Received: 26 July 2017 | Revised: 7 September 2017 | Accepted: 10 September 2017
DOI: 10.1002/ajpa.23322
CENTENNIAL PERSPECTIVE
Bioarchaeology in perspective: From classifications of the dead
to conditions of the living
Clark Spencer Larsen
Department of Anthropology, The Ohio State University, Columbus, Ohio 43220
Correspondence
Dr. Clark Spencer Larsen, Department of Anthropology, 4034 Smith Laboratory, Ohio State University, Columbus, OH 43210-1106.
Email: Larsen.53@osu.edu
Funding information
National Science Foundation; National Endowment for the Humanities; National Geographic Society; Edward John Noble Foundation; St. Catherines Island
Foundation
K E Y W O R D S : bioarchaeology, osteology, biocultural, revolutions
1 | INTRODUCTION
At no other time in either the history of physical anthropology or its
flagship journal, the American Journal of Physical Anthropology, has there
been such strong interest in the central role that human remains from
archaeological settings play in developing an understanding of the
remarkably dynamic last 10,000 years of the history, evolution, and
contextual circumstances surrounding the human condition. Over the
course of my association with bioarchaeology, I have watched the
expanding interest in the study of the remains of post-Pleistocene
humans as expressed in the number of articles in the pages of the
AJPA, increasing class enrollments and student interest generally, num-
ber of faculty positions, summer field schools around the globe, gradu-
ate student interest and recruiting, volume of presentations at the
annual meetings of the American Association of Physical Anthropolo-
gists, growth in other related professional societies (e.g., Paleopathol-
ogy Association, Dental Anthropology Association, British Association
for Biological Anthropology and Osteoarchaeology), new peer-
reviewed journals, book series, and presence in electronic news media.
I knew that bones and teeth were amazingly interesting when I took
my first osteology course taught by William Bass in my freshman year
at Kansas State University, but I had no idea of the broad extent of the
fund of data archaeological human skeletons provide for addressing
hypotheses and questions about the human past, especially as these
questions pertain to environment viewed broadly—diet, nutrition,
climate, social and cultural circumstances, health, lifestyle, activity, bio-
logical relatedness, and social, cultural, and population dynamics. As
bioarchaeology has shown, a person’s skeleton contains their cumula-
tive record of lived experiences and conditions, and skeletons from
community cemeteries provide a record over many generations.
Am J Phys Anthropol. 2018;165:865–878. wileyonlinelibrary.com/journal/ajpa
Moreover, the individual and community genetic history gives us a
baseline for addressing questions of how population history—from
marriage patterns to large scale mobility—has shaped human biology
through time and space.
Bioarchaeology has a long history around the world (O’Donnab-
hain & Lozada, 2014). Most exciting for me today is watching its spread
to areas not especially visible in the literature until relatively recently.
For example, Southeast and East Asia have become a central focus of
bioarchaeological inquiry, producing an impressive and expanding vol-
ume of problem-oriented research, fueled by long-standing as well as
newly arising questions relating to health, lifestyle, and population his-
tory (e.g., various in Oxenham & Buckley, 2016; Pechinkina & Oxen-
ham, 2013). In addition, while bioarchaeology is largely population-
oriented, other related areas of investigation pertaining to osteobiogra-
phy, biohistory, and the individual have also developed (e.g., Stodder &
Palkovich, 2012; Stojanowski & Duncan, 2016).
In this perspective, I present a personal view of major develop-
ments, turning points, and themes since the American Journal of Physi-
cal Anthropology began publication in 1918. This perspective is not
intended to be a comprehensive treatment of the history of bioarch-
aeology, the women and men who molded the field, or content (see
various in Buikstra & Beck, 2006; Larsen, 2015). Rather, I focus on
some of the research trends in building today’s integrative, contextual
study of human skeletal remains derived from archaeological settings. I
emphasize from the onset that bioarchaeological study today places
skeletons derived from archaeological settings in context, including, but
not limited to, environment, stressors present, availability of particular
resources (especially dietary), social and cultural influences, and other
factors that shape life conditions and the life course, many of which
are accumulated directly or as indirect outcomes, a process
V
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866 | LARSEN
commencing when skeletal and dental tissues begin to form in utero

rez,
and concluding only at death (Agarwal, 2016; Martin, Harrod, & Pe
2013). Placement of humans (and other primates) in context is a hall-
mark of physical anthropology.
In my view, the increasing interest in archaeological human
remains stems from the growing understanding that bones and teeth
present a multifaceted data array from which to reconstruct and inter-
pret the record of life conditions, health outcomes, and lifestyle. Bio-
archaeology’s multiscalar approach—individual, community, regional,
and global—provides a means for developing an understanding of the
human experience in the past. The record is especially strong in regard
to insights gained about behavior in multiple contexts and challenges,
adaptations, lifestyles, and health circumstances. In this regard, there is
a very strong platform for the development of an informed understand-
ing of the impacts and circumstances of fundamental transitions in
population size, mobility, diet, and other issues that are well repre-
sented in the AJPA and the growing scientific literature.
2 | BACKGROUND TO BIOARCHAEOLOGY
Although the term “bioarchaeology” as it applies to human remains was
not introduced until the 1970s (see below), the study of its core matter
—human remains derived from archaeological settings—has been
around for a long time. Much of this history stems from a period when
the groundwork for the founding of the discipline of physical anthro-
pology was being laid, including the roots of its professionalization in
the early twentieth century. In looking back over the last century or so,
especially at a time when the AJPA was being launched, it is clear that
without a strong profession behind it, there would be no support for or
direction in physical anthropology. High on the list of the develop-
ments of professional activities was the founding of the AJPA and its
sponsoring scientific association, both owing to the leadership, force-
of-will, and dogged perseverance of Ales Hrdlička. He viewed the study
of human skeletal remains—from all places and all times—as central to
understanding human biological variation. To address this variation,
thousands of these remains were either excavated or studied by him or
his associates over the course of his remarkable career, and especially
beginning with his appointment as head of the Division of Physical
Anthropology at the Smithsonian Institution’s National Museum in
1903. Although Hrdlička did not act alone in professionalizing the field,
it would have been a very different century for physical anthropology
had there been no Ales Hrdlička.
In the first decade or so of his employment at the Smithsonian,
Hrdlička began pulling together the necessary resources (including out
of his own pocket) to publish research results in his brainchild, the
AJPA (Giles, 2010). Hrdlička was keenly interested in starting a profes-
sion, modelled at least in part on his experiences in the laboratory
founded by Paul Broca in Paris, where human remains were central to
the study of human biological variation (Stewart & Spencer, 1978).
From the beginning of his quest for the development of a profession,
he envisioned three integrated parts—the peer-reviewed journal, a
scientific society, and an institute for training future professional
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physical anthropologists. For all three parts, human skeletal remains
from archaeological and other contexts (e.g., anatomical, paleontologi-
cal) were of essential importance. Although history shows that
Hrdlička’s plan for a developing an institute was a nonstarter, the
establishment of a professional journal and a professional society were
ultimately highly successful, and a century later, the journal and society
as we know them today have grown beyond his wildest dreams. In
addition, the setting Hrdlička created at the Smithsonian for research
and study has engendered growth in the science and leadership in
research, both on his part, as well as nearly a century of leaders in
physical anthropology, commencing with his recruitment of T. Dale
Stewart and Stewart’s problem-oriented research, such as regarding his
classic study on dental caries (Stewart, 1931) and other research in
skeletal biology spanning decades (see Angel, 1976; Ubelaker, 2006).
In 1918, while human remains were recognized as a significant
record of growth and development, pathology, and other issues
(Hrdlička, 1918, 1919), they were studied largely within a racialized,
typological framework, deriving from paradigms begun centuries before
and well in place by the nineteenth century (Cook, 2006; Wolpoff &
Caspari, 1997). As applied to archaeological human remains, Samuel
George Morton (1839), who Hrdlička (1914) credited with being the
founder of physical anthropology, was keenly interested in cranial vari-
ation and its classification, and concluded at least in his earlier studies
that Native Americans have a common morphology and, therefore, a
shared ancestry (and see Buikstra, 2006). As with mainstream human
biology in the nineteenth century, the typological perspective influ-
enced Hrdlička’s interpretation of human cranial variation. For example,
he classified cranial morphology categorically, including, “brachyce-
phals” (individuals and populations having relatively short cranial
vaults) and “dolichocephals” (those with relatively long cranial vaults).
Hrdlička’s other work and comprehensive vision for physical anthropol-
ogy (Hrdlička, 1918, 1919) made clear that this approach to cranial
morphology is fundamental to the science of physical anthropology.
Interestingly, though, Hrdlička (1934) observed that femoral diaphyses
form during growth and development and throughout older adulthood
are quite plastic, with morphological changes resulting mainly from
mechanical forces. Those views on at least the postcranial skeleton
were prescient of things to come nearly a half a century later regarding
bone as dynamic tissue involving cumulative developmental processes.
The typological approach to the study of human remains carried
considerable heft in physical anthropology laboratories and classrooms
in the United States and elsewhere well into the second half of the
twentieth century. Underlying the approach was a heavy focus on pop-
ulation history, to tell how people are related to each other—to families,
groups, and whole populations. Fundamental to this knowledge was an
understanding of biological relationships at all levels (Cook, 2006).
While Hrdlička was concentrating on the development of discipli-
nary infrastructure, Earnest Albert Hooton, founder of physical anthro-
pology at Harvard University, was singularly responsible for training
most of the first generation of physical anthropology’s academic lead-
ers (Giles, 2010). The majority of Hooton’s students developed exper-
tise in the study of human remains. Many of the individuals went on to

LARSEN
found or develop physical anthropology at academic institutions (e.g.,
Alice Brues, University of Colorado; Frederick Hulse, University of Ari-
zona; William Howells, University of Wisconsin; Paul Baker, Pennsylva-
nia State University; Joseph Birdsell, University of California at Los
Angeles; James Spuhler, Ohio State University; Charles Snow, Univer-
sity of Kentucky; Sherwood Washburn, University of Chicago, Univer-
sity of California at Berkeley).
Like Hrdlička and others in the first half of the twentieth century,
Hooton characterized cranial morphology in the typologically-based
tradition dominant at the time, applying the racial, categorically-based
paradigms to crania recovered from various locations (e.g., Hooton,
1918, 1920, 1930). Hooton’s typological influence is seen in the
approaches to the study of cranial morphology taken by his students in
various projects involving human remains from archaeological settings
(e.g., Angel, 1946a; Hulse, 1941; Marshall & Snow, 1956; Newman &
Snow, 1942; Snow, 1948, 1974; Webb & Snow, 1945). These
approaches made assumptions about population history, namely that
groups have specific cranial morphology that could be used to track
population history, such as in Eastern North America (e.g., Neumann,
1952; Robbins & Neumann, 1972; and see Smith, 1993). On the other
hand, Hooton approached the study of human remains in the context
of regionally-based questions, and his interest in health and demogra-
phy helped pave the way for future productive directions in bioarch-
aeology and paleopathology (Buikstra, 2006). Moreover, his book
dedicated to Hrdlička, The Indians of Pecos Pueblo: A Study of Their Skel-
etal Remains (Hooton, 1930), stands as a remarkable contribution to
the early development of bioarchaeology, including especially his inclu-
sion of contextual matters, description, and attention to paleodemogra-
phy, pathology, and morphology in temporal space. The book also
provided an important basis for the extensive follow-up research on
the Pecos Pueblo skeletal series (various in Morgan, 2010). Hooton’s
interest in regional context clearly influenced future growth in bioarch-
aeology, especially as shown in the work and career of his student, J.
Lawrence Angel, and Angel’s long-term study of populations in ancient
Greece, in particular, and the Mediterranean Basin, in general (e.g.,
Angel, 1944b, 1946b, 1966, 1971). His early focus on social context
foreshadows developments in bioarchaeology decades later.
Typological approaches to population history are deeply flawed in
part because they mistakenly assume that skeletal biology is a static
entity. While today’s view that skeletal tissues are as dynamic as any
other living tissue was mostly missed at the time, others well before
Hooton were aware of this fundamental characteristic of bone. For
example, in the late nineteenth century when Julius Wolff’s (1892) pio-
neering work, The Law of Bone Remodeling, was published, there was lit-
tle interest by those moving in the direction of what was to become
physical anthropology in the twentieth century. Decades later, another
of Hooton’s students, Sherwood Washburn (1947), pioneered experi-
mental studies on mastication in rats, a study revealing the important
role of masticatory muscles on craniofacial morphology. Clearly, the
time was not yet ripe for physical anthropologists to engage experi-
mental research having relevance to skeletal biology of past human
populations (and see Larsen, 2015; and below).
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| 867
There were a few emerging professional physical anthropologists in
the first half of the twentieth century that were identifying patterns of
skeletal variation that were not typologically motivated. Rather, these
professionals were encountering variation in archaeological skeletons
when viewed in social, cultural, and behavioral contexts. The big boost
that would lay important groundwork for what would become bioarch-
aeology is due in important ways to the establishment of federally man-
dated work relief programs in areas of the United States hardest hit by
the Great Depression of the 1930s. As part of Roosevelt’s New Deal
and putting the unemployed back to work, many thousands of Ameri-
cans were hired in a wide range of federally funded jobs, including as
excavators and field technicians in state-of-the-art archaeological exca-
vations at high-profile sites, mostly in the American Southeast where
the Depression had hit hardest. For at least some of the projects—espe-
cially those involving the excavation and study of large numbers of
human remains—graduate students and newly produced PhDs from
leading institutions (e.g., University of Chicago, Harvard University)
were recruited. Hooton’s past and current graduate students (e.g.,
Charles Snow, Frederick Hulse, and Marshall Newman) had training in
physical anthropology and skeletal anatomy. Study of human remains,
including skeletal and dental pathology, revealed a collective record of
elevated infectious disease, dental caries, and other evidence of morbid-
ity in late prehistoric farming societies (e.g., Lewis & Kneberg, 1946;
Newman & Snow, 1942; and see discussions in Larsen, 2012; Smith,
1993). In addition, other aspects of analysis were undertaken, some of
which focused on extensive documentation of large series of skeletal
remains, such as those recovered at Indian Knoll (Snow, 1948; see
Smith, 1993). Largely owing to historical events involving the end of the
Great Depression and the onset of World War II, very little of the post-
excavation research continued, and the volume of bioarchaeology in
general tapered off considerably (although see Hoyme & Bass, 1962).
Nevertheless, discussions of health and lifestyle stemming from work
relief projects foreshadowed the remarkable expansion in bioarchaeo-
logical research that was to come. In addition, not all of Hooton’s stu-
dents viewed cranial variation in post-Pleistocene populations as
sorting out into discernable types. William Howells’s (1973) Cranial Vari-
ation in Man and his subsequent analyses based on many thousands of
individuals globally revealed no evidence for racial categories, past or
present, discrediting the notion that there are identifiable types.
3 | THE BIRTH OF BIOARCHAEOLOGY
Many of the developments and innovations resulting in what bioarch-
aeology is today were first introduced in the pages of the American
Journal of Physical Anthropology, and played a founding role in trans-
forming the science from one of description and classification to one
focusing on understanding human remains as the record of once-living
people. This change in perspective gives the context for the transfor-
mation from typological, descriptive research to the investigation of
processes associated with the interplay between the complex natural,
cultural, and social circumstances that shape skeletal and dental struc-
tures in life and preserved in death.

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The growing understanding of the varied and complex circumstan-
ces that shape the human skeleton and dentition opened up a new era
of transformative research in the 1970s. The term bioarchaeology was
first introduced as it applies to human remains from archaeological con-
texts by Jane Buikstra (1977) in her pioneering “bioarchaeological”
study of human remains from the lower Illinois River valley. During a
time when classification and typology were still a part of the study of
past populations, a pivotal symposium was organized by Robert Blakely
(1977), aimed at discovering processes of biocultural integration and
adaptation as documented by the study of human remains from
archaeological settings. In that now-legendary occasion, Jane Buikstra
(1977:69) proposed a study of archaeological human remains that
was broadly contextualized, specifically, one that was comprised of
“. . .regionally based, interdisciplinary research in mortuary site archae-
ology and human osteology.”
Simultaneously, George Armelagos and his colleagues were making
the case for a skeletal biology that would shift from focus on descrip-
tion to focus on the biological study of human remains in the context
of culture, behavior, and environment, and concurrently addressing
hypotheses and research questions (Armelagos, Carlson, & Van Gerven,
1982; Carlson & Van Gerven, 1977, 1979; Lovejoy, Burstein, & Heiple,
1976). Their collective work resulted in a number of important theoret-
ical innovations, including the application of the Selye’s stress model in
the larger discussion of biocultural approaches to the study of health,
disease, and skeletal biology (e.g., Goodman, Martin, Armelagos, &
Clark, 1984) and the application of the Barker hypothesis regarding
early stress and early mortality (e.g., Armelagos, Goodman, Harper, &
Blakey, 2009). These developments laid the groundwork for research
programs investigating patterns of health and lifestyle, including large
integrative studies such as in the context of the foraging to farming
transition in prehistory (e.g., various in Cohen & Armelagos, 1984;
Cohen & Crane-Cramer, 2007), human adaptation and biological his-
tory within large regions (e.g., American Great Plains: Bass, 1981; Ows-
ley & Jantz, 1994), paleodemographic patterning (Buikstra, Konigsberg,
& Bullington, 1986), health transitions in the Western Hemisphere
(e.g., Steckel & Rose, 2002), and the African diaspora in North America
(e.g., Blakey & Rankin-Hill, 2009).
As a graduate student at the University of Michigan in the mid-
1970s, I well recall the string of “ah-ha” moments for me, especially in
reading Buikstra’s and Armelagos’s work and other newly-emerging
approaches to the study of archaeological human remains (Buikstra,
1976, 1977; Carlson, Armelagos, & Van Gerven, 1976; Ubelaker,
1974). At the time, I was just beginning a long-term research collabora-
tion with David Hurst Thomas and the American Museum of Natural
History on the U.S. Georgia Atlantic coast (Thomas, Jones, Durham, &
Larsen, 1978), hoping that the project would provide the basis for
my Ph.D. dissertation. The winds of change involving the shift from
old-style skeletal studies to the new wave of bioarchaeology, a field
built on understanding of skeletal biology, were clearly in development,
especially with Buikstra’s, Armelagos’s, and other researchers’
emphases on regionally-based, population-oriented studies (see Larsen,
2015). Their collective work and the students and professionals they
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were influencing led to a sea-change in how human remains from
archaeological contexts were being studied, what has shaped the field
as we know it today, and what has promoted new developments that
make the field so vibrant and engaged with other disciplines. I was also
excited about these new developments because in a number of key
ways, they were consistent with the innovative and interdisciplinary
approaches espoused by Sherwood Washburn (1951) in his proposed
“New Physical Anthropology,” advocating more meaningful ways of
understanding biology and links to behavior, testing hypotheses, mov-
ing from technique-oriented to problem-oriented research, and giving
new direction and relevance to physical anthropology.
The following discussion highlights selected key developments in
bioarchaeology, building on the above advances in the field. I discuss in
the remainder of this perspective what I think of as “revolutions” and
the depth of understanding we have gained about past populations
from new and innovative perspectives, tools, and ways of looking at
past human biology in context. In particular, the following focuses on
the dynamic nature of bone as it applies to masticatory and ambulatory
functions; dietary reconstruction and nutritional inferences based on
biogeochemistry of stable isotopes; the recovery and analysis of bio-
molecules, especially ancient DNA (aDNA), and its contribution to bio-
distance analysis; and the record of aDNA in documentation of ancient
pathogens.
I refer to these developments as “revolutions,” largely because of
the remarkable opportunities they offer in understanding recent human
evolution and variation. Moreover, these areas are emblematic of mod-
ern problem-oriented research that has engaged interdisciplinary
theory and method, and that have benefitted from technological and
methodological developments that I view as having significant informa-
tive power and setting the stage for future research questions and
issues. In all ways, these developments have helped to fuel interdiscipli-
nary, collaborative-oriented science that the field has experienced over
the past several decades.
Most importantly, the areas described below engage behavior—a
fundamental element of anthropology in general—broadly defined to
include elements of social, cultural, and economic behaviors that influ-
ence the access that individuals and aggregates of individuals have to
resources, and how humans function, perform, and thrive in these com-
plex landscapes. Moreover, the cumulative record of these influences
on the individual over the course of their lifetime and over the course
of aggregations of individuals over generations within a community or
regional setting give us a powerful picture of past life conditions and
adaptive networks. So, to be clear, when I refer to behavior, I am refer-
ring not to physical activity, such as movement of muscles, bodies, and
parts of bodies. Rather, I am referring to the broad suite of behavioral
elements that make us what we are in life and how the signature of
that complexity is manifested in the skeleton. In some ways, bioarch-
aeology offers a way of addressing issues relating to cause, especially
regarding the global transition from foraging to farming, arguably one of
the most dramatic changes in lifeway—on health, lifestyle, living circum-
stances over the life course, mobility, and other terms that comprise the
human condition as based on the cumulative record of life experiences.

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4 | REVOLUTION 1: SKELETAL
FUNCTIONAL ADAPTATION VIA
BIOMECHANICAL ANALYSIS
In their study of a temporal series of crania from Nubia in the Nile
River valley, David Carlson and collaborators Dennis Van Gerven and
George Armelagos documented distinctive morphological changes in
comparison of earlier hunter-gatherers and later farmers. Like other
anthropologists, they observed that hunter-gatherer populations tend
to have relatively long cranial vaults and robust faces and jaws,
whereas later farmers tend to have shorter, rounder cranial vaults with
gracile faces and jaws. The earlier, typological approaches to the study
of crania from the region had concluded that the morphological differ-
ences between earlier and later periods reflected population replace-
ment—simply, racial groups having round, gracile crania invaded and
replaced the earlier racial groups having long, robust crania. However,
Carlson and Van Gerven (1977, 1979) took issue with the racialized,
classification paradigm and applied their newly developed “biocultural”
approach that culturally and behaviorally contextualizes skeletal biol-
ogy. Viewed in this new paradigm involving cultural contexts coupled
with craniofacial biology, they made the strong case that the shift from
harder-textured foods consumed by earlier hunter-gatherers to softer,
less textured foods associated with grains boiled into soft mushes
would result in a reduction in mechanical demand on the face and jaws
during mastication. Therefore, owing to a shift from hard-textured to
soft-textured foods, craniofacial morphology underwent functional
adaptation to changing masticatory behavioral circumstances. Impor-
tantly, the testing of their masticatory-functional hypothesis offered a
new way of interpreting craniofacial morphology in this setting and
potentially other settings around the globe. Simply, while cranial mor-
phology may be one element of population relatedness, other behav-
ioral factors influence the muscle-bone interaction during the growth
and development of the cranial vault, face and jaws.
The introduction of a new way of documenting and interpreting
craniofacial morphology and functionally related adaptations, set into
motion a reinterpretation of old data and development of research in a
range of settings in local, regional, and global contexts that underwent
the foraging-to-farming transition, especially where cooking technology
and food consistency changed (reviewed in Larsen, 2015; and see von
Cramon-Taubadel, 2017; Katz, Grote, & Weaver, 2017).
This is not to say that cranial morphology is determined by exter-
nally induced factors alone. Indeed, new developments in biological dis-
tance analysis are also providing perspective on population history
having little to do with masticatory function and more to do with evo-
lutionary dynamics, such as gene flow and the degree to which popula-
tions are isolated or represent movement of population (Galland &
Friess, 2016; Herrera, Peart, Hernandez, Spradley, & Hubbe, 2017;
Jantz & Spradley, 2014; Pilloud & Hefner, 2016). As expected, bio-
logically informed research reveals that there are clear patterns of
morphological variation linked with specific populations and regions.
Using multivariate statistical methods, analyses of cranial and dental
characters documents a record of population dispersion, gene flow,
and intra- and interpopulation relatedness at various levels (e.g.,
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| 869
Hemphill & Mallory, 2004; Konigsberg, 2006; Pietrusewsky, 2013;
Stojanowski, 2013; von Cramon-Taubadel & Pinhasi, 2011; various
in Pilloud & Hefner, 2016).
As with the skull, the postcranial skeleton presents a record of
activity-based lifestyle and behavioral patterns, including especially
those relating to mechanical function involving workload and mobility.
In the nineteenth century, Julius Wolfe considered the role of long
bones in their functional context, especially with regard to adjusting
size, amount, and distribution of bone in response to mechanical forces.
This concept, Wolff’s Law (or “bone functional demand”), states that
bone tissue places itself where needed to maintain mechanical integrity
(Ruff, Holt, & Trinkaus, 2006). We are now well aware that bone tis-
sues adapt to and are shaped by the mechanical environment.
Although the mechanical environments of the cranium and postcranial
skeleton are very different—the former in regard to mechanical forces
associated with mastication, and the latter to ambulatory functions
(lower limbs) and grasping, carrying, throwing, and other behaviors
(upper limbs). Nonetheless, the skeletal tissues comprising the cranium
and postcranium operate in the context of physiological functions
involving bone modeling and remodeling during the years of growth
and development.
The breakthrough in the recent history of physical anthropology
was the application of engineering principles that influence the devel-
opment of some aspects of skeletal morphology—especially in the way
that bone tissue is distributed in the diaphysis perpendicular to the
long axis of limb bones (e.g., humerus, femur). Lovejoy, Burstein, and
Heiple (1976) were the first to apply engineering-based beam theory
to the analysis of cross-sections of long bones in archaeological con-
texts. This framework and its application to archaeological contexts
offered an important means for addressing long-standing issues regard-
ing labor, activity, and mobility in a broad comparative perspective.
Lovejoy’s prescient AJPA article signaled an interest in a number of
us regarding engineering-based behavioral models and interpretations
of the postcranial skeleton. A long history of study of postcrania
revealed generally greater skeletal size-based robusticity in hunter-
gatherers compared to farmers, likely reflecting greater mechanical
loading in the former compared to the latter. In my dissertation
research based on a regional study of the southeastern US Atlantic
(Georgia) coast, I had documented a clear reduction in skeletal robustic-
ity based on external size of femora and humeri (Larsen, 1982). In par-
ticular, the earlier hunter-gatherers had greater midshaft robusticity
than the later maize-based farmers. I presented the research at a sym-
posium organized by Christopher Ruff at the AAPA meetings in 1980.
He also presented his research on cross-sectional geometric analysis of
the famous Pecos Pueblo series. We put into place a plan to document
behavioral changes relating especially to workload and lifestyle in the
context of economic, behavioral, and cultural changes relating to the
agricultural transition on the southeastern US Atlantic coast. Our
resulting analysis opened up a window of opportunity to understand
the dynamics of a setting undergoing population size increase, eco-
nomic intensification, and major social and behavioral changes in the
rise of complex societies and colonialism (Ruff, Larsen, & Hayes, 1984;

870 | LARSEN
Ruff & Larsen, 2001). The research began a long trajectory of biome-
chanical analyses expanding to other contexts globally in a diverse
range of local, regional, and other contexts (see Larsen, 2015; Pinhasi &
Stock, 2011; Ruff et al., 2015).
The case remains, however, that without being able to observe
behavior directly, the record for skeletal robusticity and its relationship
to mechanical demand, including mobility, are limited. In this regard,
newly emerging human experimental and bioethnographic evidence
reveals important findings that are consistent with the bioarchaeologi-
cal record regarding mobility and behavioral regimes (e.g., Shaw &
Stock, 2009a, 2009b; Stieglitz, Trumble, Kaplan, & Gurven, 2017).
Moreover, the developmental changes, including bone mass and robus-
ticity in earlier life are maintained in later life well after the loading
regime ceases (e.g., Warden et al., 2014). That is, for both living and
past populations, activity confers long-term benefits—greater balance
of bone—continuing well into adulthood. Importantly, these experimen-
tal and observational studies of living humans play a key role in inform-
ing our understanding of behavioral reconstructions in past populations
as well as long-term benefits that may be selectively advantageous,
both in the past and the present.
Although the living and experimental sources discussed here are
crucial for interpreting morphology and behavior, the contribution of
bioarchaeology in the broad scope of time, culture, and context gives
the discipline tools for understanding the activity repertoire of past
populations, which of course is not possible with the narrow window
of time and change demonstrated via the study of living humans. An
important caveat remains regarding any bioarchaeological investigation
of morphological change. Using mice models, there is growing experi-
mental evidence to show that skeletons of mice from different inbred
strains express different responses to mechanical loading (Robling &
Turner, 2002). This finding underscores the point that there are
likely different mechanical-responsive patterns. However, patterns of
mechanical loading and skeletal adaptations almost certainly reflect
responses to loading (Wallace, Demes, & Judex, 2017).
5 | REVOLUTION 2: STABLE ISOTOPE
RECORD FOR RECONSTRUCTING DIET,
MOBILITY, POPULATION HISTORY, AND
HEALTH
Nutrition scientists, physiologists, and biogeochemists have long under-
stood that along with many other constituents bones and teeth contain
stable isotopes of various elements, including for example, the stable
12
isotopes of carbon ( C, 13 14
C) and nitrogen ( N, 15
N). Moreover, the
plants and animals we eat contain isotopic variations that are repre-
sented in our tissues, including the skeletal and dental tissues. For
most of the history of the field, archaeologists have relied largely on
dietary reconstruction of past populations based on the documentation
of remains of plants and animals recovered from archaeological midden
deposits and tools used to extract or prepare foods. Although these
archaeological records provide meaningful dietary context, mere identi-
fication of animal and plant remains can be a notoriously inaccurate
record with regard to assessing the relative importance of specific foods
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or nutrients, thus drawing limited inferences about a population’s or
community’s nutritional status.
In the late 1970s, J.C. Vogel, a geochemist, and Nicholas van der
Merwe, an archaeologist, applied the principles of photosynthesis—C3
(most temperate-adapted plants) vs. C4 plants (plants adapted to hot/dry
climates)—and the characteristics of stable isotope ratios to the bioarch-
aeological record of the human consumers of the plants (Vogel & van der
Merwe, 1977). In this regard, C3 plants have a stable isotope ratio
13
C/12C (d13C) that is more negative than C4 plants (e.g., maize). Vogel
and van der Merwe’s analysis of a North American temporal series of
prehistoric archaeological skeletal remains from earlier hunter-gatherers
with later maize farmers produced the predicted result, namely that the
earlier group consuming largely non-domesticated C3 sources had more
negative d13C values in their skeletons than those later groups consuming
domesticated (maize) C4 sources. Their hypothesis proved to be strongly
supported by the results, setting off a revolution in reconstruction of past
diets in a wide range of settings globally. Owing to the precision of the
method, stable isotope analysis also offered a stronger record from which
to infer nutrition and nutritional changes than the aforementioned tradi-
tional methods based on plant and animal remains alone.
Soon after their remarkable discovery, our Georgia coastal project
provided van der Merwe and Vogel’s research group bone samples
from sites representing earlier prehistoric foragers and later prehistoric
farmers. To our excitement, their results were fully consistent with a
shift from C3 to C4 diets, indicating a strong presence of maize in diets
of late prehistoric populations, and lack of maize in earlier groups.
On the other hand, other stable isotope research was showing
that consumption of marine foods (and other animal-sources of pro-
tein) will produce d13C values that overlap with those produced in
maize consumers. We subsequently began working with Margaret
Schoeninger, who like van der Merwe and Vogel, was pioneering new
developments in isotope biochemistry and whose experimental evi-
dence revealed that marine foods produce a stable isotope ratio of a
nitrogen (15N/14N expressed as d15N) signature that when plotted with
d13C values in bivariate plots allowed us to tease apart relative contri-
butions of marine-based sources vs. maize (Schoeninger, van der
Merwe, Moore, Lee-Thorp, & Larsen, 1990). Subsequently, we worked
with Schoeninger’s lab and more recently with Laurie Reitsema’s lab to
produce the robust picture of dietary change in earlier prehistoric, later
prehistoric, contact-era, and mission-era dietary transitions, mostly
involving the adoption and increased focus on maize and simultaneous
decrease in marine foods (Larsen, Schoeninger, van der Merwe, Moore,
& Lee-Thorp, 1992; Reitsema, 2016). The beauty of large temporal and
regional studies is that they identify patterns of variation across time
and across landscapes, including identification of key shifts in diet and
where foods are derived, such as those acquired via long distance (rare)
or locally (common) both in the Georgia coastal setting (e.g., Hutchin-
son, Larsen, Schoeninger, & Norr, 1998) and elsewhere (e.g., Holder,
Dupras, Jankauskas, Williams, & Schultz, 2015; Papathanasiou, Rich-
ards, & Fox, 2015; and many other settings).
Biogeochemical analysis of the hard tissues presents a remarkably
sensitive record of key life course shifts as they are linked with diet. A

LARSEN
fundamental area of physical anthropology in general and bioarchaeol-
ogy in particular is the record of growth and development, addressing
questions relating to dietary reconstruction and nutritional inference
during early development in a far less speculative manner than prior to
the isotope revolution. A central issue is the duration—from initiation
to completion—of the weaning process. There is now an impressive
record showing variation in weaning patterns in bioarchaeological con-
texts (e.g., Tsutaya & Yoneda, 2015; Wright & Schwarcz, 1998; and
many others). Based on the premise that at birth the infant skeleton
has a stable isotope signature that is nearly identical to the mother, but
as soon as breast-feeding begins shortly after birth the values rise to a
peak within the first year. This pattern reflects the fact that the infant
is a consumer of the mother’s protein product. Depending on cultural
and social factors, the peak values gradually decline as the mother
removes the infant’s access to breastmilk. With the considerable num-
ber isotopically-informed bioarchaeological weaning studies now avail-
able in the literature, it is clear that there is substantial variation on the
timing of weaning in the past. In this regard, weaning practices were—
as in today’s world—locally determined by an array of cultural, social,
and environmental factors.
At this point, in the absence of written records, bioarchaeology
comes up short on why some populations have relatively long duration
of weaning, whereas others have short weaning periods. Clearly, the
decisions are culturally and socially mediated owing to local and other
circumstances. Dupras and collaborators’s (Dupras, Schwarcz, & Fair-
grieve, 2001) analysis of the Roman-period Kellis series from Dakleh
Oasis, Egypt, outline the circumstances, explaining at least in part why
complementary (adult) foods were introduced to infants by 6 months
with breast-feeding ending by 3 years. Regardless of the duration, the
terminal access to breast milk is often associated with the appearance
and elevation in prevalence of pathological conditions reflecting growth
disruption, owing to poor postweaning nutrition and exposure to infec-
tious disease and compromised immune function, among other issues
facing infants and young children (Sandberg, Sponheimer, Lee-Thorp, &
Van Gerven, 2014; Temple, 2016).
The record of weaning and its duration is fundamental for address-
ing a wide range of demographic outcomes. For example, in general,
human population size has increased in many settings that saw a shift
in subsistence from foraging to farming. The question then becomes,
did shortened duration of weaning result in increased birthrates? The
ethnographic record indicates that shortened duration of weaning
results in an increase in number of infants born to a mother. This gen-
eral pattern suggests that decrease in birth spacing and increased
velocity of population growth are closely related (Katzenberg, Herring,
& Saunders, 1996; and see Buikstra et al., 1986). The link between lac-
tation and female fecundity speaks to the high energetic investment
associated with producing milk for the infant’s consumption (Ellison,
Bogin, & O’Rourke, 2012). Ellison et al. (2012) argue that natural selec-
tion may have influenced the mother’s reproductive capacity during
the period when her investment in the young infant is most elevated.
Today, skeletal and dental tissues from many thousands of
archaeological individuals globally have been sampled for carbon and
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| 871
nitrogen stable isotopes, revealing patterns of food consumption prac-
tices that were not available prior to the stable isotope revolution. In
the larger picture, the combination of sedentism and population
increase explains at least in part the rise in infectious disease in later
Holocene populations, including some of the key diseases that can be
documented in archaeological contexts (e.g., tuberculosis; and see
below). It comes as little surprise, then, that many of the late prehistoric
societies supported by maize-based or otherwise carbohydrate-
enriched diets, living in sedentary, densely crowded communities
around the world experienced an increase in prevalence of infectious
diseases.
The powerful nature of stable isotope analysis of carbon and nitro-
gen—and other biogeochemical markers—is evidenced by the explosion
in number of articles published, especially in the pages of the AJPA.
This expanding record gives us a remarkably rich and growing picture
of the evolution of diet, allowing an informed understanding of nutri-
tional outcomes and social impacts on a grand temporal and global
scale (Larsen, 2015).
The isotopic record offers a compelling tool for addressing broader
and fundamental social questions regarding diet and nutrition in past
societies, such as the implications of the shift from foraging C3-based
diets to agricultural (e.g., maize or millet) C4-based diets, and accompa-
nying changes in social organization, often involving a shift to more
complex, hierarchical societies having differential access to resources
among their constituents (e.g., Dong et al., 2017; Winkler et al., 2016;
various in Klaus, Harvey, & Cohen, 2017). For example, in the central
Plains of China, when millet (a C4 plant) is replaced by wheat (a C3
plant) during the Bronze Age, females saw an increased consumption
of wheat and reduced consumption of animal sources of protein (Dong
et al., 2017). The dietary record and the archaeological context point to
a pronounced social shift involving male-biased inequality. This finding
is consistent with increased sexual dimorphism in stature and greater
display of wealth in male burials compared to female burials in this
setting.
The expansive application of stable isotope analysis to past popu-
lations has resulted in the development of other windows onto the
structuring of past societies, but central to the reconstruction of past
mobility and migration patterns, areas of study having a long history of
interest by anthropologists. Biogeochemistry reveals distinctive isotope
ratios that clarify level and patterns of mobility in past societies. The
87
Sr/86Sr ratios of foods consumed by humans reflect the 87
Sr/86Sr
ratios of the underlying geology (Bentley, 2006). The presence of sig-
nificant 87Sr/86Sr differences in isotope ratio values assessed for a fully
mature adult’s first molar crowns and their femoral diaphyses provide a
strong indication that the person lived in a different setting in child-
hood than in adulthood. For example, the analysis of 87Sr/86Sr ratios in
the heartland capital of Tiwanaku (Bolivia) and the outlying community
of Chen Chen (Peru) reveals the presence of persons in the latter who
had originally come from the heartland capital (Knudson & Price, 2007).
87
Variability in the Sr/86Sr ratio in comparison of adult females and
adult males in the Neolithic farmers of central Europe reveals consider-
able variation in adult females—far more than adult males, and beyond

872 | LARSEN
expectation for the biogeochemistry (Bentley et al., 2012). The pattern
is consistent with patrilocal residence—males, but not females, remain
in the natal setting following marriage. These and other biogeochemical
signatures provide essential context for interpreting a range of social
and behavioral issues well beyond the scope of diet alone (Grupe &
McGlynn, 2006).
6 | REVOLUTION 3: ASSESSING
POPULATION HISTORY FROM ANCIENT
BIOMOLECULES: AN EXPANDING
BIODISTANCE AGENDA IN THE GENOMIC
AGE
Population history, or biodistance, has deep historical roots in the disci-
pline. For many of the areas pertaining to bioarchaeology, the ability to
document biological continuity is central in the identification and inter-
pretation of a range of issues, such as temporal variation in cranial mor-
phology or stable isotope ratios. Although the methods are different now
than they were a century ago, the general problem remains for under-
standing biological relatedness—to each other at levels of family, commu-
nity, or population. Patterns identified in the archaeological record
facilitate investigation of larger evolutionary issues, such as the direction
and amount of gene flow, social organization, and historical connectivity.
The key difference in theory and method behind today’s biodistance
analysis with that of 1918 is that the former is informed by the range of
intrinsive and extrinsive factors that influence skeletal and dental biology,
whereas the latter was informed by racial paradigms based on misunder-
standing of the biological basis of cranial morphology. Thus, the questions
asked today by bioarchaeologists are in many ways similar to those asked
a century ago, but the tools, data sources, and interpretive frameworks
used to answer the questions are substantively different.
The genetic record as represented by ancient DNA is one of a num-
ber of obvious sources of biodistance data not available in 1918. It
would not be until DNA fragments were first identified by Pääbo (1985)
from tissue samples taken from a 2,000-year-old Egyptian mummy that
some of the recent breakthroughs in understanding local, regional, and
continental patterns of migration and gene flow began to be identified.
The development of polymerase chain reaction, next generation
sequencing technology, and continuing technological advances have
made it possible to document mitochondrial and nuclear DNA variation,
to the point where the entire genome and genomic variation of modern
and archaic humans (e.g., Neandertals) are now being explored and, in
some instances, clarified. As a result, paleogenetics has developed to
the point where it is becoming central to addressing issues of individual,
family, community, regional, and phylogenetic relationships.
The aDNA record, in combination with dental and cranial sources
of morphological and morphometric evidence is beginning to address
key areas of concern that go back decades in the discipline. This record
is especially powerful when it is integrated with the bioarchaeological
and archaeological evidence, a context-related practice that is continu-
ing to develop but has much further to go (and see Johannsen, Larson,
Meltzer, and Vander Linden, 2017). This is an important point because
even if the extraction and characterization of aDNA are successfully
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performed, without the archaeological context and the biological his-
tory in cultural, social, and behavioral perspective, the use of aDNA to
reconstruct population history is similar to what the nineteenth- and
early twentieth-century physical anthropologists were attempting in
comparing artificial cranial types. The aDNA record has considerable
promise, but its potential for understanding social and cultural dynam-
ics is only now being developed. On the other hand, the record is
beginning to provide evidence of hereditary relationships, community
structure, and residential variation (e.g., Kennett et al., 2017) that is
complementing earlier craniometric biodistance analysis (Stojanowski &
Schillaci, 2006).
Collaboration of bioarchaeologists, paleogeneticists, and archaeolo-
gists collectively addressing old or otherwise long-standing issues and
questions is a key objective now in process and in planning stages world-
wide. One of the famously long and ongoing discussions is the debate
regarding the origins of farming and its spread from primary centers.
This discussion is centered around the following question: Was the
spread of crop technology from these primary centers due to the cultural
diffusion of the practice to groups peripheral to these centers, or did it
come about by demic diffusion involving actual movement of people
from one place to another, bringing the practice and technology with
them and either fully or partly replacing indigenous hunter-gatherers?
Assessment of the genetic (ancient and modern), archaeological,
and bioarchaeological literature suggests no clear answer to the ques-
tion of cultural vs. demic diffusion. Indeed, the literature presents con-
flicting evidence or a mosaic of cultural and demic diffusion, especially
regarding the origins of agriculture in Europe (e.g., Brace et al., 2006;
von Cramon-Taubadel & Pinhasi, 2011). The genetic evidence so far is
intriguing, showing presence of haplogroups (e.g., N1a) found rarely in
modern Europeans but in a substantial numbers in central-European
Neolithic populations, suggesting at least for that setting a population
diffusion/migration scenario (e.g., Deguilloux, Leahy, Pemonge, & Rot-
tier, 2012; Haak et al., 2005). Similarly, Linderholm (2011) makes the
case from the aDNA record that Mesolithic foragers in Sweden were
replaced by a later group of farmers. I suspect that simple continuity/
replacement scenarios will become rare as more research develops in
the coming years. Rather, as the aDNA record expands, it will likely
reveal abundant evidence of demic diffusion, resulting in considerable
admixture and, perhaps, some replacement. The one conclusion that is
emerging is that humans (as today) moved and mated a lot, a pattern
that extends well back into the Pleistocene.
7 | REVOLUTION 4: HEALTH HISTORY
FROM ANCIENT BIOMOLECULES
The fourth revolution has been heavily influenced by the aforemen-
tioned remarkable advances in extraction and amplification of aDNA,
including aDNA that has proven useful in identifying pathogen-specific
aDNA from bones and teeth. That said, a considerable volume of the
published research is based on insufficiently authenticated aDNA,
including especially infectious diseases, such as those caused by patho-
genic mycobacterial DNA (for tuberculosis and leprosy) and treponemal

LARSEN
DNA (Wilbur & Stone, 2012). On the other hand, much has been
learned in the recent history of aDNA research, presenting investiga-
tive opportunities not even dreamed of by most of us when bioarch-
aeology was getting off the ground in the 1970s. Importantly, the
growing aDNA record being developed in laboratories in the United
States, England, Germany, and elsewhere offers a fund of information
on specific pathogens, both their evolution and their association with
human populations (Wilbur & Stone, 2012).
Successes in identifying the aDNA record of M. tuberculosis as
documented by aDNA analyses began in the mid-1990s with the publi-
cation of the successful amplification and identification of DNA frag-
ments specific to the bacterium in Peruvian mummies (Arriaza, Salo,
Aufderheide, & Holcomb, 1995; Salo, Aufderheide, Buikstra, & Hol-
comb, 1994). Since then, other evidence has been reported in archaeo-
logical remains in the Americas (e.g., Klaus et al., 2010) and across the
Atlantic in a range of contexts (e.g., Mays, Taylor, Legge, Young, &
Turner-Walker, 2001). While the evidence has derived from various tis-
sues, the record from South American settings, especially desiccated
tissues from mummies, has proven successful.
Where, when, and how did the pathogenic bacteria evolve and
transmit to humans? Or, did the pathogen first evolve in humans and
subsequently transfer to animals? Experience in study of a range of
infectious diseases today points to animals as the original source of path-
ogenic bacteria. With regard to tuberculosis, it has long been known that
M. bovis is the pathogenic mycobacterium that causes the disease in cat-
tle. Humans are highly susceptible to the pathogen, involving spread
from cattle to humans via air or consumption of meat and dairy prod-
ucts. Therefore, it seemed that that during the Neolithic, when domesti-
cation involved close proximity between cattle and humans, zoonotic
transmission from the former to the latter occurred involving an evolu-
tionary transition from M. bovis to M. tuberculosis. However, genomic-
based phylogenetic analysis of the bacilli representing a wide range of
strains of mycobacterial species reveals that M. tuberculosis clearly did
not evolve from M. bovis. Moreover, the human-associated M. tuberculo-
sis pathogen likely has considerable antiquity, predating the arrival of
humans in the Western hemisphere (Bos et al., 2014). For the New
World specifically, Bos and collaborators (2014) successfully identified
M. tuberculosis genomes in three Middle Horizon/Late Intermediate
period sites (ca., AD 750–1350) in the Osmore River drainage of Peru.
While the paleopathology record demonstrates strong evidence for the
presence of tuberculosis, their analysis confirms that it has been endemic
and present for considerable time, certainly well before European con-
tact and colonization. Importantly, the strain identified in precontact indi-
viduals is most similar to the strains documented in seals and sea lions
rather than strains seen in today’s human populations. Rather, phyloge-
netic analysis suggests that M. tuberculosis originated in sea mammals
and was subsequently transmitted to humans.
8 | LOOKING BACK, MOVING FORWARD
Bioarchaeology has been built by many players, including all of the pro-
fessionals and the students they have taught in classrooms, seminars,
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| 873
laboratories, and field settings over the last several decades. There is of
course much more to learn, but the base to build on is well in place, to
anticipate new revolutions, to embrace new developments, to take
chances, and to expect changes that will continue to inform and pro-
mote even better understanding of the human condition and the con-
texts that shape it. In addition to the other characteristics of modern
bioarchaeology, I believe that successes of the science have been due
to it being far more informed about the biology of bone than in the
past (see Crowder & Stout, 2012; Gosman, Stout, & Larsen, 2011) and
the essential social and cultural contexts associated with life conditions,
adaptation, and challenges facing humans in the last 10,000 years,
arguably some of the most dynamic years in all of the last six to seven
million years of hominin evolution. Key to the future success of the
field will be the continued developments in articulating the social, cul-
tural, and behavioral contexts for interpreting the human skeletal
record discussed in this perspective. Unlike those who studied ancient
remains in 1918, many of today’s bioarchaeologists study human
remains that are closely linked with the archaeological record. In many
research programs, they are linked with the archaeologists document-
ing and interpreting that record. The collaborative approach is not uni-
versal, especially in areas of the world where those who excavate
skeletons and those who study them do not necessarily collaborate in
an integrative fashion. Nonetheless, the developing bioarchaeology dis-
cussed in this perspective is far different than the one in the early to
middle twentieth century where much of the research was disassoci-
ated from context, was based on race models, and was descriptive
(Buikstra, 2006).
Several years ago, a student asked me at the AAPA meetings if I
was a “scientific” bioarchaeologist or a “social” bioarchaeologist. After
giving that a bit of thought, I concluded with an answer that sums my
response: We often use the scientific method for tackling problems,
but the “social” element is only one part of the whole that helps con-
textualize the study of human remains. In my view, that is the big pic-
ture approach to what we do—we place the remains of past people
into the complex contexts of their past lives. In doing so, bioarchaeolo-
gists address fundamental changes in who we are as people and how
very successful human beings can be in adapting to and mitigating cir-
cumstances that challenge their survival.
The emphasis on context in bioarchaeology has stimulated many
new investigations concerning a range of topics that are explicitly
focused on the lived experience and social process, such as childhood,
colonialism, ethnicity and ethnogenesis, identity, gender, violence, and
inequality (e.g., Agarwal & Glencross, 2011; Dong et al., 2017; Geber,
€ sel, 2006; Knudson & Stojanowski, 2009; Martin,
2015; Gowland & Knu

rez, 2012; Murphy & Klaus, 2017; Tung, 2012; Stojanow-
Harrod, & Pe
ski, 2013; Temple & Goodman, 2014; Thompson, Alfonso-Durruty, &
Crandall, 2014; Vercellotti et al., 2014; Winkler et al., 2016). This
emphasis, coupled with the growth of techniques such as stable iso-
tope analysis, cross-sectional geometry, and next generation sequenc-
ing, has created new opportunities to study the past, with potential for
understanding life and living conditions in today’s world (cf., Armelagos,
2003; Martin et al., 2013).

874 | LARSEN
There are other developments currently taking place in bioarch-
aeology that are expanding our ability to understand “life not death”
(Milner & Boldsen, 2017), especially in regard to community and
regional patterns of health and lifestyle. This approach requires a
population-oriented perspective involving collections of archaeological
skeletons derived from single cemeteries representing individuals from
a community and groups of cemeteries representing individuals from
multiple communities. Given that a collection of skeletons is not a true
population owing to the inherent biases present in archaeological skel-
etal series (Wood, Milner, Harpending, & Weiss, 1992), there are
important factors requiring assessment for any population-oriented
study. However, in settings where contexts are well documented, there
is a strong case to be made for testing hypotheses regarding outcomes
of health and lifestyle. It is in these settings where people shared simi-
lar environments and landscapes in common geographic contexts. This
approach provides more compelling explanations of variation in health
and life conditions than the traditional biomedical approach involving
comparisons of populations having highly disparate histories and living
circumstances (e.g., comparison of “Whites” vs. “Blacks”).
One important issue relating to inequality and health is differential
ability of the immune system to mitigate infectious pathogens. How-
ever, newly developing experimental research on immune responses
demonstrates that cells exposed to mycobacterial pathogens for lep-
rosy and tuberculosis elicit inflammatory responses to other pathogens,
including those that are associated with periodontal disease (Crespo,
Klaes, Switalo, & DeWitte, 2017). The implications of this work are
that these inflammatory shifts may be detectable in archaeological
remains, reveal the larger burden of exposure to pathogens in seg-
ments of populations experiencing reduced access to quality nutrition
and predisposition to infection, and provide insights into the dynamics
of immunological responses. Identification of pathogens associated
with periodontal disease provides complementary opportunities to
investigate the oral cavity environment and associated immunological
circumstances. Recent research on aDNA derived from dental calculus
shows the presence of periodontal pathogens (Warinner et al., 2014). I
can well envision the development of insights into infectious disease
and variation in immunological competence in new and innovative
ways, including in their social, cultural, behavioral, and environmental
contexts.
In summary, the study of archaeological human remains today has
little in common with their study prior to the 1970s. Prior to the
1970s, human remains were studied in a largely descriptive manner in
the context of race identification. Beginning in the 1970s, owing to a
number of influences—such as the New Physical Anthropology move-
ment, greater emphasis on the biological processes of skeletal and den-
tal tissue growth and development, the increased attention to
problem-based research addressing questions and hypotheses, viewing
skeletal remains as individuals and populations once living, and the
development of new analytical methods—has transformed the manner
and focus of study of archaeological human remains. The century of
1918–2018 has seen exciting developments, most of which are pre-
sented in the pages of the American Journal of Physical Anthropology.
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The growing community of scientists and their discoveries will no
doubt continue to transform bioarchaeology over the course of the
next century.
ACK NOWLE DGME NT S
The preparation of this perspective facilitated considerable reflection
about my involvement in and the growth of bioarchaeology. I am
grateful to William M. Bass III, who introduced me to the record of
the human remains from archaeological contexts, and my graduate
school professors, especially Milford H. Wolpoff (advisor), C. Loring
Brace, Frank B. Livingston, Stanley M. Garn, David S. Carlson,
Michael Zimmerman, Philip D. Gingerich, and A. Roberto Frisancho
for their collective knowledge, leadership in the discipline, and sup-
port. David Hurst Thomas and Douglas H. Ubelaker mentored me
and provided wonderful opportunities to learn in fieldwork and at
their lab settings at the American Museum of Natural History and
the Smithsonian Institution, respectively. The experiences working
with them were life-changing. I am indebted to George Armelagos
and Jane Buikstra for their influence on my perspectives. I acknowl-
edge my many associates, collaborators, and students in research
programs that I have directed or codirected in the American South-
east and Great Basin, Turkey, and Europe. Support for these projects
is from the National Science Foundation, National Endowment for
the Humanities, National Geographic Society, Edward John Noble
Foundation, and the St. Catherines Island Foundation. For their help-
ful comments on drafts of the manuscript, I thank Daniel H. Temple,
Fabian A. Crespo, Douglas H. Ubelaker, Barbara Betz, and the anon-
ymous reviewer.
OR CID
Clark Spencer Larsen http://orcid.org/0000-0002-6905-8417
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How to cite this article: Larsen CS. Bioarchaeology in perspec-
tive: From classifications of the dead to conditions of the living.
Am J Phys Anthropol. 2018;165:865–878. https://doi.org/10.
1002/ajpa.23322