to humans today and to a wide range of nonpri-
mate mammals (Fashing and Nguyen, 2011). Our interest
in “care,” therefore, ought to extend beyond the assertion
that care was provided in the past, and explore the plastic-
ity of complex, context-dependent social interactions and/
or strategies adopted (whether successful or not) that
influence the lives of others.
The paleopathological focus on recognizing “compas-
sion” in the archeological record further complicates the
skeletal interpretation of “care.” Like the terms
“impairment” and “disability,” the terms “compassion” and
“care” are often treated as synonyms. They are not. While
the term “care” denotes “attending to” and provisioning, the
term “compassion” involves emotional responses including
sympathy, empathy, and an awareness of suffering. In her
astute and unsettling critique of paleopathological research
supporting evidence of prehistoric compassion, Dettwyler
(1991) lays bare a number of assumptions that underlie
inferences of compassion. She argues that one assumption,
that most people within a population are productive and
self-sufficient, overlooks the changing roles of children and
the elderly, and ignores that “illness and injury probably
incapacitate most members of a population occasionally. . .”
(Dettwyler, 1991: 380). Disability is not, therefore, abnor-
mal or unusual. Assuming that the survival of an incapaci-
tated individual indicates the presence of compassion is
also misleading, as “cruelty and indifference leave few
traces in the archaeological record” (Dettwyler, 1991: 382).
Similarly, Dettwyler directly refutes the assumption that
caring for or facilitating the survival of an incapacitated
individual is always a “compassionate” act, since cultures
that practice infanticide of impaired or deformed children
emically frame compassion as sparing their child from
hardship.
Another key component of impairment, disability, and
care research involves the need for clinical correlates.
Paleopathologists rely (or certainly ought to rely) on the
integration of clinical research in order to develop differ-
ential diagnoses of archeological specimens (Mays,
2011). Often neglected, however, is the use of clinical
data to interpret the ramifications of bone changes. There
is a tendency to equate lesions appearing macroscopically
severe with the in vivo presence of pain or debilitation.
This association is problematic. Roberts (1988) and
Grauer and Roberts (1996) point out that long bone frac-
tures, some displaying considerable angulation (up to 35
degrees) or overlap (up to 35 mm) impacting alignment
and creating shortening, are clinically considered “suc-
cessful” and can thus be used to interpret the presence of
healing and the presence of treatment in the archeological
record. Visually, a femur displaying 35 degrees angula-
tion is alarming. The impact on the individual might intui-
tively appear great, as mobility is likely impacted. But the
association between visual deformity (variation from nor-
mal) and the psychosocial impact on the individual is not
straightforward. Clinical research into the correlation
between tissue damage and pain finds varying association
in, for instance, patients with osteoarthritis of the knee
(Torres et al., 2006). Torres et al. found that pain severity,
measured in 143 patients, was not statistically correlated
with the presence of osteophytes and bone cysts, but was
statistically significant when bone attrition and bone mar-
row lesions were observed.
Further complicating the association between patho-
logical conditions and pain, Summers et al. (1991), and
later Wollaars et al. (2007) found positive and statistically
significant associations between pain in patients with spi-
nal cord injuries and “negative cognitions” such as anger
or isolation, as well as in those “less accepting” of their
disability. “Level of lesion, completeness of injury,

surgical fusion and/or instrumentation. . . were not associ-
ated with pain severity” (Summers et al., 1991: 183).
More recently, Neogi (2013: 1147) argued that “pain is a
subjective experience, influenced by a number of factors,
including genetic predisposition, prior experience. . . cur-
rent mood, coping strategies and catastrophizing, and
sociocultural environment . . ..” Hence, without taking
into account psycho-social-economic and other factors
that contribute to personal differences, assessments of the
relation between pathological structure and symptoms
will be confounded. Our take-away is that assertions of
impairment or disability in the archeological record that
are based on the presence of pathological lesions invari-
ably associated with pain are naive, and that, indeed,
impairment and disability are best evaluated as inextrica-
bly linked to all aspects of human life.
Asserting the presence of biomechanical incapacitation
or functional outcomes of pathological conditions from
skeletal remains is being increasingly addressed by paleo-
pathologists. Morphological changes to joints, for instance,
can have predictable consequences for mobility. Joint
fusion caused by arthroses or trauma leads to changes in
gait, posture, and movement, depending on the location
and etiology of the condition. The clinical record bears
witness to these associations. Correlations between joint
change and functional outcomes of osteoarthritis have been
a particular focus of the National Institutes of Health, as an
enormous initiative known as the Osteoarthritis Initiative,
seeks to document and make public osteoarthritis status
and outcome measures in patients throughout the United
States (http://www.oai.ucsf.edu/). Young and Lemaire
(2014, 2017) offer an ordinal grading system of severity of
osteoarthritis based on these data. Their resulting COAS
(Clinical Archaeological Osteoarthritis Scale) is intended
to allow researchers to model functional outcomes of OA
based on dependent variables such as age, sex, and type of
movement. Functional limitations of OA patients, such as
climbing stairs, bending, etc., might have archeological
correlates such as traversing mountainous terrain or squat-
ting, allowing paleopathologists to explore links between
lesions and impairment. Hence, the integration of the clini-
cal record with archeological or mummified specimens has
much to offer.
OSTEOBIOGRAPHY IN
PALEOPATHOLOGY
In what might appear to be an ironic twist in skeletal anal-
yses, researchers are rethinking current emphases on pop-
ulation approaches. Discussed in detail in Chapter 1,
paleopathological research fomenting from social theory
of the 1960s, redirected attention away from case studies
toward population analyses. Concentration on skeletal
Themes in Paleopathology Chapter | 3 29
populations, advocated by “new archeology,” allowed
broad evolutionary and epidemiological questions to be
posed. These cast light on changing human pathogen
interactions, and the effects of variables such as environ-
ment, subsistence strategies, time, and inclusion into bio-
social groups (whether defined geographically or as
status, sex, or age). Lost, however, was the appreciation
for highly contextualized studies of individuals.
To be clear, case studies are not the same as osteobio-
graphies. Case studies center attention on particular
lesions, diagnostic issues, or on finite details of a circum-
scribed population. On the other hand, osteobiographies, a
term first coined by Frank Saul (1976), and later rede-
fined by Robb (2002), currently seek to emphasize multi-
dimensional aspects of the life of an individual. As such,
osteobiographies draw heavily on archeological and his-
torical context and interpretation (see Stodder and
Palkovich, 2012), as well as identity theory and the life
course approach in order to understand the life of an indi-
vidual. Stodder (2012) offers a compelling segue into
osteobiographical research in her evaluation of data and
data analysis in paleopathology. Our decades-old empha-
sis on building large databases and statistically testing
hypotheses with large sample sizes, she argues, compels
us to ignore outliers (Stodder, 2012: 352). It might also
lead to disregarding statistically insignificant results.
Outliers, however, provide us with pertinent information.
If, as Stodder points out, stature is being investigated
(which might be used as a proxy for the presence of child-
hood stressors in the population), the statistical mean
along with measures of population differences become the
tools of analysis. Individuals who fall outside the circum-
scribed standard deviation are rendered inconsequential to
the study. But studying these individuals might be key to
our mapping population dynamics such as migration,
genetic and epigenetic bases for phenotypic variation,
environmental change, and interpersonal interactions.
These aspects of human life profoundly impact host
pathogen relationships, and thus warrant attention. The
geographical movement and social interactions of a single
individual can influence the spread of disease and influ-
ence the evolutionary course of a disease.
Social and identity theory has greatly impacted the
osteobiological approach, as individuals stand at the core
of evaluation. The individual is evaluated and understood
in light of the presence and implications of paleopatholog-
ical lesions, which gain meaning within complex histori-
cal and mortuary contexts. As Sofaer (2011: 285 286)
points out, “the need to better integrate knowledge from
the natural and social sciences, as well as the humanities,
is of increasing importance if we wish to understand the
challenges facing human existence in the past, present
and future.” “The skeletal body is employed as a means
of underpinning interpretations rather than as a source for
30 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
generating them” (Sofaer, 2006: 2). This call to reflect on
the intersectionality of human life created by time, place,
and interpersonal relationships serves as the foundation
for Watkins and Muller’s (2015) initiative to closely inte-
grate documentary evidence with the African American
remains in the W. Montague Cobb skeletal collection.
They assert that repositioning the Cobb Human Archive
involves assessing the repercussions of using a skewed
skeletal sample when exploring population distributions,
as the dearth of infants and very old individuals might be
supplemented by documentary sources. Limitations cre-
ated by the statistical construct of cohort evaluation, in
the presence of a chronologically skewed skeletal sample
that includes individuals who lived in Washington, DC
for a few days or up to a lifetime, may be eased in part by
the inclusion of census data and other records that provide
temporal perspectives. And very personal and very politi-
cal concepts of “ethnicity” and “race” might be used to
“disrupt narratives that position the collection as a
resource for exploring ‘unique’ features of African
descendant populations that can be taken out of context”
(Watkins and Muller, 2015: 49). Hence, the integration
“opens up analytical and interpretive possibilities that are
reflected in the most recent scholarship that focuses on
anatomical collections as well as skeletal remains from
cemeteries and other burial sites. This includes studying
individual experiences of health and disease in the larger
context of social issues, experiential interpretations of
skeletal remains that may include fictional narrative, dif-
ferent ways of theorizing connections between the past
and the present, and denaturalizing and complicating
notions of gender and age” (Watkins and Muller, 2015:
46 47).
Bringing complexity to the evaluation and life of the
individual has led some researchers to contest the bound-
aries of scientific inquiry. Boutin (2016), for instance,
situates a skeleton of a young female from c.2050 to 1800
BCE Bahrain, displaying idiopathic coxa valga with fem-
oral anteversion, and humerus varus deformity with
reduced bone length firmly within archeological and mor-
tuary contexts by offering a fictive osteobiographical nar-
rative of the individual’s life. Through the eyes of the
mother, and writing in the first person, Boutin recreates
the life events of the impacted young woman. The
impaired child is given a name, Beltani. Her life is appre-
ciated through a model of personhood where constructs of
identity and individuality are not seen as universals, are
deeply complex and comprised of many entangled facets,
and is informed by the life course paradigm. Her mother’s
thoughts and actions are thus offered based on known
clinical repercussions of the deformities and nuanced
assessments of the mortuary context within which the
individual was recovered. “. . .as she grew,” Boutin pens
in the voice of Beltani’s mother, “she helped as much as
my other daughters, never complaining about the awk-
ward positions into which she sometimes had to contort
her upper body, or the stares she had to endure as her
twisted legs carried her down the street. . .” (Boutin, 2016:
25).
Readers of these new directions in paleopathological
research are often struck by the subjective interpretations.
Some might discount them. But the approaches present
two important points: close and careful synthesis of paleo-
pathological lesions within archeological, mortuary, and
documentary context is essential, and all science is sub-
jective. Attending to the first point, it is clear that inter-
preting the etiology along with the repercussions of
pathological conditions requires the integration of as
many data points and pieces of information as possible.
Traumatic lesions result from human actions and reac-
tions, whether they be physical, physiological, or patho-
genic. Similarly, the etiological complexity of metabolic
disorders might best be appreciated alongside complex
social interactions between humans, humans and their
environments, and varying physiological needs throughout
the life course. Hence, understanding human skeletal
responses requires an appreciation for the complexity of
human life.
The second point, that all science is subjective, has
deep roots in intellectual and philosophical inquiry (Reiss
and Sprenger, 2016). Published scientific articles from the
early to mid 20th century comfortably employed the first
person or subjective narrative to report data, analyses, and
conclusions, only to become more sterile and impersonal
in later decades (Sheffield, 2010). Growing numbers of
reports written in the passive and third person have led to
readers’ interpretations that the words on the page are
truth; that there is a single understanding or interpretation
to be shared by all. But, argue Jahn and Dunne (1997:
201), “over the greater portion of its scholarly history, the
particular form of human observation, reasoning, and
technical deployment we properly term ‘science’ has
relied at least as much on subjective experience and inspi-
ration as it has on objective experiments and theories.”
The new themes driving paleopathology in exciting direc-
tions, such as social science and identity theories, feminist
and gender theories, and disability and impairment theo-
ries, support this contention. They converge on the point
that human experiences are unique and multidimensional
and are understood through many lenses, none more
important or “real” than the next. Denying the subjectivity
of our work within paleopathology removes the humanity
from the humans we seek to understand. As Zee (1975:
417) so poignantly points out, “. . . the question we asked
ourselves still remains: what is it that so readily impels us
to gather facts from a suffering person to diagnose a dis-
ease rather than into a more mutual exploration to learn
more about the personal meanings and causes of what a

person experiences?” These are the new directions bring-
ing new understandings to human health and disease.
Multiscalar studies hold great promise in offering
nuanced understandings of the past and the complex rela-
tionship between human biological and social lives. For
instance, Torres-Rouff and Knudson (2017: 381) combine
multiple lines of evidence that reflect “identities in the
past” —individual and group, mutable and immutable, in
order to offer insights into a period of marked change in
the Andes. Focusing upon the oases of San Pedro de
Atacama and the Loa Valley, they investigate the “tumul-
tuous” period of transition between the Middle Horizon
(AD 500 1100) and the Late Intermediate Period (AD
1100 1400). Mutable aspects of identity include those
inscribed upon the body and cultural practices related to
funerary ritual. Immutable are those relating to geo-
graphic origins and heredity. The changes inscribed upon
the body include cranial modification, discussed here in
Chapter 9. Biological age and sex are used as proxies for
social age and gender. Inherited measurable features of
the skeleton and nonmetric traits assist in interpreting
ancestry inferring ethnicity, while biogeochemistry pro-
vides indications of geographic origins. Their research,
and that of a growing number of other paleopathologists
and bioarchaeologists, clearly supports the promise of
incorporating numerous lines of evidence and social the-
ory with skeletal analyses.
REFERENCES
Agarwal, S.C., Beauchesne, P., 2011. It is not carved in bone:
Development and plasticity of the aged skeleton. In: Agarwal, S.C.,
Glencross, B.A. (Eds.), Social Bioarchaeology. Wiley-Blackwell,
Malden, MA, pp. 312 332.
Agarwal, S.C., 2012. The past of sex, gender, and health: bioarchaeology
of the aging skeleton. Am. Anthropol. 114 (2), 322 335.
Agarwal, S.C., 2016. Bone morphologies and histories: life course
approaches in bioarchaeology. Yearb. Phys. Anthropol. 159 (Suppl.
S61), 130 149.
Armelagos, G.J., Goodman, A., Harper, K., Blakey, M., 2009. Enamel
hypoplasia and early mortality: bioarchaeological support for the
Barker Hypothesis. Evol. Anthropol. 18, 261 271.
Barker, D.J., 2002. Fetal programming of coronary heart disease. Trends
Endocrinol. Metab. 13, 364 368.
Barrett, A.R., Blakey, M.L., 2011. Life histories of enslaved Africans in
colonial New York. In: Agarwal, S.C., Glencross, B.A. (Eds.), Social
Bioarchaeology. Wiley-Blackwell, Malden, MA, pp. 212 251.
Baxandall, R., Gordon, L., 2005. Second-wave feminism. In: Hewitt, N.
A. (Ed.), A Companion to American Women’s History. Blackwell
Publ, Malden, MA, pp. 414 431.
Bengtson, J., O’Gorman, J., 2017. Women’s participation in prehistoric war-
fare: A Central Illinois River Valley case study. IJO 27 (2), 230 244.
Binford, L.R., 1971. Mortuary practices: their study and their potential.
Mem. Soc. Am. Archaeol. 25, 6 29.
Themes in Paleopathology Chapter | 3 31
Blakely, R.L., Harrington, J.M., 1997. Bones in the Basement:
Postmortem Racism in Nineteenth-Century Medical Training.
Smithsonian Institution Press, Washington, DC.
Boraschi, D., Aguado, M.T., Dutel, C., Goronzy, J., Louis, J., Grubeck-
Loebenstein, B., et al., 2013. The gracefully ageing immune system.
Sci. Transl. Med. 5 (185), 1 9.
Boutin, A.T., 2016. Exploring the social construction of disability: an
application of the bioarchaeology of personhood model to a patho-
logical skeleton from ancient Bahrain. Intl. J. Paleopath. 12, 17 28.
Buikstra, J.E., 1995. Tombs for the living . . . or for the dead: The
Osmore Ancestors. In: Dillehay, T. (Ed.), Tombs for the Ancestors.
Dumbarton Oaks Research Library and Collection, Washington, DC,
pp. 229 280.
Buikstra, J.E., Scott, R.E., 2009. Key concepts in identity studies.
In: Knudson, K.J., Stojanowski, C.M. (Eds.), Bioarchaeology and
Identity in the Americas. University Press of Florida, Gainesville,
pp. 25 55.
Buikstra, J.E., Roberts, C.R., 2012. The Global History of Paleopathology:
Pioneers and Prospects. Oxford University Press, Oxford.
Byrnes, J.F., Muller, J.L., 2017. Bioarchaeology of Impairment and
Disability: Theoretical, Ethnohistorical, and Methodological
Perspectives. Springer, New York.
Carr, C., 1995. Mortuary practices: their social, philosophical-religious,
circumstantial, and physical determinants. J. Archaeol. Method
Theory 2 (2), 105 200.
Charles, D.K., Buikstra, J.E., 1983. Archaic mortuary sites in the central
Mississippi drainage: distribution, structure, and behavioral implica-
tions. In: Phillips, J.L., Brown, J.A. (Eds.), Archaic Hunters and
Gatherers in the American Midwest. Academic Press, New York,
NY, pp. 117 145.
Cohen, M., Bennett, S., 1993. Skeletal evidence for sex roles and gender
hierarchies in prehistory. In: Miller, B.D. (Ed.), Sex and Gender
Hierarchies. Cambridge University Press, Cambridge, pp. 273 296.
Cross, M., 1999. Accessing the inaccessible: disability and archaeology.
Archaeol. Rev. Cambridge 15 (2), 7 30.
Cybulski, J.S., 1977. Cribra orbitalia, a possible sign of anemia in early
historic native populations of the British Columbia coast. Am. J.
Phys. Anthropol. 47, 31 39.
de la Cova, C., 2011. Race, health, and disease in 19th-century-born
males. Am. J. Phys. Anthropol. 144, 526 537.
de la Cova, C., 2017. Fractured lives: structural violence, trauma, and
recidivism in urban and industrialized 19th-century-born African
Americans and Euro-Americans. In: Tegtmeyer, C.E., Martin, D.L.
(Eds.), Broken Bones, Broken Bodies: Bioarchaeological and
Forensic Approaches for Accumulated Trauma and Violence.
Lexington Books, London, p. 153.
Dettwyler, K.A., 1991. Can paleopathology provide evidence for “com-
passion”? Am. J. Phys. Anthropol. 84 (4), 375 384.
DeWitte, S.N., Stojanowski, C.M., 2015. The osteological paradox 20 year
later: past perspectives, future directions. J. Archaeol. Res. 23, 397 450.
Farmer, P.E., Nizeye, B., Stulac, S., Keshavjee, S., 2006. Structural vio-
lence and clinical medicine. PloS Medicine 3 (10), e449. Available
from: https://doi.org/10.1371/journal.pmed.0030449.
Fashing, P.J., Nguyen, N., 2011. Behavior toward the dying, diseased, or
disabled among animals and its relevance to paleopathology. Int. J.
Paleopath. 1, 128 129.
Galtung, J., 1969. Violence, peace, and peace research. J Peace Res 6
(3), 167 191.
32 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
Giefing-Kröll, C., Berger, P., Lepperdinger, G., Grubeck-Loebenstein,
B., 2015. How sex and age affect immune responses, susceptibility
to infections, and response to vaccination. Aging Cell 14, 309 321.
Gilchrist, R., 2000. Archaeological biographies: realizing human life
cycles, -courses and histories. World Archaeol. 31 (3), 325 328.
Gilligan, J., 1996. Violence: Reflections on a National Epidemic.
Vintage Books, New York, NY.
Goldstein, L., 1980. Mississippian mortuary practices: a case study of
two cemeteries in the lower Illinois Valley. Northwestern University
Archaeology Program Scientific Papers #4.
Gowland, R., Knüsel, C., 2006. Social Archaeology of Funerary
Remains. Oxbow Books, Oxford.
Gowland, R.L., Penny-Mason, B., 2018. Overview: archaeology and the
medieval life-course. In: Gerrard, C., Gutierrez, A. (Eds.), Oxford
Handbook of Later Medieval Archaeology in Britain. Oxford
University Press, Oxford, pp. 759 773.
Grauer, A.L., McNamara, E., Houdek, D., 1999. A history of their own:
Health and disease in a 19th-century poorhouse. In: Grauer, A.L.,
Stuart-Macadam, P. (Eds.), Sex and Gender in Paleopathological
Perspective. Cambridge University Press, Cambridge, pp. 149 164.
Grauer, A.L., 2018. A century of paleopathology. Am. J. Phys.
Anthropol. 165, 904 914.
Grauer, A.L., 2019. Paleopathology: from bones to behavior.
In: Katzenberg, M.A., Grauer, A.L. (Eds.), Biological Anthropology
of the Human Skeleton, third ed. Wiley-Blackwell, New York,
pp. 447 465.
Grauer, A.L., Roberts, C.A., 1996. Paleoepidemiology, healing, and possible
treatment of trauma in the medieval cemetery population from St. Helen-
on-the-Walls, York, England. Am. J. Phys. Anthropol. 100, 531 544.
Guatelli-Steinberg, D., Lukacs, J.R., 1999. Interpreting sex differences in
enamel hypoplasia in human and non-human primates: developmen-
tal, environmental, and cultural considerations. Yearb. Phys.
Anthropol. 42, 73 126.
Halcrow, S.E., Tayles, N., 2008. The bioarchaeological investigation of
childhood and social age: problems and prospects. J. Archaeol.
Method Theory 15 (2), 190 215.
Halcrow, S.E., Tayles, N., 2011. The bioarchaeologial investigation of
children and childhood. In: Agarwal, S.C., Glencross, B.A. (Eds.),
Social Bioarchaeology. Wiley-Blackwell, Chicester, pp. 333 360.
Harrod, R.P., Martin, D.L., 2014. Signatures of captivity and subordina-
tion on skeletonized human remains: a bioarchaeological case study
from the ancient Southwest. In: Martin, D.L., Anderson, C.P. (Eds.),
Bioarchaeological and Forensic Perspectives on Violence: How
Violent Death in Interpreted from Skeletal Remains. Cambridge
University Press, Cambridge, pp. 103 119.
Hodder, I. (Ed.), 1982. Symbolic and Structural Archaeology. University
Press, Cambridge.
Hodder, I., 1984. Burials, houses, men and women in the European
Neolithic. In: Miller, Tilley (Eds.), Ideology, Power and Prehistory.
Cambridge University Press, Cambridge, pp. 51 68.
Hollimon, S., 2006. The archaeology of nonbinary genders in Native
North American Societies. In: Nelson, S.M. (Ed.), Handbook of
Gender Archaeology. AltaMira Press, Walnut Creek, pp. 435 450.
Hollimon, S., 2011. Sex and gender in bioarcheological research.
In: Agarwal, S., Glencross, B. (Eds.), Social Bioarchaeology. Wiley
Blackwell, New York, pp. 149 182.
Jahn, R.G., Dunne, B.J., 1997. Science of the subjective. J. Sci. Explor.
11 (2), 201 224.
Kieffer, C.L., 2015. Sacrifice of the social outcasts: two cases of
Klippel-Feil Syndrome at Midnight Terror Cave, Belize. IJO.
Available from: https://doi.org/10.1002/oa.2456.
Kirkpatrick, R.C., 2000. The evolution of human homosexual behavior.
Curr. Anthropol. 41, 385 398.
Klaus, H.D., 2012. The bioarchaeology of structural violence: a theoreti-
cal model and a case study. In: Martin, D.L., Harrod, R.P., Perez, V.
R. (Eds.), The Bioarchaeology of Violence. University of Florida
Press, Gainesville, FL, pp. 29 62.
Knüsel, C., 2011. Men take up arms for war: sex and status distinctions
of humeral medial epicondylar avulsion fractures in the archaeolog-
ical record. In: Baadsgaard, A., Boutin, A.T., Buikstra, J.E. (Eds.),
Breathing New Life into the Evidence of Death: Contemporary
Approaches to Bioarchaeology. School for Advanced Research
Press, Santa Fe, pp. 221 251.
Lebel, S., Trinkaus, E., Faure, M., Fernandez, P., Guérin, C., Richter, D.,
et al., 2001. Comparative morphology and paleobiology of Middle
Pleistocene human remains from the Bau de l’Aubesier, Vaucluse,
France. Proc. Natl. Acad. Sci. U.S.A. 98, 11097 11102.
Lewis, M.E., 2007. The Bioarchaeology of Children: Perspectives from
Biological and Forensic Anthropology. Cambridge University Press,
Cambridge.
Martin, D.L., Harrod, R.P., Fields, M., 2010. Beaten down and worked
to the bone: bioarchaeological investigations of women and violence
in the ancient Southwest. Landscapes Violence 1 (1), 1 19.
Mays, S., 2011. The relationship between paleopathology and the clinical
sciences. In: Grauer, A. (Ed.), Companion to Paleopathology. Wiley,
New York, pp. 285 309.
Mays, S., Gowland, R., Halcrow, S., Murphy, E., 2017. Child bioarch-
aeology: perspectives on the past 10 years. Child. Past 10 (1), 38 56.
Mitchell, P., 2012. Anatomical Dissection in Enlightenment England and
Beyond: Autopsy, Pathology, and Display. Ashgate, Farnham.
Morris, I., 1991. The archaeology of ancestors: the Saxe/Goldstein
hypothesis revisited. Cambridge Archaeol. J. 1 (2), 147 169.
Neogi, T., 2013. The epidemiology and impact of pain in osteoarthritis.
Osteoarthritis Cartilage 21, 1145 1153.
Nystrom, K.C., 2014. The bioarchaeology of structural violence and dis-
section in the 19th-century United States. Am. Anthropol. 116 (4),
765 779.
Nystrom, K.C., 2017a. Introduction. In: Nystrom, K.C. (Ed.), The
Bioarchaeology of Dissection and Autopsy in the United States.
Springer, New York, pp. 1 22.
Nystrom, K.C., 2017b. The Bioarchaeology of Dissection and Autopsy
in the United States. Springer, New York.
Ortner, D.J., 1998. Male/female immune reactivity and its implications
for interpreting evidence in human skeletal paleopathology.
In: Grauer, A.L., Stuart-Macadam, P. (Eds.), Sex and Gender in
Paleopathological Perspective. Cambridge University Press,
Cambridge, pp. 79 93.
Ortner, D.J., 2003. Identification of Pathological Conditions in Human
Skeletal Remains, second ed Academic Press, New York.
Oxenham, M.F., Tilley, L., Matsumura, H., Nguyen, L.C., Nguyen, K.T.,
Nguyen, K.D., et al., 2009. Paralysis and severe disability requiring
intensive care in Neolithic Asia. Anthropol Sci. 117. Available from:
https://www.jstage.jst.go.jp/article/ase/117/2/117_081114/_html/-char/ja.
Penny-Mason, B.J., Gowland, R.L., 2014. The children of the reforma-
tion: childhood palaeoepidemiology in Britain, AD 1000 1700.
Medieval Archaeol. 58, 162 194.

Perry, M., 2008. Redefining childhood through bioarchaeology: toward
an archaeological and biological understanding of children in antiq-
uity. In: Baxter, J.E. (Ed.), Children in Action: Perspectives on the
Archaeology of Childhood, Number 15 (1). Archaeological Papers
of the American Anthropological Association, pp. 89 111.
Rakita, G.F.M., Buikstra, J.E., 2005. Introduction. In: Rakita, G.F.M.,
Buikstra, J.E., Beck, L.A., Williams, S.R. (Eds.), Interacting with
the Dead: Perspectives on Mortuary Archaeology for the New
Millennium. University Press of Florida, Gainesville, FL.
Reis, J., Sprenger J., 2016. Scientific objectivity. In: Zalta, N., (Ed.), The
Stanford Encyclopedia of Philosophy, Summer 2016 edition. ,http://
plato.stanford.edu/archives/sum2016/entries/scientific-objectivity/..
Robb, J., 2002. Time and biography. In: Hamilakis, Y., Pluciennik, M.,
Tarlow, S. (Eds.), Thinking Through the Body. Springer, Boston,
MA, pp. 153 171.
Roberts, C.A., 1988. Trauma and Its Treatment in Medieval British
Antiquity. PhD dissertation. Bradford University, Bradford.
Roberts, C.A., 2000. Did they take sugar? The use of skeletal evidence
in the study of disability in past populations. In: Hubert, J. (Ed.),
Madness, Disability, and Social Exclusion: The Archaeology and
Anthropology of “Difference”. Routledge, Abingdon, pp. 46 59.
Sanders, V., 2006. First wave feminism. In: Gamble, S. (Ed.), The Routledge
Companion to Feminism and Postfeminism. Routledge, London,
pp. 15 24.
Saul, F.P., 1976. Osteobiography: life history recorded in bone.
In: Giles, E., Friedlander, J.S. (Eds.), The Measures of Men.
Peabody Museum Press, Cambridge, MA, pp. 372 382.
Saxe, A., 1970. Social Dimensions of Mortuary Practices. University of
Michigan.
Sharlach, L.B., 2009. First wave feminism. In: Ness, I. (Ed.), The
International Encyclopedia of Revolution and Protest. https://doi.
org/10.1002/9781405198073.wbierp0556.
Sheffield, N., 2010 2011. Duke Graduate School Scientific Writing
Resource. ,https://cgi.duke.edu/web/sciwriting/index.php..
Sofaer, J.R., 2006. The Body as Material Culture: A Theoretical
Osteoarchaeology. Cambridge University Press, Cambridge.
Sofaer, J.R., 2011. Towards a social bioarchaeology of age. In: Agarwal,
S.C., Glencross, B.A. (Eds.), Social Bioarchaeology. Wiley-
Blackwell, Malden, MA, pp. 285 311.
Solecki, R.S., 1971. Shanidar: The First Flower People. Knopf Publ,
New York.
Stinson, S., 1985. Sex differences in environmental sensitivity during
growth and development. Yearb. Phys. Anthropol. 28, 123 147.
Stirland, A.J., 1997. Care in the medieval community. Int. J. Osteoarch.
7, 587 590.
Stodder, A.L.W., 2012. Data and data analysis: Issues in paleopathology.
In: Grauer, A.L. (Ed.), Companion to Paleopathology. Wiley-
Blackwell, New York, pp. 339 356.
Stodder, A.L.W., Palkovich, A.M., 2012. The Bioarchaeology of
Individuals. University of Florida Press, Gainesville.
Stuart-Macadam, P., 1987. Porotic hyperostosis: new evidence to support
the anemia theory. Am. J. Phys. Anthropol. 74 (4), 521 526.
Summers, J.D., Rapoff, M.A., Varghese, G., Porter, K., Palmer, R.E.,
1991. Psychosocial factors in chronic spinal cord injury pain. Pain
47 (2), 183 189.
Themes in Paleopathology Chapter | 3 33
Tegtmeyer, C.E., Martin, D.L., 2017. Broken Bones, Broken Bodies:
Bioarchaeological and Forensic Approaches for Accumulated
Trauma and Violence. Lexington Books, London.
Tilley, C., 1993. Interpretive Archaeology. Berg Publishers, Providence,
RI.
Tilley, C., 1999. Metaphor and Material Culure. Wiley Publishing,
Chicester, UK.
Tilley, L., 2015. Theory and Practice in the Bioarchaeology of Care.
Springer International, New York.
Tilley, L., Schrenk, A.A., 2017. New Developments in the
Bioarchaeology of Care. Springer International, New York.
Torres, L., Dunlop, D.D., Peterfy, C., Guermazi, A., Prasad Hayes, P.K.
W., Song, J., et al., 2006. The relationship between specific tissue
lesions and pain severity in persons with knee osteoarthritis.
Osteoarthritis Cartilage 14 (10), 1033 1040.
Torres-Rouff, C., Knudson, K.J., 2017. Integrating identities: An innova-
tive bioarchaeological and biogeochemical approach to analyzing
the multiplicity of identities in the mortuary record. Current
Anthropol 58 (3), 381 409.
Trinkhaus, E., Zimmerman, M.R., 1982. Trauma among the Shanidar
Neandertals. AJPA 57 (1), 61 76.
Tung, T.A., 2014. Gender-based violence in the Wari and Post-Wari era
of the Andes. In: Knusel, C., Smith, J.J. (Eds.), The Routledge
Handbook of the Bioarchaeology of Human Conflict. Routledge,
New York, N, pp. 333 354.
Watkins, R., Muller, J., 2015. Repositioning the Cobb Human Archive:
the merger of a skeletal collection and its texts. Am. J. Hum. Biol. 27,
41 50.
Watts, R., 2013. Childhood development and adult longevity in an
archaeological population from Barton-upon-Humber, Lincolnshire,
England. Int. J. Paleopathol. 3, 95 104.
Webb, S., 1984. Intensification, population and social change in south-
easter Australia: the skeletal evidence. Aboriginal Hist. 8 (1/2),
154 172.
Wollaars, M.M., Post, M.W., van Asbeck, F.W., Brand, N., 2007. Spinal
cord injury pain: the influence of psychologic factors and impact on
quality of life. Clin. J. Pain 23 (5), 383 391.
Wood, J.W., Milner, G.R., Harpending, H.C., Weiss, K.M., 1992. The
osteological paradox - problems of inferring prehistoric health from
skeletal samples. Curr. Anthropol. 33, 343 370.
Yaussy, S.L., DeWitte, S.N., Redfern, R.C., 2016. Frailty and famine:
Patterns of mortality and physiological stress among victims of
famine in medieval London. Am. J. Phys. Anthropol. 160 (2),
272 283.
Young, J.L., Lemaire, E.D., 2014. Linking bone changes in the distal
femur to functional deficits. IJO 24 (6), 709 721.
Young, J.L., Lemaire, E.D., 2017. Using population health constructs to
explore impairment and disability in knee osteoarthritis. In: Byrnes,
J.F., Muller, J.L. (Eds.), Bioarchaeology of Impairment and
Disability. Springer International, Cham, pp. 159 182.
Zee, H.J., 1975. Subjectivity in science. Ann. Intern. Med. 82 (3),
417 418.
Zuckerman, M.K., Crandall, J.J., unpublished manuscript. Reconsidering
sex, gender, and sexuality in relations to health and disease in
bioarchaeology. J. Anthropol. Archaeol.
Chapter 4
Fundamentals of Human Bone and Dental
Biology: Structure, Function, and
Development
Niels Lynnerup1 and Haagen D. Klaus2
1
Department of Forensic Medicine, University of Copenhagen, Copenhagen, Denmark, 2Department of Sociology and Anthropology, George Mason
University, Fairfax, VA, United States
The average adult human skeleton contains 206 bones and
32 teeth; in subadults, there are more than twice that num-
ber of bones, in addition to a set of deciduous teeth. A
range of intrinsic and extrinsic parameters and biologic
states guide the normal growth, maintenance, repair, met-
abolic functioning, and optimization of bone form and
strength. Yet, these tissues are subject to a diverse spec-
trum of developmental, metabolic, infectious, reticuloen-
dothelial, hematopoietic, degenerative, metastatic, and
endocrine influences. These conditions may disrupt or
manipulate underlying bone physiology. Accordingly,
detailed knowledge of bone and tooth biology is a funda-
mental baseline in the practice of paleopathology. This
chapter provides a relevant overview concerning current
understandings regarding how human bone and dental tis-
sue form, grow, and function—crosscutting gross anat-
omy, bone microstructure, and cellular and molecular
levels.
SKELETAL STRUCTURE, FUNCTION, AND
CELLULAR BASIS OF BONE BIOLOGY
Evolution of the Vertebrate Skeleton
Bone was once thought of as a relatively simple tissue
type when compared to the overt complexity of the ner-
vous system, for example. This view of bone was rein-
forced further by the fact that skeletal phenotypes are
varyingly biologically and evolutionarily constrained,
such that a diverse range of pathophysiological processes
can evoke only a specific range of possible forms of
abnormal bone phenotypes. Additionally, bone is charac-
terized by a rather slow tissue turnover rate compared to
Ortner’s Identification of Pathological Conditions in Human Skeletal Remains. DOI: https://doi.org/10.1016/B978-0-12-809738-0.00004-1
© 2019 Elsevier Inc. All rights reserved.
other tissue types, resulting in much longer time intervals
between disease onset and morphological alterations. Yet,
such simplistic notions of the skeletal system have been
increasingly shown to be deeply flawed. Human bones
are a product of approximately three-quarters of a billion
years of evolution (Donoghue and Aldridge, 2001; Ota
and Kuratani, 2009). An evolutionary view is the starting
point to understand the skeleton today.
Vertebrates are a relatively small branch on the tree of
life and appear in the fossil record relatively recently.
Fossil data and comparative morphologic, genomic, and
embryological evidence suggest the long-term evolution-
ary pathway likely involved a common ancestor that was
invertebrate chordate—either a complex annelid-like
worm or a more basic enteropneusta-like marine worm
(Holland et al., 2015). Ancestral chordates appear to have
been filter feeders that possessed a notochord, gill slits,
an endostyle, and a post-anal tail (Gee, 1996; Lowe et al.,
2015). In time, prismatic mineralized cartilage developed
around the notochord and brain and a bilateral body plan
emerged. Crown-group vertebrates (the last common
ancestor to all living vertebrates and their fossil ancestors)
emerged in the Cambrian epoch and were jawless, fish-
like animals (Janvier, 2015). Interestingly, various lines
of evidence suggest more than 80% of the basic relation-
ships and components of the vertebrate skeleton were
established within 15 million years after the emergence of
true vertebrates (Hall, 2002, 2015)—underscoring the
degree of adaptive advantage that bones conferred as well
as the conserved nature of underlying bone form and
function.
All vertebrates possess a bony vertebral column and
joints. The vertebral column is key to maintaining specific
35
36 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains

postures, and joints permit forms of locomotion beginning carpal and tarsal bones. This tripartite approach to classi-
with movement of the early vertebral column. Subsequent fication not only reflects a straightforward way to start
evolution of a cranium, mandible, and appendicular skele- sorting and identifying commingled bones, but it also
ton allowed vertebrates an opportunity to develop a more directly reflects differences in how bones are vascularized,
complex central nervous system, face, limbs, and tissues form and grow, and highlights varying biomechanical
regulating hematopoiesis and mineral homeostasis. The properties, metabolic functions, and locomotor functions.
first vertebrates lacked jaws, but one branch of this For example, flat bones are often enriched with red mar-
diverse and now mostly extinct group, known as gnathos- row, generating new blood cells, and they are therefore
tomes, indeed developed a mandible and paired appen- also highly vascularized. In turn, they represent ideal
dages during the late Ordovician period or early Silurian microenvironments for various kinds of bloodborne cancers
period (Brazeaum and Friedman, 2015). Between 423 and or iron-loving pathogens to metastasize and colonize (e.g.,
419 million years ago (mya), mandibulated bony fish and Wilbur et al., 2008; Klaus, 2017). Conversely, the tubular
tetrapod ancestors began to undergo remarkable diversifi- long bones possess load-bearing capabilities that reflect not
cation. By the close of the Devonian at 360 mya, adapta- only long-term patterns of physical activity and adaptation,
tions for terrestriality were present (Clack, 2012; also see but also risk specific kinds of fractures.
Dial et al., 2015). Today, more than 99% of all living ver- Long bones share a common gross anatomy (Fig. 4.1).
tebrates are gnathostomes. Most of their mass is comprised of a relatively long, cylin-
Over this history and within the deeply conserved ele- drical tube- or shaft-like structure called the diaphysis.
ments of bone biology, significant complexity developed
and accumulated. In the last 200 years of scientific study,
many of the biological complexities of the skeleton have
come to light, but it seems that for every new fact gained,
two more questions arise. Bone has been revealed to be a
highly complex organ system, involving specialized inter-
cellular signaling and tightly coordinated systems of
information exchange that precisely regulate turnover,
growth, repair, and disease response, offering direct
bridges between bone and the central nervous system and
neuropeptides, the deep synergisms between bone and the
immune system, endocrine and paracrine system influ-
ences, the gut microbiome, and deeply embedded life his-
tory traits (e.g., Gosman et al., 2011; Baldock, 2013;
Lorenzo et al., 2015; Yan et al., 2016; Okamoto et al.,
2017; Martin and Sims, 2018; Wei and Karsenty, 2018).

Gross Function and Anatomy

Functionally, the skeleton offers protection for vital soft
tissue structures, such as the brain, spinal cord, heart, and
lungs. The senses of sight, smell, hearing, and taste reside
within the skull. Bone also provides the underlying archi-
tecture for ligament and muscle attachments to facilitate
locomotion. It is a principal organ maintaining mineral
homeostasis, including its role as a reservoir for bioavail-
able calcium and phosphate, and bone is the factory for
the production of blood cells and cells of the innate
immune system (Favus and Goltsman, 2013; Frisch and
Calvi, 2013).
On a gross anatomical level, the bones of the skeleton
can be classified as: (1) long, tubular bones of the extrem- FIGURE 4.1 Gross anatomy of the exterior of a long bone (in this case
a right adult femur) denoting the diaphysis (primary center of ossifica-
ities, (2) flat bones, which form parts of the “walls” of
tion), the proximal distal epiphyses (secondary centers of ossification, an
adjacent body cavities (e.g., braincase, thoracic and pelvic apophysis (relatively rare tertiary centers of ossification), and both proxi-
cavities), and (3) irregular bones, such as the geometri- mal and distal metaphyses and articular surfaces (29-year-old female,
cally complex vertebrae, bones of the facial skeleton, and Terry Anatomical Collection, NMNH P000171R; photo: HDK).
 Fundamentals of Human Bone and Dental Biology: Structure, Function, and Development Chapter | 4
A diaphysis is the product of the primary center of ossifi-
cation (see section on Embryological and Developmental
Processes below) and internally, it features a medullary
cavity, or marrow space. Long bones tend to gradually
increase in width toward the ends in the region termed the
metaphysis. At the end of a long bone is its epiphysis,
which represents a secondary center of ossification. During
the process of bone growth, epiphyses are separated from
metaphyses by a cartilaginous growth plate that ossifies
and unites the two following the completion of bone
growth. Tertiary centers of growth are termed apophyses
and form at the site of tendinous insertions, such as the
greater and lesser trochanters of the femur.
The external surface of every bone is covered in a
thin, wax-like membrane called the periosteum. The space
between the periosteum and the bone itself is termed the
periosteal envelope. The periosteum is highly vascular-
ized and highly osteogenic (Ragsdale and Lehmer, 2012).
Arterial vessels penetrate the periosteum and the surface
of the bone through numerous microscopic nutrient
foramina (Fig. 4.2). In long bones, a major artery supplies
Articular cartilage
Epiphyseal
artery
Epiphysis
Metaphyseal
artery
Metaphysis
Periosteal
arteries
Diaphysis
Nutrient artery
Compact bone
FIGURE 4.2 Diagram showing the blood supply of an adult long bone.
The nutrient artery and the epiphyseal arteries enter the bone through
nutrient foramina. These openings in the bone arise developmentally as
the pathways of the principal vessels of periosteal buds. Metaphyseal
arteries arise from periosteal vessels that become incorporated into the
metaphysis as the bone grows in diameter. Reproduced with permission
from Ross, M.H., Pawlina W., 2011. Histology: A Text and Atlas with
Correlated Cell and Molecular Biology. Wolters Kluwer/Lippincott
Williams & Wilkins, Philadelphia, p. 222.
37
the bone marrow, and the nutrient foramen is correspond-
ingly much larger and can be seen macroscopically. This
major artery, penetrating the diaphysis, divides into two
branches traveling toward each epiphysis, with a further
division between a primary epiphyseal and a metaphyseal
branch. Veins follow the arteries, although the vertebrae
have distinct venous foramina on the lateral aspects of
vertebral bodies.
Periosteal blood vessels are very important for
bone growth (see below), as well as for bone repair
after fracture and responses to infection. If the perios-
teum is removed, the underlying bone will die and
necrotize. The high degree of vascularization also
means that bloodborne pathogens, emanating from
elsewhere in the body, may be spread with relative
ease to bone, resulting in periosteal inflammatory
reactions (Weston, 2012; Klaus, 2014). The periosteum
also carries the greatest number of peripheral nerves
associated with bone, such that pain associated with
fracture is mostly due to periosteal nerve signaling.
In contrast, a deeper, chronic infection of bone may
result in less physical discomfort. The periosteum is
comparatively thick and also serves as an anchor for
fibers that facilitate muscle attachments (Sharpey’s
fibers). These fibers penetrate the outer surface of the
bone and thus form a very strong anchor for the mem-
brane. Bone morphology at these sites of attachment
may be influenced further by mechanical loads, result-
ing in new bone formation if there is an increased strain
on the muscle attaching at the site. Macroscopically,
these form roughened and topographically rugose areas
called entheses.
Bone Tissue: Composition and Organization
Bone is one of the hardest materials synthesized by any
biological process. Bone is a composite material—part
mineral, part collagen—and therefore possesses the opti-
mal qualities of both—simultaneously rigid and flexible
and lightweight (amounting to only about 30% of the
weight of the human body). As a composite, the skeleton
is made up of inorganic mineral content (B60%), organic
components (B25%), and water (B15%). Chemically,
the inorganic component of bone tissue is hydroxyapatite
(Ca10 (PO4)6 (OH)2), which is a specialized crystalline
form of calcium phosphate. These plate-shaped crystals
have a very large surface area (about 100 m2 per 1 g of
hydroxyapatite crystals). Bone matrix also incorporates
organic tensile collagen fibers (type I), which are
alkaline-soluble polypeptides. The combination of organic
collagen fibers and inorganic crystals is suspended in an
intercellular matrix of proteoglycans giving bone unique
mechanical and physical properties. Bone possesses a
38 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains
high tensile strength—1100 kg/cm2 parallel to the bone’s
axis—more than twice that of porcelain and other ceramic
fabrics. Due to the crystalline nature of bone mineral, the
longitudinal compressive strength is also very high,
exceeding 2000 kg/cm2. Indeed, bone tissue is often com-
pared to fiberglass plastic: a flexible but strong fiberglass
weave (collagen) surrounded by hard but fragile, epoxy
resin (mineral).
Bone is comparable to all connective tissues in that it
consists of cells and an intercellular substance. In bone,
the intercellular matrix consists of collagen fibers and
mineral salts, mostly calcium phosphate (85%), but also
calcium carbonate (10%), in addition to magnesium and
alkali salts (5%). Almost the entire body’s calcium store
is maintained within the skeleton, which therefore serves
as a calcium reservoir (Favus and Goltsman, 2013). There
is a tightly regulated exchange of calcium between blood
and the skeleton in order to maintain an allostatic balance.
Calcium is important for muscle function, and even small
deviations in blood calcium can result in spasmodic
cramps. Due to its importance, the body may be forced to
draw upon calcium deposits in the skeleton if there is
insufficient dietary calcium intake.
For any single bone there are two types of tissue: an
external compact lamellar tissue, which forms a smooth
surface (or “cortex,” hence the alternate term cortical
bone) and an internal spongy or cancellous substance (tra-
becular bone). Compact bone covers all parts of the skele-
tal surface. Macroscopically, it has a homogeneous
appearance and is most pronounced in the diaphyses of
the largest long bones, where it surrounds the marrow
space in a cylindrical wall up to several millimeters thick.
A layer of compact bone also covers joint surfaces,
although they are considerably thinner. Cancellous bone
is most prolific in irregular and flat bones, such that it
occupies the space between the external and internal sur-
faces of flat bones, including the parietal bones where it
is termed diploë. Cancellous bone forms a fine network of
thin bone beams (trabeculae). The total internal cancel-
lous bone surface area of the skeleton therefore is very
large. Trabeculae also play a biomechanical role, perhaps
most clearly seen in longitudinal sections of the femoral
neck, where the orientation of trabecular networks reflects
the optimal directional transmission of force (including
stretching, torsional, and compressive forces) and under-
scores how bone architecture elegantly maximizes
strength with a minimum of material.
The internal surface of a long bone diaphysis is com-
posed of the medullary cavity and the endosteal envelope
that is best described as cellular membrane (Hall, 2015).
The medullary cavity contains bone marrow that fills the
internal bone cavity beneath the cortical surface. In adult
long bones, this cavity is cylindrical. Marrow space is also
found continuing onward into the metaphyses where it is
interwoven into trabecular space. Marrow spaces are filled
with both red and yellow bone marrow. The red marrow is
hematopoietic and is the main blood-forming organ after
birth. Hematopoietic marrow red blood cells, white blood
cells, and platelets are produced in marrow tissue that is
nourished from the blood vessels that pass through the
nutrient foramen (Kumar et al., 2014; also see
Chotinantakul and Leeanansaksiri, 2012). The term red
bone marrow refers to the mainly hematopoietic cells, or
blood stem cells, responsible for forming the different cel-
lular components of blood. In children, all bone marrow is
red due to the demands of blood cell formation in a grow-
ing body. From about the fifth year of life onwards, areas
of red marrow (for instance, in the cranial vault) are gradu-
ally replaced by yellow fat cells. This process begins at the
center of the bone and expands outward. After skeletal
growth has ended (from the age of 20 25), there is usually
only red bone marrow in the proximal aspects of femora
and humeri and in various flat bones (sternum, ribs, clavi-
cles, scapulae, and the pelvic bones). In older adults, these
bones may eventually contain only yellow bone marrow.
In rare cases of unusually severe or chronic anemia or
another increased need for blood cells, yellow bone mar-
row can be converted to red bone marrow.
Cartilaginous Tissue
The cartilage of the joint surfaces consists of hyaline car-
tilage, which covers them with no intervening periosteal
membrane (for excellent overviews, see Resnick, 2002;
Lories and Luyten, 2018). Hyaline cartilage possesses a
very smooth, strong, and elastic quality. This smoothness,
combined with synovial joint fluid, minimizes friction
in the joints. These biomechanical properties of hyaline
cartilage make a joint highly resistant to wear despite the
enormous strain and compressional forces to which is
exposed (e.g., up to 300 kg in the hip joint). Cartilage
may vary in thickness from 1 to 7 mm. On convex
joints, the cartilage is thickest in the middle, whereas on
concave surfaces, the thickest portions are distributed to
the joint edges.
As with bone, hyaline cartilage consists of cells sur-
rounded by an intercellular matrix. These cells are called
chondrocytes, and produce a proteoglycan matrix as well as
collagen fibers. The fibers are usually oriented in ways that
maximize their biomechanical efficiency vis-à-vis joint
movements. A simple and consistent fiber orientation is eas-
iest to achieve if the joint has one axis of motion (e.g., a
simple hinge joint between phalanges). Unsurprisingly,
joints with more axes of movement are also more prone to
 Fundamentals of Human Bone and Dental Biology: Structure, Function, and Development Chapter | 4
cartilage damage (e.g., shoulder, elbow, hip, knee). Hyaline
cartilage (as elastic and fibrous cartilage) lacks blood ves-
sels and nerves, and thus regenerates slowly or not at all if
the tissue is damaged. This may result in the underlying
joint bone surface being directly exposed in severe cases of
osteoarthritis. If adjacent joint surfaces are thus denuded,
they may interact directly, leading to pathological eburna-
tion and surface grooving.
The two other kinds of cartilage are elastic cartilage,
found in the epiglottis, pharynx, and the outer ear.
Fibrous cartilage (fibrocartilage) is found in the vertebral
discs, the intraarticular components of major joints (e.g.,
the menisci of the knee joints), and symphyses.
Interestingly, the sacroiliac joint has hyaline cartilage on
the sacral joint surface and fibrocartilage on the iliac
joint surface. Overall, the three cartilage types differ in
relative amounts of collagen, elastic fibers, and proteogly-
can matrix. Elastic cartilage is yellowish in color due to a
higher content of elastic fibers, while the latter is more
whitish and less elastic (Ross and Pawlina, 2011;
Standring, 2015).
Bone Cells
The growth, regulation, repair, and other functions of skele-
tal tissue are driven by the behavior of just three kinds of
cells: osteoblasts, osteoclasts, and osteocytes. Osteoblasts
are bone-producing cells derived from osteoprogenitor cells
that are products of mesenchymally derived cell lines.
Osteoblasts are mononuclear cells and possess copious ribo-
somes and mitochondria that reflect their intensive protein
synthesis duties in the production of new bone. Osteoblasts
were once envisioned solely as bone-producing cells, but
we now know that their activities range from the synthesis
and deposition of an unmineralized collagen mass (osteoid),
to the secretion of calcium phosphate, osteoclast regulation,
hematopoiesis, and immune functions (Gosman, 2012).
In particular, there appear to be fundamental interactions
between blood and osteoblasts as they continuously
“talk” to each other as osteoblasts provide critical
regulatory support of the hematopoietic stem cell line
(Wu et al., 2009).
Osteoblasts are plump and polygonal in shape and
genetically are a sophisticated derived fibroblast (Ducy
et al., 2000). Osteoblasts produce a mucoprotein matrix
mainly consisting of highly structured layers of collagen
fibers (type I fibrils). This is followed by deposition of
solid particles of calcium phosphate as hydroxylapatite
crystals are embedded upon and between the collagen
fibers. Their ultimate fate resides in becoming either
osteocytes or bone-lining cells (Fig. 4.3).
Osteoclasts are cells that remove bone. They are large,
multinucleated cells that arise from the fusion of usually
39
Osteoblast lineage
Stem cell
?
Preosteoblast
Osteoblast
Osteocyte Bone-lining cell
FIGURE 4.3 A schematic representation of the origin and fate of
osteoblasts. Osteblasts share a stem cell source in common with fibro-
blasts. When the stem cell differentiates into a preosteoblast, its daughter
cells are committed to osteoblasts development. Ultimately, they may
undergo apoptosis, serve as BLCs, or become trapped in matrix and fur-
ther develop as an osteocyte. Reproduced with permission from Garner,
S.C., Anderson J.J.B., 2012. Skeletal tissues and Mineralization. In:
Anderson, J.J.B., Garner S.C., Klemmer P.J. (Eds.). Diet, Nutrients, and
Bone Health. CRC Press, Boca Raton, FL., p. 36.
between 10 and 20 mononuclear phagocytes, which them-
selves are macrophage-related hematopoietic precursor
cells in bone marrow (Boyle et al., 2003) (Fig. 4.4).
Osteoclasts possess a ruffled border that facilitates a
tight seal between cell and bone surface (Teitelbaum,
2000; Ross, 2013). Under this seal, the osteoclast creates
its own microenvironment. Osteoclasts possess electro-
genic proton pumps (H1-ATPase) and a Cl channel
that allows the cell to secrete hydrochloric acid and
collagen-degrading proteolytic enzymes. Under the seal,
local pH drops to around 4.5 and is sufficient to dissolve
bone matrix (Ross, 2013). Exposed collagen fragments
are then broken down by the enzyme cathepsin K and
the mineral and protein remains are expelled into the
surrounding intracellular fluid (Stenbeck and Horton,
2004). Just like osteoblasts, osteoclasts are mobile and
can move along a surface between 95 and 115 μm/h
(Hall, 2015) (Fig. 4.5).
Osteocytes are osteoblasts that became trapped in
bone matrix deposited by other surrounding osteoblasts.
The trapped cell then differentiates into an osteocyte,
decreasing in size and organelle volume. They occupy
ovoid-shaped cavities in the bone matrix called lacunae.
40 Ortner’s Identification of Pathological Conditions in Human Skeletal Remains

Osteoclast lineage Osteocytes appear to be proverbial “control freaks”

CFU-GM
(Gosman, 2012) that are the primary regulators of
bone mass as they integrate mechanical and hormonal
signals (Dallas and Bonewald, 2008; Bonewald, 2013;
(+) Hall, 2015; and see below). An emerging consensus
GM-CSF
portrays osteocytes as the master mechanosensory cell
Promonocyte in bone. Osteocytes retain thread-like cytoplasmic
IL-1; IL-6; (+) TGF-β extensions from their earlier lifetime as osteoblasts.
TGF-α (+)
These pass through small channels in the bone matrix
called canaliculi. These specialized extensions connect
Early to one another via gap junctions that allow osteocytes to
Monocyte
pre-osteoclast form a functional and communicative syncytium that
links all cells within bone. Together, they all function in
(+) a way that can be seen as a single mineral regulatory
PTH; endocrine gland (Bonewald, 2013; Qing et al., 2008).
1,25D
Tissue
More than 90% of all cells within mature bone are
Late
pre-osteoclast macrophage osteocytes.
Most critical to an understanding of bone cells is that
they do not operate independently of each other.
Osteocytes appear to first assess mechanical signals and
(+) then regulate bone shape and density by targeting
PTH;
1,25D osteoblast and osteoclast functions. Basically, it is the
Osteoclast Giant cell
balanced actions of osteoblasts and osteoclasts that
IL-6 (+) secure bone growth and maintenance. The balance
between osteoblastic and osteoclastic activity is equally
Osteoblast (–)
(+) CT fundamental for understanding the pathophysiology of
all diseases affecting bones (see Chapter 5). Some dis-
(+) Active osteoclast ease processes give rise to greatly enhanced osteoblastic
PTH; 1,25D; activity, while others are characterized by an elevated
IL-1; TGF-α osteoclastic response. Others feature mixed responses
or begin with elevated osteoclast activity but are later
eclipsed by new bone formation. Ultimately, these
FIGURE 4.4 A schematic representation of the origin and fate of
osteoclasts. Osteoclasts orginate from a stem cell population of colony- cells are not behaving autonomously, but are responding
forming-unit-granulocyte-macrophages (CFU-GM) that can either give to a yet deeper level of control: molecular signaling
rise to osteoclasts or monocytes/macrophages. CFU-GMs are stimulated mechanisms.
by the granulocyte-macrophage colony stimulating factor and differenti-
ate into premonocytes. This cell will then commit to either a osteoclastic
or monocytic pathway under the stimuli from varying local factors
including the influence of various interleukins (e.g., IL-1, IL-6) along Molecules and Signaling Pathways: Master
with transforming growth factor-α (TGF-α) for the osteoclast linage and
transforming growth factor-β (TGF-β) for the monocyte linage(s).
Control Mechanisms
Additional influences of parathyroid hormone (PTH) and 1,25-dihydrox- Since the late 1980s, advances in molecular biology and
yvitamin D (1,25D) and other cytokines, either acting alone or via osteo-
genetics have progressively revealed the major molecules,
blast mediation, further develops the cell and regulates the behavior of a
mature osteoclast throughout its life. Reproduced with permission from cell signaling pathways, and genes that drive and regulate
Garner, S.C., Anderson J.J.B., 2012. Skeletal tissues and Mineralization. bone cell behavior. These insights hold myriad implica-
In: Anderson, J.J.B., Garner S.C., Klemmer P.J. (Eds.). Diet, Nutrients, tions for paleopathological problem-solving (e.g., Crespo
and Bone Health. CRC Press, Boca Raton, FL., p. 38. et al., 2017).
Osteoblast-mediated bone production is fundamen-
tally controlled by a group of proteins called Wnt
It was once commonly held that osteocytes were simply (named after the wingless gene in Drosophila) via
unlucky osteoblasts that found themselves at the wrong Wnt cell surface coreceptor LRP5 (low-density
place at the wrong time and when trapped in the matrix lipoprotein-related protein 5). The Wnt/b-catenin
they entered into a passive state, retired from all meaning- pathway is commonly called the canonical pathway, as
ful and functional roles. Currently emerging understand- it activates gene expression in cell nuclei to incite
ing demonstrates the opposite. osteoblasts to form new bone (Fig. 4.6). Osteoblasts
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marriage of long ago, and what had come of it. Had they had
children, did their blood still run on the far worlds? But even if it did,
what was that now to those two of long ago?
There had been a poem about flowers at the end of the old book on
flowers Haller had lent him, and he remembered some of it.

"All are at one now, roses and lovers,

Not known of the winds and the fields and the sea,
Not a breath of the time that has been hovers
In the air now soft with a summer to be."

Well, yes, Kellon thought, they were all at one now, the Rosses and
the Jennies and the things they had done and the things they had
thought, all at one now in the dust of this old planet whose fiery final
summer would be soon, very soon. Physically, everything that had
been done, everyone who had lived on Earth, was still here in its
atoms, excepting the tiny fraction of its matter that had sped to other
worlds.
He thought of the names that were so famous still through all the
galactic worlds, names of men and women and places.
Shakespeare, Plato, Beethoven, Blake, the old splendor of Babylon
and the bones of Angkor and the humble houses of his own
ancestors, all here, all still here.
Kellon mentally shook himself. He didn't have enough to do, that was
his trouble, to be brooding here on such shadowy things. He had
seen all there was to this queer little old place, and there was no use
in coming back to it.

But he came back. It was not, he told himself, as though he had any
sentimental antiquarian interests in this old place. He had heard
enough of that kind of gush from all the glittering phonies in the ship.
He was a Survey man and all he wanted was to get back to his job,
but while he was stuck here it was better to be roaming the green
land or poking about this old relic than to have to listen to the
endless babbling and quarrelling of those others.
They were quarrelling more and more, because they were tired of it
here. It had seemed to them a fine thing to posture upon a galactic
stage by helping to cover the end of Earth, but time dragged by and
their flush of synthetic enthusiasm wore thin. They could not leave,
the expedition must broadcast the final climax of the planet's end,
but that was still weeks away. Darnow and his scholars and
scientists, busy coming and going to many old sites, could have
stayed here forever but the others were frankly bored.
But Kellon found in the old house enough interest to keep the waiting
from being too oppressive. He had read a good bit now about the
way things had been here in the old days, and he sat long hours on
the little terrace in the afternoon sunshine, trying to imagine what it
had been like when the man and woman named Ross and Jennie
had lived here.
So strange, so circumscribed, that old life seemed now! Most people
had had ground-cars in those days, he had read, and had gone back
and forth in them to the cities where they worked. Did both the man
and woman go, or just the man? Did the woman stay in the house,
perhaps with their children if they had any, and in the afternoons did
she do things in the little flower-garden where a few bright, ragged
survivors still bloomed? Did they ever dream that some future day
when they were long gone, their house would lie empty and silent
with no visitor except a stranger from far-off stars? He remembered
a line in one of the old plays the Arcturus Players had read. Come
like shadows, so depart.
No, Kellon thought. Ross and Jennie were shadows now but they
had not been then. To them, and to all the other people he could
visualize going and coming busily about the Earth in those days, it
was he, the future, the man yet to come, who was the shadow. Alone
here, sitting and trying to imagine the long ago, Kellon had an eery
feeling sometimes that his vivid imaginings of people and crowded
cities and movement and laughter were the reality and that he
himself was only a watching wraith.

Summer days came swiftly, hot and hotter. Now the white sun was
larger in the heavens and pouring down such light and heat as Earth
had not received for millennia. And all the green life across it
seemed to respond with an exultant surge of final growth, an act of
joyous affirmation that Kellon found infinitely touching. Now even the
nights were warm, and the winds blew thrilling soft, and on the
distant beaches the ocean leaped up in a laughter of spray and
thunder, running in great solar tides.
With a shock as though awakened from dreaming, Kellon suddenly
realized that only a few days were left. The spiral was closing in fast
now and very quickly the heat would mount beyond all tolerance.
He would, he told himself, be very glad to leave. There would be the
wait in space until it was all over, and then he could go back to his
own work, his own life, and stop fussing over shadows because
there was nothing else to do.
Yes. He would be glad.
Then when only a few days were left, Kellon walked out again to the
old house and was musing over it when a voice spoke behind him.
"Perfect," said Borrodale's voice. "A perfect relic."
Kellon turned, feeling somehow startled and dismayed. Borrodale's
eyes were alight with interest as he surveyed the house, and then he
turned to Kellon.
"I was walking when I saw you, Captain, and thought I'd catch up to
you. Is this where you've been going so often?"
Kellon, a little guiltily, evaded. "I've been here a few times."
"But why in the world didn't you tell us about this?" exclaimed
Borrodale. "Why, we can do a terrific final broadcast from here. A
typical ancient home of Earth. Roy can put some of the Players in
the old costumes, and we'll show them living here the way people
did—"
Unexpectedly to himself, a violent reaction came up in Kellon. He
said roughly,
"No."
Borrodale arched his eyebrows. "No? But why not?"
Why not, indeed? What difference could it possibly make to him if
they swarmed all over the old house, laughing at its ancientness and
its inadequacies, posing grinning for the cameras in front of it,
prancing about in old-fashioned costumes and making a show of it.
What could that mean to him, who cared nothing about this forgotten
planet or anything on it?
And yet something in him revolted at what they would do here, and
he said,
"We might have to take off very suddenly, now. Having you all out
here away from the ship could involve a dangerous delay."
"You said yourself we wouldn't take off for a few days yet!" exclaimed
Borrodale. And he added firmly, "I don't know why you should want
to obstruct us, Captain. But I can go over your head to higher
authority."

He went away, and Kellon thought unhappily, He'll message back to

Survey headquarters and I'll get my ears burned off, and why the
devil did I do it anyway? I must be getting real planet-happy.
He went and sat down on the terrace, and watched until the sunset
deepened into dusk. The moon came up white and brilliant, but the
air was not quiet tonight. A hot, dry wind had begun to blow, and the
stir of the tall grass made the slopes and plains seem vaguely alive.
It was as though a queer pulse had come into the air and the ground,
as the sun called its child homeward and Earth strained to answer.
The house dreamed in the silver light, and the flowers in the garden
rustled.
Borrodale came back, a dark pudgy figure in the moonlight. He said
triumphantly,
"I got through to your headquarters. They've ordered your full
cooperation. We'll want to make our first broadcast here tomorrow."
Kellon stood up. "No."
"You can't ignore an order—"
"We won't be here tomorrow," said Kellon. "It is my responsibility to
get the ship off Earth in ample time for safety. We take off in the
morning."
Borrodale was silent for a moment, and when he spoke his voice had
a puzzled quality.
"You're advancing things just to block our broadcast, of course. I just
can't understand your attitude."
Well, Kellon thought, he couldn't quite understand it himself, so how
could he explain it? He remained silent, and Borrodale looked at him
and then at the old house.
"Yet maybe I do understand," Borrodale said thoughtfully, after a
moment. "You've come here often, by yourself. A man can get too
friendly with ghosts—"
Kellon said roughly, "Don't talk nonsense. We'd better get back to the
ship, there's plenty to do before take off."
Borrodale did not speak as they went back out of the moonlit valley.
He looked back once, but Kellon did not look back.

They took the ship off twelve hours later, in a morning made dull and
ominous by racing clouds. Kellon felt a sharp relief when they
cleared atmosphere and were out in the depthless, starry blackness.
He knew where he was, in space. It was the place where a
spaceman belonged. He'd get a stiff reprimand for this later, but he
was not sorry.
They put the ship into a calculated orbit, and waited. Days, many of
them, must pass before the end came to Earth. It seemed quite near
the white sun now, and its Moon had slid away from it on a new
distorted orbit, but even so it would be a while before they could
broadcast to a watching galaxy the end of its ancestral world.
Kellon stayed much of that time in his cabin. The gush that was
going out over the broadcasts now, as the grand finale approached,
made him sick. He wished the whole thing was over. It was, he told
himself, getting to be a bore—
An hour and twenty minutes to E-time, and he supposed he must go
up to the bridge and watch it. The mobile camera had been set up
there and Borrodale and as many others of them as could crowd in
were there. Borrodale had been given the last hour's broadcast, and
it seemed that the others resented this.
"Why must you have the whole last hour?" Lorri Lee was saying
bitterly to Borrodale. "It's not fair."
Quayle nodded angrily. "There'll be the biggest audience in history,
and we should all have a chance to speak."
Borrodale answered them, and the voices rose and bickered, and
Kellon saw the broadcast technicians looking worried. Beyond them
through the filter-window he could see the dark dot of the planet
closing on the white star. The sun called, and it seemed that with
quickened eagerness Earth moved on the last steps of its long road.
And the clamoring, bickering voices in his ears suddenly brought
rage to Kellon.
"Listen," he said to the broadcast men. "Shut off all sound
transmission. You can keep the picture on, but no sound."
That shocked them all into silence. The Lee woman finally protested,
"Captain Kellon, you can't!"
"I'm in full command when in space, and I can, and do," he said.
"But the broadcast, the commentary—"
Kellon said wearily, "Oh, for Christ's sake all of you shut up, and let
the planet die in peace."

He turned his back on them. He did not hear their resentful voices,
did not even hear when they fell silent and watched through the dark
filter-windows as he was watching, as the camera and the galaxy
was watching.
And what was there to see but a dark dot almost engulfed in the
shining veils of the sun? He thought that already the stones of the
old house must be beginning to vaporize. And now the veils of light
and fire almost concealed the little planet, as the star gathered in its
own.
All the atoms of old Earth, Kellon thought, in this moment bursting
free to mingle with the solar being, all that had been Ross and
Jennie, all that had been Shakespeare and Schubert, gay flowers
and running streams, oceans and rocks and the wind of the air,
received into the brightness that had given them life.
They watched in silence, but there was nothing more to see, nothing
at all. Silently the camera was turned off.
Kellon gave an order, and presently the ship was pulling out of orbit,
starting on the long voyage back. By that time the others had gone,
all but Borrodale. He said to Borrodale, without turning,
"Now go ahead and send your complaint to headquarters."
Borrodale shook his head. "Silence can be the best requiem of all.
There'll be no complaint. I'm glad now, Captain."
"Glad?"
"Yes," said Borrodale. "I'm glad that Earth had one true mourner, at
the last."
THE END
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