Professional Documents
Culture Documents
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Bioarchaeology
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Bioarchaeology
Interpreting behavior from the
human skeleton
Newcastle-upon-Tyne
R. A. Foley Department of Biological Anthropology, University of
Cambridge
A/so in the series
Anthropo/ogy
The University of North Carolina, Chapel Hill
J. E. Lindsay Carter and Barbara Honeyman Heath Somatotyping Development and applications
Roy J. Shephard Body Composition in Bio/ogical Anthropology
Ashley H. Robins Biological Perspectives on Human Pigmentation
C. G. N. Mascie-Taylor and G. W. Lasker (editors) Applications of
Biological Anthropology to Human Ajfairs
Alex F. Roche Growth, Maturation, and Body Composition - The Fels
Longitudinal Study 1929-1991
Eric J. Devor (editor) Molecular Applications in Biological Anthropology
Kenneth M. Weiss The Genetic Causes of Human Disease - Principies
and evolutionary approaches
Duane Quiatt and Vernon Reynolds Primate Behaviour - Information,
social knowledge, and the evolution of culture
G. W. Lasker and C. G. N. Mascie-Taylor (editors) Research Strategies
in Biological Anthropology - Field and survey studies
S. J. Ulijaszek and C. G. N. Mascie-Taylor (editors) Anthropometry: the
individual and the population
C. G. N. Mascie-Taylor and B. Bogin (editors) Human Variability and
Plasticity
S. J. Ulijaszek Human Energetics in Bio/ogical Anthropology
CAMBRIDGE
UNIVERSITY PRESS
Contents
Acknow/edgments
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XI
lntroduction
2.1
2.2
2.3
2.4
2.5
2.6
Introduction
Introduction
Dental caries
Periodontal disease (periodontitis) and tooth 1oss
Nonspecific infection
Specific infectious diseases: treponematosis, tuberculosis,
and leprosy
3.6 Summary and conclusions
lntroduction
Accidental injury
Intentional injury and interpersonal violence
Medical care and surgical intervention
Interpreting skeletal trauma
6
8
23
29
56
61
64
64
65
77
82
93
107
109
109
110
119
152
154
159
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Introduction
Articular joints and their function
161
161
162
162
Contents
5.4
5.5
5.6
185
188
193
195
195
197
219
220
224
226
226
227
242
247
268
270
270
271
290
300
300
302
302
305
310
324
331
333
333
334
340
341
10
References
343
General index
432
Site index
459
Acknowledgments
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xii
Acknowledgrnents
Introduction
l~an?us
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I~stitution al;~;
~ecogniz~d b~
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~n ~k~:1;r:::~;s;~~::!:~' s~:t~;;~ni:T~e;~~:~es~
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C 1 1 0d
omsrnna, exas, Oklahoma Kansas
in~o~~a~~~~~o~;:P~;:!~~a~;~:i~~::~~~~~~~~~/:~;r si:~~ d~scriptiv;
98
. or
~~o:sanlds ~f squbare miles, over 300 mortuary site; have been repo~::~'~yg
c aeo& og1sts,
fro m_ on1Y 50 0 f t hese s1tes
.
(Steele
Olive ut skeletons
) H
are described
1989
and interpreted in deta~lm~n r~mams from ~nly eight sites were described
observed. 'ma
. . n t e sa~e rev1ew, Owsley and coworkers
t f 1. f ny A~encan archeolog1sts have not appreciated the full
pbeohen .ia o doshteolog1cal research as a source of information on biocultural
adapt a f ton. M any ofthese v1ews
.
. av10r an uman
.
persist as reflected
an_ ar~h~olog1st'~. statement to a reporter visiting a field schol excavaln m o orado: Human bones don't provide that much information
.
fter ali, w~ know that they are Indians."' (1989:122).
:?
~r~~~l~~::~~~:
c:~:ions
that 'burials on ?istorical sites are much m".:'r~
kl worth . Unless the ctrcumstances are very special I
.
~
~a;e~'t
exampl~p~~:~tk
Introduction
2
Introduction
.
h
excavatin is time-consuming
when encountered in situ as obiects ~-t o;e "real" archaeology' (Bush &
and which somehow does not cons 1 u e
.
h
.
owing evidence to suggest that
Zuelebil, 1991:5) ... '
osil!ve note t ere 1s gr
On a more p .
, . skeletal studies into their research dearchaeolo~is~s are 1~corpo~:t1~:se for testing hypotheses and drawing
:
.
-Ashmore et al., 1982; Roberts,
1991; Chamberlam,
Ford, 1979; _Gilbert~ ~i~lkd i~~~- ~~~g& Brown, 1979). Additionally,
1991; Sobohk, 1994a, Sttr an '
'
. is a dominant area of inquiry
within biological anthropology, oste~log71983 to 1992 about 20% of ali
malo te 1sc1pm
- the pnmary JOU
f!'
:tu
pretive role of human remains, these works .serve primarily as 'how to'
guides to bone identification and skeletal analysis and not as resources for
the investigation of broader issues in biological anthropology and sister
disciplines. The present book focusses on the relevance of skeletal remains
to the study of the human condition and human behavior generally;
namely, how skeletal and dental tissues from archaeological settings revea!
life history at both the individual and the population levels. The goal of th.is
book is to provide a synthesis of bioarchaeology, an emerging discipline
that emphasizes the human biological component of the archaeological
record. Although first applied to archaeozoology (Clark, 1972), the study
of animal remains in archaeological contexts, it has become convention to
use the term bioarchaeology in reference to the study of archaeological
human remains exclusively.
The enormous potential of bioarchaeology for understanding the past
has only recen ti y become realized. This is the case for severa! reasons. First,
most human osteological analysis has been descriptive and oriented around
case studies. Even for large assemblages of skeletons, osteological reports
tended to overlook pattern and tendency in a population perspective. This
descriptive orientation reflects the historical role of medica! practitioners
and their emphasis on diagnostic approaches to the study of ancient human
remains, especially in regard to paleopathology and disease. This is
especially true in the older paleopathology literature, in which diagnostic
case studies predominate. Second, most well documented, large collections
ofhuman remains havebeen excavated only within the last few decades. A
book like this would not ha ve been possible prior to the last decade or so.
Finally, theoretical and methodological developments underlying the
studies presented here are also quite recent.
This book takes a population perspective. Individual-based case studies
are discussed, especially because collectively they help to build a picture of
biological variability in earlier societies. The population approach is critica!
for characterizing patterns ofbehavior, lifestyle, disease, and other aspects
that forro the fabric of the human condition. The discussion in the following
pages also underscores the importance of culture in interpreting population
characteristics. Dietary behavior, for example, is highly influenced by
culture. If an individualis taught that aspecific food is'good' to eat, then the
consumption of that food tem becomes fully appropriate in that cultural
context. Other factors media te the consumption of a food or foods within a
society (e.g., environment, local plants and animals). However, cultural
behavior plays an essential role in determining diets of a group of peo ple.
Unlike many ofthe aforementioned guides to osteological analysis, this
book is not methodologically driven, although methodological develop-
lntroduction
4
Introduction
c_omparingdatasetsfromskeletal~~s a~~
at
f
nea events' (1976 454
.our o the vast holdings in this s
. ). Th1s bookprovides
gamed about earlier peoples based tor~house, d1splaying the knowledge
on t e study of their mortal remains.
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2.1
Introduction
Physiological disruption resulting from impoverished environmental circumstances - 'stress' - is central to the study of health and well-being and
the reconstruction of adaptation and behavior in earlier and contemporary
human societies (Goodman et al., 1988; Huss-Ashmore et al., 1982). Stress
is a product of three key factors, including (!) environmental constraints;
(2) cultural systems; and (3) host resistance. Goodman and coworkers
(Goodman, 1991; Goodman & Armelagos, 1989; Goodman et al., 1984,
1988) have modeled the interaction ofthese factors at both the individual
and the population levels (Figure 2.1 ). This model emphasizes the environment in providing both the resources necessary for survival and the
stressors that may affect the health of the population. Cultural systems
serve as protective buffers, and they provide behaviors necessary for
extraction of importan! nutrients and resources from the environment. Ali
stressors can never be fully buffered; sorne slip through the fi!ter of the
cultural system. In these instances, the individual may exhibit a biological
stress response observable at the tissue leve! (bones and teeth). Physiological disruption feeds directly back into environmental constraints and
cultural systems. This model makes clear that health is a key variable in the
adaptive process.
Stress has significant functional consequences. Elevated stress can lead
to a state of functional impairment, resulting in diminished cognitive
development and work capacity. The reduction in work capacity can be
detrimental if it impedes the acquisition of essential resources (e.g., dietary)
for the maintenance of the individual and the population. If individuals of
reproductive age are affected by poor health, then decreased fertility may
be the outcome. Ultimately, the success or failure of a population to
mitigate stress has far-reaching implications for behavior and the functioning of the society (see also Martin et al., 1991).
Biological anthropologists employ a variety of skeletal and dental stress
indicators which can be measured empirically. Use of multiple indicators
gives a comprehensive understanding of stress and adaptation in the past
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2.2.1
Growth rates
Although generally continuous, growth from birth through a~olescence is
punctuated by two intensive periods of activity. The. first penod shows a
great increase in growth velocity during infancy, fallmg o~ soon after ~he
first year of life. The second involves another marked mcreas~ dunng
adolescence, then declines and. reduces to zero growth wh~n ep1physeal
fusion of the long bones (femur, tibia, fibula, humerus, radms, and uln~)
and other skeletal elements is complete in early adulthood. Growth r.ate is
widely recognized as a highly sensitive indicator of health and well-bemg of
a community or population (Crooks, 1995; Eveleth & Tanner, 1990,
Gracey, 1987; Gray & Wolfe, 1996; Huss-Ashmore ~ !ohnston, 1985).
Growth is affected by various factors, such as genet1c mfluences, growth
hormone deficiencies, and psychological stress (Eveleth & Tanner, 1990;
olfe 1996) but the preponderance of evidence underscores the
G ray & W
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influence of environment - especially nutrition - on the growm~ c 1 .
Infectious disease, such as episodic diarrheal disease, can also con~r.1bute to
poor growth (e.g., Jenkins, 1982; Martorell et al., 1977). Nutn~1on ~nd
disease have a synergistic relationship wh~reby poorl~ nour.ished iuvemles
are more susceptible to_ infection, and disease and mfect10n r~duce the
ability of.the body to absorb essential nutrients (Keusch & Farthmg, 1986;
10
11
similarity between the two groups from birth to six months and from four
years to 10 years. F or juveniles aged six months to four years, Libben tibiae
are shorter than Bt-5 tibiae. The growth period between six months and
four years- the period differing most between Btc5 and Libben populations
- is highly sensitive to metabolic disruption. During this period, the infant
undergoes weaning, involving the shift from a relatively stable, nutritious
food source (mother's milk) to a potentially less stable, less digestible, and
less nutritious food (e.g., maize). Passive immunities derived from consumption of breast milk are lost during weaning during this period of life
(Popkin et al., 1986). These immunities are crucial for early health and
well-being sin ce the child's immune system is not fully developed until after
five years of age (Newman, 1995). Mensforth (1985) found a high
prevalence of nonspecific periosteal infections in the Libben infants,
suggesting that high levels of infectious disease in infancy and young
childhood contributed to growth retardation. Although both groups
apparently enjoyed adequate nutrition, Mensforth argues that the Libben
population had a subsistence economy with a relatively greater diversity of
resources that were immediately available withoutneed of extensive travel
for acquisition ofresources. Thus, in comparisonwith the Bt-5 population,
the Libben population experienced greater sedentism and size which
fostered poor sanitation, elevated infectious disese, and poor health.
Comparison of Libben with a modero reference population (Denver,
Colorado) confirms the presence of growth suppression in the first three
years oflife in the former, after which the growth rates are similar between
the two groups (Lovejoy et al., 1990). Lovejoy and coworkers ( 1990) argue
that massive infection was the cause of growth retardation. They suggest
that inflammation would result in an increased production of cortisol, the
major natural glucocorticoid, which results in limitation of growth and
availability of amino acids. Thus, elevation of infection in the Libben
population may have hada strong influence on growth generally (Lovejoy
et al., 1990).
Historie-era skeletal series furoish important insights into stress in the
recent past. Saunders and coworkers (1993, 1995) analyzed growth data
available from a large series of juvenile remains from the St. Thomas'
Anglican Church cemetery in Belleville, Ontario. The cemetery was used by
a predominantly British-descent population during the period of 1821 to
1874. Comparisons of femur length from juveniles buried in the cemetery
with a tenth century Anglo-Saxon series from Raunds, England, and
modero growth data from Denver, Colorado (Maresh, 1970), indicate a
strong similarity in overall pattero of growth between the three groups
(Figure 2.2). The two cemetery samples are temporally separate, but share
12
400
350
E
g
300
.e
&_250
gi 200
~ 150
100
5o L.~~~~~ry~'4~5~-5r'7
-~7~,8~~9~1co ~1~1~12~13
2 3
Age (years)
Figure 2.2. Fitted curves for femoral diaphyseal length for. the
nineteenth-century St. Thotnas' Church cemetery (dotte~ hne), tenth-century
R.aunds Anglo-Saxon-sketetons (dashed line), and twentleth-century Den~er,
Colorado, livingpopulatiOn (solid line). (From Sau~ders & Hoppa, 1993,
reproduced with pennission of authors and John W1ley & Sons, lnc.)
general ethnic origins with the modern U.S. population. Figure 2.2 sh~ws
that the St. Thomas' series is slightly shorter for age than the modern senes.
That the Raunds .series is considerably shorter than either of the other
groups is to be expected given the inferior living standards of tenth century
England. With regard to the St. Thomas' skeletons, Saunders an.d
coworkers suggest that juveniles died from ~cute causes. ~nd not chromc
conditions (e.g., chronic infections or chromc undernutntlon) that would
result in a.decrease in skeletal growth. Children less than two years of age
had slightly lower growth rates than modern twentieth centur_Y popu~a
tions. They regard this as perhaps representing stresses assoctated w1th
poor maternal health and prenatal growth. .
.
Analysis of juvenile cortical bone growth v1a measurement of cortical
thickness provides a complementary source of information to the length of
.long bones. In living populations, defici~~cies in cor'.ical bone mass ar~
present in groups experiencing undernutnt10n (e.g., Fnsancho et al., 1970,
Garn 1970; Garn et al., 1964; Himes, 1978; Himes et al., 1975). Garn and
cowo~kers (1964), for example, showed that malnourished Guate~alan
children have reduced cortical bone in comparison with well no~nshed
reference groups. Although bone lengths increased dun~g p~nods of
growth.recovery, cortical thickness continued to show defic1enc1es dueto
13
2.2.2
Stature
Substantial evidence drawn from the study ofliving populations reveals the
strong relationship between growth suppression in childhood and attainment of adult body size, including terminal height: Growth~retarded
children should be short-statured adults. Study of living populations
provides sorne support for this conclusion. Comparison of growth of
undernourished Thai children with American (U.S.) children reveals that
despite a longer period of growth in the former (by about one year) the
reduction in growth over their lifetimes resulted in shortened terminal
height (Bailey et al., 1984; see also Bogin & MacVean, 1983; Frisancho et
al., 1970; Satyanarayana et al., 1980).
The close ties between stress- especiaHy poor nutrition- and stature are
abundantly documented in research developingout ofa growing interest in
anthropometric history (Floud et al., 1990; Komlos, 1989, 1994, 1995;
Steckel, 1995). Originally inspired by controversy over the health and
14
15
Stature (cm)
Males
Crossn
Fort William Henry"
Harvie''
Prospect HiJJd
Colonial U.S.t
Mt. Gileadf
Clifts Plantationg
Belleviewg
Ft. Laurensh
Snake Hill;
Bradford's Company;
Old Quebedl
West Point cadetsk
Modero U.S.1
Fema/es
Cross
Harvier
Prospect Hilld
Colonial U .S."
Mt. Gileadf
Modero U.S.'
175
177
171
173
173
172
169
170
174
176
174
173
172
174
163
161
161
160
162
161
16
17
McCaa & Mrquez Morfin, 1995; Wright & White, 1996). Comparisons of
agricultura! populations with other settings indicate relatively short
statures in Mesoamerica (Storey, l992a) and Ecuador (Ubelaker, 1994)
which are linked with chronic malnutrition.
Other archaeological settings show reduction in stature in the shift to
agricultura! economies. Preliminary evidence indicates that late Pleistocene
foragers in South Asia were taller and more robust than their farming
descendants (Kennedy, 1984). Similarly, comparisons of skeletal series
from the Upper Paleolithic through the Neolithic in western Europe
indicate a general reduction in average stature, which is especially pronounced in the comparison of Mesolithic with Neolithic subsamples
(Meiklejohn et al., 1984; although see Jacobs, 1993). Finally, in Sudanese
Nubia, reduction in stature coincided with agricultura! intensification,
especially in comparison of the earlier A-group (3400-2400 se) and Jater
X-group (AD 350-550) populations (Van Gerven et al., 1995). These studies
point to the possibility of increasing dietary stress as a causal factor in
stature reduction.
Much of the research on body size in children and adults in archaeological settings is oriented toward tracking the consequences of adaptive
transformations, primarily from foraging to farming; relatively little is
known about other dietary transitions. The consequences of change in
dietary focus not involving agriculture are manifested in temporal comparisons of native populations from the Santa Barbara Channel region of
southern California (Lamber!, 1993, 1994). In this region, populations
shifted their dietary emphasis from terrestrial resources - especially plant
foods - to marine resources after 500 BC (Glassow, 1996). Over the period
of 6000 BC to AD 1782, stature decreased by about 10 cm. Lamber! (1993,
1994) argues that stature reduction was fostered by decline in health, dueto .
the combined effects of declining nutrition and elevated infectious disease:
Protein, mostly derived from fish, was abundan!, but other importan!
nutrients may have been lacking in the diets of later prehistoric populations. During the latest prehistoric period, for example, island populations
traded beads and other manufactured products with mainland populations
for plant foods, especially roots and seeds, suggesting that islanders lacked
immediate access to key plant resources. In addition, the later populations
were more sedentary, and they consumed a narrower range of foods than
the earlier populations. Environmental evidence indicates periodic and
lengthy periods of drought, which would ha ve reduced the availability of
potable water and plan\ foods (e.g., acorns and other seeds). The worsening
of nutritional quality was probably compounded by other stressors,
particularly infectious disease. In addition to increases in other. stress
18
The influence of nutritional deprivation on human growth and terminal height is revealed in the study of components of past ~roups t~at
may have been.differentially buffered against str.ess. Compans~n of ehte
and.nonelite adults from Middle Bronze Age s1tes (2000 ne) m Greece
shows that elites are about 6cm taller than nonelites (Angel, 1975, 1984).
Similarly, the tallest adults in the Etruscan period in Tarquinia, Italy'. are
associated with high-status chamber tombs (Becker, 1993). In a Ma1tasChiribaya (ca. 2000 BP) population from northern Chile, shaman males
are taller than other, nonelite males, which may indicate better health
and resources in the former (Allison, 1984). High-status adult males in
sorne Mesoamerican populations. appear to be taller than low-status
19
individuals or the general population (Haviland, 1967; Helmuth & Pendergast, 1986-1987; but see Wilkinson & Norelli, 1981). Likewise, elite
males are taller than nonelite males in severa! contexts in the prehistoric
southeastern and midwestern United States (Buikstra, 1976a; Cook,
1984; Hatch, 1976; Hatch & Willey, 1974; Powell, 1988). These apparent
status differences in attained height suggest that elite males may have had
nutritional advantages resulting in greater height than nonelite individuals. There are no clear differences in stature between elite and nonelite
adult females in any of these New World settings. This suggests that the
burden of stress may be on adult males in ranked societies, at least as it is
exhibited in attained height.
2.2.3
20
causal factor in.determining cranial base form is incorrect. The phenomenon of cranial base flattening is largely unexplained.
2.2.4
'
Pelvic morphology
Severe vitamin D deficiency (rickets) caused by inadequate protein consumption weakens growing bone during early childhood, because the
rapidly forming protein matrix <loes not mineralize sufliciently. This results
in pelvic deformation, dueto the forces created by body weight and gravity
(Angel, 1975, 1978a, 1982, 1984;Angel & Olney, 1981; Greulich & Thoms,
1938; Nicholson, 1945; Thoms, 1947, 1956; Thoms et al., 1939; Walker et
al., 1976). Pelvic inlet deformation is. characterized by a reduction in
anterior-posterior diameter relative to the medial-lateral diameter (called
'platypellism'). Flattening of the pelvis is well documented in clinical
populations (e.g., Greulich & Thoms, 1938; Nicholson, 1945; Thoms, 1947)
and in modern anatomical samples in comparison of lowei; and middle
class groups from the United States (Angel, 1982). For example, British
women who were young children during the waryears of 1914 to 1918 have
flattened pelvic inlets (Nicholson, 1945). Presumably, these women had
relatively poor nutrition during these years. Consisten! with the relationship between growth and nutritional status, women with flattened pelves
tend also to be short-statured.
Comparisons of pelvic inlet form between earlier and later (or modern
reference) populations suggest improvements in nutritional health in
several settings, including the eastern Mediterranean (Angel, 1984; Angel
& Olney, 1981), North America (Angel, 1976), and Sudanese Nubia(Sibley
et al., 1992). Preliminary evidence shows differences in pelvic shape by
status group.Low-status adult females from the Middle Woodland (Klunk
and Gibson Mound groups)"period in the lower Illinois River valley have
flatter pelvic inlets than high-status adult females (Brinker, 1985). These
differences appear to reflect better nutrition in the high-status women than
in Jow-status women.
Other aspects of pelvic morphology may also be linked to negative
environmental factors. Sciatic notch widths (innominate bone) are appreciably larger in nutritionally stressed eighteenth and nineteenth century
British from the St .. Bride's Church, London, than in better fed twentieth
century Americans (Walker, unpublished manuscript). Rickets. was a
severe health problem in industrial England, and is well documented in the
St. Bride's Church population. Archival documents indicate that the births
of St. Bride's individuals with wide sciatic notches occurred during cold
months of the. year, the period when. rickets. was especially prevalen!..
21
2.2.5
22
nutritional stress (e.g., Adams, 1969; Angel, 1984; Buxton, 1938). Buxton
(1938) asserted that less bone is required in the construction ofa diaphysis
ifit is flattened ratherthan round. He viewed the temporal trend ofrounder
diaphyses as representing an increase in amount ofbone, inferring a decline
in nutritional deficiency in recen! 'civilized' populations. Structural analysis of long bone diaphyses reveals that flattening is related not to the
amount ofbone present, but rather to the manner in which it is distributed
when viewed in cross section. Mechanical loading, not nutritional stress, is
the primary determinantofflatness oflong bone diaphyses (see Chapter 6).
Nutritional deprivation or other physiological stressors certainly have an
influence on amount of bone, but the relationship between nutritional
status and diaphyseal shape is unsubstantiated.
2.2.'6
The effects of catch-up growth on stature and long bone. lengths are
problematic for documenting stress history of an individual during their
growth years. An individual may be stressed early in life, but amelioration
of negative conditions (e.g., improvement in nutritional status) during later
juvenile years may result in obliteration of evidence of growth disruptions
that had occurred earlier in life. In the Dickson Mounds senes, for
example, although juvenile growth became stunted in the transitio.n to
intensive farming for the period of AD 950 to 1300, no appreciable
reductions occurred in adult height (Lallo, 1973). Thus, adult heights in
this population are uninformative about juvenile stress.
The. similarity of stature in Dickson Mounds may be simply due to
growth recovery. Vertebral growth provides a means of addressing the
problem of growth stress identification not possible with attained height.
At the time of birth, vertebral neural canal size is approximately 65%
complete; ful! size is reached by about four years of age (Clark, 1988; Clark
et al., 1986). Vertebral body height continues to grow into early adulthood,
well after the third decade of life. Thus, early and late stress in the life
history ofthe individual is represented in the respective size ofthe vertebral
neural c<tnal and vertebral body height in adult skeletons. If there is a
reduction fo canal size but not in vertebral height, then catch-up growth
probably occurred following early stress (prior to four years of age). If?oth
neural canal size and vertebral body height are small, then stress was hkely
to be present throughout most of the years of growth and development,
certainly after four years of age and possibly into adulthood (Clark, 1988;
Clark et al., 1986).
Analysis of thoracic an.d lumbar vertebrne from the Dickson Mounds
23
si te reveals that growth of neural canal size was completed prematurely, but
growth in vertebral body height continued throngh the juvenile years into
adulthood (Clark et al., 1986). This growth pattern suggests that stress
amelioration in the later juvenile years accounts for the similarity in adult
long bone lengths and stature in the earlier and later populations from
Dickson Mounds.
Young adults (15--25-year age group) in the Dickson Mounds series ha ve
significantly smaller vertebral neural canal size than older adults (25 +
years) (Clark eta/., 1986). This finding suggests that small neural canal size
is linked with a reduced lifespan. Additionally, vertebral wedging, the
condition whereby anterior body height is reduced compared to posterior
body height, is associated with small vertebral neural canal size. Vertebral
wedging is symptomatic of adult and postmenopausal bone loss (osteoporosis). The association between smallerueural canal size and wedging
suggests that stress occurring during an individual's juvenile years may be a
predisposing factor for poor health during adulthood.
2.3
2.3.J
24
eruption timing shows sorne correlation with body size (Garn et al., 1960;
McGregor et al., 1968).
It is not possible to identify delays in dental eruption timing in
archaeological seties based on teeth alone, since age-at-death must be
determined by comparing the archaeological dentitions with sorne standard based on individuals of known age (e.g., Moorrees et al., 1963).
Relative differences between dental and skeletal development may provide
sorne insight in to growth stress. Comparison of skeletal age and dental age
in Medieval period skeletons from Sudanese Nubia reveals that most
individuals (70.5%) have skeletal ages younger than their dental ages
(Moore et al., 1986). These relative differences indicate that skeletal growth
may have been retarded. Dietary reco.nstruction suggests that growth
retardation was due to nutrition.al deprivation, a finding that is consisten!
with other skeletal indicators ofstress (e.g., iron deficiency anemia; Moore
et al., 1986).
2.3.2
Tooth size
Like bone size, tooth size involves a complex interplay between environment and heredity. Unlike skeletal elements, tooth crowns do not remodel
once they are fully formed. Therefore, teeth provide an unchanging record
of size well in advance of the adult years. Tooth size appears to be highly
heritable, indicating that variation between and within human populations
can be explained mostly by genetic differences (Chapter 7; and see Kieser,
1990). Twin studies revea! that as much as 80% to 90% of observed
covariation in tooth size is due to additive genetic factors; the remaining
10-20% is attributed to environment (Townsend et al., 1994). Other
estimates of heritability vary widely (see Kieser, 1990), but most workers
agree that environmental influences on tooth size are significan!, albeit
small (e.g., Dempsey et al., 1995; Garn et al., 1965; Garn, Osborne et al.,
1979; Potter et al., 1983; Townsend, 1980, 1992; Townsend & Brown,
1978). Therefore, tooth size represents a measure of deviation from genetic
growth potential in response to sorne stressor or stressors (Bailit et al.,
1968, 1970; Evans, 1944; Garn, Osborne et al., 1979; Garn et al., 1980;
Goose, 1967). Placenta( insufficiency, maternal health status, nutritional
status, anda variety of genetic and congenital defects (Down's syndrome,
cleft palate, prenatal rubella, congenital syphilis) are linked with reduced
tooth size (Cohen et al., 1979; Garn & Burdi, 1971; Garn, Osborne et al.,
1979; Goodman et al., 1989). Understanding the influence of nutrition on
tooth size is hampered by the paucity of .data holding genetic factors
costant. in situations of variable nu.tritional quality. Goodman and
25
coworkers (1989) tested the hypothesis that poor nutrition will result in
small permanent tooth size by comparison of dietary supplemented and
nonsupplemented individuals in a Nahuatlcommimity from Tezonteopan,
Mexico. Overall, supplemented individuals have larger tooth size than
nonsupplemented individuals, with statistically greater differences occurring for the mate first incisor, second incisor, and first molar buccolingual
dimensions. These findings are consisten! with experimental research on
laboratory animals showing tooth size reduction ih response to developmental disruptions and nutritional deprivations (e.g., Bennett et al., 1981;
Holloway et al., 1961; Paynter & Grainger, 1956; Riesenfeld, 1970;
although see Murchison et al., 1988).
Prehistoric maize agriculturalists from coastal Georgia post-dating AD
1150 had smaller teeth than did their foraging predecessors (Larsen, 1982,
1983a). Tooth size was reduced in both the permanent and deciduous
dentitions, which may reflect increase in physiological stress dueto declines
in dietary quality and health status generally. Tooth size reduction in the
primary dentition suggests a negative change in maternal health status and
placenta! environment, since deciduous teeth form in utero. Given the
relatively narrow temporal window of tooth size reduction in this and other
populations with the shift from foraging to farming (e.g., Coppa et al.,
1995; Hinton et al., 1980; Meiklejohn & Zvelebil, 1991; y'Edynak, 1989),
these changes probably indicate an increase in stress that accompanied this
transition. In contras!, Lunt ( 1969) documented a temporal increase in
permanent tooth size from Medieval times to the present in Denmark,
attributed to improved dietary conditions in later times (and see Lavelle,
1968).
Dental size decrease or increase in Holocene populations cannot be
explained fully by nonevolutionary factors. In prehistoric Nubian populations, there is a relatively greater reduction in posterior tooth size than
anterior tooth size, which Calcagno (1989) attributes to a selective
advantage for smaller posterior teeth in caries-prone agriculturalists. These
findings underscore the complexity of tooth size, requiring consideration of
both extrinsic and intrinsic circumstances in specific settings.
The hypothesis that members of a population who suffer most from
illness and physiological stress are more likely to die at an earlier age than
other (healthier) members of a population has been tested by the comparison of permanent tooth size of juveniles and adults in different settings in
the American Southeast, namely in the late prehistoric Averbuch series
from the Middle Tennessee River valley (Guagliardo, !982a) and the
Spanish mission Santa Catalina de Guate from St. Catherines lsland,
Georgia (Simpson et al.; 1990). Both populations were sedentary mai'e
26
Juvenile
Tooth
SD
Mean
SD
16
23
28
34
25
38
21
7.66
6.94
8.59
I0.12
9.77
11.93
12.09
0.56
0.39
0.66
0.59
0.56
0.68
0.67
33
37
55
70
72
77
85
7.48
6.91
8.64
I0.09
9.89
12.14
12.01
0.40
0.36
0.47
0.49
0.64
0.51
0.68
-2.4
-0.4
0.6
-0.3
1.2
1.7
-0.7
20
27
32
37
33
45
31
:S.84
6.23
7.51
8.09
8.42
11.11
10.76
0.38
0.40
0.57
0.46
0.52
0.49
0.57
22
47
77
95
95
72
87
5.89
6.34
7.85
8.30
8.63
11.24
I0.76
0.33
0.38
0.53
0.44
0.47
0.52
0.61
0.8
1.7
Maxillary
11
12
PMI
PM2
MI
M2
Mandibular
ll
12
PMI
PM2
MI
M2
/o Difference"
Mean
4.3'"
2.Sb
2.4b
l.2
o.o
27
28
29
past groups should provide importan! insight into stress and developmental instability in earlier societies.
2.4
2.4.J
30
endemic. Schistosomiasis ('snail fever') triggers an immunological response after the eggs of blood-vessel inhabiting worms (genus Schistosoma) become lodged in body organs (e.g., liver, intestinal wall,
urogenital tract)., The disease has a tropical worldwide distribution
(Farley, 1993). Hookworm disease results from the ingestion or inhalation
of infective larvae of the hookworm (Ancy/ostoma duodena/e, Necator
americanus). The worm extracts blood by grasping the host's intestinal
wall with its sharp teeth (Despommier et al., 1995; Hotez & Pritchard,
1995). The consequences can be especially severe owing to losses of large
amounts of blood when severa! hundred or more worms are simultaneously feeding on the same host. Hookworms are also geographically widespread, mostly in tropical settings, and their presence is linked with iron
deficiency anemia in a range of settings (e.g., Layrisse & Roche, 1964; and
see below).
Various genetic diseases also cause iron deficiency, including thalassemia, sickle cell anemia, nonspherocytic hemolytic anemia (e.g., glucose6-phosphate dehydrogenase deficiency [favism], pyruvate kinase deficiency), spherocytosis, and rarely, hereditary elliptocytosis.
Skeletal changes associated with chronic anemia that are identified
radiographically include perpendicular orientation of trabeculae in the
cranial diploe (called 'hair-on-end'), expansion (hyperostosis) of the
diploe, thinning of compact cranial bone, and orbital roof thickening
(Stuart-Macadam, 1987). Postcranial changes have also been observed,
such as in metaphyses of long bones, but they are generally less severe and
reduced in prevalence in acquired anemias than in genetic anemias
(Stuart-Macadam, l 989a). Skeletal changes result from the hypertrophy of
the blood-forming tissues (marrow) in order to increase the production of
red blood cells in response to the anemia. The increase in marrow
production results in a replacement of the o uter table of compact bon e with
exposed diploic bone, which gives the appearance of raised and hypervascularized areas of skeletal tissue.
The skeletal changes associated with iron deficiency anemia are part of a
generalized syndrome called porotic hyperostosis, a term introduced by
Angel (1966a; see also Hill & Armelagos, 1990) that he u sed to describe
pathology involving the outer table of cranial vault bones (Figure 2.3).
Similar lesions found in the roof areas of the eye orbits, called cribra
orbitalia, are also frequently observed in archaeological remains (Figure
2.4). Sorne argue that cribra orbitalia is one of the earliest manifestations of
anemia, with changes on the ftat bones of the cranial vault appearing
subsequently (Carlson et al., 1974; Lallo et al., 1977; Walker, 1985). There
is wide variation in frequency of orbital versus nonorbital lesions across
31
32
Figure 2.4. Cribra orbitalia; Santa Catalina de Guale de Santa Maria, Amelia
increase in red blood cell production and marrow expansion does not
involve the use of ali available marrow space (Stuart-Macadam, 1985). The
restriction of active porotic hyperostosis to youngjuveniles indica tes that
the effects of anemia on the adult componen! of past populations cannot be
33
34
35
The most abundan! data on porotic hyperostosis are available from the
New World, especially North America. In the American Southwest,
porotic hyperostosis is highly prevalen! (e.g., Akins, 1986; El-Najjar et al.,
1975, 1976, 1982; El-Najjar& Robertson, 1976; Hooton, 1930; Kent, 1986;
Lagia, 1993; Martin et al., 1991, 1995; Palkovich, 1980, 1987; Stodder,
1994; Stodder & Martin, 1992; Walker, 1985; Zaino, 1967, 1968). Among
mostly late prehistoric Puebloan samples studied by El-Najjar and collaborators (e.g., El-Najjar et al., 1976) from Canyon de Chelly, Chaco
Canyon, Inscription House, Navajo Reservoir, and Gran Quivira, porotic
hyperostosis was found in 34.3% ofindividuals. At Chaco Canyon alone,
sorne 71.8% of individuals display the characteristic lesions. Similarly, high
prevalences have been reported from late prehistoric and contact-period
siles, including San Cristobal (90%), Hawikku (84%), Black Mesa (88%),
Mesa Verde (70%), Dolores (82%), Cases Grandes (46%), and La Plata
Valley (40%) (Martin et al., 1995; Stodder, 1994; Weaver, 1985). There are
sorne southwestern samples that have relatively low prevalences (e.g., 16%
for Navajo Reservoir children; see Martin et al., 1991). Martin and
coworkers (1991) note that comparisons of data collected by different
researchers is problematical, because of the varying methods used in
identification and recording of porotic lesions. For example, sorne researchers may include slight pitting when analyzing their data sets, whereas
others may not. Unfortunately, this distinction is only rarely noted in
bioarchaeological reports, regardless of geographic or cultural setting.
El-Najjar ( 1976) links the elevated levels of porotic hyperostosis in the
American Southwest and other regions ofthe New World to the effects of
over-reliance on maize in conjunction with food processing techniques that
may contribute to iron deficiency. Specifically, he regards the presence of
phytate - an iron inhibitor - as well as lime treatment as decreasing the
nutritional value of maize.
Analysis of archaeological samples from other maize agriculturalists in
the New World provides mixed support for El-Najjar's dietary hypothesis.
Relatively high prevalences of porotic hyperostosis ( > 15-20%) are
present in agriculturalists in the American Midwest (e.g., Cook, 1984;
Garner, 1991; Goodman et al., 1984; Lallo et al., 1977; Milner, 1983, 1991;
Milner & Smith, 1990; Perzigian et al., 1984; Rose et al., 1984), Southeast
(e.g., Boyd, 1986; Eisenberg, 1986a, 199la, 1991 b; Hancock, 1986; Parham
& Scott, 1980), and Northeast (e.g., Magennis, 1986; Pfeiffer & Fairgrieve,
1994), as well as a range of other settings in Mesoamerica and South
America (e.g., Cohen et al., 1994; Hodges, 1989; Hooton, 1940; Hrdlicka,
1914; Saul, 1972; Trinkaus, 1977; Ubelaker, 1984, 1992a; White et al.;
1994). For sorne regions where skeletal remains of foragers (or less
36
intensive agriculturalists). have been compared with those of agriculturalists, there are clear temporal increases in porotic hyperostosis prevalence
(e.g., Cook, 1984; Lallo et al., 1977; Perzigian et al., 1984; Rose et al., 1984;
although see Hodges, 1989).
Skeletal series from large, late prehistoric Mississippian centers in the
American Southeast (e.g., Blakely, 1980; Larsen, Ruff et al., 1992; Powell,
1988, 1989), contact era part-time maize agriculturalists in the Great Plains
(Miller, 1995), a large urban center in Mesoamerica (Storey, 1992a), and
the coastal desert of Peru and Chile (Allison, 1984) ali display low
prevalences. These findings are not consisten! with the dietary hypothesis,
suggesting that other factors underlie the etiology of poro tic hyperostosis.
The dietary hypothesis <loes not account for the relatively high frequencies of porotic hyperostosis in sorne foraging populations. A number of
Pacific coastal foraging groups with access to iron-rich marine resources
have high prevalences of porotic hyperostosis. Moderate levels of porotic
hyperostosis are present in precontact and contact era Northwest coas!
populations (13-14%; Cybulski, 1977, 1992, 1994). In this setting, European-introduced diseases may have prevented adequate iron metabolism
during the contact period (Cybulski, 1994). The presence of porotic
hyperostosis prior to contact indicates that there may have been other
importan! factors, such as blood loss and parasitism (see Cybulski, 1994).
Late prehistoric foragers from the islands and mainland of the Santa
Barbara Channel Island region ofCalifornia have higher prevalences than
earlier foragers, increasing from 12.8% in the Early Early period to 32. l %
in the Late Middle period (Lamber!, 1994; Lamber! & Walker, 1991;
Walker, 1986a). Late period populations living on islands located furthest
from the mainland coast havean extraordinarily high prevalence of poro tic
hyperostosis (73.1% on San Miguel Island). Walker and Lambert suggest
that water contamination explains the elevated prevalence of the condition.
High prevalence of iron deficiency anemia in island populations coincides
with a period of increasing sedentism and population size, anda shift from
terrestrialto marine diets. In the Late period, groups became concentrated
around a limited number of water sources. As a result, diarrhea-causing
enteric bacteria may have contaminated these water sources. Ethnographic
evidence indica tes that island populations preferred eating raw (vs. cooked)
fish (see Walker, 1986a), thus also increasing their chances of acquiring
parasitic infections.
Prevalences of porotic hyperostosis in prehistoric Australian foragers
are consistently high in tropicaJ/subtropical environments and low in
desert environments (Webb, 1995). For example, in southeastern Australia, prevalences range from 62.5% ( < 21 years) in the Rufus Valley to
1,
37
30.0% ( < 21 years) in the desert. Half of the juvenile crania from the
tropics of northeastern Australia are porotic. Various factors appear to
have contributed to iron deficiency anemia in Australia, but parasitism is
primary. The Murray Valley, southeastern coast, and tropics provide well
suited environments for support of various intestinal parasitic organisms,
including Trichuris trichuris, Ascaris /umbricoides, Strongyloides stercoralis, and Enterobius vermicularis. In the tropics, hookworm infection
may have been a principal cause. In living populations occupying the
tropics of Australia, sorne 40% of children are infected with this helminth.
Although it is unknown whether hookworm parasites were present in this
region prior to contact by Europeans (in 1788), had they been, they would
have caused the same types ofhealth problems seen in living groups toda y.
The cumulative evidence showing a patchwork distribution of porotic
hyperostosis independent of diet in past populations' makes El-Najjar's
dietary hypothesis unlikely. Samples studied by El-Najjar and others in the
American Southwest tend to be from maize-dependent populations inhabiting canyon bottomlands. Perhaps the greater prevalence of porotic
hyperostosis in the canyon sites was due to problems arising from poor
drainage and contaminated water and generally more restricted diets
rather than to maize consumption (Walker, 1986a).
El-Najjar and coworkers underplay the role of parasitism in iron
deficiency anemia in the prehistoric American Southwest. They assert that
parasitic infections (e.g., from hookworm) were 'extremely rare in Southwestern American Indians' (1975:921). However, recen! analysis of coprolites from archaeological sites in the American Southwest indicates the
presence of disease-causing parasites (e.g., Enterobius vermicularis,
Moniliformis c/arki, Strongy/oides spp.; Cummings, 1994; Reinhard, 1992).
Comparison of foragers and farmers indicates a dramatic increase in
helminth parasitism in the latter, especially in E. vermicularis, the organism
that causes pinworm infection. This finding is consisten! with a decrease in
sanitation and increase in population crowding (Reinhard, 1992). Additionally, the dark and crowded living conditions in prehistoric Southwestern Pueblos of the Anasazi would have exacerbated these conditions;
promoting infection and anemic responses (and see Kent, 1986).
A limited number of settings in South America show high prevalence of
porotic hyperostosis (Peru and Ecuador: Hrdlicka, 1914; Ubelaker, 1981,
1992a). Porotic hyperostosis is low in prevalence in mountainous regions,
and appears to be restricted primarily to late prehistoric coastal occupations. The penchant for coastal settings may reflecta more restricted access
by native populations to fresh, parasite-free water sources in these areas.
Ubelaker (1992a) contends that the coastal pattern of elevated porotic
38
39
40
41
2.4.2
Figure 2.5. Harris lines on juvenile tibia (left) and femur (right); anatomical
specimens. The dashed lines indicate thc contours of the growth disr.uption
(From Garn et al., 1968; reproduccd with permission of authors and
Eastman-Kodak Company.)
42
43
maize.
The use of transverse lines for documenting stress in past populations is
clouded by the fact that lines have a tendency to fade or vanish with
advancing age, dueto bone remodeling. In a study of living populations
composed of individuals ofknown stress history, lines showed a decrease in
width with advancing age; sorne lines disappeared, whereas others were
inexplicablyretained well into adulthood (Garn & Schwager, 1967; Garn et
al., 1968).
Study of formation rates, persisten ce, and loss of lines in tibiae from the
Medieval period Kulubnarti si te in Sudanese Nubia revealsa clear history of
growth disruption, but in the context of complexitiesaddressed by Garn and
coworkers on line disappearance and persistence (Hummert & Van Gerven,
1985). Juveniles in this setting display high prevalence ofHarris lines; adults
have few lines. The reduction in frequency of lines in older individuals
suggests that lines denoting previous stress events had disappeared, dueto
bone remodeling. Therefore, although lines present in adults certainly refiect
episodes of metabolic stress, their absence represents either the lack of stress
or simply the resorption oflines. Thus, transverse lines are relatively more
representative of stress history in juveniles than in adults.
Another difficulty of using Harris lines for assessing stress is the high
degree of frequency variation in relation to individual health history. Study
of living individuals of known stress history reveals the presence of
numerous lines in clinically normal children with uneventful health
histories (Garn et al., 1968), and few lines in children who are well below
weight-for-age (Walimbe & Gambhir, 1994). These findings and the lack of
close association between transverse lines and disease episodes in archaeological populations (e.g., Mensforth, 1981) and in living populations (e.g.,
Marshall, 1968) suggest that this stress indicator should be interpreted
cautiously in bioarchaeological analysis, especially in consideration of
health status and its relationship to specific behavioral, environmental, and
dietary adaptations.
2.4.3
44
Cementoenamel
junction
45
Dentin
Root
l
~
from small pits or furrows to large, deep grooves or even large areas of
missing enamel. Typically, these defects are horizontal grooves that are
called chronological or linear enamel hypoplasias (Figure 2. 7). The color
and hardness of hypoplastic enamel is normal.
Hypocalcifications are enamel defects involving change in color or
opacity, reflecting variation in enamel quality or hardness. The enamel
surface is usually smooth and appears intact. Hypoplasias occur when
ameloblasts fail to produce the normal thickness of enamel matrix during
enamel development. Hypocalcifications appear to result from a disruption
of the mineralization process during the maturation stage of enamel
development. This dichotimy may not be so clear cut, since hypocalcifications ha ve been experimentally documented in the initial stage of enamel
formation (Suckling, 1989).
Hypoplasias result from three potential causes, including hereditary
anomalies, localized traumas, and systemic metabolic stress (Goodman &
Rose, 1991 ). Defects arising as hereditary anomalies or as localized
traumas are rare in human populations, indicating that the vast majority of
hypoplasias seen in contemporary and archaeological populations are
linked to systemic physiological stress. The causal stressors assciated with
46
hypoplasias are numerous and varied. Clinical and epidemiological investigations in living populations document associations with systemic diseases,
neonatal disturbances, and nutritional deprivation (reviewed by Hillson,
1996; Pindborg, 1982). Experimentally induced stress in laboratory animals has also shown the direct link between enamel deficiency and stress
(e.g., Kreshover, 1944; Kreshover & Clough, 1953a, l 953b; Suckling et al.,
1983, 1986; Suckling & Thurley, 1984). Studies of non human primates with
known life histories revea! links between enamel defects and life events,
including birth, parturition, poor physical growth, social stress, and the
stresses associated with capture (Bowman, 1991). Thus, enamel defects are
a nonspecific indicator of physiological stress (Goodman & Rose, 1990;
Kreshover, 1960; Pindborg, 1982).
Ecological factors are critica! far understanding the prevalence and
pattern of enamel defects in human populations. Studies ofliving populations with dietary deficiencies show the primacy of nutrition in the
development of normal enamel. Analysis of individuals born during the
starvation famine of 1959-1961 in the People's Republic ofChina reveals
that enamel farmed during the famine is highly defective, unlike the enamel
that farmed either befare or after the famine (Zhou, 1995). Rural
individuals have more defects than urban individuals, a pattern consisten!
with records indicating that the rural population was subjected to more
stress than the urban population (Zhou, 1995).
Enamel hypoplasias show a predilection far anterior teeth and far the
cervical and middle thirds of tooth crowns, suggesting that specific teeth
and regions of crowns are differentially susceptible to growth disruption
(Condon & Rose, 1992; Goodman & Armelagos, 1985a, 1985b; Hutchinson & Larsen, 1988; Li et al., 1995; Pedersen & Scott, 1951; Zhou, 1995).
Susceptibility to growth disruption in specific teeth and specific areas of
tooth crowns may vary according to the deposition rate of enamel matrix:
47
position of the active ameloblasts at the time of the insult (Figure 2.8). They
typically begin at the cuspa]/incisal slope of the hypoplasia and terminate
at the dentoenamel junction (Condon & Rose, 1992; Goodman & Rose,
1990). A common association between these incremental structures and life
history is the birth event, resulting in a distinctive 'neonatal line' on the
farming teeth, namely the deciduous teeth and the permanent first molars
(Schour, 1936; Whittaker & Richards, 1978). Wide Wilson bands are
associated with trauma tic births (Eli et al., 1989), suggesting that the width
of Wilson bands may represen! an iudicator of stress severity.
Wilson bands are not always associated with a surface macrodefect such
as a hypoplasia (Bullion, 1986; Condon, 1981; Condon & Rose, 1992;
Danforth, 1989; Goodman &Rose, 1990; Rose, 1977; Wright, 1990). The
lack of a consisten! association between macro- and microdefects suggests
that their etiologies are different. Wilson bands appear to represen! brief
periods of stress lasting from one to five days, whereas hypoplasias appear
to represen! long-term str~ss lasting from weeks to severa] months
(Condon, 1981).
48
49
Stress chronology
Because metabolic insults leading to growth disruption affect only the part
ofthe tooth that is in the process offorming, location ofthe disturbance on
the tooth crown provides a precise chronological indicator of stress history.
Toothenamel begins to form at about four months in utero, beginningwith
the deciduous first incisors, and is co)Ilplete when the crowns of the
permanent third molars are fully farmed at about age 12 (Smith, 1991; Ten
Cate, 1991). The location of the position of an enamel defect (e.g.,
hypoplasia) relative to the cementoenameljunction can be used to plot the
age of disturbance (Goodman et al., 1980; Sciulli, 1992; Swiirdstedt, 1966).
Earlier researchers suggested that there is a preprogrammed stress clock in
humans (e.g., Massler et al., 1941; Sarna! & Schour, 1942). However, the
body of evidence that has built up over the last five decades indicates little
support far a universal model of the timing of growth disruption (see
Goodman, 1989; Goodman & Armelagos, 1985a, 1985b; Zhou, 1995; and
below).
Although the relationship between enamel defects and age has been
recognized since the nineteenth century (e.g., Talbot, 1898), this chronological approach has only recently been applied to archaeological remains.
The pioneering investigation of Medieval-era dentitions from Westerhus,
Sweden, by Swiirdstedt (1966) revealed that hypoplasias peaked in the two
to faur year period, a pattern that has been identified in many other
archaeological samples (e.g., Corruccini et al., 1985; Goodman et al., 1980;
Hillson, 1979; Hodges, 1989; Hutchinson & Larsen, 1995; Martn et al.,
1991; Powell, 1988; Storey, 1992a, 1992b) (Figure 2.9).
The tendency far hypoplasias to occur after the first year in archaeological samples suggests that stresses may be due to the negative effects of
weaning (e.g., Cappa et al., 1995; Corruccini et al., 1985; Lanphear, 1990;
Lillie, 1996; Moggi-Cecchi et al., 1994; Ogilvie et al., 1989; Simpson et al.,
1990; Ubelaker, 1992b; Webb, 1995; and many others). Weaning may not
be the most appropriate explanation in ali circumstances. A test of the
weaning hypothesis based on the study of historical records and archaeological dentitions from enslaved African-American populations living in
Maryland and Virginia reveals that the peak frequencies ofhypoplasias are
in the 1.5--4.5-year age intervals, whereas weaning took place only nine
months to one year afier birth (Blakey et al., 1994). This discrepancy
between age pattern ofhypoplasias and weaning led Blakey and coworkers
(1994) to conclude that weaning was not the primary causal factor leading
to enamel defects. Rather, other stresses of enslavement, including nutritional problems, poor hygiene, and illness, were likely to be responsible far
30
!!l.
120
~
...
10
1\
~\
o
o
3
4
Age (years)
hypoplasia frequencies (o/o per half-year age group) compared. with Georgia
coastal foragers (unbroken line) and farmers (dashed line). (Frm Hutchinson
& Larsen, 1995; reproduced with permission of the American Museum of
Natural History.)
the age pattern of physiological perturbation in this setting. The predilectlon far the two to four year period reflects at least in part the greater
susceptibilityofthe regan ofthe tooth crown associated with this period of
cnamel deposition and tooth farmation. Therefare, the general acceptance
of weaning as a cause far the age profile of enamel defects in archaeological
scttings, usually between two and faur years, is incorrect (and see
Katzenberg et al., 1996). Weaning may certainly be a cause of stress leading
to poor enamel. However, the link between enamel defects and weaning is
coincidental rather than real in many circumstances.
Duration/severity of stress
50
Sarna! & Schour, 1941, 1942; Simpson et al., 1990). Suckling and
coworkers ( 1986; Suckling, 1989) concluded on the basis of experimental
work with laboratory sheep that severity of stress plays a vital role in
determining the width of individual hypoplasias. Therefore, hypoplasia
size may reflect either stress duration or severity or perhaps sorne unknown
combination of both.
Stress histories in human populations
The connection between poor living conditions and enamel defect prevalence is well supported by epidemiological studies of contemporary human
populations. In general, individuals fron developed nations tend to have
far lower prevalences than individuals from underdeveloped nations (see
Goodman & Rose, 1991). In tbis regard, less than 10% ofindividuals from
developed nations ha ve one or more hypoplasias, whereas hypoplasias are
commonplace in many underdeveloped settings or in disadvantaged
subgroups of populations with poorer diets, more disease, or sorne
combination of undernutrition and disease (e.g., Anderson & Stevenson,
1930; Baume& Meyer, 1966; Dobney& Goodman, 1991; Enwonwu, 1973;
Goodman et al., 1987, 1991; Goodman, Pelto et al., 1992; Infante, 1974;
Infante & Gillespie, 1974, 1977; Li et al., 1995; Zhou, 1995; Lukacs &Joshi,
1992; Massler et al., 1941; May et al., 1993; Pedersen & Scott, 1951; Sawyer
& Nwoku, 1985; Sweeney et al., 1971 ).
Severa! case studies are especially informati ve regarding the link between
stress, socioeconomic status, and life history. Children from villages in the
Solis Valley of the Temascalcingo region ofthe Mexican highlands display
prevalences of hypoplasias that document the relationship between poor
growth status and physiological disruption (Goodman, Pelto et al., 1992).
Children with enamel defects have reduced body weights and heights-forage in comparison with children who lack defects. Predictably, children
with hypoplasias tend to be from families of lower socioeconomic status
living under conditions of malnutrition and poor sanitation. The association between negative environments and defect prevalence is well illustrated in settings involving selected dietary supplementation (Dobney &
Goodman, 1991; Goodman et al., 1991; May et al., 1993). In rural Mexico
and Guatemala, for example, children receiving dietary supplements have
far fewer linear enamel hypoplasias than their nonsupplemented peers. In
Guatemala, children who were ill more than 3.6% of the time had more
hypoplasias than other children (May et al. 1993). Comparable patterns of
morbidity differences are reported far Wilson band prevalence in living
populations. Children with histories of chronic systemic disease in Shef-
51
52
53
1996b; Rose et al., 1978; Simpson et al., 1990; Stodder, 1995; Swiirdstedt,
1966; White, 1978). These findings suggest that individuals experiencing
stress during childhood are predisposed to early death. This may indicate
that individuals who are stressed in childhood continue to be stressed as
adults, resulting in weaker constitutions and earlier death. Alternatively,
individuals experiencing stress early in their Iives may somehow lose the
ability to deal with stress la ter in life. Finally, higher social position of sorne
juveniles in stratified societies (e.g., Dickson Mounds) may buffer them
from stresses experienced by lower social ranks (Goodman & Armelagos,
1988). Unfortunately, with archaeological samples, it is diflicult to determine the likelihood of the alternative explanations (see also Goodman,
1989; Goodman & Armelagos, 1988). The similar pattern of age differences
in vertebral neural arch size, tooth size, and enamel defect frequency,
especially in comparison of juveniles and adults from the same population,
strongly suggests that individuals surviving to adulthood enjoyed relatively
better health than !hose members of the population who expired prior to
reaching adulthood.
Wilson bands in archaeological remains also provide an importan!
avenue for investigating other major transitions in human populations that
might compromise health status and stress levels. Wright (1990) determined prevalences of bands in precontact and contact-era mandibular
canines from Lamanai, Belize, a Maya center occupied from the Preclassic
period through the Historie period. Following an initial period of abandonment after contact by Europeans, the si te was re-occupied as a Catholic
mission by Mayan Indians until the mid-seventeenth century. Diet changed
relatively little in the prehistoric to historie transition, but archiva! records
indicate that other stressors (e.g., European-introduced diseases) compromised health .in native populations following contact. Comparisons of
Postclassic and Historie dentitions show a dramatic increase in physiological stress: 84% of bands observed in the samples combined are from the
Historie dentitions. Historie individuals also show more bands than
precontact individuals (2.4 vs. 0.88 per individual, respectively). Given the
lack of major dietary changes, Wright (l 990) argues that the differences in
microdefect prevalence can be attributed to changing disease patterns with
contact, such as the introduction of malaria and other Old World parasitic
infections, diseases leading to acute health crises.
A contrasting temporal trend in microdefect prevalence in comparison
of prehistoric and contact era native populations is identified at another
mission locality in Belize. Microdefect prevalence in mission Indians at
Tipu decreases in comparison with precontact lndians (Cohen et al., 1994;
Danforth, 1989). The different temporal trends at Lamanai and Tipu may
54
55
o.o
0.5
1.0
1.5
2.0
2.5
3.0
3.5
4.0
4.5
5.0
5.5
6.0
6.5
7.0
10.0-16.0
SI
o
o
4
Sil
25
13
13
14
38
31
37
28
53
47
39
40
18
32
21
25
25
20
19
29
28
32
lO
15
o
o
24
SIII
29
53
75
90
65
76
52
74
57
8
5
61
17
17
35
100
56
Adult stress
1.5.J
Bone mass
Adult stress
57
at about age 35 (Garn, 1970; Heaney, 1993; Pfeiffer & Lazenby, 1994). At
about age 40, bone commences resorption endosteally but continues to be
deposited periosteally. The imbalance ofbone loss and bone gain on these
respective surfaces dueto relatively greater endosteal losses results in a net
reduction ofbone tissue during and following the fifth decade oflife (Bales
& Anderson, 1995; Garn et al., 1967, 1992; Smith & Walker, 1964).
Adult bone loss leads to increased risk of fracture in older adulthood
owing to a complex disorder called osteoporosis (Anderson, 1995; Anderson & Pollitzer, 1994; Heaney, 1993; Stini, 1990, 1995). Two types ofbone
loss dueto osteoporosis are identified clinically, including that arising from
reduction in estrogen levels following menopause (Type I), and gradual
age-related reduction in borre mass in adult females and males (Type II)
(Drezner, 1995; Stini, 1990). Women lose relatively more borre mass than
men, due to the combined affects of Type 1 and Type II osteoporosis.
Estrogen is critica! in borre maintenance (Drinkwater, 1994). Even in
younger women undergoing overvigorous exercise regimens and accompanying loss of menstruation (secondary amenorrhea), the reduction in
estrogen results in significan! borre losses (Anderson, 1995; Kreiner, 1995).
The rate of borre loss in human populations is variable, and environmental factors such as nutritional status are significan! influences (Arnaud
& Sanchez, 1990; Martin et al., 1985; Pollitzer & Anderson, 1989;
Schaafsma et al., 1987). Clinical evidence indicates that individuals with
low calcium intakes are more prone to adult borre loss, and other dietary
factors such as high protein consumption are also implicated (Arnaud &
Sanchez, 1990; Nordin, 1984; Pfeiffer & Lazenby, 1994; Stini, 1990, 1995).
Body weight, heredity, and lactation status are also importan! risk factors
(Arnaud & Sanchez, 1990; Evers et al., 1985; Heaney, 1993; Kreiner, 1995;
Pollitzer & Anderson, 1989; Schaafsma et al., 1987; Stini, 1990, 1995).
Comparisons of active vs. sedentary populations or athletes vs. nonathletes
indicate the strong influence of physical activity on borre maintenance:
simply, active individuals have stronger, denser borre than sedentary
individuals (Anderson & Pollitzer, 1994; Drinkwater, 1994; Lacey et al.,
1991; Marcuset al., 1992; McMurray, 1995; Y ano et al., 1984). Owing to a
decrease in physically demanding lifestyles in a number of countries (e.g.,
Sweden, United States, United Kingdom, China), there appears to be a
secular increase in osteoporotic fracture (Allander, 1995).
Adult borre mass is documented in human remains from a variety of
archaeological settings, including Sudanese Nubia, and eastern, southwestern, and high-latitude North America (see Pfeiffer & Lazenby, 1994);
Much of this research shows either a general similarity or accelerated
patterns of borre loss in archaeological samples and in living populations
58
(e.g., see Carlson et al., 1976; Cook, 1984; Dewey et al., 1969; Van Gerven,
1973; Van Gerven et al., 1995). Variation in relation to differing lifestyles
and subsistence strategies has been examined in sorne detail with the use of
alternative data ollection protocols, including raw measures ofbone mass
(cortical thickness [CT], cortical area [CA], bone mineral content [BMC])
or size-standardized measures (per cent cortical area [%CA or PCCA] or
per cent cortical thickness [%CT or PCCT]) (Ruff, 1992). Comparisons of
femoral cortical thickness in X-group (AD 35~550) intensive agriculturalists from the Wadi Halfa area ofSudanese Nubia with modern Euroamericans and Native Americans revea! similar trends of initial gains in bone
mass from the third to fourth decades, followed by losses (Martin &
Armelagos, 1979; Martin et al., 1985). In Nubian females bone mass
decreased after age 20. Martin and coworkers (1985) speculate that
premature osteoporosis was due to nutritional inadeqqacies associated
with an over-reliance on a single dominan! crop (millet) ~ such as
protein--calorie malnutrition or imbalance of calcium/phosphorus ratios or the influence of disease. Perhaps the bone losses in childhood in this
setting (Hummert, 1983; and see above) predisposed adults, especially
females, to premature bone loss.
Similarly, bone mass (%CA from second metacarpals) in late prehistoric
maize agriculturalists from southern Ontario is below what has been
documented in living populations (Pfeiffer & King, 1983; cf. Garn, 1970).
Although various factors may be involved, the reliance on maize and attendant protein-calorie malnutrition may have contributed to low bone m~ss.
With the assumption that cortical thickness and Nordin's Index (cor!Jcal
thickness/total subperiosteal area) are useful measures of bone mass and
nutritional quality, Owsley (1991) compared femoral bone mass in a
temporal series of Great Plains Arikara dating from ca. AD 1600 to 1832.
These comparisons reveal an increase in bone mass in the transition from
Adult stress
59
Histomorphometry
60
the Haversian canals. Each !acuna contains cells that maintain the bone tissue
and are arranged in layers (lamellae). (From Larsen, I987; illustration by
61
2.6
ratios characteristic of maize-based diets (Stout, 1983), or perhaps a
variation in skeletal tissue maturation rates in different populations (Stout
& Lueck, 1995).
The rate of mineralization ofosteoid (unmineralized bone matrix) in the
Haversian canal can be influenced by stress. Under conditions of normal
bone development, osteons mineralize uniformly, but under conditions
involving slower growth - such as with nutritional stress or disease delayed osteoid mineralization results in the creation of hypermineralization zones. Viewed in cross section, one or more hypermineralized zones
62
deficiencies often resulting from the synergy between poor nutrition and
infection. If these conditions are maintained throughout the years of
growth and development, then the affected individuals are likely to be
short-statured as adults. Overall, the early environment has long-lasting
effects on health status over the lifespan. Juveniles experiencing elevated
stress can have poor health and shortened lifespans as adults (and see
Henry & Ulijaszek, 1996).
.
Do small-bodied humans have an adaptive advantage over large-bodied
humans living under circumstances of reduced resource availability or
adequate nutrition? Seckler (1980, 1982) proposed that shortness in body
height in developing nations is an adaptation to reduced food supplies.
This reduction, he argues, results in ind_ividuals who are small but healthy.
If reduced body size is adaptive, then reduced height should have no
associated functional costs, In fact, small body size is linked with various
negative factors, including increased disease and poor nutrition. Poor
growth status is associated with a range of functional co~ts and consequences, including decreased activity and poorer learning (Crooks, 1995;
Dasgupta, 1993; Goodman, 1991, 1994; Stinson, 1992). Although smaller
body size appears to enable individuals to perform sorne activities with
lower energy requirements, the efficiency is reduced (Stinson, 1992).
Clearly, there are negative consequences of small body size in disadvantaged settings, indicating that this reduction is maladaptive.
Sorne also suggest that there may be an adaptive advantage for iron
deficiency, especially in light ofthe apparent link between iron withholding
and microbial invasion (Kent et al., 1990, 1994; Stuart-Macadam, 1992a,
l 992b). Clinicians observe that decreases in serum iron reduce the
availability of iron for microbial growth, thus inducing a 'nutritional
immunity' (Weinberg, 1974, 1992; see discussion and review by Keusch &
Farthing, 1986). Weinberg (1992) notes that animal and human studies
show that hosts not withholding iron are at increased risk for infection
(bacteria!, fungal, protozoan), and conversely, risk of infection decreases
with increased iron withholding (although see Berger et al., 1992; Keusch &
Farthing, 1986). Thus, iron deficiency anemia - at least in the mild to
moderate form - may be an adaptive response to chronic pathogen loads
(Kent et al., 1990, 1994; Stuart-Macadam, 1992a, 1992b).
Goodman (1994) evaluated the pathogen load model, and suggests that
instead ofbeing an 'adaptation' to stress, iron deficiency anemia should be
considered an 'adjnstment' to stress. Like reduced growth, iron deficiency
has a series of direct functional costs for a population. For example, even
when an individual is only slightly deficient in iron, a number of key
enzymes involved in vital functions (e.g., DNA synthesis) are affected. At
1
1'
1
1
i
1
1
l
1
1
1
1
63
the organism level, iron deficiency anemia has profound negative effects on
work capacity, cognition, and the maintenanceof a healthy immune system
(Goodman, 1994). Additionally, dietary iron has a very importan! influence on iron status. Low iron intake is the leading cause of iron deficiency
anemia in the United States (children and adult women) and Argentina
(children) (see Goodman, 1994). Thus, in archaeological settings, although
porotic hyperostosis may be an indication of the body's attempt to adjust
to increased pathogen loads, it nevertheless reflects an increased health
burden.
In the study of past societies, it is possible to test the 'adaptive' vs. 'stress'
models for the growth and nutrition stress indicators discussed in this
chapter. Significantly, various indicators are linked with decreased survival
as determined by mean age-at-death of individuals with and without (or
with relative differences in prevalence of) the indicator. Enamel defects
(macro- and microdefects), vertebral neural canal size, and tooth size show
clear links with lifespan. Where age-at-death and stress indicators are
examined concurrently, individuals without enamel defects, with larger
neural canal size, and with larger tooth size appear to have died later in life.
These findings suggest that skeletal stress indicators are related to quality
of life and do not represent adaptations.
Dental caries
Exp~sure
3.1
Introduction
to iefectious pathogens
65
3.2
Dental caries
3.2.I
66
Meiklejohn et al., 1992; Milner, 1984; Molnar & Hildebolt, 1987; Molnar &
Molnar, 1985b; Newbrun, 1982; Powell, 1985; Rowe, 1982; Woodward &
Walker, 1994).
Intrinsic characteristics offood and the consistency and mannerin which
it is prepared strongly influence cariogenesis in human populations. For
example, in the American Southeast, more than three times the number of
late prehistoric Caddoan farmers have carious teeth than the FourcheMaline foragers that predate them (Powell, 1985). The higher prevalence in
the later population is due to two factors: (!) consumption of maize, a
highly cariogenic food (see also below); and (2) reduced occlusal surface
wear, which increases the probability that cariogenic bactetial colonies will
aggregate in caries-prone areas, such as in grooves between cusps of
Dental caries
67
premolars and molars (Powell, 1985). Maat & Van der Velde (1987) also
found a negative correlation between frequency of occlusal surface caries
nnd degree of dental wear in molars from sailors recovered from a
seventeenth and eighteenth century Dutch whaling station in the Spitzbergen Archipelago (Svalbard). In this series, increased wear appears to be
ussociated with fewer carious lesions. They concluded that '(t)hese findings
strongly suggest a competitive relationship between progress in caries and
attrition' (Maat & Van der Velde, 1987:281). The studies linking reduced
wear to increased caries prevalence are convincing, especially because
common sites of plaque and cariogenesis are the grooves and fissures of
unworn crowns (and see Christopherson & Pedersen, 1939; Corbett &
Moore, 1976; Milner, 1984).
The relationship between low caries rates and high-wear dental environments should not be overgeneralized. In their study of the Mesolithic
dentitions from Cabeco da Arruda and Moita do Sebastiiio, Portugal,
Meiklejohn and coworkers (1992) found a positive correlation between
caries and wear in molars; the most heavily worn crowns are the most
carious. In this setting, the Mesolithic Portuguese individuals consumed
figs and carob, foods high in sugar content that also produce high rates of
wear. Occlusal surface wear is also excessive in Archaic-period foragers
from the lower Pecas region of southwestern Texas, resulting in pulp cavity
exposure and tooth loss (Hartnady & Rose, 1991; Sobolik, !994b). Caries
prevalence is high (14% of teeth), and is indistinguishable from many
agricultura! groups (see also Bement, 1994; Marks et al., 1988).
Coprolite analysis reveals that various highly abrasive materials were
included in foods consumed, including phytoliths, seeds and small bones,
and calcium-oxylate crystals from succulents and cacti (Hartnady & Rose,
1991; Sobolik, !994b). Historie accounts also document the introduction
of abrasives to food, including ash for baking of sotol cactus and dirt to
'sweeten' meals. High-carbohydrate foods such as succulent fibers, prickly
pear fruits, pecans, and mesquite resulted in active cariogenesis. Thus, like
the Portuguese Mesolithic foragers, Pecas region Indians show a positive
relationship between tooth wear and caries.
3.2.2
68
Dental caries
69
20
- 15
ff.
10
Period
Figure 3.2. Percentage of teeth affected by dental caries in eastern North
America. {Based on data from Milner, 1984; Larsen et al., 1991.)
70
River valley, the Ohio River valley, and the mid-South generally, the first
agricultura! development involved use of starchy seed-bearing plants (e.g.,
chenopod, sunflower, and cucurbit), coinciding with social and economic
transformations during the Middle Woodland period from ca. 250 BC to AD
200 (Fritz, 1990; Smith, 1992). For this time period, Rose and coworkers
(1991) identified an increase in carious lesion frequency that is probably
Iinked with this reorientation of diet well before the adoption of maize in
later prehistory (ca. AD 900).
No comparisons have been made with regard to relative severity or
prevalence differences that might be present between consumers of different domesticated plants. No studies have compared caries prevalence
values for populations dependen! on maize in the New World vs. wheat in
the Old World. Thus, there is no understanding ofthe relative differences of
the impact ofagriculture on populations consuming different types of plant
domesticates. Lubell and coworkers (1994) remark that European Neolithic populations have generally lower caries prevalence values than
pro vides an importan! meaos of assessing the severity of the disease process
(see Hillson, 1996). Comparisons ofprehistoric foragers with la ter farmers
in the Tennessee River valley and on the Georgia coast indicate that,
through time, lesions increased in size and affected more areas of tooth
crowns (Larsen, 1982; Smith, 1982). Carious lesions in prehistoric foragers
are mostly restricted to the cervical region of the posterior dentition, which
may be dueto food impaction between teeth (Smith, 1982). In contras!, late
prehistoric maize farmers display lesions on multiple locations of tooth
crowns.
Dental caries
71
72
3.1.3
Dental caries
73
al., 1982; Lee, 1968; Meehan, 1977; Walker & Hewlett, 1990; Woodburn,
1968).
For the most part, sex differences in caries prevalence in archaeological
samples are identified in populations practicing agriculture to sorne degree.
However, female and male foragers from the Santa Barbara Channel
Island region display differences in caries prevalence for much of prehistory
(Lamber! & Walker, 1991; Walker, 1986b; Walker & Erlandson, 1986).
Ethnographic observations indicate a distinctive sexual division of labor
whereby men hunted and fished and women collected plants. Historie
accounts indicate that men ate more of the game they hunted than did
women. Thus, it appears that greater consumption of plants by women - a
factor related to their subsistence responsibilities - is reflected in their
greater caries prevalence.
Walker and Hewlett (1990) have investigated dental caries in severa!
groups of central African pygmy foragers (Aka, Mbuti, Efe) and Bantu
farmers in order to build support for a behavioral interpretation of
prevalence variation. Comparisons of subsistence patterns and food
consumption practices in these groups revea! clear intra- and interpopulation differences in caries prevalences that provide an importan! perspective
on dietary behavior and cariogenesis. Aka and Mbuti foragers practice
net-hunting, and the Efe foragers bow-hunting. The sources of animal
protein available to these groups vary depending on season (e.g., meat from
seasonally available animals), and is mostly obtained through hunting and
collecting. Manioc and other cultigens acquired through trade with Bantu
agriculturalists form a significan! componen! of their diets. The relative
consumption of meat to cultigens is highly variable, ranging from a high
proportion for the Mbuti (al leas! par! of the year) to a much lower
proportion for the Efe. The Aka apparently consume similar amounts of
meat to Mbuti foragers, since both are net-hunters. On the basis of
comparisons of amount of time spent hunting, it appears that Mbuti and
Aka foragers consume more meat than do Efe foragers. In addition to meat
and cultigens, honey is an importan! part of the diet for part of the year.
For Mbuti foragers, this highly cariogenic food contributes nearly 80% of
calories for a one-month period (lchikawa, 1981 ).
Virtually ali foods consumed by Bantu villagers are plant domesticates,
including manioc, maize, rice, peanuts, and plantains. The little amount of
meat that is consumed - approximately 2.5% of foods - is acquired from
pygmy foragers.
Field observations indicate differences in food consumption between
females and males in the foragers, but not in the farmers. Diets of forager
women contain more plants than do those of forager men. Aka men
74
Teeth (n)"
'Yo carious
Aka
Total
Males
Fema!es
(3099)
( 1706)
(1393)
5.2
4.2
6.6
Mbuti
Total
Niales
Females
(1773)
(1048)
(753)
6.0
5.1
7.3
(277)
(277)
6.0
6.0
(630)
(308)
(322)
8.1
9.1
7.1
Efe
Total
Males
Fema les
Ban/11
Total
Males
Females
consume more meat than do women, and it is acquired on the hunt prior to
their return to the home village; sorne of the choicest cuts of meat are shared
among the roen who participated in the hunt.
Bantu farmers ha ve the highest dental caries prevalences in comparison
with foragers (Table 3.1), which appears to reflect thcir greater consumption ofplant carbohydrates. Although foragers ha ve relatively lower caries
prevalence, the values are nevertheless appreciable (5-6%; Table 3.1),
which points to the significantcomponent ofplant carbohydrates acquired
frorn Bantu farmers via trading relationships. Caries prevalence is also
higher for pygmy women than for men. These differences are related to
food consumption variation between male and fema!e pygmy foragers. The
frequency ofbetween-meal eating in pygmy women is an additional factor
that provides partial explanation for sex differences. In these foragers, men
concentra te theireatingin several large mea Is, and women snack frequently
during the day. Clinical evidence froni Western populations indica tes that
snacking between meals (especially carbohydrates) results in elevated
Dental caries
75
caries rates (e.g., Burt et al.; 1982, 1988; Gustafsson et al., 1954; Konig,
1970; Konig et al., 1969; Nizel, 1973; Rowe, 1982; Weiss & Trihart, 1960;
although see Rugg-Gunn et al., 1984).
Ethnographic documentation of dietary practices in native groups in
South America provides additional insight into differences between females and males in dental health (P. L. Walker et al., unpublished
manuscript). In the three groups studied - the Yanomamo of Venezuela,
the Yora of southeastern Peru, and the Shiwiar (Achuar) of Ecuador mea! and fish provide a significan! par! of the diet, and most carbohydrates
are from plan! crops, such as manioc and bananas. Ali three groups have
significan! caries rates, in part dueto the consumption of cultigens, but also
dueto the reduction in their isolation and greater access to processed foods.
Yora and Shiwiar women spend many hours processing manioc in their
mouths for production of beer (chicha). For these two groups, women
display a somewhat higher frequency of carious teeth than roen. Their
relatively greater exposure to cariogenic manioc explains higher caries
rates.
Given the finding that females are frequently more carious than males, it
seems possible that there may be sorne underlying, nonbehavioral reasons
for these differences. Permanent teeth erupt slightly earlier in females than
in males, exposing their teeth at an earlier age to caries-promoting factors
(e.g., Carlos & Gittlesohn, 1965; DePaola et al., 1982; Dunbar, 1969;
Walker, 1986b). However, tooth eruption differences between males and
females show either weak or no correlation with dental caries prevalence
(e.g., Moorrees, 1957; Toverud et al., 1952; Ziskin, 1926). Alternatively,
there is a long-held popular notion that pregnancy results in poor dental
health, including tooth loss and caries. There is evidence for increasing
gingival inflammation in sorne pregnant women (e.g., Arafat, 1974; Loe,
1965; Loe & Silness, 1963; although see Jonsson et al., 1988; Maier &
Orban, 1949). No evidence exists for increase in tooth loss or dental caries
due to gingivitis or other factors during pregnancy (Larsen et al., 1991;
Walker, 1986b; Walker & Hewlett, 1990). Therefore, sex differences in
eruption timing or pregnancy cannot explain variation in dental caries
between males and females. If dental caries differences could be explained
by sorne underlying physiological or developmental reason, then the
pattern of greater prevalence of caries in women than roen should be
universal, or nearly so. In fact, there are exceptions indicating that other
factors are involved (e.g., a number ofinvestigations show either the same
prevalences or greater prevalence in males than infernales in archaeological
and clinical settings: Barmes, 1962; Burns, 1979, 1982; Clarkson &
Worthington, 1993; Moore & Corbett, 1973; Pietrusewsky, 1988; Powell,
76
1988; Rowe, 1982; Sutter, 1995; Wells, 1980; White, 1994). Therefore,
variation in caries prevalence in females and males is behaviorally
mediated.
3.2.4
77
African foragers. Walker & Hewlett (1990) found tbat high-status pygmy
leaders bave fewer carious lesions than low-status nonleaders. They suggest
that greater access to meat by leaders (from gifts and tribute), combined
with reduced consumption of carbohyrate-rich plants, best explains the
discrepancy between social ranks.
3.3
3.3.1
78
79
Figure 3.3. Periodontal disease; central European Bronze Age. Note large gap
between the alveolar bone and cementoenamel junction for all teeth. (From
Hildebolt & Molnar, 1991; reproduced with permission of authors and
Wiley-Liss, Inc., a division of John Wiley & Sons, Inc.)
checked, the skeletal support for the teeth diminishes, and, ultimately,
exfoliation occurs (Figure 3.4). Once gone, the soft tissue heals, and the
tooth socket is completely remodeled. Although the progression of
periodontal disease is well documented in human populations, ancient and
modern, it remains an etiological conundrum. There is general agreement
that bacteria- perhaps as many as 40 dilferent taxa- may be involved in the
onset and progression ofthe disease (Drake et al., 1993; Enwonwu, 1995;
Hillson, 1986). Other importan! intluencing factors include poor oral
hygiene, cariogenesis, malocclusion, nutritional status and to a lesser
extent pregnancy, puberty, and psychological stress (see Enwonwu, 1995;
Hildebolt & Molnar, 1991; Hillson, 1986).
3.3.2
80
81
82
3.4
Nonspeciflc infection
3.4.1
Skeletal lesions of infectious origin represen! a continuum initially involving the periosteum, followed in severity by involvement of cortical bone
generally, and, at the extreme end, extension of infection into the medullary
cavity. Periostitis (or periosteal reaction) is the least severe, and osteitis and
Nonspecific infection
83
84
Nonspecific infection
1
1
85
86
Nonspecific infection
87
unpublished manuscript; and see above for more detailed description of the
samples).
Unlike the Dickson Mounds study, these analyses focus on comparisons
of populations consuming exclusively wild plants and animals with
populations utilizing maize to varying degrees. For most ofthe time prior
to the arrival of Europeans in the sixteenth century, native groups were
nonsedentary foragers who obtained most foods from a combination of
hunting, gathering, and fishing. Archaeological and isotopic documentation of subsistence economy indicates that marine resources from estuarine
and marine contexts provided most of the protein in the na ti ve di et (Larsen,
Schoeninger et al., 1992; Reitz, 1988, 1990). During the twelfth century AD,
maize rapidly took on an increasingly importan! dietary role. With the
arrival of Europeans and the establishment of Catholic missions, there was
a subsistence reorientation whereby maize became highly significant
Coupled with these dietary changes were alterations in population size,
density, and sedentism. Prehistoric foragers in the region appear to ha ve
been sparsely settled and highly mobile. Prehistoric farmers, though, were
living in larger, more densely occupied villages, probably for longer periods
oftime (Larsen, 1982). ln the mission period, populations wereencouraged
to live in and around mission settlements. Although population size was
reduced dramatically during the contact period, the settlements were
permanent and villages were crowded.
Prevalence comparisons for periosteal reactions in adult tibiae indicate
that there is an increase in frequency prior to contact from 9.5% to 19.8%.
In the early mission population, the prevalence dedined slightly to 15.4%,
but then greatly increased to 59.3% in the late mission period. These general
. increases affected both adult males and females. The moderate increase in
the precontact populations is well under that observed at Dickson Mounds:
In ali likelihood, the relatively lower prevalence in the Georgia Bight
reflects considerably smaller population size and less of a commitment to
maize agriculture in the later prehistoric period in comparison with interior
Mississippianpopulations (e.g., Dickson Mounds). The marked increasein
the late mission period is probably tied to the relocation and increased
concentration of native populations around mission centers and the
introduction of new diseases, including possibly venereal syphilis. The
change in settlement provided conditions conducive to the maintenance
and spread of chronic infectious diseases and other factors that lead to an
increase in bone lesions (Larsen & Harn, 1994; C. S. Larsen et al.;
unpublished manuscript). The effects of increased infection rates would
probably have been exacerbated by the increase in emphasis on nutrition-
ally poor foods, especially maize.
88
Nonspecific infection
89
Larsen & Harn, 1994; Milner, 1991).Some ofthis variability is undoubtedly due to interobserver differences in recording methods. The variable
pattern of infection prevalence also points to a high degree of divers~ty
between human groups occupying very different landscapes and phys10graphic zones, ranging from highly fertile river bottoms (e.g., Lallo et al.,
1978) to marginal uplands (see Eisenberg, 1986a) or coastal regions (e.g.,
Larsen, 1982). Detailed analysis of population trends indicates that
population histories fluctuated dramatically, with regard to both size a_nd
distribution (Milner, 1990). Living in peripheral settings did not prov1de
freedom from disease - sorne of the highest prevalences of bone infection
are in the so-called 'marginal' habitats (e.g., Eisenberg, l986a, I99la,
1991b).
Sorne evidence suggests that high population density and disease
burdens in combination with other factors (e:g., warfare), may have
contrib~ted to cultural terminations during later prehistory well befare the
arrival of Europeans (e.g., Eisenberg, !986a; and see Larsen, 1994).
Improved survivorship, coupled with a decline in prevalence of skeletal
infection, in Ontario suggests that populations may have adjusted to high
density in this setting (Katzenberg, I 992a). This suggestion contrasts
sharply with other contact-era settings where periosteal infections have
been shown to increase in a dramatic fashion (e.g., Larsen & Harn, 1994;
Stodder, 1994; Ubelaker, 1994).
90
Nonspecific infection
91
92
93
3.5
3.5.1
Treponematosis
Treponematosis is represented today by four ~ecogniz~~ disease syndromes: venereal syphilis, nonvenereal (endem1c) syph1hs (also called
bejel), yaws, and pinta '(Hudson, 1965; Mandell et al., 1990; Ortner &
Putschar, 1985). Ali but pinta result in hard tissue responses. Unfortunate'
94
!y, the skeletal lesions of the other syndromes are so similar that it is
virtually impossible to distinguish among them. The pathogens responsible
for the disease are bacteria! spirochetes of the genus Treponema, including
T. careteum (pinta), T. pal/idum pertenue (yaws), T. pal/idum pallidum
(venereal syphilis), and T. pallidum endemicum (endemic syphilis) (Mandell
et al., 1990). Because of the high degree of morphological and immunological similarity, sorne argue that thesemay not represen! different species, but
rather reflect differences in expressions due to cultural and environmental
factors that affect the mode of infection. This conclusion is borne out by
recen! DNA hybridization studies (see Ortner et al., 1992).
Treponema! infection is introduced via the skin or mucous membranes.
For venereal syphilis, the spirochete typically enters the body during sexual
contact from lesions on the genitals. F or endemic syphilis and yaws, the
pathogens are spread from nongenital lesions on the arms, legs, or trunk
during nonsexual contact between individuals (e.g., physical contact
between children playing). Foral! three syndromes, the infection spreads
throughout the bodyvia the circulatory system. Congenital transmission of
venereal syphilis involving passage of the spirochete from the mother
transplacentally to the fetus is well documented (Hudson, 1965; Ortner &
Putschar, 1985). In living populations, pinta, endemic syphilis, and yaws
have been found to be especially prevalen! in rural settings with poor
sanitation. Additionally, in tempera te to hot clima tes, individuals typically
wear relatively little clothing, which facilitates the spread of infection
through direct contact with infected minor wounds, such as abrasions and
cuts. Venereal syphilis characteristically appears in populations with
higher levels of sanitation, such as in urban settings in Western countries.
These groups tend to be fully clothed, which provides fewer opportunities
for the spread of infection via skin contact in the manner typical of
nonvenereal syphilis, pinta, and yaws.
The skeletal manifestations of the treponematoses are described in the
paleopathological literature in detail (Hackett, 1976; Ortner & Putschar,
1985; Powell, 1988). Yaws is most commonly represented by the inflammatory response of the periosteum surrounding the bones of the forearm,
hand, and lower leg bones. In the most severe form, repeated episodes of
periosteal reaction and remodeling may result in hypertrophy of the
anterior crests of the tibia, presenting an appearance of bowing called
'saber-shin'. Bone surfaces with el ose proximity to skin- such as the cranial
vault and the anterior tibia - also may express active lesions or pitted
defects from gummatous granulomas. Destructive nasal and hard palate
changes may occur, but are less prevalen! than in venereal or endemic
syphilis.
95
96
and the far more extensive study ofNorth American skeletal remains than
in Europe or other areas of the Old World, the issue is unresolved.
Researchers generally agree that treponematosis was present in both the
Old World and fhe New World well prior to European contact (see Baker &
Armelagos, 1988). Cases of treponematosis are only sparsely documented
in Europe and Asia (e.g., Dutour et al., 1994; Henneberg et al., 1992;
Roberts, 1993; Rothschild & Rothschild, 1995; Suzuki, 1982-1984).
Treponematosis is widespread in Australia (Hackett, 1936, 1976; Webb,
1995) and in sorne areas of the Pacific (e.g., Marianas: Hanson, 1988;
Stewart & Spoehr, 1952; Stodder et al., 1992). The evidence for
treponematosis from the New World is quite abundan!, it having been
identified both with regard to a range of case studies and population
differential diagnoses in North American contexts. In addition to earlier
studies (see review in Baker & Armelagos, 1988), examples of treponematosis are associated with highly diverse settings, including southern coastal
California (Cybulski, 1980; Lamber!, 1994; Walker & Lambert, 1989),
Northwest Coast (Cybulski, 1990), Great Plains (Schermer et al., 1994),
Southwest (Lahr & Bowman, 1992; Stodder, 1994; Stodder & Martin,
1992), and coastal Chile (Arriaza, 1995). In many contexts, the skeletal
manifestation appears somewhat intermedia te between those of the two
modern endemic syndromes. Considering the evolutionary nature of
human infectious disease over centuries and across host populations of
differing genetic composition, these departures from the modern pattern
are not unexpected.
The preponderance ofNew World treponematosis data is from precontact American Midwest and Southeast human remains. In the lower Illinois
River valey, a pattern of proliferative bone infection that is strongly
suggestive of endemic treponematosis rather than venereal syphilis is
present (Cook, 1976). There is a high frequency oftibial periostitis affecting
adult males and females alike, progressively increasing with age. The
prevalence is especia!ly elevated in later prehistoric, maize-dependent
groups with high population density. Although the pattern is variable
across populations, it has been described in (mostly late) prehistoric human
remains from other groups in Illinois (Garner, 1991; Milner, 1983, 1992;
Milner & Smith, 1990), Arkansas (Powell, 1989), Louisiana (Lewis, 1994;
Robbins, 1978), Mississippi (Ross-Stallings, 1989), Kentucky (Cassidy,
1984), Tennessee (Eisenberg, 1986a, 199la, 199lb}, Alabama (Powell,
1986, 1988, 199la, 199lb, 1992a), North Carolina (Bogdan & Weaver,
1992; Monahan, 1995; Reichs, 1989), Georgia (Powell, 1990, 1991 a, 1992b,
1994), and Florida (Bullen, 1972, 1973; Dickel, 1991; Hutchinson, 1993;
Hutchinson & Mitchem, 1996; i~can & Miller-Shaivitz, 1985; Miller-
97
Shaivitz & i~can, 1991). Severa! of these case studies provide details on
lesion morphology and other characteristics that strongly suggest the
presence of sorne form of endemic treponematosis.
Hutchinson (1993) found extreme proliferative periosteal apposition on
long bones (especially tibiae) and stellate lesions in crania from late
prehistoric and early contact contexts in the central Florida Gulf coast. F or
example, in the sixteenth century componen! of the Tatham Mound
sample, three crania exhibited healed stellate lesions (Figure 3.7). Similar
lesions are present in crania from the postcontact Weeki Wachee Mound
and precontact components of the Safety Harbor and Tierra Verde sites.
Although saber-shin tibiae are also present in these samples, stellate scars
appear to be the best single criterion for endemic treponematosis in the
Florida Gulf coast region (Hutchinson, 1993; and see Milner & Smith,
1990, for a discussion ofNorris Farms, Illinois). Similarly, detailed study of
the Mississippian-period Irene Mound series from north coastal Georgia
indicates the presence ofthe disease. In this series, lesion morphology and
prevalence is consisten! with the presence of treponematosis (endemic
syphilis)(Powell, 1990, 199 la). These include cranial stellate scars, destruc-
98
99
John Lawson noted that natives in theeastern part ofthe colony ' ... have a
sor! ofRheumatism or Burning ofthe Limbs, which tortures them grievously, at which time their Legs are so hot, that they employ the young People
continually pour water down them ... This not seldom bereaves them of
their Nose. I have seen three or four of them render'd most miserable
Spectacles by this Distemper. Yet, when they have been so negligent, as to
let it run on so far without curbing of it; at last, they make shift to patch
themselves up, and Iive forman y years after .. .' (Lawson, 1967:231 ). These
descriptions correspond well with modern clinical descriptions of deep leg
pain and orofacial lesions (see Hackett, 1951; Hudson, 1958).
Sorne examples of nonspecific periosteal reactions observed in archaeological skeletons are probably oftreponemal origin, especially in individuals who also display the classic lesions (e.g., stellate scars on crania and/or
highly proliferative bone on tibiae diaphyses). In cases where distinctive
symptoms are absent, the delineation between nonspecific infection (periostitis) and endemic treponematosis is difficult to determine.
The well documnted paleopathological diagnoses of treponematosis in
North American archaeological settings make it clear that the disease was
well established in the New World prior to the arrival ofEuropeans. In the
lower American Midwest and Southeast, regions that are subtropical or
that experience seasonal high hilmidity and temperatures, there appears to
be a cline of skeletal expression from the hotter, more humid coas tal regions
to the somewhat drier interior regions, resembling modern inter-regional
treponema! clines spanning 'classic' yaws and 'classic' endemic syphilis in
central and southern Africa (Basset et al., 1994; Froment, 1994; Grin, 1956).
Identification of Treponema DNA in prehistoric Chilean mummies (Rogan
& Lentz, 1994; Grin, 1956) is consistent with this interpretation.
Variation by sex or status in skeletal populations is difficult to assess,
especially given the vagaries of diagnosis. Where probable cases of
treponematosis have been identified, the prevalences in adult males and
females are broadly similar (e.g., Cook, 1976; Powell, 1988; although see
Powell, 1990). Powell's (1988, 1992b) investigation of the Moundville
skeletal series indicates no clear distinction between status groups. These
findings point to the widespread nature of the disease, but, as with
prevalen ce determination, it is not possible to determine precise frequencies
in subgroups of populations.
tion of portions of the nasal margin and maxillary palate, and numerous
tibiae with grossly expandeddiaphyses and bone proliferation (Figure 3.8).
Just how life-threatening or debilitating treponema! disease was in these
earlier societies is unknown. Due to the apparent endemic nature of the
disease, it may not have been a primary cause of mortality. Based on the
high degree ofhealing, Powell (1988) argues that populations at Moundville
had more or less successfully adapted to the disease. The presence of the
characteristic lesions indicates that the disease did impose a health burden
and resulted in no small amount of discomfort. In his description ofnative
populations in North Carolina in the early eighteenth century, the explorer
3.5.2
Tuberculosis
100
culo sis progressively destroys bone tissue, and is most commonly expressed
as erosive vertebral lesions of the lower back (lower thoracic and lumbar
vertebrae) and resorptive and slight proliferative changes of the pleural
(internal) surfates of ribs (Buikstra, 198la; Ortner & Putschar, 1985;
Pfeiffer, 1991; Powell, 1988; Roberts et al., 1994; Steinbock, 1976). The
disease is caused by the acid-fast, gram-positive bacillus Mycobacterium
tuberculosis (Cotran et al., 1994). The primary mode of transmission is by
breathing airborne microbes, usually in droplets introduced by sneezing or
coughing (Smith & Moss, 1994).
The infection pathway is usually through the respiratory tract, resulting
in a primary infection in lung tissue and subsequent secondary infection in
regional hilar lymph nodes (Hopewell, 1994; Ortner & Putschar, 1985).
Overa period ofyears, the bacilli may then spread to skeletal tissues via the
circulatory system, with a propensity for hematopoietic marrow_ and
cancellous bone. The vertebrae, ribs, sternum, and (for subadults) long
bone metaphyses are especially favored sites of secondary infection,
because of the presence of a rich blood supply and the scarcity of
phagocytic cells (Hopewell, 1994; Ortner & Putschar, 1985; Thijn &
Steensma, 1990). Any bone or joint can be involved in tubercular infections
(Berney et al., 1972). The process can result in extensive destruction of
cancellous bone, most commonly in vertebral bodies (Figure 3.9). With the
loss of bone mass, vertebral bodies may collapse; the resulting severe
kyphosis is called 'Pott's disease' after the original description by Sir
Percivall Pott (1779). Only rarely are transverse processes, pedicles,
laminae, or spinous processes of vertebrae affected.
The proliferative lesions on the pleural surfaces of ribs are presumed by
sorne to be associated with tuberculosis, but the cause remains unclear
(Kelley & Micozzi, 1984; Pfeiffer, 1991; Roberts et al., 1994). Examination
of skeletal remains from individuals with known cause of death shows that
tuberculosis sufferers are likely to possess these rib lesions, but this is not
always the case. lndividuals with nontubercular pulmonary disease sometimes have similar rib lesions. Therefore, ali rib lesions of this type should
not be interpreted as diagnostic oftuberculosis (Pfeiffer, 1991; Roberts et
al., 1994).
Tuberculosis has been traced to at least 5000 years ago in the Old World
(Daniel et al., 1994; Steinbock, 1976). Most early authorities argued that
the disease was absent in the New World prior to European contact (e.g.,
Hrdlicka, 1909). The identification of acid-fast bacilli and soft-tissue
tubercular lesions (Allison et al., 1973, 1981 ), and especially M. tuberculosis
DNA in precontact Peruvian and Chilean mummies (Arriaza et al., 1995;
Salo et al., 1994), demonstrates its early prehistoric presence in the New
101
:i
World. The global distribution of the disease via detection of M. tuberculosis DNA is also indicated in archaeological bone from Old World sites in
Turkey and northern Europe (Dixon et al., 1995; Spigelman & Lemma,
1993).
Destructive vertebral or proliferative rib lesions are identified in archaeological skeletal series representing a diversity of groups \Vorldwide, '
including the Middle East (Baker, 1997; Buikstra et al., 1993; Morse, 1967;
Ortner, 1979; Strouhal, 1991 ), Denmark and northern Europe (Bennike,
1985; Formicola et al., 1987; Inglemark, 1939; Manchester, 1991; Waldron,
1993), Greece (Angel, 1984), Japan (Suzuki, 1991), and elsewhere (see
Ortner & Putschar, 1985). ln the New World, a spate ofreports documents
102
103
104
3.5.3
Leprosy
105
(a)
(b)
Figure 3.10. (a) Alveolar atrophy (leprosy); St. J0rgensg8.rd, Nrestved,
Denmark. (From M0ller-Christensen, 1978; reproduced with permission of
Odcnse University Press.) (b) Metatarsal atrophy of left fnot; St. J0rgensgrd,
Odense, Denmark. (Photograph by Kirsten Anderson; reproduced with
permission.)
been admitted into the leprosy hospital in order to isolate them from other
members of the population at large (Meller-Christensen, 1978). MellerChristensen 's extensive studies of this series revealed a distinctive 'facies
leprosa' skeletal syndrome involving atrophy of the nasal and maxillary
regions, alveolar resorption, and anterior tooth loss (Figure 3.1 O). Additionally, hand and foot elements are atrophied and shortened (Andersen et
106
107
.&
al., 1992) (Figure 3.10). Similar pathology has recently been identified in a
survey of a large skeletal series recovered from the St. forgensgard leprosy
cemetery near Odense, Denmark (K. Anderson, personal communication).
Other skeletal pathology found in leprous individuals incldes cribra
orbitalia, periostitis on tibiae and fibulae, and maxillary sinusitis (Andersen, 1969; Boocock et al., 1995; M0ller-Christensen, l 978), although these
conditions are no! symptomatic of leprosy by themselves. Increased
prevalence of maxillary sinusitis, for example, appears to accompany
elevation in air pollution and the confines ofurban living in la ter Medieval
England (Lewis et al., 1995). Sorne individuals display dental changes
whereby the crown bases of maxillary incisors are concentrically constricted (Roberts, 1986). The presence of malformed teeth indicates that the
infection occurred early in childhood.
Other probable examples of leprosy from archaeological contexts are
from mostly isolated skeletons from Great Britain, Poland, Nubia, the
Near East, and possibly other localities (see Boocock et al., 1995; Lewis et
al., 1995; Manchester, 1991; Manchester & Roberts, 1989; M0llerChristensen & Inkster, 1965; Ortner & Putschar, 1985; Steinbock, 1976;
Zias, l 99 l ), but the bioarchaeological evidence is far less profuse in these
regions !han in Denmark. Manchester and Roberts (Manchester, 1991;
Manchester & Roberts, 1989) assessed skeletal, archaeological, and ar-
X
~
the disease was endemic. They argue that following the introduction of the
endemic form of leprosy, perhaps by the late Reman period, the disease
increased in prevalence, peaking during the thirteenth century or somewhat
later. Leprosy then declined and disappeared by the end of the fifteenth
century, but remained in Scandinavia much later than in other regions of
3.6
108
highly feared, and individuals with the disease were considered as living
dead, who were to be isolated, forgotten, and removed from society
(Moore, 1987).
Various causal factors discussed in this chapter underscore the fact that
infectious disease has a varied etiology. To be sure, specific pathogens are
linked with nonspecific and specific infectious diseases, and are identified as
their 'causes'. However, even when hosts are infected by these pathogens,
actual disease transpires only when pathogen virulence coincides with host
susceptibility in a conducive environment.
The prevalence of skeletal lesions in an archaeological population <loes
not show a direct one-to-one correlation with actual prevalence in a living
population. Far example, tuberculosis was highly prevalen! in sorne
preantibiotic groups, but it only rarely spread to the skeleton (Roberts et
al., 1994), generally reported as 3 to 7% of cases (reviewed in Milner &
Smith, 1990). Sorne archaeological series show somewhat higher prevalences (e.g., Buikstra & Williams, 1991; Eisenberg, 1986a; Milner & Smith,
1990). Additionally, it is importan! to distinguish between disease and
infection. Disease prevalence - whether drawn from living or extinct
populations - may represen! only a small part of the total picture of
infection. Similarly, the risks outlined in this chapter for infection may
differconsiderably from the factors that ultimately influence and determine
whether disease will develop from the initial infection.
Diseases involving skeletal tissues must have contributed significantly to
the burden of ill health in many earlier societies, jusi as they do today.
Various segments of populations may have been affected by infectious
disease differently, but the exact experience was always mediated by local
4.1
Introduction
110
Accidental injury
111
4.2
Accidental injury
Elemental patterns
Figure 4.1. Medial vicw ar distal right radii: healed Colles's fracture (right);
normal (left); anatomical specimens. (Photograph by Paul Braly.)
112
Accidental injury
113
114
Accidental injury
115
adult male. The presence of a large callus at the fracture site, but with
exposed trabecular bone, indicates that healing was incomplete at the time
of death. Finally, like Shanidar !, another adult male (Shanidar 5) has
frontal bone trauma. The injury was completely healed at death.
The Shanidar cranial injuries are part of an overall pattern of head and
neck injuries in European and western Asian late archaic Horno sapiens
(Berger & Trinkaus, 1995). Nearly one-third of these hominids have head
and neck trauma, which is more than twice the prevalence of a recen!
clinical sample from New York (Berger & Trinkaus, 1995) (Table 4.1).
Survey of traumas associated with a variety of occupations in recen!
humans indicates that American rodeoathletes also have a high prevalence
of head and neck injuries relative to other regions of the body (Berger &
Trinkaus, 1995). The pattern in rodeo athletesreflec(s the dangers of riding
highly irritated animals (e.g., Bos taurus, Equus caballus); head and neck
injuries in rodeo athletes result from impacts after being thrown from these
animals. By logical extension, the high prevalence ofhead and neck injuries
in Neandertals relates to hunting activities, especially involving encounters
with medium-sized ungulates (Berger & Trinkaus, 1995). Use of spears
would have necessarily placed hunters in close proximity to, and hence in
danger of bodily injury from, their enraged prey.
Sudanese Nubians
Figure 4.2. Atrophy and healed fracture o right humerus; Shanidar, Iraq. The
left humerus is nonnal. (Photograph and copyright by Erik Trinkaus;
reproduced with permission.)
man, 1982). Shanidar 1, a mature adult male, has multiple injuries. The
bones of the right upper arm and shoulder are less than half the size of the
bones of the left upper arm and shoulder (Figure 4.2), which may ha ve been
caused by either childhood nerve damage or adult disuse atrophy following
a severe injury (Trinkaus & Zimmerman, 1982). The diminutive right
humerus has two fractures, both at the distal end. One ofthe fractures may
be one side of a false joint (pseudarthrosis) or evidence of an amputation
(Stewart, 1977; Trinkaus & Zimmerman, 1982). This individual also
displays evidence of severe cranial trauma involving extensive scarring of
the frontal and a crushing fracture of the left eye orbit. Ali postcranial and
cranial injuries were completely healed at the time of death.
Other accidental trauma includes a rib fracture in Shanidar 4, a mature
Temporal comparisons revea! importan! trends in accidental injury patterns in recen! humans (and see below). In the Wadi Halfa region of
Sudanese Nubia, fracture prevalence increased in the Christian period (AD
550-1300) relative to earlier periods (Armelagos, 1969; Armelagos et al.,
1981). Within the Christian period in Kulubnarti, Nubia, there is a general
increase in fracture prevalence from 18% to 23% in the early (AD 550-750)
to late (AD 750-1550) periods, respectively (Burrell 'et al., 1986; Jurmain et
al., 1994; Kilgore et al., 1997; Van Gerven et al., 1995). Upper limb
fractures show an especially pronounced increase (30%) (Van Gerven et al.,
1995).
Increase in skeletal trauma in juveniles and old adults is more pronounced than in other age groups in the late Christian period at Kulubnarti. These age-specific increases in fracture prevalence may be due to
elevated risks ofliving in two-storey houses in the late period vs. one-storey
houses in the early period (Burrell et al., 1986). Access to the living area on
the second storey in late'period houses was gained by use of a retractable
ladder, which may have caused falls and other types of accidents (Burrell et
al., 1986).
ll6
Accidental injury
Head/
Group
neck
Trunk
arm
Hand
Neandertal (n = 17)
Bt-5 (n =223)
Libben (n =94)
Nubia (n= 160)
London (n = 1730)
Ncw York (n =11959)
New Mexico (n = 792)
Rodeo (n= 181)
29.6
1.8
14.8
51.1
21.3
6.9
25.9
22.4
29.7
53.1
31.6
25.3
23.I
25.9
6.4
10.6
6.2
13.7
1.6
39.2
7.0
12.3
12.5
9.9
Pelvis
Leg
Foot
3.7
6.3
3.7
3.1
o.o
o.o
11.l
6.3
3.2
1.9
24.4
3.8
0.2
11.1
9.0
39.4
22.6
21.9
23.6
6.1
0.5
2.1
3.3
23.6
20.6
11.1
6.1
1.3
7.0
5.6
25.9
9.4
Aside from gross comparisons of fracture prevalence in human populations, there is surprisingly little information on temporal trends in recen!
humans. Angel (1974) attempted to document temporal trends by comparison of post-Mesolithic archaeological skeletons (n = 2125) and modern
117
Euroamerican samples. This comparison reveals severa! general characteristics of accidental injury: adult males have more injuries than adult
females; older adult females have more fractures than older adult males (see
also Buhr & Cooke, 1959); and recent populations have far more fractures
than earlier populations. The higher frequency of skeletal injury in adult
males than females in Angel's samples probably reflects a greater exposure
of mento trauma. The reversa! in older adults indicates the effects of bon e
loss, especially in postmenopausal women. Reduced bone mass predisposes an individual to fracture, especially in the femoral neck region, where
there is relatively little cortical bone to begin with (cf. Garn, 1970; Stini,
1990). Angel (1974) contends that the higher fracture prevalence in the
modern samples results from the hazards associated with twentieth century
technology and urban living, such as the reliance on automobiles, walking
on staircases, and urban crowding.
Other findings drawn from the study of skeletal series in the North
American Eastern Woodlands, Southwest, and Texas indicate a general
decrease in individuals affected by postcranial fracture in comparison of
preagricultural hunter-gatherers and later agriculturalists (Steinbock,
1976). These findings suggest that, in general, forager lifestyles may have
been more dangerous than agricultura! lifestyles. However, the huntergatherer sample is largely drawn from the Archaic period (4000--2000 BC)
ludian Knoll site, thus limiting comparisons to mostly a single locality. The
samples are reported by the ratio of number of fractures to number of
individuals, making it difficult to assess fracture patterns and prevalence.
Thus, the low prevalence in the later populations may simply reflect the
incompleteness of skeletons rather than the true prevalence of traumatic
injury.
Consisten! with Steinbock's (1976) assessment is a significan! decrease in
forearm trauma (from 4.0% to 0.6%) in a comparison of early prehistoric,
Archaic period foragers with late prehistoric maize agriculturalists in the
Tennessee River valley (Smith, 1990). These findings suggest that foragers
in this setting led a more hazardous lifestyle than did their agricultura!
descendants, at least with respect to the kinds of conditions resulting in
forearm trauma (Smith, 1990).
Other Southeastern prehistoricmaize agriculturalists show generally low
prevalence of accidental injuries. At Moundville, for example, the total
frequency of bones affected is only 0.7% (Powell, 1988). Many of these
fractures are associated with lower-status individuals; no high-status adult
females have accident-related traumatic injuries (Powell, 1988). Therefore,
at least in this setting, sedentary populations had a relatively accident-free
lifestyle, and high-status elite women may have been spared from activities
118
Intentional injury
equally affected by traumatic injury, indicating that the elevated rates were
due to accidents associated with activities such hunting forays and travel
(in adults) or play (in juveniles).
4.3
Age pattern.<
119
interaction.
120
Intentional injury
121
.1\
adaptive shifts.
4.3.J
American Midwest
Norris Far1ns
Figure 4.3. Cutmarks on adult frontal (scalping); Norris Farms, Illinois. (From
Milner & Smith, 1990; reproduced with permission of authors and Illinois
State Museum.)
122
Intentional injury
123
than males (n = 5) with cranial injuries suggests that women were the
preferred target ofviolence. For sorne ofthese women, violent encounters
may have occurred on more !han one occasion - five female crania have
multiple healed depressed fractures. One of these individuals has a severe
depressed fracture and an accompanying large incision on the occipital as
well as multiple fractures on the left and right parietals.
Although other women may have been responsible for the cranial
injuries in this population (cf. Burbank, 1994), the demographic characteristics of the injured group suggest that maleswere the primary aggressors.
Males and females show a very different age pattern of injury: the fema!e
peak age of trauma is the early adu!t years, male trauma is evenly
distributed across age groups. Ma!e trauma is oriented toward the front of
the vault, and fema!e trauma is present throughout the cranium. Wilkinson
& Van Wagenen (1993) suggest that violence directed at women by menor
women against women or co-wives in polygamous societies fits well with
ethnohistoric accounts of Eastern Wood!ands native popu!ations (Wilkinson & Van Wagenen, 1993).
4.3.2
At the late prehistoric (AD 1000-1300) Riviere aux Vase site in southern
Michigan, nonlethal cranial injuries consisting of round or elliptical vault
depressions produced by wood or stone clubs has been identified (Wilkinson & Van Wagenen, 1993). A higher frequency of adult females (n = 14)
Crow Creek
Study ofhuman remains from various late prehistoric siles in the Missouri
River valley located in the present states of South Dakota and North
Dakota reveals evidence for violen! confrontations between tribal groups
124
competing for overlapping resources and territory (e.g., Bovee & Owsley,
1994; Hollimon & Owsley, 1994). The proto-Arikara (ca. AD 1325) skeletal
series from the Crow Creek site supplies importan! details on prehistoric
conflict (Gregg & Gregg, 1987; Willey, 1990; Willey & Emerson, 1993;
Zimmerman et al., 1981 ). Analysis of sorne 500 individuals buried in a
single pit indicates that nearly ali members of a village were massacred
during the course of a single raid. The presence of carnivore tooth marks
and weathering reveals that, following the attack, the deceased were
exposed for a period of time prior to their burial by returning survivors or
allies (cf. Milner & Smith, 1989). Analysis of the human remains suggests
that although ali of the deceased were victims of a single attack, violence
had a well-established history in the Crow Creek villagers; a number of
massacre victims had healed violence-related injuries, including scalping
(Willey, 1990; Willey & Emerson, 1993).
Virtually ali individuals in the series have unhealed cutmarks on frontals
and other cranial elements, indicating removal of the scalp with a stone
knife al or following the time of death. Forty per cent of the victims ha ve
era nial depressed fractures, mostly located on the top and si des of vaults. In
addition to scalping trauma, various other mutilations are common,
including tooth evulsions, alveolar and tooth fracture, nose removal,
decapitation, and dismemberment of both upper and lower limbs. The
presence of cutmarks on mandibles - especially on ascending and/or
inferior borders of rami - suggests that mutilation also involved tangue
exc1s1ons.
Demographic assessment of the Crow Creek population indicates that
young adult (15-24 years) males outnumber young adult females by a
factor of two. Additionally, there are twice the number of older adult
(45-59 years) females than older adult males in the series. The absence of
young women may reflect captive taking, escape, or actual demographic
composition of the population from which victims are drawn (Willey &
Emerson, 1993). Similarly, the paucity of elderly men reflects their escape
or actual demographic composition. lt is unlikely that older males were
captured, because they would ha ve represented a relatively low economic
return for the raiders. Perhaps older males are missing because of earlier
raids and endemic warfare (Willey & Emerson, 1993).
Larson Village
During the historie period, the Arikara occupied a series of temporally
successive villages as the tribe migrated northward up the Missouri River
valley. One such villa ge was decimated by violence. At the seventeenth
lntentional injury
125
126
4.3.3
American Southeast
Koger's !stand
lntentional injury
127
15
z 10
--.
....- - .
"
10
20
30
40
50
60
Age (years)
1985). Adult ( > 15 years) males and females have very different age-at"
death profiles (Figure 4.4). Female deaths peak slightly during the late
teens and early twenties, progressively fall to a low point in the thirties, and
again rise in the forties. This pattern also characterizes other archaeological
series (e.g., Buikstra & Konigsberg, 1985). Male deaths are few in number
during the late teens, but high in the thirties. Presumably, many of these
deaths resulted from violen! intergroup encounters. The age composition
of the small burials (containing fewer than five individuals) is different from
the age composition ofthe multiple burials (five or more individuals). The
former is dominated by infants and younger juveniles. The latter has an
un usual peak in the thirties, and most of these deaths are adult males. The
pattern is distinctive in that there are relatively few women or children in
the multiple burials, suggesting that women and children had either
escaped or been captured; males may ha ve died while protecting the village
from aggressors:
The loss of adults, and especially men who were responsible for
protection of the group and acquisition of resources not acquired by
women (e.g., animal protein), would have had far-reaching consequences
for the population's ability to mitigate stress in a hostile setting. In later
prehistory, political systems and population size declined in this and sorne'
other regions of the American Southeast and Midwest (e.g., seeAnderson,
1994; Milner, 1990; Steponaitis, 1991). This hostile environment may have
128
129
(a)
(b)
(e)
()
(e)
Figure 4.5. Depressed fractures in adult crania: ellipsoidal parietal injury (a),
deep circular injury (b), multiple circular injuries (e), cllipsoidal occipital injury
(d), circular parietal injury (e; arroW points to residual fracture line); Santa
Barbara Channel islands, California; (From Walker, 1989; reproduced with
permission of author and John Wiley & Sons, Inc.)
130
40 years of age. This age-specific pattern suggests that adults not involved
in behaviors and activities resulting in cranial trauma had somewhat
greater longevity in comparison with victims (Lambert, 1994).
Most of the ihjured adults are males. In these individuals, roughly
two-thirds oftheinjuries occur on the left side ofthe frontal, indicating that
conflicts were face-to-face encounters with a right-handed perpetrator
striking the left side of the head of the victim (Lambert, 1994). This
patterning of nonlethal trauma is remarkably similar to trauma observed in
the Yanomamo foragers ofVenezuela (Chagnon, 1992). Yanomamo men
attack other males with heavy wooden clubs. Although numerous casualties result from these encounters, they are rarely lethal.
Adult female cranial injuries are haphazardly distributed on the face and
vault - only about one-third of the injuries are on the frontal bone
(Lambert, 1994). The random distribution of depressed fractures indica tes
that, although females were occasionally involved in face-to-face attacks,
most were from other directions, including from behind (e.g., while fleeing
an attacker). Additionally, sorne of these nonfrontal injuries could be from
accidental causes (Walker, 1989).
The presence of projectile injuries and associated projectile points in
sorne individuals reveals appreciable numbers of deaths ca u sed by violen ce:
of 1744 individuals, 3.3% had been killed or wounded by single or multiple
projectile injuries. Unlike the cranial trauma, the majority of projectile
victims (at least 71%) died from their wounds, indicating the lethal
intentions of the attacker. The demographic composition of individuals
with either lethal (projectile wounds) or nonlethal (cranial depressed
fractures) injuries is similar in at least two respects: first, young and mature
adult males are the most affected; and second, children, older adults, and
women are the least affected. Like cranial trauma, aggression leading to
injury and death involved primarily adult males under 40 years of age
(Lambert, 1997), which is consistent with patterns observed for many
nonindustrial communities globally (e.g., Chagnon, 1992; Meggitt, 1977;
and see discussion in Lambert, 1994).
The temporal patterns of nonlethal and lethal skeletal injury are
distinctive in prehistoric southern California coastal populations. Nonlethal cranial depressed fractures are common in the Early Middle period
(1490 BC-AD 580), whereas lethal projectile injuries are far more common in
the Late Middle period (AD 580-1380)(Lambert, 1994, 1997). The increase
in frequency of projectile injuries may be tied to the adoption of the
bow-and-arrow in California during the sixth century AD (Lambert &
Walker, 1991). Perhaps the rapid adoption of the bow-and-arrow at this
time may ha ve been fostered by competition and conflict between popula-
Intentional injury
131
tions in North America generally (Blitz, 1988). Regardless of the motivation for increased lethal violence, the later peak fa projectile injuries
signifies a shift to more serious - and deadlier - forms of conflict in Iater
prehistoric southern California populations.
The availability of abundant bioarchaeological, archaeological, and
climatological data provides a more comprehensive understanding of
factors that may have motivated violent behavior in the Santa Barbara
Channel region. Analysis of tree ring and other climatological data
suggests that the .Middle period saw an increase in environmental instability, periodic droughts, and decreased terrestrial resource productivity.
These changes, coupled with warming of the Pacific Ocean during this time,
reduced marine productivity (Lambert, 1994, 1997). In light of these
changes, Lambert ( 1994) speculates that elevated competition for increased
resource stress may have engendered an increase in violence in these
populations. This hypothesis is consistent with other skeletal data showing
a decline in quality oflife and increase in health stress (e.g., Lambert, 1993,
1994; Walker, 1986a; Walker & Lambert, 1989; see Chapters 2 and 3).
Along with the increase in trauma and disease during this time, there is
evidence for increasing social complexity. For example, the burial of
high-status grave goods (e.g., beads) with infants is more common during
the peak in drought in later prehistory, suggesting ascribed rather than
achieved social rank. Perhaps increased intergroup competition '-' and;
hence, increased violence - for scarce resources during episodes of
environmental degradation fostered more complex social organization in
Iater prehistory (Fischman, 1996).
4.3.5
The Arctic
Like California, native groups living in the Arctic are often perceived as
living in a state of quiet repose and nonviolence (see Fienup-Riordan,
1994). Early documentation of Eskimos often presents them as 'passive to
the point oflethargy' (Fienup-Riordan, 1994:322). Severa! bioarchaeological investigations are providing new data that indicate the need for revision
of these perceptions. Findings from these studies suggest that, although
violence may have been rare, it erupted on occasion and had dire
consequences for sorne groups.
Saunaktuk
Study of human remains from the Saunaktuk site located east of the
Mackenzie Delta inthe CanadianNorthwest Territories provides compell-
132
Intentional injury
ing evidence for violen! confrontation between native groups (Melbye &
Fairgrieve, 1994; Walker, 1990). Human remains from this locality
represen! a minimum of 35 Inuit Eskimo villagers, sorne of whom died
violently in the !'ate fourteenth century AD. Evidence for violen! death and
body treatment in this Arctic setting is indicated by extensive perimortem
skeletal modifications, including knife cuts, slashing, piercing, gouging,
and splitting of long bones (Melbye & Fairgrieve, 1994). None of the
remains had been purposefully buried. Most of the victims are juveniles
(68.6%), suggesting that adults- particularly males - may have been away
hunting, leaving a relatively defenseless group vulnerable to attack.
Hundreds of knife cuts, especially around articular joints and the neck
(e.g., occipital condyles and upper cervical vertebrae), indica te the practice
of dismemberment and decapitation. Numerous cuts on facial bones on
most victims identify widespread facial mutilation or disfigurement. Many
other cuts, such as on clavicles and scapulae, reflect an overall pattern
associated with purposeful dismemberment, removal of muscle and other
soft tissues, and intentional mutilation. Unique to the Saunaktuk skeletal
series is the presence of gouges at the ends of long bones. Adult distal
femara from two individuals display large perimortem mediolateral gouges
passing completely through the cortical and cancellous tissue. These
modifications are consisten! with oral tradition describing a type of torture
whereby the victim's knees were pierced and the individual was dragged
around the village by a cord passed through these perforations.
With few exceptions, long bones had been split in to seores of longitudinal sections. The surfaces of split bones are smooth and display tiny step
fractures identical to borre modifications in butchered animal bones.
Presumably, this breakage pattern was produced by extraction of the
nutritionally rich marrow for consumption (Melbye & Fairgrieve, 1994; cf.
White, 1992). The striking similarity between butchered animal remains
found in archaeological siles and the Saunaktuk human skeletal assemblage suggests that the deceased had been cannibalized. In sum, at least
sorne members of this late prehistoric Saunaktuk population had been
tortured, and ali had been murdered, mutilated, and cannibalized.
There is a rich historical record that provides a context for understanding violence between native groups living in this region of the Arctic. In
areas where Inuit and Amerindians carne into frequent contact - such as
the Mackenzie Delta region - violen! interactions between the groups were
commonplace. Oral traditions and historical accounts describe the horrific
nature of intergroup violence. For example, Samuel Hearne, who explored
the region in the late eighteenth century for the Hudson Bay Company,
observed the murder and mutilation of Inuit villagers by a group of
133
4.3.6
The Anasazi were one of severa! complexes oflate prehistoric societies who
were ancestral to sorne of the modern native populations curren ti y living in
the 'Four Corners' region (present-day states ofUtah, Colorado,Arizona,
and New Mexico) of the American Southwest. The region has been the
focus of intensive archaeological investigation for more than a century,
producing an abundance of human remains. Most human remains are
intentional interments, ranging from isolated burials found in house floors,
trash and storage areas, and large cemeteries. Individuals are generally
singular and are accompanied by grave goods (Turner, 1993a).
A small - but highly visible - proportion of graves are multipleindividual interments containing many disarticulated and broken skeletal
134
Intentional injury
135
136
4.3.7
Ritualized violence and offering of remains- such as heads and other body
parts - as a mediator between the living, their ancestors, and the
supernatural world has been documented in a wide range of settings in
precontact Latin America (e.g., Benson & Boone, 1984; Carneiro, 1990;
Fowler, 1984; Redmond, 1994; Verano, 1995). Numerous iconographic
descriptions and ethnohistorical accounts provide information on sacrificial death and mutilation, and discoveries of missing heads and limbs in
archaeological settings provide verification for this behavior (e.g., Fowler,
1984). Little systematically collected skeletal evidence exists of these
activities (Verano, 1995). lmportant recent research in Peru and Mexico is
beginning to allay this shortcoming.
In coastal Peru, a series of mutilated human remains and contextual data
lntentional injury
137
138
with a single hole and the base of the skull shows damage, ranging from
foramen magnum enlargement to removal of most of the cranial base.
Although the reasons for these alterations are not known, the frontal
perforation probably supported a rope handle. The foramen magnum may
have been enlarged for removal of brain tissue, perhaps as a part of the
mortuary ritual.
In precontact Mexico, skeletal evidence has accrued on body processing,
including cutmarks, decapitation, dismemberment, burning, and intentional borre breakage in a limited number of different temporal and cultural
contexts (Pijoan & Mansilla Lory, 1990, 1997). As has been discussed for
the American Southwest, cutmarks and other cultural modifications of
skeletal remains by themselves do not provide information on the motivations for body processing (and see White, 1986). However, a rich
historical and iconographic record indicates that ritual was the key
motivation for cannibalism in early contact period Mexico.
ted by single and multiple interments ofmostly adults from the Temple of
the Feathered Serpent dating to the early occupation of the site and
construction ofthe temple (ca. AD 100-250) have been analyzed (Cabrera
Castro, 1993; Cabrera Castro et al., 1991; Serrano Snchez, 1993;
Sugiyama, 1989). Sacrificial victims associated with the temple number
sorne 120 individuals (Cabrera Castro, 1993). The skeletal remains have
not been systematically studied for cutmarks or other skeletal modifications, but the position of the hands behind the backs for sorne individuals
suggests that they had been bound at the time of death. The general lack of
disarticulation as well as the inclusion of co-mingled individuals in single
burial pits (e.g., burial 190 included 18 adult males in a linear pit at the
midpoint between the southeast and southwest corners of the temple)
suggest that death probably took place at one time for all individuals
(Sugiyama, 1989). The identity ofthe victims are not known. However, in
reference to one of the multiple burials on the margins of the temple (burial
190), the inclusion of numerous obsidian points and blades and other
offertory items suggests that they 'were military men, priestly soldiers, or
men disguised as military personages' (Sugiyama, 1989:98).
Ritualized violence, sacrifice, and related behaviors are well documented
in the iconography of numerous precontact Latin American cultures. Mass
burials, mutilation, decapitation, and the presence of trophy heads in
Intentional injury
139
4.3.8
Easter lsland
140
Jntentional injury
(247)
(122)
(138)
(132)
(92)
(133)
13.4
26.2
14.5
16.7
6.6
23.3
3.6
7.4
4.4
6.8
1.5
(151)
(83)
(123)
(51)
(62)
(86)
19.9
27.7
31. 7
33.3
24.2
32.6
4.0
8.4
10.6
11.8
9,7
10.5
2.4
4.9
5.9
4.8
3.5
Males
Central Murray
Rufus River
South Coast
Desert
Tropics
East Coast
3,8
Fen1ales
Central Murray
Rufus River
South Coast
Desert
Tropics
East Coast
,N
o
0,8
0.7
1.5
1.3
were embedded in the frontal and left parietal of a historie period adult.
The bone tissue surrounding the entry wounds is fully healed, suggesting
that the victim survived for a period of time following the attack. Most
other victims offirearms were probably notas fortunate as this individual.
In summary, the skeletal evidence of trauma on Easter Island is
consisten\ with folklore documenting endemic interpersonal conflict in the
sense that violence was frequent. Contrary to this record, bioarchaeological analysis indicates that violence resulted in frequent nonlethal injuries
rather than widespread death (Owsley et al., 1994). Additionally, the
practice of cannibalism is not confirmed by study of skeletal remains.
Therefore, the folklore of Eastern Islanders overemphasizes warfare,
violence, and cannibalism.
4.3.9
Australian foragers
141
Labren & Bertyman, 1984; Owsley, 1994; Robb, 1997; Walker, 1989; and
many others; although see Wilkinson & Van Wagenen, 1993), which
probably reflects the central role of men in the violen! resolution of
conflicts in most human societies. This sex-specific pattern of head injury
contrasts with that in Australia. In virtually ali samples throughout the
continent, adult females show a consistently higher prevalence of cranial
injury than adult males, thus contributing to the greater prevalence in
females than males overall (Table 4.2). Sorne of the sex differences in
specific localities are slight, but many series show a remarkably strong
disparity between sexes. F or example, in the south coastal Swansport
sample, 39.6% (21/53) and 19.3% (11/57) of females and males exhibit
cranial trauma, respectively. For the few skeletal series (4/22) where males
have more cranial injuries than females, the differences are statistically
indistinguishable. The disparity in cranial injury between males and
females is not restricted to prevalence alone. F or virtually ali regions of
Australia - regardless of ecological or cultural setting - women show a
predominance of depressed fractures on right parietals and occipitals.
This pattern suggests that attacks carne from behind the victim, perhaps
while she was fleeing the attacker. Adult males show a different pattern of
injury location: for the entire series, more left parietals are fractured than
right parietals. This pattern suggests that male violence usually involved
facial confrontations.
It is not uncommon for an individual in the Australian samples to have
two, three, or even four cranial bones that display depressed fractures
(Table 4.2). Consisten! with the sex differences in prevalence of crania,
affected, more women than men have multiple injuries. The general
pattern, then, indicates that violence and aggression were directed more at
women than at men in prehistoric foraging societies throughout Australia.
Violence was not limited to prehistoric Australian Aboriginal societies.
Ethnographers observe that violence is a common occurrence anda part of
everyday discourse (Burbank, 1994). Unlike Western societies, such as in
the United States, where fighting - and especially aggression against
women - is viewed as a deviant behavjor, physical aggression in native
Australians is considered an accepted if not legitimate form of social
interaction (e.g., Burbank, 1994; Myers, 1986). Burbank (1994) provided
detailed observations on physical aggression in men and women in an
aboriginal group living in Arnhem Land (Northern Territory). Her study
shows that both men and women were heavily involved in confrontations.
However, the majority of aggressors and their victims are adult females.
These observations of both deceased and living native Australians revea! a
striking consistency in behavior between prehistoric and contemporary
142
4.3.10
The historical record ofviolence and warfare is abundan\ for northern and
western Europe. Systematic studies of violence have been produced for
severa! areas of northern Europe, including Scandinavia, and especially
prehistoricand early historie Denmark. Analysis ofhuman remains reveals
evidence of trauma tic injury, decapitation, and mutilation. Like much of
the history ofpaleopathology, these studies are largely descriptive, having
focussed on single ora few individuals (Bennike, 1987, 1991).
Relying primarily on remains dating from the Mesolithic (ca. 83004200 se) to the Middle Ages (to AD 1536), Bennike (1985) identified
patterns of injury in Denmark. These patterns can be characterized as
involving a predominance of cranial trauma in mostly adult males
(mostly depressed fractures) on anterior cranial vaults, indicating face-toface violent interactions. Folklore and historical accounts emphasize the
high prevalence of violence during the Viking period (AD 800--1050).
Bennike's (1985) assessment clearly indicates that the Mesolithic and
Neolithic periods were far more violent than the Viking period: Mesolithic crania display the highest prevalence of cranial trauma (43.8%),
which markedly declines in the Neolithic (9.4%), !ron Age (4.7%), Viking
period (4.3%), and Middle Ages (5.1%). Violence is well illustrated by the
presence of projectile injuries, sword and axe cuts, cranial depressed
fractures, and decapitation (Bennike, 1985; Ebbesen, 1993; Kannegaard
Nielsen & Brinch Petersen, 1993). At the Mesolithic site of Bogebakken, a
borre projectile was found lodged between the second and third thoracic
vertebrae of an individual. In fact, all projectile wounds at this and many
other Danish sites are f9und in the thoracic and head regions, revealing
the lethal intentions of the attackers.
In interments dating to the Middle Ages, the heads ofvictims had been
removed and placed between their legs. The reasons for this unusual
treatment are unclear, but during the Middle Ages the practice was
associated with criminals in order to prevent their return following death
(Bennike, 1985; and see below). Decapitation and other forms ofhead and
neck trauma were probably more common than is indicated by the skeletal
evidence alone. A number of Neolithic and !ron Age bog corpses show
evidence of decapitation and strangulation; the latter may have involved
lntentional irijury
143
hanging (e.g., Bennike, 1985; Glob, 1971). Owing to the relatively small
number of projectile- and weapon-related deaths, it is not possible to
identify a pattern of decrease or increase in violence-related death in
Denmark (see Bennike, 1985). However, there is a shift from use of
projectile weapons to axes and swords in the Iron Age (Bennike, 1985). The
lethal nature of this new weaponry for the enemies of Danes is demonstrated in at least one battle site (see below).
Violence in western Europe during the Mesolithic may have been highly
regionalized, with a relatively high prevalence in sorne regions (e.g.,
Denmark) but not in others. For western Europe as a whole, violen!
trauma was relatively infrequent during the Mesolithic (ConstandseWestermann & Newell, 1984). An increase in population density, reduced
territories occupied by ethnic groups, increase in social complexity, and
resource circumscription during the Mesolithic suggests the potential for
an increase in hostilities. However, trauma prevalence does not change in
comparison to the succession of periods during this time frame (Constandse-Westermann & Newell, 1984).
Battle o/ Wisby
144
wounds are variable, ranging from scratches and nicks on individual bones
to dismemberment. The latter, for example, is illustrated by the presence of
severed hands and feet, partial limbs, and complete limbs. One individual
expresses the intensity of battle: the lower halves of both left and right
tibiae and fibulae are completely severed. The lower legs are affected more
than any other area of the body: about two-thirds (65%) of cutting trauma
involved tibiae. Ingelmark observed that the focus on the lower limbs
probably reftects the use of shields and protective clothing for the body
trunk, leaving the legs especially vulnerable to injury. Sword blows directed
at the lower legs typically resulted in the slicing and chipping of bone on the
tibia anterior crest.
Poor protection of heads of individuals from the Gotlander army is
indicated by the large number of cranial injuries, sorne of which involve
extremely deep cuis. The heads of sorne groups of Gotlander soldiers may
ha ve been better protected than others. For example, only 5.4% of crania
are injured in common grave no. 3; this frequency contrasts with the
prevalence of cranial injuries of 42.3o/o and 52.3o/c, in common graves no. 1
and no. 2, respectively. The majority of cranial wounds are on the left side
of the head - this fits the expected pattern of injury sustained by a weapon
held by a right-handed individual during a face-to-face encounter (and see
Courville, 1965). Sorne crania have injuries on posterior vaults, suggesting that these victims were struck from behind while fteeing their attackers.
The presence of ali ages of males suggests that the majority of the male
population in and around Wisby were recruited for the defense of the city.
Analysis of pathological conditions suggests that virtually anyone who
could walk (and even those who could not) were drafted. Ingelmark
(1939: 192) remarked on the 'good many morbid processes' present in the
skeletal assemblage of battle victims. Many vertebrae have pronounced
osteoarthritis, and al leas! four individuals have extensive vertebral
tuberculosis. One individual displays a completely ankylosed (fused)
knee. The angle of ftexion (about 55) greatly disabled the individual:
running was an impossibility for this victim. A number of individuals
show wellhealed femoral neck fractures which limited their ambulatory
capabilities. These observations, combined with other health problems :including skeletal infections and numerous healed, but poorly aligned
limb fractures (n = 39) - also contributed to reduction in efficiency on the
battlefield. The defending army, then, was not composed of a group of
robust males who were in their peak years of fighting prowess. Like many
of the skeletal samples discussed in this chapter from both New and Old
World contexts, these victims of the massacre were members of a
Intentional injury
145
population not unfamiliar with violence during their lifetimes (cf. Milner
et al., 1991; Willey, 1990). A number ofbattle victimshave healed cranial
trauma (e.g., depressed fractures).
Beheading in ancient Britain
146
Intentional injury
147
1993). The narrowness of the cuts indicates that the decapitation was
completed with a narrow blade administered as blows to the neck.
4.3.11
When Europeans began exploration of the New World in the late fifteenth
century, they brought with them a weapons technology that facilitated
their conques! of native populations. The early expeditions were violen!
affairs at times, resulting in brutal treatment ofnatives (see Weber, 1992).
Although these tactics seem repulsive now, they were well .within the
bounds ofbehavior for European males during the Middle Ages. Historical
literature and accounts of violen! confrontations (e.g., Wisby) indicate that
con:flict behavior between European males was excessively cruel, at least at
times (see also Weber, 1992). The study of native remains dating to the
early period of contact with Europeans has provided a new dimension to
understanding the nature of the interactions between these groups.
(a)
La Florida
(b)
Figure 4.7. Cut adult scapula (a) and humerus (b); Tatham Moun.d, Florida.
(From Hutchinson & Norr, 1994; reproduced with permission ofauthors and
Wiley-Liss, Inc., a division of John Wiley & Sons, Inc.)
148
Intentional injury
American South1vest
Like many regions discussed in this chapter, relatively little is known about
violence and conflict in the American Southwest prior to or at the time of
contact (see Stodder & Martn, 1992). The presence of fortifications and
other defensive architecture and high frequencies of traumatic injury
(including injury resulting from violence) in sorne later prehistoric siles
suggests that confrontations were frequent (e.g., Stodder & Martn, 1992).
Cranial trauma in the American Southwest showsan increase in prevalence
during the late prehistoric period, which continued into the historie,
mission period (Stodder, 1990, 1994; and see Stodder & Martn, 1992).
Archaeological and historical evidence shows that the high frequency of
cranial trauma in the historie period can be attributed to confrontation
between Spaniards and Indians as well" as among Pueblos and between
Pueblos and non-Pueblo native groups. Study of skeletal remains reveals
that most cranial injuries are in adult males (Stodder, 1994). At San
Cristobal and Hawikku, sites with significan! contact period skeletal
assemblages, very high frequencies of cranial injuries have been reported
(Stodder, 1994). Twenty per cent and 17% of males have cranial trauma
from the San Cristobal and Hawikku sites, respectively. Paleopatholog1cal
markers of stress (e.g., dental defects) indica te that physiological perturbations were generally high during the late prehistoric and contact periods;
this may have contributed to fostering intra- and intergroup hostilities
during this time (Stodder, 1994).
1
1
1
4.3.12
149
During the North American French and Indian War (called the Seven
Years War in Europe), France and Great Britain fought overcontrol ofthe
vast territories of the Northeast and Canada. During the summer of 1757,
the British garrison surrendered Fort William Henry at the southern end of
Lake George, New York, to French and Canadian troops and their Native
Americans allies (Starbuck, 1993). As part of the conditions of surrender,
British soldiers and dependents were allowed to leave the fort and return to
British-controlled territory under French protection. The French-ally
Native American warriors felt slighted that scalps and other prizes of
warfare would not be forthcoming. In retaliation, the French-ally lndians
killed the remaining British troops at the fort. Warriors then proceeded to
kili or capture the hundreds of civilians and soldiers under the care of the
French troops.
Analysis of the remains of five adult males buried in a mass grave within
the fort indicate clear evidence of violence-related injuries, bearing testimony to the historical and fictionalized (e.g., Cooper, 1919) accounts ofthe
battle (Liston & Baker, 1996). Four ofthe five show premortem trauma to
the legs, refiecting injuries received duringthe siege ofthe fort. None ofthe
injuries are healed, suggesting that they died prior to or during the siege.
The trauma was not lethal, but serious enough to preven! their departure
from the fort. These skeletons display a range of perimortem trauma that
probably represents injuries resulting in death. One individual shows a
series of four cutmarks on the posterior surface of the odontoid process of
the second cervical vertebra. The pattern of trauma suggests that the
soldier had been beheaded from behind. Another individual expresses a
series of radiating fractures through the face and frontal, indicating a
crushing ofthe skull with a blunt object. Ali five individuals show notches,
slices, and gashes in skeletal elements of the anterior and posterior trunk
(e.g., scapula, ribs) and pubic region. Themorphology of cutmarks evinces
the use of both knives and axes in the mutilation of victims.
Snake Hill
1
1
!
Sorne ofthe most vigorous fighting between British and Americans during
the War of 1812 took place in the frontierregion between Lake Ontarioand
150
Lake Erie along the Niagara River (Whitehorne, 1991). During the
four-month period in 1814 when American troops successfully captured
and held Fort Erie on the Canadian side of the river, heavy siege and
combat resulted in the deaths ofhundreds of soldiers from both British and
American armies. Archaeological excavations at the battle site of Snake
Hill, Ontario, resulted in the recovery of the skeletal remains of American
soldiers from burial and medical waste pits (Owsley et al., 1991; Thomas &
Williamson, 1991). Demographic assessment of the complete or nearly
complete skeletons (n = 26) indicates that most individuals were young
adult males, aged 15 to 29; seven soldiers were older than 30 years at death.
Half (50%) of the individuals in the sample had fractures caused by damage
from firearm projectiles. The general lack ofhealing in most cases indicates
that thcse wounds were usually fatal. The highest percentage of fractures
in volved ribs (28%; 7/25), followed by femara (25%; 7/28), and crania
(9.1%; 2/22).
Locational assessment of skeletal wounds indicates that most injuries
(69.8%) were above the waist. In regards to the total number ofnoncranial
and nonvertebral trauma (n = 53), twice the number of fractures occurred
on the left side (56.6%; 30/53) than on the right side (26.4%; 14/53) ofbattle
victims (excluding skull and vertebrae [17 .0%; 9/53]). This pattern may
reflect handedness or body postores during the battle (Owsley et al., 1991).
Cause of death is especially apparcnt for several victims. For example, a
young adult male died of a massive head injury in which a firearms
projectile had passed through the left and then right parietals. This
individual also had a large, completely healed cranial depressed fracture
from an earlier injury. Other individuals had fractured facial bones from a
firearms projectile and shattered long bones.
Battle of the Little Bighorn
In present-day South Dakota in June of 1876, General George Armstrong
Custer and 267 soldiers and civilians were overwhelmed and massacred by
a superior force ofNative Americans (see Scott & Fox, 1987). Reminiscent
of prehistoric and historie conflicts between nati ve groups in the region (cf.
Crow Creek, Larson Village site; and above), this battle was part of an
overall pattern of political domination and control of lands and resources
in the Great Plains by opposing groups.
Within two days of the battle, eyewitness accounts described mutilation
(including scalping) and dismemberment not unlike patterns observed in
other Plains samples (e.g., Crow Creek, Larson Village site; see above)
(Scott & Fox, 1987; Snow & Fitzpatrick, 1989). Temporary graves were
Intentional injury
151
hastily prepared at the locations where individuals were killed. Sorne ofthe
bodies were identified, but owing to decomposition and mutilation, many
were not. Skeletal fragments of battle victims from erosion and limited test
excavations provide the basis for detailed study of battle injuries (Snow &
Fitzpatrick, 1989).Analysis of375 partial and complete bones and 36 teeth
from a minimum of 34 individuals indicates three primary types of
perimortem trauma, including blunt-force trauma, cutmarks, and bullet
wounds (Snow & Fitzpatrick, 1989; Willey & Scott, 1996). Blunt-force
trauma involved massive fragmentation of crania, and, to a lesser extent,
postcranial elements. All 14 partial crania showed massive injuries due to
heavy blows. Additionally, the presence of cutmarks on various skeletal
elements indicates widespread perimortem mutilation. Severa! different
forms of cutmarks, ranging from fine incisions to pronounced incisions,
reflect the use of metal arrows or knives.
Cutmarks on a variety of skeletal elements indica te the high degree of
mutilation ofbattle victims. One individual, for example, has cutmarks on
a humerus head and sternum. The use ofheavy metal-edged weapons (e.g.,
hatchets) is clearly indicated in severa! instances. Far example, a completely transected cervical vertebra indicates decapitation by a single blowto the
neck. One individual shows distinctive sets of chopping blows to the proxi-
mal ends of the left and right femara indicating purposeful dismember"
ment.
In addition to traditional native weaponry, the presence of gunshot
wounds in six individualsindicates the useoffirearms by Native Americans
at the battle site (Willey & Scott, 1996). Individual M85, for example, had
at least two upper body gunshot wounds, including an entran ce wound on a
rib margin and shattered ribs from another wound. A third wound is
represented by a bullet or bullet fragment embedded in the distal left radius.
This individual also displays cutmarks on his clavicle. Three gunshot
wounds are located in the crania, one entering from the back and two
entering from the right side and exiting from the left.
In summary, based on the study ofhuman remains from the battle site, a
sequence of events can be reconstructed: namely, soldiers were wounded,
killed (frequently with blunt-force trauma to the skull), and mutilated
(Snow & Fitzpatrick, 1989). As would be expected, except for the use of
firearms, the pattern ofkilling and mutilation of vctims is strikingly similar
to that observed in other North American native populations from the
Great Plains and Midwest discussed in this chapter (e.g., Crow Creek,
Larson Village site, Norris Farms). A consideration of the direction of
entry wounds is consistent with historical records indicating that the battle
was chaotic (Willey & Scott, 1996).
152
4.4
153
154
crania from North America are associated with cranial injury (e.g., Ortner
& Putschar, 1985; Stone & Miles, 1990).
Evidence for treatment of dental disease has been identified in the form
of alterations ofteeth, namely drilling in tooth roots (e.g., Bennike, 1985;
Schwartz et al., 1995) and crowns (e.g., Koritzer, 1968). These boles are
usually found in association with carious or otherwise diseased teeth,
indicating a therapeutic intention.
. . .
In summary, the study of samples worldwide reveals that mJunes
sustained either by accidental means (e.g., limb fractures) or during violent
confrontations (e.g., cranial depressed fractures) were treated in sorne
earlier societies. These findings indicate that many past societies were
aware that proper restoration of function could be brought about only by
appropriate treatment protocols (see Roberts, 1988).
4.5
155
156
ments, demographic, cultural and social infl.uences, and individual behavior. Therefore, although these apparent similarities are informative, it is
importan! to identify specific causal factors in specific settings before
making conc,lusions regarding the relationship between population distribution and injury mortality in humans generally.
In sorne regions, clear patterns of violence have begun to emerge. For
example, an increasingly hostile landscape in the Eastern Woodlands
generally is corroborated by the archaeological evidence of increase in
defensive construction in later prehistory (e.g., Milner et al., 1991 ). Similar
patterns of increasing conflict and aggression in the final centuries before
European conques! are indicated in the southern California Santa Barbara
Channel islands, the American Southwest, Great Plains, and Arctic. These
regional investigations suggest that resource productivity and environmental stability may have hada strong influence on the presence or degree
of conflict in the past. Similar patterns of increase in evidence of violence
are also documented in 01d World settings (e.g., Kennedy, 1994).
Overall, bioarchaeological studies indica te that violene and conflict are
not random events, but are strongly influenced by extrinsic factors, such as
resource depletion. Dietary deprivation may have been a motivation for
cannibalism. Ca_nnibalism is poorly understood in archaeological settings
where historical -evidence is limited, such as in the prehistoric American
Southwest. Historical records in a number ofsettings are informative. For
example, cannibalism may have been symptomatic of a larger pattern of
animosity and aggression between groups (e.g., Saunaktuk) ora key part of
ri.tualized violence (e.g., Teotihuacan).
Study of trauma in skeletal remains reveals that the areas of the body
attacked are highly patterned in comparisons of different societies. Walker
(1997b) observed that in modero industrial Western societies (e.g., United
States) the head and neck are highly favored targets ofattack, probably for
both strategic and symbolic reasons. He argues that the face is an appealing
target, because the injuries are especially painful. The face and head
generally bleed profusely and bruise. easily, and this may symbolize the
aggressor's dominance (Walker, 1997b). This probably explains why the
most highly traumatized focal point of the body in recent urban populations is the nose, followed by the zygoma and the mandible (e.g., Hussain et
al., 1994)..
Many past societies show a penchant for head injury, but these injuries
are usually directed at the vault and not the face or dentition. Dental
fractures are present in archaeological remains, but they are rare (Alexandersen, 1967; see Leigh, 1929; Lukacs & Hemphill, 1990; al so J. R. Lukacs,
unpublished manuscript, for regional studies documenting dental trauma).
11
i
i!
1
157
The location of the injury on the body provides insight into sorne of the
details of conflict between the individuals involved. For example, many
cranial injuries are found on the left side of the frontal or other anterior
elements, indicatingthat a right-handed attacker successfully engaged their
weapon while facing their victim (e.g., native Australian males). A more
haphazard pattern of cranial injury (e.g., prehistoric Michigan) or higher
frequency of trauma on the right side or posterior vault indicates that
injuries were sustained while the victim was fleeing their attacker or
perhaps while lying prone (e.g., Wisby). This pattern is more common in
women than in men in sorne settings (e.g., Australia, Michigan), suggesting
that aggression was directed at women. Ethnographic evidence reveals
that, although the aggressor was often an adult male, attack by adult
females on other females (and on males) occurred with no small amount of
frequency in sorne settings (Burbank, 1994).
Historical documentation suggests that children have long been a target
ofviolent injury and death. deMause (1974), for example, regarded child
abuse as widespread in Europe prior to the eighteenth century. Nevertheless, examination of thousands of archaeological skeletons reveals no
evidence of skeletal trauma - localized trauma-induced subperiosteal
lesions in multiple stages of healing, and perimortem fracture - associated
with battered-child syndrome (Walker, 1997a; Walker et al., 1997).
Certainly, juveniles in earlier societies were victims of homicide and
violence (e.g., Crow Creek, Norris Farms). Juvenile skeletons, however;
lack the injuries associated with long-term abuse. This suggests that, like
the pattern of facial injuries in twentieth century Western societies, child
abuse resulting in severe skeletal trauma is primarily a modero phenomenon. Walker (1997a) suggests that the rise of childhood abuse is dueto the
loss of social controls of violen! behavior in recen! urban settings.
Technological factors are importan! in interpreting patterns and types of
skeletal injuries. The introduction of the bow-and-arrow is linked with an
increase in lethal conflict (e.g., southern California). Prior to the invention
offirearms, violence-related injuries were caused primarily by projectiles,
cutting, and blunt force.
Study of a number of skeletal samples possessing both lethal and
nonlethal forms of trauma provides insight into the previous history of
interpersonal aggression both within and between past societies. The
Wisby, Crow Creek, and Norris Farms victims display numerous healed
injuries (e.g., cranial depressed fractures) that reflect a long and well
established history of conflict well prior to the event or events resulting in
widespread death. In a sense, then, these injuries foreshadowed a future act
(e.g., a major battle) that later resulted in more widespread violence.
158
159
of the victim was sometimes placed between the legs (Denmark, England),
perhaps as the ultimate insult. Other unique and highly localized forms of
body treatment ofthe living victim were probably'practiced. For example,
gouging of the knees at Saunaktuk in the Arctic may have been associated
with a practice of dragging the victim through the village prior to their
death.
Traumatic injury data are importan! for dispelling prejudices and
assumptions about past societies. For example, hunter-gatherer societies
around the world are often characterized as peaceful inhabitants of
stress-free environments living in a state of blissful repose (e.g., Lee &
DeVore, 1968; Service, 1966; and see discussions by Burbank, 1994;
Fienup-Riordan, 1994). This characterization may reflect the fact that
anthropologists doing fieldwork among these societies are guests. What
guest is going to go back home and write about the unpleasant things they
observed, especially with regard to violen! encounters between individuals
(and see Keeley, 1996; Erchak, 1996)? Many anthropologists underplay the
negative or offensive, avoiding realism in social beliefs. In fact, a number of
cnltures described as nonviolent or peaceful have homicide rates far
exceeding !hose of sorne Western nations (Knauft, 1987). Evidence drawn
from the study of prehistoric remains from a number of settings presents a
very different picture, both about foragers and about other types of
societies. The point is not to replace peaceful characterizations of earlier
societies with violen! ones. Rather, these findings underscore the importance of substituting an incorrect image with one that fits the evidence. This
approach is critical for informing our perspective on past groups as
functioning societies rather than as images of what earlier social behavior
mus! have been like. This new-found precision adds a more complete
historical context for the study ofrecen! human behavior. Anthropologists
and others seem to employ - either consciously or unconsciously - their
own cultural and social assumptions about earlier societies in order to
'remember' the past. We project these assumptions into a past that seems to
refiect curren! and highly biased perspectives on the condition of humankind, be they peaceful or violen!. These skeletal data help us to reconstruct
and interpret trauma and violence in a more comprehensive and accurate
manner.
4.6
160
behavior in earlier societies. Human populations living in difficult circumstances have an elevated prevalence of skeletal injuries due to accidental
causes. The skeletal record of conftict in the past is highly visible in a
number ofpopulations worldwide. Study of remains from a wide variety of
contexts helps to provide better understanding of the circumstances of
violence whether dueto intra- or intergroup confticts. Sorne conftict may
result in' cannibalism, but, from the study of human remains alone, it is
difficult to identify the cause ofthis practice. Ritualized violence, including
sacrifice and elaborate body treatment, is also highly visible in sorne
archaeological settings. Although limited in scope, its study provides an
importan! link between culture and treatment ofthe body, both in life and
in death. Regardless of the circumstances of death, contextual data are
essential for its interpretation, including resource availability, the history
of intra- and intergroup social relationships, and weapons technology.
5.1
Introduction
162
5.3.1
Osteoarthritis
Osteoarthritis is a multifactorial disorder representing a pattern of responses to various predisposing factors (Hoffman, 1993; Hough &
163
Sokoloff, l 989; Rogers & Waldron, 1995). Epidemiologists and anthropologists observe a great <leal of worldwide variation in osteoarthritis in
relation to age. For example, young adults and older juveniles in sorne
human populations express a relatively high frequency of the condition
(e.g., Chapman, 1972; Chesterman, 1983; Larsen, Ruff et al., 1995). In
urbanized industrial societies, osteoarthritis rarely occurs before age 30
(Dekker et al., 1992).
There is sorne variation in osteoarthritis prevalence and severity with
climate (see Moskowitz, 1989); symptoms appear to be somewhat less
severe in warm climates, and this may be related to temperature, sun
exposure, and amount of clothing worn (Moskowitz, 1989). Body weight
can also be an importan! factor. Clinical and epidemiological studies
indicate a greater incidence of osteoarthritis in obese individuals, especially
in the weight-bearingjoints (e.g., Kellgren, 1961; Kellgren & Lawrence,
1958; Leach et al., 1973; Moskowitz, 1989). Obese women have rn
especially high incidence of knee osteoarthritis, which is related to adde-d
mechanical stresses on this major weight-bearingjoint (Cooper et al., 1994;
Moskowitz, 1989; Spector et al., 1994).
Jurmain (1977a) documents an increase in female osteoarthritis in later
middle-age, more so than in adult males of the same age. In twentieth
century United States groups, more women than men have osteoarthritis
after age 55 (Roberts & Burch, 1966). Similarly, the disorder is more severe
and more generalized in women than in men in Great Britain (Kellgren et
al., 1963). These studies suggest that there may be a hormonal influence in
osteoarthritis. Other factors such as metabolism, nutrition, bone density,
vascular deficiencies, infection, trauma, and heredity can influence the
disorder (see Duncan, 1979; Engel; 1968; Jurmain, 1977a; McCarty &
Koopman, 1993; Moskowitz, 1989; Ortner & Putschar, 1985; Rogers &
Waldron, 1995).
The primary contributing factor to osteoarthritis is mechanical stress
and physical.activity (Duncan, 1979; Hough & Sokoloff, 1989; Jurmain,
1977a, 1977b; McKeag, 1992; Nuki, 1980; Peyron, 1986; Radin, 1982,
1983; Radin et al., 1972). Asan individual ages, a stressedjoint or joints will
probably exhibit the biological concomitants of osteoarthritis. Radin
( 1982:20) defined osteoarthritis as 'the result of a physiological imbalance
between the mechanical stress in the joint tissue and the ability of the joint
tissues to withstand ... stress'.
The mechanical stress argument is supported by various findings. For
example, industrial laborers show patterns of articular degeneration in
relation to particular physical activities in the workplace. Strenuous lifting
by miners causes articular change in the hips, knees, and vertebrae
164
(Anderson et al., 1962; Kellgren & Lawrence, 1958; Lawrence, 1977); use of
pneumatic tools by ship builders and others results in similar modifications
(e.g., Lawrence, 1955, 1961); lifting of long tongs to move hot metals by
foundry workers results in degenerative changes in the elbow (Hough &
Sokoloff, 1989); and repetitive activity involving the hands in cotton mili
workers results in different patterns of osteoarthritis (Hadler, 1977; Hadler
et al., 1978; and see Merbs, 1983). Other findings for manual laborers,
farmers, ballet dancers, various types of athletes, and !hose who engage in
rigorous exercise generally support these observations (see Cooper et al.,
1994; Croft, Coggon et al., 1992; Croft, Cooper et al., 1992; Forsberg &
Nilsson, 1992; Jurmain, 1977a; Lawrence, 1977; McKeag, 1992;
Nakamura et al., 1993; Stenlund, 1993). New epidemiological findings are
providing importan! corroboration for these conclusions linking mechanical demand and osteoarthritis. Comparisons revea! markedly higher
prevalence of knee and hip osteoarthritis in North Carolina rural populations than the U.S. (primarily urban) population as a whole (hip: 25.1%vs.
2.7% in the 55-64 age cohort; Jordan et al., 1995). These differences reflect
the greater physical demands of the rural lifestyle in the United Sta tes.
The linkages between physical activity and osteoarthritis are not
straightforward. The hand bones ofweavers from the Spitalfields, London,
skeletal series have no more osteoarthritis than hand bones from the
general sample (Waldron, 1994). Manual laborers in this series have no
more or less osteoarthritis than the population as a whole. These findings
anda survey of inconsistencies found in the epidemiological literature led
Waldron to conclude 'that there is no convincing evidence of a consistent
possible.
Epidemiological studies provide an importan! baseline for interpreting
osteoarthritis in past human populations. Bioarchaeological and epidemiological studies are not strictly comparable, as the !alter are almos!
always based on clinical contexts, either radiological examinations or
palien! interviews, which do not identify subtle degenerative changes seen
in actual skeletal specimens that bioarchaeologists study. Hard tissue
changes observed in the clinical setting are not strictly comparable to those
observed in archaeological or other types of skeletal collections.
The pathophysiology of osteoarthritis is poorly understood regarding
the relationship between hyaline cartilage and borre changes. Sorne have
165
166
167
5.3.2
168
169
Sadlermiut Eskimos
170
Pecos
Black
White
Joint
Males
Knee
Hip
Shoulder
Elbow
27.0
51.0
47.3
12.5
3.0
2.9
1.1
5.8
38.2
47.3
50.9
19.8
4.5
1.8
3.8
5.2
29.3
20.7
33.3
11.4
1.7
2.3
1.5
3.8
32.4
35.2
53.6
31.1
18.0
35.6
37.4
44.3
12.7
10.9
13.1
8.2
1.0
31.9
47.8
53.6
21.7
18.6
7.8
8.9
0.9
16.I
20.7
22.2
I0.4
o.o
o.o
o.o
32.0
22.4
23.I
22.0
4.0
1.7
2.6
7.3
Fema/es
Knee
Hip
Shoulder
Elbow
3.0
13.5
2.8
o.o
171
demands for the Pecos Pueblo agriculturalists may be mostly limited to the
growing season, whereas Eskimos are subjected to high levels of activity
throughout the entire year (Jurmain, 1977a; see also Merbs, 1983).
Assessment of osteoarthritis prevalence in prehistoric adult males and
females from the American Great Basin contributes to an ongoing debate
about workload in harsh environmental settings (Larsen, Ruff et al., 1995).
Archaeologists suggest two alternative models for characterizing native
subsistence strategies in the Great Basin (Thomas, 1985). One model states
that prehistoric native populations pursued a limnosedentary exploitive
strategy whereby food and other resources were obtained primarily in
ecologically rich circumscribed wetland areas that punctuated the desert
landscape, thus resulting in a sedentary lifeway. Alternatively, the limnomobile model contends that these wetlands do not provide sufficient
resources for the support ofnative populations, at least on a full-time basis.
These wetlands are subject to occasional resource crashes arising from
droughts and floods. From this point ofview, native populations relied in
part on marsh resources, but spent significan! amounts oftime in collection
and transport of foods recovered from upland settings in the nearby
mountains and elsewhere. The implications of the former model is that the
more sedentary wetlands-focussed adaptation involved less mechanical
stress than the nonsedentary lifestyle; the limnomobile model is built on the
premise that the carrying of supplies and long-distance travel was physically demanding, requiring strength and endurance (Larsen, Ruff et al., 1995).
In order to determine which of the two models best characterizes the
adaptive strategies of native populations in the Great Basin, Larsen and
coworkers (1995) assessed the pattern and prevalence of osteoarthritis in
prehistoric human remains recovered from the Stillwater Marsh region, a
large wetland area in western Nevada. Analysis of these remains revealed
an abundance of osteoarthritis. Most adults, including ali individuals over
age 30, have osteoarthritis in at least one, and usually multiple, articular
joints. Articular pathology for older adults involves severe proliferative
lipping on joint margins, eburnation, vertebral compression fractures, and
Schmorl's nodes. Contrary to expectations of the limnosedentary model,
these findings suggest that hunter-gatherers in this setting led extremely
demanding lives. The high prevalence of osteoarthritis suggests elevated
mechanical demand, such as in heavy lifting and carrying. These findings
also imply that prehistoric groups may not have been tethered to the marsh,
and they exploited a wide range of resources from both the marsh and the
surrounding uplands. Beyond concluding that the Great Basin lifeway was
physically demanding, however, it is not possible to state whether these
populations were sedentary or mobile from osteoarthritis evidence alone.
172
173
174
Iations from the Pickwick Basin, Alabama. She suggested that only one
arm and just the elbow joint are involved in the use of the atlatl (sensu
Angel, 1966b), whereas both left and right arms and the elbow and
shoulder joints of each are involved in the use of the bow-and-arrow. Thus,
the joints of the upper limb should show different distributions of
osteoarthritis reflecting either an atlatl pattern (unilateral, elbow) or
bow-and-arrow pattern (bilateral, elbow and shoulder). Because males in
most human societies are responsible for hunting, they should show a
higher prevalence of osteoarthritis than females. Furthermore, as expected,
early prehistoric males have a higher prevalence of elbow osteoarthritis
than late prehistoric males, a pattern that probably reflects the use of the
atlatl in the earlier group and the bow-and-arrow in the later group.
Contrary to expectations, both temporal groups display slight right
dominance of osteoarthritis. Early prehistoric Jemales ha ve the highest
frequency of the right-dominant elbow osteoarthritis. These findings
provide mixed support in this setting for the link between weapons use and
degenerative articular pathology.
The Angel and Bridges studies indicate that sorne groups using the atlatl
ha ve a distinctive pattern of elbow osteoarthritis (e.g., Eskimos), whereas
others do not (e.g., Pickwick Basin). These differences may reflect the
relative importance or intensity of specific activities (Bridges, 1990). For
example, traditional Eskimo diets are heavily dominated by mea!, and they
relied exclusively (or nearly so) on hunting over the course of the entire
year. Thus, their atlatl use was highly intensive. Early prehistoric lndians
living in the Pickwick Basin hada far more di verse di et, which was acquired
only partially by hunting. For much of the summer and spring, native
populations utilized riverine resources (e.g., fish) and various flood-plain
plants (e.g., edible seeds); hunting was practiced mostly during the winter.
Therefore, the very different pattern of elbow osteoarthritis in Tennessee
Indians cannot be attributed solely to use of the atlatl or the bow-andarrow. Rather, a range of activities probably contributed to the patterns of
upper limb osteoarthritis (Bridges, 1990).
In contras! to the pattern of right-side dominance of osteoarthritis in
upper limbs (e.g., Merbs, 1983; Webb, 1995), sorne groups display bilateral
symmetry. Elbow osteoarthritis in native populations from Chavez Pass,
Arizona, is highly prevalen! and bilaterally symmetric (Miller, 1985; Nagy
& Hawkey, 1993). In this setting, mechanical loading of both elbows while
processing maize with grinding implements - pushing manos against
metates with the hands - involves equal use of the left and right arms
(Miller, 1985; and see Merbs, 1980). In traditional Southwestern native
societies, females are responsible for this activity. Thus, the relatively
175
higher frequency in adult females in the Chavez Pass series reflects the role
of women in food preparation.
Horseback riding
The vertebral column has been studied in a large number of settings in the
Americas (summarized by Bridges, 1992) and elsewhere. For prehistoric
North America, these comparisons revea! a number of tendencies. First,
prevalence is always greatest in the articular region between the fifth and
sixth cervical vertebrae; second, there is a tendency for the lower thoracic to
be affected more than the upper thoracic vertebrae; third, the second to
fourth lumbar vertebrae usually show the greatest degree of marginal
lipping in comparison with other vertebrae; and finally, the region
176
177
pattern of increase in sexual dimorphism has been documented in prehistoric northwest Alabama (Bridges, 1991a). In this setting, differences in
osteoarthritis prevalence between Archaic period males and females are
not statisticaUy significan!, whereas later Mississippian period males have
more severe osteoarthritis than females (Bridges, 1991a). These patterns in
Georgia and Alabama do not specificalty define behaviors associated with
either sex, but they are suggestive of contrasting patterns of physical
activity (see also below). The presence of more significan! differences
between agriculturatist mates and females in both settings suggests the
possibitity that sex differences in tabor demands were greater in later than
ineartier prehistory. Simitarly,comparisons offoragers from Indian KnoU,
Kentucky, with maize agriculturatists from Averbuch, Tennessee, indica te
a different prevalence of osteoarthritis between adult males and females
(Pierce, 1987). For example, !odian Knotl males have a significantly greater
frequency of shoulder, hip, and knee osteoarthritis than femates; Averbuch
males have significantly greater osteoarthritis for the shoulder and hip, but
not the knee. These differences are suggestive of change in workload and
activity with the adoption of agriculture.
Untike males, agriculturatist females from the lower Ittinois River valley
have a higher prevalence of vertebral osteoarthritis than forager females
from the same region (Pickering, 1984). These differencesare especialty
pronounced in cervical vertebrae; this may be related to an increase in
mechanical demand in this region of the skeleton with the shift to
agriculture (Pickering, 1984).
Fahlstrom (1981) identified an unusually high prevatence and severity of
shoulder osteoarthritis in adult males in the Medieval skeletal series from
Westerhus, Sweden. Historical analysis ofthis population suggests that the
high frequency in males reflects work and activity practices that are
exclusive to men, including parrying in sword fighting, spear throwing,
timber cutting, and other activities associated with repetitive, heavy
toading of the shoulder joint (Fahlstrom, 1981).
Sorne analyses revea! no appreciable differences between males and
femates in archaeological settings. For example, males and females in the
Dickson Mounds, Itlinois, series show no differences in prevalence of
appendicular osteoarthritis (Goodman et al., 1984; Lalto, 1973). The
simitarity between sexes infers that mechanical loading of most articular
joints in this setting was broadly the same in adults regardless of sex, in
contras! to most other prehistoric Eastern Woodlands populations (cf.
Bridges, 1992).
Two clear trends emerge when sex differences are examined (e:g.;
Bridges, 1992). First, where there are statisticaUy significan! differences
178
between males and females, males show a higher prevalence of osteoarthritis than females. Second, in specific regions of the New World, maize
agriculturalists tend to display more sexual dimorphism in degenerative
pathology than foragers. This suggests a difference in behavior leading to
degeneration of articular joints in agriculturalists but not in earlier
foragers. The change in pattern of sexual dimorphism suggests that there
was a fundamental shift in the division of labor once agriculture was
adopted (and see Bridges, 1992).
5.3.4
Age variation
5.3.6
5.3.5
179
l1
1!
180
Males
Mississippian
Archaic
Joint
Shoulder
(n)
36.8
(19)
Elbow
Wrist
(n)
27.3
(22)
9.5
(21)
Hip
(n)
(20)
(n)
Knee
(n)
Ankle
(n)
Right
Left
42.l
(19)
40.9
(22)
15.8
(19)
5.0
(20)
31.8
(22)
5.0
27.3
(22)
23.8
(21)
1-(
o.o
(22)
Left
30.0'
(20)
28.0h
(25)
O.O
(23)
Left
Right
Left
Right
30.4
(23)
24.0
(25)
7.7
(13)
26.4
(19)
JO.O
(20)
17.4
(23)
(13)
28.6
(14)
37.6
(16)
6.7
(15)
17.6
( 17)
20.0
(20)
O.O
(14)
Right
o.o
o.o
(21)
(21)
8.6
(23)
21.7
(23)
O.O'
(24)
M ississippian
Archaic
4.8/J
(21)
o.o
o.o
o.o
( J3)
(10)
22.3
(18)
5.9
(17)
15.8
(19)
0.0-1
(18)
15.8
(19)
5.6
(18)
7.1
(14)
21. l
(19)
o.o
( 17)
23.5
(\ 7)
o.o
o.o
(\ 6)
(19)
These la ter groups were largely sedentary and lived primarily in villages on
river floodplains, although smaller temporary uplands habitations were
utilized far hunting <leer and other animals (e.g., small mammals, turkey,
waterfawl) on a seasonal basis. In sum, although sharing sorne features, the
subsistence strategies and settlement patterns in the earlier and later
periods were very different. Because faraging and farming involved very
different kinds of physical activity, the respective populations should
.
display different prevalence and patterns of osteoarthritis.
Comparisons of shoulder, elbow, wrist, hip, knee, and ankle osteoarthntis show a number of importan! temporal trends in the Alabama series
(Bridges, 199la). Far individuals 30 to 49 years of age-at:d~ath, the
Archaic group has generally more osteoarthritis than the M1ss1ss1pptan
group, and the increases are especially consisten! far males (Table 5.2).
Statistically significan! differences between periods are presentin only a few
of thejoints. However, the overall greater prevalence in the Archaic sample
is clear. The severity of osteoarthritis tells the same story: Archaic
populations have generally greater severity ofthe disorder than Mississip-
181
182
service under the repartimiento labor system and farced to make longdistance trips to various localities in the province (Hann, 1988; Lyon,
unpnblished manuscript; Worth, 1995). These trips involved carrying
heavy burdens over great distances (Hann, 1988), which placed demands
on the lower limbs in walking and on upper limbs and trunks. The general
increase in mechanical demands on native populations in the seventeenth
to the increasing labor demands placed on native groups during the mission
period (and see Chapter 6).
In summary, from the osteoarthritis evidence, there was an apparent
decline in physical demands with the transition to agriculture in northwestern Alabama and coastal Georgia. It is importan! to point out that the
Alabama and Georgia populations may not be directly comparable. There
is a large temporal gap between the early (faragers) and late (farmers)
prehistoric groups in northwestern Alabama. The skeletal series used for
making comparisons are separated in time by sorne 2200 years ( 1000 BC-AD
1200). The osteoarthritis profile during the period of time immediately
prior to the Mississippian agriculturalists is unknown. It is possible that a
period of less intensive maize agriculture prior to AD 1200 produced levels
of osteoarthritis similar to those of either the foragers or the farmers.
Larsen's (1982) study of osteoarthritis in the prehistoric Georgia Bight also
involved the analysis of skeletal remains from populations spanning a
lengthy period; the precontact preagricultural group comprises ali human
remains predating AD 1150. However, most prehistoric farager remains are
drawn from the 450-year period immediately preceding the adoption of
maize agriculture (ca. AD 700-1150). Unlike the Alabama series, there is a
183
cultural and biological continuum with Iittle orno time gap separating the
population groupings. The temporal differences between the Alabama and
Georgia skeletal assemblages may not be significan!, because subsistence
reconstruction based on stablecarbon isotopes far the Eastern Woodlands
indicates that the shift to maize agriculture was widespread in the region
after about AD 900 (e.g., Ambrose, 1987; Smith, 1989). Thus, the two
regions are broadly comparable, at least with respect to the timing of the
introduction of maize agriculture and underlying socioeconomic factors.
The adaptive systems representing the Alabama and Georgia populations are different in sorne other importan! aspects. The Alabama skeletal
series represents populations of intensive terrestrial maize agriculturalists,
whereas the Georgia series represents populations of maize agriculturalists,
but who also engaged in fishing and collecting marine resources from local
estuarine and ocean contexts. Activity differences rellecting these adaptive
contrasts are suggested by long bone structural analysis (see Chapter 6).
It would be over simplistic to say that the shift to or intensification of
agriculture in prehistoric North America involved a reduction in workload
in native populations. For example, comparisons of less intensive with
more intensive agriculturalists from the Dickson Mounds site, Illinois,
show a general increase in prevalence and severity ofvertebral osteoarthritis in the more intensive agriculturalists (Lallo, 1973). In adults (sexes
combined), the frequency increases from 39.7% in the Late Woodland
period to 65.8% in the Middle Mississippian period. Similarly, there is a
much greater frequency of osteoarthritis in Mississippian period agriculturalists from the A verbuch site compared to foragers from the Indian
Knoll (Pierce, 1987; and see Hodges, 1989).
Much of the facus on the temporal comparisons of osteoarthritis deals
with the shift from foraging to farming. The study of osteoarthritis in
relation to other changes in economic systems has also proven highly
infarmative. In the Santa Barbara Channel islands and mainland Pacific
coas!, the facus on hunting and gathering of terrestrial resources was
replaced by intensivefishing in la ter prehistory (Walker & Hollimon, 1989).
The latter adaptation is especially well documented by early explorers and
others who first arrived in the region. These observations provide an
importan! perspective on the types of activities undertaken by native
populations that may have potential inlluence on articular pathology.
Early accounts of native groups note the presence of an elaborate fishing
technology and material culture, including such items as harpoons, fish
traps, nets, and fishhooks. In addition to fishing, shellfish were collected
from rocks by the use of prybars constructed from wood or bone. Boats
made from carved planks were used far travel between islands and between
184
islands and the mainland. Plant foods (e.g., acorns, chia and other seeds)
werecollected in large quantities, especially on the mainland. Various roots
and bulbswere extracted from the ground with digging sticks. Economic
tasks followed a strict division oflabor by sex- men hunted and fished, and
women collected plant foods and shellfish. These differences in work
activities are reflected in dietary differences between adult females and
males (Walker & Erlandson, 1986). For example, early prehistoric women
have higher caries rates than early prehistoric men, which reftects the
greater consumption of cariogenic plant carbohydrates by women (and see
Chapter 3).
Comparison of osteoarthritis in early and late prehistoric Indians from
this setting reveals temporal changes that are suggestive of alterations in
activity and workload with the transition to a marine focussed economy
(Walker & Hollimon, 1989). Severity of osteoarthritis, ranging from slight
articular surface porosity to extensive marginal lipping and eburnation,
increases in the late prehistoric period, especially in the lower limb (Walker
& Hollimon, 1989). On the basis of archaeological evidence, Walker &
Hollimon (1989) speculate that the increase in osteoarthritis may be dueto
increased work that involved trade, exchange, and more pedestrian travel.
The severity of elbow and wrist osteoarthritis increased in Late period
adult males, but not in adult females in the Santa Barbara Channel Islands
region. Osteoarthritis severity declined in the shoulder and hand. These
changes may be linked to an alteration in weaponry (replacement of the
atlatl by the bow-and-arrow) and fishing equipment (shift from harpoons
to nets). The temporal increase in forelimb osteoarthritis is perhaps related
to the increased use of canoes and fishing neis in the Late period (Walker &
Hollimon, 1989). Overall, the increase in osteoarthritis was greater for
males than females. Although there are severa! possible explanations for
this trend, the greater role of men in fishing suggests that the workload
increase was greater formales than females. The net result was a decrease in
the difference of osteoarthritis prevalence between men and women in the
Late period (Hollimon, 1991).
Viewed in a regional perspective, these North American studies show a
tremendous range ofvariation in mechanical stress loads. This high degree
of variability from one setting to another suggests that at leas! sorne factors
that intluence osteoarthritis are dependen! on local circumstances. Webb's
(1989, 1995) study of native populations from Australia serves to underscore the point that what are often perceived as uniform adaptations- even
over very large areas - are in fact highly variable. His comparative analysis
of osteoarthritis prevalence reveals very differenl'patterns and mechanical
stress levels in skeletal series from regio ns of Australia. F or example, elbow
185
5.4
Nonpathological skeletal modifications reflecting habitual postures provide a picture ofbehaviors such as squatting and kneeling(see K. Kennedy,
1989; Trinkaus, 1975). For example, the crouched posture that characterizes squatting involves extreme flexion of the hip, knee, ankle, and foot
joints. As a result, mechanical demands on lower limb articular joints may
produce distinctive joint modifications. Charles (1893) described an extension of the articular surface from the femoral head onto the superioranterior neck - called Poirier's facet - which he assumed was related to
abduction and hyperftexion of the thigh during squatting. Most later
studies show no relationship between the articular extension and squatting
(see Angel, 1964; Trinkaus, 1975). Other skeletal modifications purported
to be linked with squatting include facets on the superior-posterior margins
of the femoral condyles, rounding of the posterior aspect of the lateral tibia!
condyle, retroversion of the tibia! plateau, formation of a groove for the
posterior cruciate ligamen! (on the femoral intercondylar line), angulated
facets on the anterior aspect of distal tibia, talar neck facet surfaces, and
various other alterations on the talus and calcaneus (reviewed by Trinkaus,
1975). Trinkaus (1975) concluded that none of these features bears an
unambiguous association with squatting. For example, the presence of
articular facets on the superior-posterior femoral condyles and the groove
created by the posterior cruciate ligamen! on the femoral intercondylar line
are not necessarily associated with squatting. The presence of a relatively
high frequency of femoral condylar facets, and extensions in the knee,
ankle, and subtalar articulations in western European Neandertals is
suggestive of a habitual squatting posture. The extreme degree of skeletal
robusticity in these late archaic Horno sapiens indicates very high levels of
activity generally, which would also promote the development of these
articular joint modifications. Therefore, the combination of squatting and
great physical activity best explains these skeletal features, at least in these
late Pleistocene hominids (Trinkaus, 1975).
Trinkaus's (1975) study reminds us that it is difficult to sort out general
activity levels and specific articular joint modifications in interpreting
skeletal morphology. Study of distinctive metatarsal-phalangeal joint
articular modifications in severa! different contexts reveals that habitual
186
187
188
5.5
5.5.1
Cortic~l
defects
189
190
5.5.3
populations. In his sample from north of the Yukon River, Alaska, more
than 40% of individuals exhibit separa te neural arches, which he originally
attributed to inbreeding of isolated groups (but see Stewart, 1979). Sorne
191
192
lower back, perhaps accompanied by jarring and twisting, are linked with
spondylolysis (Merbs, 1989b). Given the broad ran.ge ofbehaviors that are
associated with these defects, no specific stresses appear to lead to
spondylolysis (Bridges, 1989a; Merbs, 1989b). Merbs (1989b) speculates
that spondylolysis may have an adaptive value in that it may engender
flexibility of the lower back (see also Snow, 1974). Bird and coworkers
(1980), for example, reported that adults with the defect considered
themselves 'more supple in youth' than adults who lacked the defect. The
defect seems to have minimal negative influence on physical performance.
F or example, college football players with the defect appear to lose neither
practice nor playing time, and they continue to play professionally in later
years (see discussion in Merbs, 1989b).
The el ose association between mechanical loading of the lower back and
spondylolysis indicates that the defect should be documented in archaeological skeletal series. In northwestern Alabama, early prehistoric huntergatherers have a higher prevalence of spondylolysis than late prehistoric
agriculturalists (Bridges, l 989a). This finding is consistent with moderate
increases in osteoarthritis (see above) in this series. Statistical analysis of
the co-occurrence of spondylolysis and osteoarthritis reveals only a weak
relationship between the two conditions. Osteoarthritis prevalence is
broadly similar between individuals with and without spondylolysis.
Structural analysis of long bones reveals a general increase in mechanical
demand in the agricultura] groups, which seems to contradict the findings
based on spondylolysis prevalence (see Chapter 6). Bridges (l 989a)
speculates that spondylolysis may be associated with unusual postures or
specific activities affecting the lower spine rather than overall activity levels
(cf. Hoshina, 1980).
Adult males in populations with appreciable frequencies of spondylolysis
ha ve a higher prevalence than adult females (e.g., Arriaza, 1995; GunnessHey, 1980, 1981; Merbs, 1983, 1995; Stewart, 1979; Trembly, 1995;
Waldron, 1993). Presumably, mechanical demands causing the defect were
greater for men than for women in these societies. There are sorne notable
exceptions to this pattern, which emphasizes the role of mechanical demand
and the influence of culture cross-cutting gender lines. Contact era Ornaba
and Ponca women from northeastern Nebraska, for example, ha ve a higher
prevalence of spondylolysis and spinal osteoarthritis than men (Reinhard et
al., 1994). Highly demanding activities affecting the female spine are well
documented ethnohistorically; these include hide scraping undertaken in a
stooped posture (Reinhard et al., 1994). In addition, women were responsible for a range of other physically demanding activities not shared by men,
including house construction and firewood gathering.
1
%
1
1'
*
1
1
1
!
1
1
1
l
1
1!
193
5.6
Study of articular joint modifications relating to mechanical demand especially osteoarthritis- offers insight in to the stresses of different activity
194
patterns and lifestyles in past populations. Generally, the more mechanically demanding the lifeway, the greater the prevalence of osteoarthritis
and other degenerative pathological conditions related to activity (e.g.,
spondylolysis). Conversely, less demanding work repertoires result in a
relatively lower prevalence ofthese conditions. Temporal comparisons of
6.1
trends. For example, although there are sorne exceptions, la ter prehistoric
agriculturalists tend to show a reduction in osteoarthritis and degenerative
pathology than earlier foragers. This provides sorne support for the
traditional point of view that foragers work harder than agriculturalists.
More importantly, local circumstances and conditions influence osteoar-
Julius Wolff, a leading nineteenth century German anatomist and orthopedic surgeon, recognized the great sensitivity of bones to mechanical
stimuli, especially with regard to their ability to adjust size and shape in
response to externa! forces. Wolff concluded that 'every particle of mature
bone is very active. Such activity must appear in the externa! shape of the
bones' (1892:78). What he called the 'law of bone remodelling' - now
known as Wolff's Law- simply states that bone tissue places itself in the
direction of functional demand.
A great deal of evidence has accrued in support of Wolff's hypothesis,
thus demonstrating the primacy of the mechanical environment in interpreting skeletal structural variation. Experimental and other research on
bone remodeling is instrumental in identifying patterns of skeletal modification under different loading regimes (see Lanyon et al., 1982; Meade,
1989; Trinkaus et al., 1994). Using laboratory dogs, Chama y & Tschantz
(1972) observed that the surgical removal of portions of radii resulted in the
hypertrophy of ulnar diaphyses. The ulnar diaphyses increased in size by
31%after just 16 days and 60-100% at nine weeks. Similarly, Lanyon and
coworkers (Goodship et al., 1979; Lanyon & Bourne, 1979) documented
increased apposition of bone on the radius following ulnar osteotomies in
pigs and sheep. Nonsurgical load alterations have also resulted in changes
in bone mass. Woo and coworkers (1981) identified significan! endosteal
. apposition in young pigs subjected to exercise. Simkin and coworkers
(1989) compared hnmeri from swimming and nonswimming rats in an
experimental setting. The swimming rats included a group trained to swim
for one hour per day and a group that underwent the same training, but
also hada lead weight (approximately 1% of the rat's body weight) tied to
their tails. Comparison of bone size and structure revealed that both
groups of swimming rats had greater periosteal apposition than the
thritis prevalence and pattern in a far more profound way. Adult males
generally show a tendency for greater degenerative pathology than adult
females in archaeological settings. Although osteoarthritis is related to
mechanical loading, the relationship to leve! or type of activity is not a
direct one. High levels of osteoarthritis in the skeleton suggest demanding
lifestyles, but do not indicate whether these demands also include longdistance or frequent travel as a cause, except possibly with regard to the
high prevalence offoot osteoarthritis. Other skeletal indicators ofmobility
are discussed in the following chapter.
1
1
1
196
Structural adaptation
Cross-sectiona/ geometry
1977; Ruff et al., 1994), rodeo cowboys (Claussen, 1982), and baseball
pitchers (King et al., 1969), exhibit marked hypertrophy of the externa!
diaphyses of long bones of the playing si de. In professional ten nis players,
for example, one study revealed that males ha ve a 35% increase in cortical
area in the distal humerus of the playing arm vs. the nonplaying arm;
females have a 29% increase (Jones et al., 1977). The effects of elevated
.. 0.. ..G~
in clinical settings. Adults who exercise tend to ha ve higher bon e mass and
size than adults who are relatively sedentary, especially in males (reviewed
by McMurray, 1995).
Removal of normal functional loading, especially involving extended
periods of bed rest, weightlessness in spacefiight and absence of gravi-
UNLOADED
decreased bone mass (e.g., Abram et al., 1988; Jenkins & Cochran, 1969;
Kazarian & Von Gierke, 1969; Kiratli, 1992; Lanyon & Rubin, 1984;
Lazenby & Pfeiffer, 1993; Meade, 1989; Morey & Baylink, 1978; Prince et
al., 1988; Sevastikoglou et al., 1969; Todd & Barber, 1934; Whalen et al.,
1988; see discussion by Trinkaus et al., 1994).
Bone is anisotropic in that it is characterized by differcnt material
properties depending upon the direction ofloading (Cowin, 1989; Currey,
1984; Nordin & Frankel, 1980; Wainwright, 1988). Long bones, for
example, are stronger in the longitudinal direction than in any other plane
(Wainwright, 1988). The primary loading forces affecting bone include
tension, compression, shear, bending, and torsion (Figure 6.1). These can
be best understood by considering a thin section, or slice ofbone. Tensile
loading occurs when equal and opposite forces are applied outwardly from
the surface of the slice. Compression, the opposite of tension, occurs when
equal and opposite loads are directed toward the two surfaces. Shear loads
involve the application of forces parallel to the surfaces under consideration. Bending forces produce two types of stresses, tension on the convex
side and compression on the concave side. Torsional loading is the twisting
197
TENSION
SHEAR
COMPRESSION
TORSION
BENDING
COMBINED LOADING
TORSION-COMPRESSION
Figure 6.1. Loading modes that affect.long bones. (From Nordin & Frankel,
1980; reproduced with permission of authors and Williams & Wilkins.)
6.2
Cross-sectional geometry
198
Structural adaptation
(a)
Figure 6.2. Cross section of bone undergoing bending (a) a~d torsion. (b), and
showing stress distribution around the neutral plane and axis, respect1vely.
Note that the magnitude of forces (indicated by heavier arrows) is greatest at
the periphery of the bone and least nearest the neutral plane or axis. (F~o?1
Nordin & Frankel, 1980; reproduced with permission of authors and W1lhams
& Wilk_ins.)
& Conroy, 1996),femoral necks (Beck et al., 1990; Phillips et al., 1975), and
second metacarpals (Roy et al., 1994). These investigations, as well as
experimental evidence based on laboratory animals (e.g., Abram et al.,
1988; Simkin et al., 1989), indicate the value of structural analys1s m
drawing inferences about physical activity and behavior patterns.
In bending and torsion of a hollow beam, such as a long bone, the
magnitud e of mechanical stresses is proportional to the distance fro':'1 the
central or 'neutral' axis of the bone (Figure 6.2). The neutral axis 1s the
plane (bending) or axis (torsion) where stress is zero. Thus, ali else ~ein.g
equal, the cross section that is strongest is that in which t?e matenal 1s
oriented furthest from the neutral axis (Currey, 1984; Nordm & Frankel,
1980; Ruff, 1992). And, by inference, the greater the distance from the axis,
the greater the magnitude of stresses (see Nordin & Frankel, 1980; Ruff,
1992; Wainwright, 1988). In long bones and in sorne other.ele.ments (e.g.,
Cross-sectional geometry
199
metacarpals), the cross-sectional area and the manner in which the bone is
distributed about an axis reflect mechanical/functional behavior.
Bending a ruler is a good analogy for demonstrating the principies
associated with beam analysis. If one attempts to bend a ruler against its
narrow edges, there is little or no give. In contras!, if forces are applied
virtually anywhere along its flat surface, especially toward the middle, the
ruler bends readily. From a mechanical perspective, the small amount of
give when applying bending forces to the narrow edges occurs because the
materials in this axis are distributed relatively far from a central, neutral
axis. Thus, in this plane of bending, the ruler has a great deal of strength.
Conversely, the ease of deformation when bending forces are applied to the
flat surface is made possible by the lack of material far from the neutral
axis; therefore there is very little strength of material when the ruler is
subjected to bending forces in this direction. A ruler is structured so as to
resist bending from one direction only. Given the tubular shape of long
bones, they are able efficiently to withstand the mechanical demands
associated with bending and torsion from multiple directions.
Beam analysis involves the measurement of geometric properties from
cross sections taken perpendicular to the long axis of a skeletal element. As
demonstrated by the ruler analogy, these properties are based on both the
amount and the distribution of bone tissue in the cross section. As suc:n,
they are direct measures of the 'strength' (more precisely, rigidity) of the
bone cross section in resisting forces or loadings. The properties measured
in archaeological human bones should, then, represen! a measure of the
cumulative forces operating on the skeletons of individuals during their
lifetimes.
Cross-sectional geometric properties measure the amount and the
distribution of skeletal tissue in a section. These properties include section
'areas' and 'second moments of area' (or 'area moments ofinertia') (Figure
6.3). Areas include total subperiosteal area (TA), endosteal or medullary
area (MA), and cortical area (CA). Measurement ofmediolateral (mi) and
anteroposterior (ap) breadths of long bone diaphyses are used to calcula te
areas with formulas for two planes of measurement (e.g., Ruff & Jones,
1981):
TA= n(Tap/2)(Tm1/2)
MA = n(Map / 2)(Mm1/2)
CA=TA-,-MA
where T is the ti:ltal externa! diameter and M is the medullary diameter.
Thus, CA is a measure ofthe amount of cortical bone in a cross section and
200
Cross-sectional geometry
Structural adapta/ion
001GPF
/\
\U
\
Cortical properties
TA
CA
X bar
Y bar
lx
ly
Theta
lmin
Imax
~,
(1 O, 1O) (30,
454.96
287.26
19.40
31.74
16468.40
12422.80
97.48
16539.40
12351.80
28891.20
10)
is also an indicator of strength of the long bone diaphysis under pure axial
loading (loading that is simultaneously applied to both ends of the bone).
Per cent cortical area (%CA= CA{f A) is an a!ternative indication of
amount of compact cortical bone. CA and %CA provide very different
representations of bone mass, owing to the fact that the latter, but not the
former, measures cortical bone relative to TA (and see Ruff, 1992; Ruff &
Larsen, 1990). TA and MA are measurements of the two major surfaces of
the bonecortex, including the outcr periosteal and inner endosteal surfaces,
respectively (Ruff, 1991). Expansion in TA and MA indicates a greater
distribution of skeletal tissue further from the neutral axis of the bone.
Bone areas, especially CA, are proportional to strength in compression
and tension when the forces are applied noneccentrically along the central
longitudinal axis ofthe bone diaphysis (axial loading). Beca use long bones
are curved and are affected by muscular forces applied off-center to a bone
axis, pure axial loadings in either compression or tension are rare; most
forces involving long bone diaphyses are eccentrically applied. Second
moments of area ha ve been shown generally to be more accurate indicators
of bone strength and mechanical function than areas alone. For example,
201
where a; is the unit area and d; is the perpendicular distance from the center
of the unit to the neutral axis. lx can refer to the bending strength in the
anteroposterior plane and !y to bending strength in the mediolateral plane
(Ruff, 1989). Other values of I expressing the maximum and minimum
bending strength in a cross section are referred to as I max and I m;n,
respectively, where Imax measures the maximum strength in resistance of
bone to bending and Imrn measures the minimum strength in resistance of
bone to bending. Torsional strength is calculated in reference to the neutral
center or 'centroid' of the cross section and is called the polar second
moment ofarea or 'J.' Jis equal to the sum ofthe values of Imax and Imrn (or
/., and !y), which are always perpendicular to each other (Ruff & Hayes,
1983a).
Values of I and J are calculated as products of very small unit areas in the
cross section and squared distances of the unit areas relative to the neutral
axis (for values of !) or the neutral center of the section (for values of J).
Therefore, second moments of area are presented as linear dimensions to a
specific power. Because ofvariability in body size or length oflong bones in
the comparison of different population samples or temporal series within a
particular setting, properties should be size standardized when comparisons are made between or within comparative groups (Ruff, 1984). Earlier
work suggested that, for the femur, dividing areas by the square of the
length and second moments of area by the fourth power of the length was
an appropriate meaos of size standardizing (e.g., Ruff, 1984; Ruff &
202
Structural adaptation
Larsen, 1990). More recent analyses utilizing additional and more extensive samples of extinct and recen! human skeletal samples suggest that
more appropriate powers for size standardization of the femur are bone
Jength cubed and bone length to the power 5.33 for areas and second
moments of area, respectively (Ruff et al., 1993).
Cross-sectional geometric analysis was first applied to samples of more
than a single bone in a variety of settings involving both human and
nonhuman primates (e.g., Jungers & Minns, 1979; Kimura, 1971, 1974;
Klenerman et al., 1967; Lovejoy et al., 1976; Martin & Atkinson, 1977;
Miller & Piotrowski, 1977; Minos et al., 1975; Piziali et al., 1976). These
studies were generally limited to fewer than 10 individuals beca use of the
lengthy and tedious process involved in manually calcuJating geometric
properties. Specifically, two problems with calculating section properties
include the determination of endosteal and periosteal boundaries and the
mathematical integration of areas that are necessary to complete the
calculations (Ruff, 1992). The development of automated protocols far
computer analysis of large numbers of cross sections (Nagurka & Hayes,
1980) and new technologies has made it possible to carry out stud1es
involving more extensive samples. These developments have fostered a
more comprehensive understanding of variability both within and between
populations from archaeological and paleontological contexts (e.g., Berget
& Churchill, 1994; Bridges, 1985, 1989b, 199la, 1995a, 1995b; Brock, 1985;
Brock& Ruff, 1988; Churchill, 1994; Kimura & Takahashi, 1982; Larsen &
Ruff, 1991, 1994; Larsen, Ruff et al., 1995, 1996; Robbins et al., 1989; Ruff,
1987, 1989, 1991, l 994b; Ruff & Ha yes, 1982, 1983a, 1983b; Ruff & Larsen,
1990; Ruff et al., 1984, 1991; Sumner et al., 1985; Trinkaus et al., 1994; Van
Gerven et al., 1985).
The method of geometric analysis involves the preparation of a section
image made perpendicular to the longitudinal axis of the bon e. This image
is obtained through severa! alternative means, from existing breaks on long
bone diaphyses (e.g., Lovejoy & Trinkaus, 1980), direct cutting (e.g.,
Larsen & Ruff, 1994; Ruff & Hayes, 1983a, 1983b; Ruff et al., 1984), or
noninvasive imaging, especiallycomputed tomography (CT) (e.g., Brock &
Ruff, 1988; Bridges, 1989b; Larsen, Ruff et al., 1995; Ruff, 1987). Other
useful noninvasive techniques include multiple plane radiography (e.g.,
Biknevicius & Ruff, 1992; Fresia et al., 1990; Runestad et al., 1993;
Trinkaus & Ruff, 1989) and photon absorptiometry (e.g., Martin & Burr,
1984; Van Gerven et al., 1985; and see discussion of imaging techniques by
Ruff, 1989; Ruff & Leo, 1986). Noninvasive imaging is useful in situations
in which the cutting of bone specimens is not possible (e.g., fossil
hominids). Unlike invasive cutting, the advantage of noninvasive analysis
Cross-sectional geometry
203
is that section properties can be determined directly from the images (e.g.,
CT sean).
The accuracy of geometric analysis is dependen\ upon the availability of
fully intact periosteal and endosteal section con tours. The con tour integrity
of the periosteum can be deterrnined through visual inspection. For
noninvasive analysis, it is advisable to examine the endosteal surface by
cutting one or two specimens or using diaphyses that have already been
broken. In addition to well preserved periosteal and endosteal surfaces, the
accuracy of analyses - including size (length) standardization as well as
precise location of sections - requires the presence of largely intact ends of
long bones. Therefore, even though large numbers of skeletons may be
present in an archaeological series, if differential preservation exists, then
only subsamples composed of well preserved specimens can be analyzed
(e.g., Ruff & Larsen, 1990; Ruff et al., 1984).
There are various ways of obtaining landmark data for section analysis.
Commonly, once the section image is obtained, a magnified photographic
slide image is projected onto a digitizer screen, and the periosteal and
endosteal borders are manually traced with a digitizer stylus directly input
to a microcomputer, recording x and y coordinates at intervals of 1 mm. An
automated computer program (e.g., SLICE; Nagurka & Hayes, 1980) then
calculates the section properties.
Studies of cross-sectional geometric properties address a range of
long-standing issues in anthropology dealing with activity and physical
behavior, especially in regard to subsistence strategies, the relationship of
sex differences to dietary and subsistence adaptation, and variability in
skeletal growth and development in response to environmental and dietary
change.
6.2.J
204
Structura/ adaptation
Midshaft
400
femur
400
350
300
"'E'
<(
350
~~
..........,~// ...............
250
200
150
100
50
300
250
~CA
200
"'
1
m e"'
E'
(L
oo
E'
(L
(L
ii:
"' Cl"'
Cl
00
roo
<D
Subtrochanteric
femur
~
'
Mid-distal
1.25
350
'\,
300
------..--.
250
'
c----nc ......,
1.30
400 humerus
200
CA
1.20
11-------0-----....,
1.00
0.95
150
150
PCCA 100
PCCA 100
...------.
'iiio <(e,
"'
ii: '!2>
o
;j
"'
Cl
50
~
e, o00 e,
<(
o
E'
(L
(L
j
~
"' "'
e"'o
e"'o
Cl
"'
Cl
"'
50
1.15
~:.., 1.10
~1.05
11-------"----....
CA
e,
205
Cross-sectional geometry
PCCA
j
33
e,
e,
"'E'
e"'
e"'o Cl"'o
Cl
"'
<(
(L
0.90 L~-~-~~-~-~e,
e,
;
'iii
1
~
e"'o
(L
Cl
Cl
(L
"'
E'
ii:
"'
"'o
""'
(L
"'
Cl
<(
e"'o
"
Cl
uniformly mobile and farmers are no!. Increasing evidence indicates that a
number of hunter-gatherer groups in the prehistoric past were relatively
sedentary(e.g., Erlandson, 1994; O'Neill, 1994). Rather, these biomechanical studies suggest tendencies in patterns of mobility from high levels in
foragers to low levels in agriculturalists.
Torsional loading bears no apparent relationship to subsistence strategy
in females in these North American groups. Rather, female torsional
Joading corresponds with degree of ruggedness of terrain - mountainous
populations (Stillwater, Pecos Pueblo) ha ve the highest values of J, coastal
populations have the Jowest values (Georgia coast), and Great Plains
populations are intermediate. The differences between males and females
are also indicated by comparison of Ruff's mobility index. Stillwater males
shows a very high degree of activity involving use of the lower limbs, such
as in long distance traveL Their values are among the highest in comparison
with other North American populations, and, like data presented for
second moments of area (]), the index value fits into a continuum from
hunter-gatherers to agriculturalists (see Figure 6.5). On the basis of this
index, it appears that Stillwater females are relatively less mobile than
206
Structural adapta/ion
males, which is true for ali other North American series. Comparisons of
females and males in the lxf I, ratio revea! that Stillwater foragers are highly
sexually dimorphic, which is a paramount characteristic of huntergatherers generally and is related to sexual division of labor and a strong
male-female dichotomy in activity patterns (Larsen, Ruff et al., 1995; Ruff,
1987; C. B. Ruff, unpublished manuscript).
In the Stillwater series, humeri show a somewhat different pattern of area
and second moments of area from that of femora. Especially striking are
thc !ow values ofhumeral CA and J (Larsen, Ruff et al., 1995). This finding
is consisten! with the hypothesis that mechanical loading is localized in the
skeleton. With regard to the Stillwater adults, the presence of high bone
strength in the fen1ur and not the humerus indicates a prin1arily mechanical
effect (ruggedncss of environment, and thus, high lower limb loadings);
whereas the !ow values of CA for both skeletal elements - femur and
humerus - are more likely to be due to so1ne systemic influence, such as
undernutrition.
Therefore, the generally high levels of robusticity in Stillwater adults especially in femara - is consistent with findings based on osteoarthritis
prevalence (and supporting a limnomobile subsistence strategy' see Chapter 5). Behaviors causing osteoarthritis and elevated robusticity result in
respective high frequencies of pathology and high second moments of area.
Comparative analysis of long bone geo1netry gives greater insight into
207
Cross-sectional geometry
Table 6.1. Humeral polar moments of area - 35%, 50%, and 65%
sections - infossi/ and recen/ human adults. (Values divided by length to
thefourth power and multiplied by HJ1. Adaptedfrom Churchi/l, 1994:
Tables 16 and 17.)
Females
Males
Group
35o/o
50%
65/o
35%
50%
65%
Euroamerican
79.9
(19)
89.7
(19)
96.8
(19)
124.3
(25)
149.3
(25)
101.9
(25)
113.5
(24)
117.8
(25)
141.2
(21)
Amerindian
102.3
(n)
(20)
Peruvians
66.1
(n)
(1)
Late Upper Paleolithic 125.2
(n)
(6)
184.7
(22)
121.4
(20)
86.5
(1)
210.0
(22)
128.3
(20)
95.6
(1)
151.4
(25)
198.6
(25)
102.1
(20)
127.6
(3)
145.5
(25)
169.6
(25)
265.6
(25)
118.5
(20)
139.7
(3)
178.8
(25)
316.5
(25)
124.8
(20)
147.3
(3)
155.0
(6)
154.3
(9)
183.8
(9)
183.0
(8)
132.8
(3)
172.I
(5)
120.0
(3)
95.9
(2)
151.0
(4)
144.0
(6)
92.2
(1)
166.7
(7)
128.8
(6)
192.7
(5)
123.5
(1)
194.9
(6)
(n)
Afroamerican
(n)
Aleut
(n)
Skhul/Qafzeh
(n)
Archaic humans
(n)
101.3
(1)
125.2
(4)
79.5
(2)
133.3
(4)
82.9
(2)
184.2
(6)
208
Structural adaptation
of per cent cortical area (%CA), suggesting that the differences in borre
structure are not dueto dietary stress, but rather to behavioral and activity
differences.
Cross-sectional geometry
Table 6.2. Percentage change with age in femoral and tibia/ crosssectional geometric properties. Calcu/ated by the formula: {[ ( 40 +
years) - (20 to 39 years) J + (20 to 39years)}x100. ( Adaptedfrom
Rujf & Hayes, 1982: Table l.)
Bon e
(section location)
Males
Tibia (20%)
Tibia (50%)
Femur (50%)
Femur (80%)
Until recently, this hypothesis was difficult to test beca use of the imprecision of radiographic measures of cortical borre remodeling (see Ruff &
Hayes, 1982). In order to examine the issue of age changes and periosteal
expansion in more detail, Ruff & Hayes (1982, 1983b) analyzed section
properties - areas and second moments of area - from the late prehistoric/
protohistoric Pecos Pueblo site, New Mexico. Analysis of femoral and
tibia! diaphyseal sections reveals that both sexes saw increases in MA and
TA and decreases in CA with advancing age (Table 6.2). Second moments
of area (/max and lm;n) increase in older adults. Thus, in support of the
compensatory hypothesis, continuous periosteal expansion in older adults
appears to maintain the mechanical integrity of the long bone despite
overall decline in bone mass.
Variation in the comparison of different sections along femoral and tibia!
diaphyses also reveals that skeletal remodeling with age is less pronounced
CA
MA
TA
0.2
0.1
-0.5
13.IY'
35.7'
25.4'
13.1 6
73n
12.4'"
6.7'
4.7"
9.4
14.6"
12.3'
7.5
7.8
19.9
6.8
4.8
-14.5..,
-13.1 6
-6.6
-2.4
9.1"
62.4'
66.6'"
4J.2r
9.1"
12.9'
11.1
13.6'"
-1.5
10.3
-9.6
16.5'
-3.0
4.1
15.6'
16.l)"
-1.1
lmax
/min
Fen1a/es
Tibia (20%)
Tibia (50%)
Femur (50o/I))
6.2.2
209
Femur (80%)
in the most distal and proximal sections. The greater remodeling in tibia!
and femoral midshaft and adjacent sections is probably due to lhe
relatively greater mechanical loads - especially bending - relative to distal
and proximal ends (Ruff & Hayes, 1982; Ruff, 1992).
Juveniles also appear to show compensatory diaphyseal remodeling.
Analysis of Medieval period juvenile (~16 years) tibiae from Kulubnarti,
Sudanese Nubia, shows increases in per cent cortical area (%CA) and
second moments of area (l.,, !y) from ages 3 to 12 (Van Gerven et al., 1985).
After age 12, areas decline, while second moments of area contiflue to
increase dramatically. Thus, despite decline in borre area, continucd
increase in second moments of area in the later juvenile years appears w
foster mechanical integrity throughout the years of growth and development.
In order to document in more precise fashion the ontogenetic age
patterns of diaphyseal remodeling, Ruff and coworkers (1994) examined
areas and second iuoments of area of humeri in professional tennis players
aged 14 to 39 years. Both males and females showed a pattern of endosteal
contraction and periosteal expansion resulting in large increases in bone
area (CA) and torsional strength (J) in the dominan! playing arm. The
resulting robusticity is primarily due to greater periosteal expansion and
not endosteal contraction. The degree of humeralrobusticity has a stroilg
210
Cross-sectional geometry
Structural adaptation
association with age: individuals who began playing tennis earlier had
greater robusticity than individuals who began later. The increased
mechanical loading in children and young adolescents has a more pronounced effect on the periosteal surface; after mid-adolescence, loading
appears to have a more pronounced affect on the endosteal surface. These
findings indica te an age pattem of sensitivity in the bone-forming surfacesperiosteal vs. endosteal - in response to increases in mechanical stimuli.
The cause of the shift in focus of sensitivity is unknown, but may be related
to changes in hormonal levels and their different effects on the two bone
surfaces (Ruff et al., 1994).
6.1.3
l
1
l
211
212
Cross-sectional geometry
Structural adaptation
213
214
Structural adaptation
Cross-sectional geometry
Therefore, these two indicators ofmechanical stress seem to yield contradictory results. Bridges ( 1991 b) suggests that osteoarthritis and long b~ne
geometry are due to different types of activities. F or example, c1tmg
findings from sports medicine and other research, she notes that so~e
normal activities - such as running - do not con tribute to osteoarthnlls
(e.g., Eichner, 1989; Panush & Brown, 1987; and. s~e Chapter 8). Less
frequent movements that lead to microtrauma and mJ~ry of JO~nts ~ay be
importan! factors in the explanation of osteoarthnt1s. 1t 1s v1rtually
impossible to distinguish traumatic from daily 'wear-and-tear' osteoarthntis, thus making it difficult to identify specific causes where behav10rs are
unknown, such as in archaeological settings. Skeletal structurnl change has
been tied to long-term repetitive forces (e.g., Lanyon & Rubm, 1984; Shaw
et al., 1987). Osteoarthritis and diaphyseal cross-sectional geometry should
not necessarily be concordant, because osteoarthritis would be expected to
develop in older adults, whereas diaphyseal remodeling is a lifelong
response to mechanical stimuli (see Ruff, 1992).
.
Analysis of two diaphyseal sections of the femur (subtr~chantenc
section taken at 80% of bone length measured from the distal end;
midshaft 50% from the distal end) and one section of the humerus
(mid-dis;al, 35% from the distal end) from Georgia Bight populations
yields a different pattern of structural change than in Alabama (Larsen &
Ruff 1991, 1994; Ruff & Larsen, 1990; Ruff et al., 1984). Structural analys1s
of p;econtact hunter-gatherers and precontact agriculturafts shows no
appreciable change in CA, whereas MA and TA declmes m both sexes.
These findings indicate little orno change in bon e mass, but rather, a t1ghter
distribution of skeletal tissue about the neutral axis in the later period.
Additionally, second moments of area decline in both males and females
(Figure 6.6). Therefore, in contras! to findings from Alabama, mechamcal
demand declined in later prehistory with the adopt10n of a hfeway
involving maize agriculture.
The results reported from two regions ofthe southeastern Umted States
are different, especially in the comparison of second moments of area.
These findings are not conflicting, especially in view of the fact that the
Alabama and Georgia Bight populations represen! very different adaptational and behavioral circumstances associated with shifts in subsistence
technology and economic focus. Both settings saw a change from foraging
to maize farming. However, the adoption ofmaize was only one factor of
many involved in adaptive shifts taking place in la ter prehistory, especially
when Alabama was compared with the Georgia Bight. Although these two
regions shared a number of resources in common (e.g., terrestrial animals),
the continued dependence on marine and estuarine resources even durmg
215
100 (a)
O Males
lm Females
# 50
Preagricultural
----Agricultura!
0'-=-L
CA MA TA lmax lmln
J
Cross-sectional property
100 (b)
@)
O Males
*"
'
'
!mFemales
50
-
Preagricultural
----Agricultura!
CA
MA
TA
lmax
Imin
Cross-sectional property
216
Structural adaptation
with the early contact period on St. Catherines Island, there is a reversa!
of the decrease in bone strength that had been documented in the
prehistoric populations from the region. Femora and humeri show
increases in MA: and TA; CA is unchanged in the early and late contact
periods. Mirroring the changes in MA and TA are successive increases in
second moments of area in the early and late contact periods. This trend
occurs uniformly for female geometric properties. In males, the trend in
the early and late contact periods is less straightforward. Geometric
properties increase in the early contact period, but decline slightly (e.g.,
subtrochanteric Im.x) or remain the same (midshaft /,) in the late contact
period.
In males humeral areas and second moments of area in crease successively in the e~rly and late contact periods. In females, there is a continued
decline in humeral val ues in the early contact period, but this reverses in the
late contact period. Thus, both females and males in the late contact period
experience a marked increase in humeral strength. In general, the changes
observed in bone strength in the contact period indicate that native
populations experienced increases in mechanical loads, probably due to
increased manual labor and physical demands placed on them by the
Spanish (and see Chapter 5).
Analysis of cross-sectional 'shape' of long bones provides insight into
variability in bending forces with regard to relative loading differences
between planes. An fx/Iy ratio close to 1.0 reftects a nearly circular shape,
and a ratio deviating from 1.0 represents an ovoid shape. This ratio for
the femur midshaft assesses the distribution of bone in the mediolateral
and anteroposterior planes; an ovoid-shaped midshaft in the anteroposterior plane (i.e., ratio > 1) represents relatively greater bone strength
and functional demand in the anteroposterior direction than in the
mediolateral direction. Ruff (1987) has shown a temporal decline in the
ratio in recen! human groups, which he interpreted to reflect a general
reduction in the amount of anteroposterior bending forces as populations
have become increasingly sedentary (cf. Lovejoy et al., 1976). Therefore,
at least with respect to Holocene populations, the lx/fy ratio represents a
mobility index.
In the Georgia Bight, early contact femora show a general reduction in
the femoral midshaft fx/Iy ratio relative to earlier prehistoric populations.
Historie sources indicate that populations during the historie period
became generally less mobile as they needed to or were coerced to live in
and around mission centers (e.g., Santa Catalina de Guale). This skeletal
indicator ofmobility is, therefore, in accord with other sources describing
population sedentism during the historie period. These structural modifi-
Cross-sectiona/ geometry
217
Structural adaptation
218
700
600
S!. 650
550
:?
"
~
N
"'e
J!!
s
::
"'
1'.i 600
~u
"O
<(
"'
!!
(/)
500
450
550
lii
8.
"'
"O
500
J!!
400 (/)
(/)
CA
450
10-7
10-6
10-5
10-4 10-3
Years BP
10-2
10- 1
350
10~
219
6.3
1
~
220
Structural adaptation
6.4
221
222
Structural adaptation
223
tissue in a section.
6.4.2
224
Structural adaptation
~o
/:t. lo Cortical thickness:
11 lo Cortical area:
6. Cortical area:
.6. Second moments of area:
-33/o
-33!o
+ 12/o
+ 100/o
(Houston & Zaleski, 1967). The orientation of the femoral neck relative to
the diaphysis appears to be responsive to the combined forces of body
weight, muscle forces, and activity generally. Additionally, the srnaller
angle provides greater hip joint rnechanical stability under increased
rnechanical loading (Trinkaus, 1993).
Comparisons of a range of human groups - foragers, agriculturalists,
and urban dwellers- support the mechanical hypothesis for variation in the
femoral neck-shaft angle (Trinkaus, 1993). Cornparisons indica te that the
foragers have the lowest femoral neck-shaft angles (mean= 125.7), urban
dwellers have the highest angles (mean= 132.3), and agriculturalists have
values between those of foragers and urban dwellers (mean= 128.2)
(meanscalculated frorn Trinkaus, 1993:Table 4). This pattern of increasing
neck-shaft angles from rnobile foragers to sedentary urban groups closely
parallels the general trends of decreasing robusticity based on whole bone
rneasurements and diaphyseal structure (Ruff et al., 1993), albeit with a
wide range of overlapping variability between groups (see Trinkaus, 1993).
6.5
225
articular joints.
Because humans do not use their upper lirnbs in arnbulatory activities,
the influence ofbody weight is very rninirnal in the overall deterrnination of
sizeand rnorphology ofarrn skeletal elernents (e.g., humeros). The study of
upper lirnb bilateral asyrnmetries in various human populations allows
inferences to be rnade about loading levels and patterns in relation to
different lifestyles and rnechanical functions.
Measurement of areas and second rnoments of area in a range ofhuman
populations, ancient and rnodern, reveals a general trend for decline in
robusticity. Although this trend is especially pronounced in the transition
frorn archaic to modern Homosapiens in the late Pleistocene (Ruff et al.,
1993), it has continued throughout the Holocene. The reduction during the
Holocene is probably tied to increasing sedentisrn associated with plaut
dornestication (Larsen, 1995). Sorne modern human populations are quite
robust (e.g., early modern Europeans, Great Basin Arnerindians), which
rnay be linked with living in marginal environrnents and the great physical
effort in the food quest in these types of settings.
Externa! shaft dimensions, rneasures of bone rnass or volurne (e;g;,
%CA), and femoral neck-shaft angles provide sorne insight into activity
patterns, but precision in behavioral interpretation is dependen! upon
analysis of skeletal tissue distribution via cross-sectional geometry. External dirnensions are lirnited, in that they do not take into account the
distribution of bone in cross section. The relative degree of flattening of
long bone diaphyses, especially in the proximal and midshaft fernur and
midshaft tibia, is related to type and leve! of rnechanical loading and not to
nutritional factors or the actions of specific rnuscle groups.
7 .1
Introduction
7.2
7.2.1
Determinants of form
227
Cranial form in the growing child and the maturing adult is determined by
a complex interaction of intrinsic (genetic) and extrinsic (environmental)
factors. The overall form is principally a product of natural selection (cf.
Herring, 1993; Maynard Smith & Savage, 1959). Animal heritability
(Atchley, 1993) and experimental and observational studies on animals and
humans (Herring, 1993; Kiliaridis, 1995) demonstrate the considerable
infiuence of environment, especially in relation to the cumulative effects of
mastication and mechanical loading of the face and jaws.
Experimental studies involving extirpations of masticatory muscles in
laboratory animals show associated craniofacial skeletal modifications,
especially with regard to a reduction in size and robusticity (e.g., Avis,
1959, 1961; Horowitz & Shapiro, 1951, 1955; Moore, 1967, 1973; Pratt,
1943; Schumacher et al., 1979; Washburn, 1947a, 1947b). Craniofacial
skeletons of animals fed soft foods tend to be smaller and less robust than
animals fed hard foods (e.g., Beecher & Corruccini, 1981, 1983; Bouvier &
Hylander, 1981, 1982, 1984; Bouvier & Zimny, 1987; Corruccini &
Beecher, 1982, 1984; Hinton, 1990; Moore, 1965; Tuominen et al., 1993;
Watt & Williams, 1951; Whitley et al., 1966; and see reviews by Herring,
1993; Kiliaridis, 1995). The profound effects of alteration of masticatory
loading are also demonstrated in the experimental transpositions of
masseter and temporalis muscles in laboratory monkeys (Hohl, 1983). The
anterior relocation of these muscles leads to a number of changes, including
superior facial tilting. These extirpation and translocation studies show
that alterations in mechanical loading produce shifts in masticatory
behavior that result in distinctive craniofacial morphological changes.
7.2.2
Contrary to the assertion that human head form is stable and highly
heritable (e.g., Dixon, 1923; Neumann, 1952; and see Gould, 1996; Marks,
1995), diachronic population studies revea! a high degree of plasticity.
Franz Boas demonstrated that head shape- based on a ratio ofhead length
to breadth (cephalic or cranial index)- of American-born immigrants was
appreciably different from that of their European foreign-born parents
(e.g., Boas, 1912, 1916; also Hrdlicka, cited in Boas, 1916:716); On the basis
of this observation of plasticity, Boas was strongly opposed to the use of
228
cranial form for tracing population history and linking past with living
groups. Rather, he argued that 'the anatomical forms of the present
population and of ancient skeletons do not allow us to draw inferen~es
regardingnatinality of the ancient inhabitants' (Boas, 1902:445). The h1gh
degree of developmental plasticity in twentieth century populations was
furtherconfirmed by Shapiro and Hulse's comparisons of Japanese born m
Hawaii and Japanese immigrants to Hawaii (Shapiro, 1939). The differences between the two were pronounced; the longer the time the immigrant
population was living in Hawaii, the greater the differences with the
ancestral population still living in Japan.
An independent approach to understanding the plasticity of cranial
shape in volved efforts to document t~e influence of culture and behavior. in
past populations. In sharp contras! to ancient British populations, Ke1th
observed that in modern Britons 'many persons have small, contracted
pala tes ... Their noses are narrow; so are their faces' (1950:402). Late Celtic
faces became smaller, reflecting in part the 'change in dietetic which has
occurred since the early years of the Christian era, cooked food and soft
cereals replacing tough meats and imperfectly ground corns' (Keith,
1916: 198). Keith (l 916) noted concomitan! changes in the occlusal surfaces
of teeth, especially with regard to a reduction in tooth wear in later
populations.
.
.
Following Keith's inchoate attempts at relatmg temporal trends m
craniofacial morphology to shifts in masticatory behavior and diet, various
researchers documented other trends in archaeological populations, both
regionally and globally. Weidenreich (1945) recognized the inappropriateness of using cranial shape for identifying racial groups, instead observing a
trend in human evolution - and especially during the Holocene - for
increasingly shorter crania, a process he called brachycephalization. Most
workers argued that long-headed ('dolichocephalic') populations had been
replaced by alien short-headed ('brachycephalic') populations, thus explaining the trend (e.g., Retzius, 1900; and review by Weidenreich, 1945).
Contrary to this consensus, Weidenreich (1945) forcefully argued agamst
invasion and replacement models, by showing the widespread trend of
cranial vault shortening taking place in earlier populations throughout
Europe (see also Sokal & Uytterschaut, 1987; Sokal et al., 1987), the
Middle East, South and Central Asia, and in more recen! settings (and see
Vladescu, 1992).
As with other regions ofthe world, studies ofnative New World groups
in the first half of this century emphasized diffusionistic interpretations of
cranial shape variation, especially arguing that earlier long-headed
'dolichocephals' were replaced by later 'brachycephals' (e.g., Dixon, 1923;
229
Hooton, 1930, 1933; Hrdlicka, 1922a; Hulse, 1941; Newman & Snow,
1942).
Coinciding with and following the publication ofWeidenreich's (1945)
classic article, other researchers began to notice a temporal change in
cranial form, especially reductions in robusticity and/or increasing vault
roundness, in comparison of earlier and later prehistoric North American
Indians (e.g., Anderson, 1967; Boyd, 1988; El-Najjar, 1981; Guagliardo,
1982b; Hoyme & Bass, 1962; Ivanhoe, 1995; Newman, 1962; Newman &
Snow, 1942; Steele & Powell, 1992; Webb & Snow, 1945) and elsewhere
(e.g., Abdushelishvili, 1984; Henke, 1984; Nakahashi 1993 Newman
1951; Rightmire, 1984; Rosing & Schwidetzky, 1984; S~gne, 1976; Smith'.
Bar-Yosef et al., 1984; Suzuki, 1969; Walimbe & Gambhir, 1994;
Walimbe & Kulkarni, 1993; Wu & Zhang, 1985; and references cited
below). In certain settings - especially in North America - cranial
shortening can also be attributed to artificial deformation in sorne
cultures, a practice largely limited to late prehistoric populations. In sorne
cases, changes consisten! with a general pattern of gracilization are also
present in areas of the skull not affected by vault deformation (faces and
jaws; e.g., Larsen, 1982).
Thus, increasing evidence indica tes that the worldwide trend of cranial
shortening and gracilization is much better understood in relation to
masticatory, dietary, and technological changes, especially those associated with the shift from foraging to food production and the consumption of softer foods by later prehistoric populations. These changes in
subsistence practices and their influence on craniofacial anatomy ha ve been
investigated in a number of regions, including the Nile Valley, central
Europe, and the American Eastern Woodlands.
The Ni/e Va/ley
230
l986b; Carlson, l976a; Carlson & Van Gerven, 1977, 1979; Van Gerven et
al., 1973, 1977).
.
Recent anal yses of crania and dentitions from Lower Nubia indica te that
the evidence for the diffusionist model of biological change is less than
compelling. Independent analyses of skeletal and dental di serete and me trie
variables and other lines of evidence suggest that the earlier and later
Nubian populations representa biological continuum with no invasion by
nonindigenous populatipns (e.g., Batrawi, 1946; Berry & Berry, 1972;
Calcagno, l986a, l 986b; Franceschi et al., l 994; Greene, l 972; Mukherjee
et al., 1955; Nielsen, 1970; Van Gerven et al., 1977). Therefore, the
differences in cranial morphology between earlier and later populations
observed by Elliot Smith & Wood Jones ( 191 O), Morant (1925), and others
are best understood in relation to factors not involving population
replacement.
For better understanding of thcse factors, espccially those rclated to
dietary and technological change, Carlson and Van Gerven and their
coworkers (Armelagos et al., 1984; Carlson, 1976a, 1976b; Carlson & Van
Gerven, 1977, 1979; Hinton & Carlson, 1979; Van Gerven et al., 1973,
1977) compared craniofacial morphology in a Nubian-based temporal
sequence, including foragers from the Mesolithic (ca. 12000 BP), initial
agriculturalists from the combined A- and C-groups (3400-1200 BC), and
intensive agriculturalists from the combined Meroitic, X-group, and
Christian horizons (AD 0-1500). These comparisons reveal that Nubian
foragers and incipient agriculturalists have ftat and elongated vaults with
well developed, protruding supraorbital tori and occipitals. In contras!,
Iater intensive agriculturalists have rounded vaults with small and more
posteriorly positioned faces and masticatory muscle attachment sites
(temporalis and masseter) and reduced temporomandibular joint size
(Figure 7.1 ).
Carlson and coworkers posit a n1astfrYlfory-functional hypothesis for
explainingcraniofacial changes in Nubia (Figure 7.2). They argue that the
primary factor influencing Nubian craniofacial anato1ny was the change in
231
The masticatory-functional interpretation of diachronic change in craniofacial morphology in Nubia offers an important means of interpreting
morphological changes elsewhere, especially where food production and
agriculture have supplanted hunting and gathering as a primary mode of
subsistence. Neolithic mandibles from Lepenski Vir, Vlasac, and Vinca in
the Balkans region of central Europe show a reduction in size in
comparison with earlier Mesolithic mandibles (y'Edynak & Fleisch, 1983).
In this region, the change in size of the mandible coincides with the shift
from foraging and fishing in the earlier period to the farming of severa!
grains (e.g., eincorn and emmer wheat); Unlike the foraging adaptation;
the latter dietary focus also involved extensive cooking of food in ceramic
vessels. The mastication of generally softer foods, therefore, resulted in
232
CULTURAL CHANGE
1
1
CHANGE!N
MASTICATORY FUNCTION
1
ALTERATION OF THE
PATIERN OF CRANIO-
FACIAL GROWTH
--'-----
, __I
~
REDUCED GROWTH OF THE
MAXILLO-MANDIBULAR COMPLEX
I_
/
1
\
DECREASED MEGHANICAL
STIMULATION OF THE
PERIOSTEAL MEMBRANE
'-,_T
REDUCED SIZE ANO MORE
INFEROPOSTERIORLY LOCATED
REDUCED ROBUSTICITY OF
THE FACE ANO JAWS
,.
1..
----------
COMPENSATORY BIOMECHANICAL
RESPONSE OY THE CRANIAL VAULT ANO BASE
--
-r
DECREASED
REDUCED GtABELLAR
CRANIAL LENGTH ~ AND OCCIPITAL REGIONS
-o-
~-------L-----~--~
... -- _J
c=J
C)
PROCESSES
MECHANISMS
233
7.2.3
The size and robusticity of the supraorbital torus has been intensively
investigated by biological anthropologists and anatomists for well over a
century (see review by Russell, 1985). The development of the torus is
highly variable in humans, and ranges from a thick, bar-like projection in
early Horno (e.g., Weidenreich, 1943) to mild expression in most recent
human populations, with sorne notable exceptions (e.g., native populatins
234
in the American Great Basin and Australia). Beca use only small muscles of
facial expression are directly associated with the supraorbital torus, the
region is often interpreted as essentially nonfunctional and nonadaptive
(e.g., Owen, 1855; and later researchers). Alternatively, the feature has been
interpreted as deriving from a wide range of plausible (and implausible)
causes, including pathological processes (Virchow, 1872) or various
nonmechanical functions - keeping the hair out of one's eyes (Krantz,
1973), protection from blows to the head (Tappen, 1973, 1979), or as
anatomical sun visors (Boule & Vallois, 1957) (see also Russell, 1985).
The most serious studies of the supraorbital region focussed on the
functional-mechanical paradigm, especially in the context of mastication
and mechanical adaptation and patterns of tooth loading (Russell, 1982,
1985). Based on her reading ofEndo's (1966) experimental and mathematical analysis of the facial skeleton, Russell (1985:343) concluded that the
supraorbital torus is analogous to a beam whereby 'supraorbital development is a function of bending stresses concentrating in the frontal .bone
above the orbits during anterior tooth loading'. The torus can be modelled
'as though it were a beam extending across the superior orbital margins.
This beam is intermittently bent by the downward pull ofthe masticatory
muscles and the upward push of the bite force.' Thus, with increased
chewing (or related behaviors}, greater bending stress on the supraorbital
torus should result in greater bony development in the glabellar region in
particular and the supraorbital torus in general.
Evaluation of Russell's argument in light of Endo's original published
discussion indicates that the supraorbital region is not highly stressedeither
in anterior incisa! loading (as interpreted by Russell, 1985) or even in
posterior Ioading (see Hylander & Johnson, 1992; Hylander et al., 1992;
Lieberman, 1995; Picq & Hylander, 1989). In fact, careful reading of
Endo's experimental results suggests that he could find little evidence of
stress in the region during isometric biting on the anterior teeth (see Endo,
1966). Additionally, biomechanical analysis ofld World monkey crania
suggests that the link between anterior dental loading and compensatory
remodeling in the browridge is unfounded (Ravosa, 1988).
Although the size and morphology of the supraorbital torus is not
understood, comparison of supraorbital development in temporal sequences of recent humans from archaeological settings suggests that
browridge size is best thought of as a general indicator of craniofacial
robusticity. Carlson (1976a) found that the torus was more developed in
the preagricultural foragers than in farmers from Nubia (and see above). In
the American northern Great Plains (South Dakota and North Dakota),
comparison of size and morphology of the adult female and male
235
236
A rich body of ethnographic evidence indica tes that very heavy mechanical demands are placed on the craniofacial complex of Eskimos, specifically involving heavy use of jaws and teeth in masticatory and extramasticatory functions. In his observations of Eskimos, De Poncins (1941 :7172) noted, 'They had long since stopped cutting the meat with their circular
knives; their teeth sufficed, and the bones of the sea] cracked and splintered
in their faces. What those teeth could do, I airead y know. When the cover of
a gasoline drum could not be pried off with the fingers, an Eskimo would
take it between his teeth and it would come easily away. When a strap of
sealskin freezes hard - and I know of nothing tougher than sealskin - an
Eskimo will pul it in his mouth and chew it soft again' (quoted by
Hylander, 1977:142).
Drawing on these various lines of evidence, biological and behavioral,
Hylander (1977) argued that Eskimo craniofacial morphology longobserved by biological anthropologists (e.g., forwardly placed zygomas) is
oriented toward maximizing the efficiency and power of chewing, especially
involving anterior tooth use: the craniofacial complex is suited to the
generation and dissipation of pronounced, vertically oriented masticatory
forces in the front of the mouth. This assessment is confirmed by analysis of
position of attachrnent sites for masseter and temporalis muscles and
incisors in a sample of prehistoric Inuit crania (Spencer & Demes, 1993).
The anteriorly placed masticatory muscles and posteriorly placed incisors
'indica te an increased efficiency for the application of either high magnitude or repeated bite forces on the anterior dentition' (Spencer & Demes,
1993:15).
Other high-latitude foragers display pronounced craniofacial robusticity. Like Eskimos, crania from Tierra del Fuego and Patagonia, South
America, bear robust supraorbital tori and anteriorly placed zygomas,
sagittal keeling, occipital tori, and pronounced attachment siles for the
temporalis muscle (Lahr, 1995). The functional-masticatory paradigm
(Hylander, 1977; Spencer & Demes, 1993) best explains the similarities
between Eskimos and Fueguian/Patagonians, especially in regard to the
common skeletal responses to highly demanding masticatory regimes in
these two different settings.
Sorne workers have argued that the round head of high-latitude
populations would be best suited for cold adaptation, as a sphere
maximizes volume for heat retention and minimizes surface area far heat
loss prevention (e.g., Beals, 1972; Beals et al., 1984; Crognier, 1981).
237
7.2.5
7.2.6
Tori located on the hard palate and lingual corpus and alveoli of the
mandible are the focus of attention regarding genetic vs. nongenetic
environmental influences on skeletal variation (Halffman et al., 1992;
Hauser & De Stefano, 1989; Morris, 1981) (Figure 7.3). Sorne workers
conclude that tori found in archaeological remains are indicative of high
mechanical demands placed on the masticatory apparatus (e.g., Halffman
et al., 1992; Hooton, 1918; Hrdlicka, 1940b; Pedersen, 1944; Scott et al.,
1991), but others argue that tori are largely genetically controlled (see
Hauser & De Stefano, 1989; Morris, 1981).
There are relatively high frequencies of palatine tori in far northern and
circumpolar populations, including Icelanders (Hooton, 1918), Lapplanders (Schreinder, 1935), and Eskimos (Hylander, 1977; and discussion
above). Torus prevalence is also high in Medieval Norse living in Iceland
and Greenland in comparison with Europeans generally (e.g., Mellquist &
Sandberg, 1939; Pedersen, 1944). In order to address the question ofwhy
Medieval Norse and indigenons Arctic groups have developed a convergence in torus expression with Eskimos and other circumpolar groups,
238
239
7.2. 7
Figure 7.3. Superior (top) and posterior (bottom) views of large palatine torus
(A) and maxillary torus (B); Norse male from Benedictine Nunncry, Eastern
Settlement, Greenland. (From Halffman et al., 1992; reproduced with
permission of authors and John Wiley & Sons, Inc.)
Halffman and coworkers (1992; Scott et al., 1991) studied torus frequency
and size in a series of Medieval Norse skeletal remains from Norway,
Greenland, and Iceland dating to the eleventh to fourteenth centuries.
Temporal comparisons revea! that later Norse from Norway, Greenland,
and Iceland have a significantly higher prevalence of tori than do early
Norse from Greenland. Tori prevalence approaches 100% in later populations, which is amongthe highest in the world (cf. Halffman et al., 1992: 156;
Hauser & DeStefano, 1989). Tori increase in size in older adults, suggesting
that the trait is strongly influenced by age. The pattern of increase in torus
frequency and size within a relatively short temporal span (severa! hundred
Growth in the skull is not quiescent once adulthood is reached. Like the
postcranial skeleton, appositional borre growth in .the skull continues
during adulthood. These changes are documented . in cross-sectional
240
241
Middle (n = 58)"
Mean
SD
Mean
SD
Mean
SD
Vault Jength
Vault breadth
Vault height
Pace height
Mid-face breadth
180.9
138.5
142.3
122.9
99.8
6.3
5.6
5.4
5.8
5.2
182.3
137.I
140.3
122.l
99.9
7.1
5.8
5.3
6.8
4.8
180.4
138.5
140.4
122.1
100.3
6.0
4.5
4.3
4.2
4.8
Fe1na/es
Vault Jength
Vault breadth
Vault height
Face height
Mid-face breadth
173.0
134.0
136.5
114.5
96.4
5.9
4.3
4.2
6.0
4.2
173.6
134.7
137.5
112.6
95.5
5.1
4.5
5.2
5.8
5.2
174.2
135.l
137.3
114.4
97.2
6.7
6.6
4.8
7.2
4.4
Measurement
Males
242
7.3
7.3.l
243
244
reduction of the jaw bone has progressed through the course of human
micro-evo1ution, resulting in disharmony between the sizes of teeth and
jaws' (Inoue et al., 1986: 164; see also Suzuki, 1969). To test this hypothesis
further, malocclusion prevalences of Jomon period foragers (divided into
an early anda late sample}, Yayoi period early farmers, and Kofun period
protohistoric farmers fro1n n1ostly southern Ja pan were compared. Factor,
7.3.2
245
246
247
reduction are based on the premise that reduction occurs in the absence of
selection (cf. Brace et al., 1991; Macchiarelli & Bondioli, 1986), but these
models exclude a broader consideration of the potential health consequences of occlusal abnormalities. Therefore, it appears that there is an
adaptive advantage to maintaining a harmonious relationship between
teeth and the skeletal structures supporting them (for an alternative
perspective, see Kieser et al., 1985).
- 1500
NE
.. 1400
~
~ 1300
o 1200
-~
*g_
1100
"O
"'E
7 .4
1000
(/)
Hominid
caries, periodontal disease, and tooth loss (Calcagno & Gibson, 1991;
Inoue et al., 1986). Caries and dental impactions can also lead to abscesses
and localized infections, which can progress to more severe types of
systemic infections (e.g., gangrene, septicemia, osteomyelitis; Calcagno &
Gibson, 1991). Beca use systemic infections are potentially life-threatening,
teeth in populations consuming soft, cariogenic foods may be under
selection for reduction in these circumstances (Calcagno & Gibson, 1991 ).
However, small teeth in highly abrasive masticatory environments may
wear too rapidly, thus resulting in premature loss of crown height and pulp
exposure. Exposed pulp is highly susceptible to bacteria! infection (pulpitis). This suggests, then, that teeth may also be under selection for large
occlusal size in abrasive environn1ents.
The above discussion highlights the extreme ends of variation involving
the costs and conseq uences of soft and hard diets in relation to mastication,
tooth size, and potential selective conditions. Most human populations lie
somewhere between these ends of the spectrum of masticatory adaptation.
Tooth size is, therefore, most likely to be a product ofan ongoing 'selective
compromise' involving the promotion of optimum dental health among
other things (Calcagno & Gibson, 1991 ). Severa] other models of tooth size
The use of the teeth in eating involves a two-stage process: first, the initial
preparation of food with the anterior teeth; and second, the reduction of
food with the posterior teeth. These activities result in the wearing of
occlusal surfaces as the upper and lower teeth come into contact with each
other and with the food being prepared or reduced. There has been a shift
away from the use of the teeth toward the use of the hands in the
manipulation of the environment and a reduction in the importance of
mastication in the consumption of processed foods in the evolution of the
Hominidae. It is also true that the presence of significant amounts and
variable patterns of dental wear in past and living human populations
indicates the continued importance of teeth in survival and adaptation.
Tooth wear is variously defined in the literature. Two forms of wear,
abrasion and attrition, are most commonly discussed (Townsend et al.,
1994). Abrasion is caused by contact between the tooth and the food or
other solid exogenous materials, especially as food is forced over occlusal
surfaces. Attrition is caused by tooth-on-tooth contact in the absence of
food or various other abrasives. Additionally, erosion - the loss of tooth
surfaces dueto chemical dissolution - is sometimes considered as a form of
wear (e.g., Davis & Winter, 1980; Linkosalo & Markkanen, 1985). Because
of the difficulty of distinguishing abrasion, attrition, and erosion, this
discussion regards 'wear' as including any combination thereof (also see
Wallace, 1974). Tooth wear varies widely between human populations.
Owing to localized behavioral characteristics, and differences in cultural
practices, age, sex, diet, and orofacial morphology, it provides enormously
important information on earlier foodways and masticatory behavior (e.g.,
Benfer & Edwards, 1991; Molnar, 1971, 1972; Molnar et al., 1983; Molnar
& Molnar, 1990; Powell, 1985; Richards, 1990; Richards & Miller, 1991;
Walker et al., 1991). The significance of dental wear in relation to diet was
succinctlysummarized by Walker, who noted that 'From an archaeological
standpoint, dietary information based on the analysis of (wear) is of
248
249
considerable value since it offers an independent check against reconstruction of prehistoric subsistence based on the analysis of floral, fauna! and
artifactual evidence' (1978:101).
Dental wear variation is reported as either severity or forro or both.
Wear can also involve substantial losses in the regions of contact between
teeth (Begg, 1954; van Reenen & Reinach, 1988; Wolpoff, 1971), resulting
in reduction in length (mesial~distal). Although wear in its severe forro
can predispose a tooth to discase (e.g., pulpitis, caries) and loss, tooth
wear is a normal physiological process rather than a disease (contra Wells,
1975).
The process oftooth wear is well understood: it commences with loss of
occlusal enamel, followed by deposition of secondary dcntin serving as a
protective zone overlying the pulp chamber. In extreme forros of wear, ali
enamel is removed from the occlusal surface, leaving a ncarly continuous
Figure 7.5. Lingual tilting of mandibular first molar; Tutu, St. Thomas, U.S.
Virgin Jslands. Note wear on superior bucea! roots and pronounced Occlusal
and interproximal wear on all teeth. (Digital photograph by R. P. Stephen
Davis.)
height (e.g., Hartnady & Rose, 1991; Larsen & Kelly, 1995). Moderate to
severe wear results in a reduction in tooth size dueto the combined effects
Sorne of the most comprehensive infarmation on the behavioral significance oftooth wear is from temporally successive series ofhuman remains.
Investigation oftooth wear and the mechanical environment reveals a high
degree of consistency with skeletal indicators of masticatory stress.
Populations with high levels of mechanical demand and/or reliance on
abrasive diets have relatively advanced wear (e.g., Barondess & Sauer,
1985; Hansen eta!., 1991; Hartnady & Rose, 1991; Hemphill, I992; Marks
et al., 1988; Molnar et al., 1983; Powell & Steele, 1994; Scott et al., 1991;
Walker, 1978). Traditional populations undergoing shifts from diets
containing tough foods to diets dominated by processed foods during the
twentieth century show reduction in wear (e.g., Davies & Pedersen, 1955;
Staz, 1938). Importan! differences in interproximal wear and specific
patterns of wear also help infarm our understanding of masticatory
behavior in human populations.
7.4.1
Macrowear
250
251
252
253
Archaic
(n)
4.9
(8)
5.4
(6)
5.3
(6)
6.1
(18)
6.4
(3l)
7.1
(22)
4.8
(33)
5.3
(11)
5.4
(9)
5.9
(22)
6.0
(8)
4.3
(43)
4.6
(22)
4.9
(lO)
4.7
(6)
Woodland
(n)
Mississippian
(n)
Archaic
4.8
(7)
5.5
(16)
6.0
(7)
6.3
(5)
6.7
( 18)
6.7
(30)
6.9
(8)
4.2
(14)
5.0
(32)
5.4
(13)
5.7
(9)
6.2
(8)
4.0
(28)
4.3
(42)
5.1
(3)
4.5
(3)
(n)
Woodland
(n)
Mississippian
(n)
254
255
(b)
Figure 7.6. Lateral views of tooth wear in Nublan A-group agriculturalist (a)
and Eskimo hunter-gatherer (b) showing greater angle of wear plane in the
former. (From Smith, 1984; reproduced with permission of author and John
Wiley & Sons, Jnc.)
(a)
(b)
Figure 7.7. Occlusal view of tooth wear in Nubian X-grcup agriculturalist (a)
and Mesolithic hunter-gatherer (b) showing cupiJed occlusal wear in the former
(first molars). (From Smith, 1984; reproduced with permission of author and
John Wiley & Sons, lnc.)
(Figure 7. 7). Overall, wear plan e angles and form reflect the kinds of foods
being eaten as well as the manner in which they are prepared. Smith drew
the general conclusion that these differences were related to greater
'toughness or fibrousness' of the diets in foragers than in farmers.
Therefore, the pattern of flat and cupped wear that Wood Jones (I 91 O)
observed in bis comparisons of Predynastic and Christian era populations
in Nubia reflects the distinctions between the respective hunter-gatherer
and agriculturalist groups documented in Smith's study. Additional
confirmation of Smith's findings has been provided in other settings
256
257
!'"
258
259
inal males than females, females in the lower Murray River valley exhibit
less anterior wear than males (Molnar et al., 1989). Further up the Murray
River valley, females exhibit much more wear than males, a pattern that is
reminiscent of the Narrinyeri and Kaurna groups (cf. Richards, 1984).
Although the reasons for these differences are unknown, they attest to the
presence of highly variable wear patterns between closely related populations in what is oftencharacterized as a homogeneous region (Molnar et al.,
1989).
Gross wear differences by social group and rank ha ve been identified in
past societies. In the Medieval Edo period of Ja pan, members of the elite
Shogun class had virtually no occlusal surface wear, unlike lower-status,
non-Shogun individuals (Suzuki, 1969). This indica tes a lcss mechanically
demanding, less abrasive diet in thc elite than the remainder of Japanese
society. The presence of narro\.v faces, re<luced size of the maxillae and
mandibles, and gracile masticatory n1uscle attachment sites in Shogun
individuals corroborates a reconstruction of the consun1ption of soft,
Extramasticatory wear
(1991) note that 'teeth can show the effects of a wide variety of activities
unrelated to eating that rcsult in unusual, and at times highly distinctive,
patterns of abrasion, crown fractures, or trau1natic tooth loss'. Until
recently, a variety ofhuman populations havc used their teeth in extramasticatory ways, and in no other group is this expressed as well as among
Eskimos (Cybulski, 1974; Larsen, 1985; Leigh, 1925; Merbs, 1983; Milner
& Larsen, 1991; Molnar, 1972; Pedersen, 1952; and many others). The
following discussion considers unintentional changes on teeth arising from
extramasticatory activities. (For a discussion of intentional mutilations of
teeth, see Milner & Larsen, 1991.)
Modifications involving transversely oriented grooves on worn occlusal
surfaces of permanent mandibular incisors and canines are especially
260
261
figure 7.9. Lingual wear on maxillary permanent left first incisor; Tut, St.
Thomas, U.S. Virgin lslands. (Digital photograph by R. P. Stephen Davis.)
262
mandibular teeth contrasts with the pattern ofhigh frcquency and prescncc
mostly in maxillary teeth in dental samples from Brazil (Turner &
Machado, 1983), Panama (Irish & Turner, 1987), the Virgin lslands (C. S.
Larsen et al., unpublished manuscript), and Texas (Hartnady & Rose,
1991). Thercfore, LSAMAT appears to be a primarily Ncw World
pheno1ncnon.
7.4.3
Microu1ear
masticatory behavior not available from the study of gross wcar alone
(Teaford, 1991 ). The superior depth of fcus and highly detailed resolution
in SEM analysis has made it an especially powerful too! for documenting
minute aspects of wear (Teaford, 1991 ). Photographs (called micrographs)
taken with the SEM instrument represen! an clcctronic map of brightness
and contrast on the tooth surface. Becausc the analysis of microwear
features (e.g., pits and scratches) requires high magnification (e.g., x 500),
only small areas can be assessed at any one time. However, the details made
visible in SEM analysis potentially allow precise determinations of tooth
use.
263
264
(a)
(b)
premaize agriculturalists and Mississippian (AD 1000-1350) maize agriculturalists in the lower Illinois River valley of western Illinois increased with
age during both periods (Bullington, 1991 ). Reflecting the difference in use
of anterior and posterior teeth, the incisors have relatively more scratches
265
than pits and the molars have more pits !han scratches. These differences
indicate the use of the incisors in initial processing and the molars in
crushing hard objects. The earlier group focussed on wild plants and
animals in combination with reliance on various starchy seeds and with
hard seed coats. The later group continued to utilize these food items, but
with partial replacement by maize. In addition, these foods were probably
boiled in ceramic vessels for long periods of time, thus reducing the
toughness of foods consumed by prehistoric populations (Bullington,
1991 ). Unlike the. changes observed in the Mississippi River valley and the
Georgia Bight, comparisons of the Middle Woodland and Mississippian
groups revealed no differences in microwear. Bullington (1991) suggests
that the lack of temporal change may reflect the fact that her study focusses
exclusively on young juveniles; other investigations focus on adult microwear. There is a strong similarity in microwear between the foragers and
farmers overall, but the two groups are distinctive in the youngest cohort
(ca. six to 12 months). Mississippian teeth have lower feature frequencies
than Middle Woodland teeth. This suggests that very young children in the
later period were consuming softer foods than very young children in the
earlier period.
Similar changes in microwear in relation to the shift from hard- to
soft-textured diets ha ve been identified in a number of settings in the Old
World. Comparison of canines and molars from two Neolithic (ca. 2000
ne) and two historie (AD 175()-1800) dentitions from western coastal
Kyushu, Japan, shows a change from generally long, wide striations to
thin, narrow striations (Hojo, 1989). Hojo (1989) suggests that the la ter diet
included smaller abrasives than the earlier diet.
Change in occlusal microwear in relation to the shift from foraging to
dependence on cereal grains in western Asia has been investigated via
comparison of dentitions from the earlier discussed Neolithic settlement at
Abu Hureyra in the Euphrates River valley in northern Syria (Molleson,
1994; Molleson & Jones, 1991; Molleson et al., 1993). Microscopic analysis
of molars provides corroborative evidence for the study of gross wear.
Comparison of pit diameter, feature frequency, pit density (percentage
determined from number and size ofpits), and ratio of striations and other
linear features to pits shows a clear shift in microwear in deciduous and
permanent teeth through time. The change in wear is especially conspicuous in the shift from foraging in the Mesolithic to early agriculture in the
pre-pottery Neolithic, being accompanied by a dramatic increase in
microwear feature density. Molleson and coworkers (1993) attribute this
trend to a shift from a relatively soft diet based on consumption of roots
and small, wild grains to a few cereal types prepared with grinding stone
266
Figure 7.11. Chipped teeth; Greenland Eskimo. (From Hansen eral., 1991;
reproduced with pennission o authors and Greenland National Museum and
Archives.)
striations than do agriculturalists. The greater frequency of bucea! striations in plant consumers may be dueto the presence of abrasive phytoliths
(Lalueza et al., l 996).
pre-pottery Neolithic, followed by a decline with the adoption of agriculture and agricultura! intensification (Pastor, l 994). In general, trends in
microwear indica te the higher prevalence of finer scratches in the huntergatherers and incipient agricu1turalists, whereas microwear features become coarser in more intensive agriculturalists. The latter characteristic
reflects an increased consumption of cereal grains and use of grinding
267
7.4.4
li
1
l
'
268
7 .5
269
Isotopic analysis
8.1
coprolites, and tools used for extracting food from the environment or for
processing it once it is acquired. These approaches do not necessarily
represen! the proportions of foods or food classes consumed by past
populations. For example, the notoriously poor preservation of plants in
many archaeological contexts can prevent the documentation of their role
8.2
Isotopic analysis
8.2.1
Background
271
Isotopes are chemical elements that share the same number of protons and
electrons, but differ in the number of neutrons. Unlike radioactive or
unstable isotopes (e.g., 14 C), stable isotopes of the same element do not
transmute over time. Most elements exist in two or more isotopic forms. Of
the severa! hundreds of stable isotopes across ali elements, 1Oelements have
at leas! two isotopes with biological significance. Two of these 1Oelements
have received the preponderance of attention by anthropologists in
reconstructing and interpreting earlier diets, namely carbon (C) and
nitrogen (N).
8.2.2
Carbon has two stable isotopes, 12C and 13 C. Field and laboratory studies
involving controlled feeding experiments have shown that stable carbon
isotope ratios in an animal's tissues, including bone, reflect the ratios of diet
(Render et al., 1981; DeNiro & Epstein, 1978; Tieszen et al., 1983). The
relative abundance of isotopes between dietary resources is quite small.
Thus, the ratios in tissues are expressed in parts per thousand (read as parts
'per mil' or %o} relative to an international standard (marine fossil,
Belemnitella, from the Peedee geological formation in South Carolina
[PDB)), as delta (J) values. Thesevalues denote differences that originate in
plant photosynthetic pathways, including either C3 (Calvin-Benson}, C4
(Hatch-Slack), or CAM (crassulacean acid metabolism). C4 plants discriminate Iess against the isotopically heavier Be isotope when using C02,
the carbon source for ali terrestrial plants, from the atmosphere. As a
result, C4 plants have less negative J 13 C values than C 3 plants. In temperate
areas, most plants are the C3 variety (e.g., sorne grasses, trees, shrubs,
272
Isotopic analysis
Collagen requires essential amino acids for its formation and, therefore,
largely reflects the 13 C of the protein componen! of diet. The apatite
signatures represen! the whole diet, which may include carbohydrates and
fats, in addition to protein (Ambrose et al., 1995; Ambrose & Norr, 1993;
Cooke et al., 1996; Krueger & Sullivan, 1984; Tieszen & Fagre, 1993).
Experimental research by Ambrose & Norr (1993) contradicts earlier
discussions and studies linking o13 C values in collagen to whole diet (e.g.,
Schoeninger, 1989; van der Merwe, 1982) and bone apatite values to
carbohydrates and fats (e.g., Lee-Thorp et al., 1989).
Marine plants ha ve '5 13 C values that are between the values ofCi and C4
terrestrial plants, owing to variation in their carbon sources, ranging from
AD
273
o -10
1lt
"'~
~ -15
"
"'"' -20
e:
"
::;;
-25
-4000
-3000
-1000
-2000
Age
(ec/o)
f f
!i1 {t
1000
-~
274
Isotopic analysis
ha ve been a greater part of the diet in the outlying areas around the late
prehistoric Mississippian center of Cahokia than in Cahokia itself
(Buikstra & Milner, 1991). These differences may represen! social distinctions between the core and hinterlands of the Cahokia chiefdom. Analysis
of o"C values in the late prehistoric Ohio River valley drainage indicates a
link between social complexity and maize production (Schurr & Schoeninger, 1995). Comparison of populations from late prehistoric tribal-society
Fort Ancient sites (Bau1n, Gartner, Feurt, Sun Watch) with contemporary
populations from organizationally 1nore co1nplex Mississippian sites
11
j
275
276
The picture of dietary ecology has changed markedly with the work on
stable isotopes in Mayan populations. Temporal and spatial comparisons
of 0 13 C values from Lamanai and Pacbitun, Belize, indica te that prehistoric
Mayans had less emphasis on maize than did sorne other Mesoamerican
groups (e.g., Tehuacan) (Whiteet al., 1993, 1994; White & Schwarcz, 1989).
Nonlinear temporal changes suggest a variable emphasis on maize in
Belize. Pre-Classic populations (1250 BC-AD 250) have Iess negative ;De
values ( - 12.4%0) at both siles, suggesting strong reliance on maize. Late
and Terminal Classic groups have increasingly negative iillC values al
Lamanai and less negative values at Pacbitun, indicating respective
decrease and increase in maize consumption at the two Iocalities (White et
al., 1993). Convergence of diet and reduced reliance on maize at the very
end ofthe Terminal Classic period is inferred by more negative .JllC values
in both populations.
Following the collapse ofthe Classic period Maya in the eighth to ninth
centuries AD, ii 13 C values at Lamanai (data are not available for Pacbitun)
show a marked reversa] in the trend of decreased maize consumption
documented for the Late and Terminal Classic periods. Post-Classie and
Historie period Maya skeletons have substantially less negative values
(-9.3%0, -9.9%0, respectively), indicating an increased reliance on maize.
/sotopic analysis
277
278
Adult males show less negative values than adult females at both Pacbitun
(White et al., 1993) and Copn (Reed, 1994). Lamanai females and males
show no difference in values (White & Schwarcz, 1989). Reed (1994) and
White and coworkers (1993) contend that sex differences in diet at Pacbitun
and Copn represent variation in socioeconomic status, with males
consuming more maize than females. With regard to Copn, Reed observes
that this difference 'parallels the observation of higher frequencies of
anemia, infection, and a statistically significan! higher rate of caries in
females than in males' (1994:216). Based on discussions presented in
Chapters 2 and 3, his findings of reduced consumption of maize in females
actually contradicts these other data sets. For example, greater caries
prevaience in females suggests more emphasis on maize in women than in
men in this setting. Therefore, although both isotopic and caries analyses
point to possible differences in diet by gender, the contradictory results do
not provide a clear picture of what these differences may have been. At
Lamanai, like the isotope evidence, dental caries prevalence data suggest
distinctive differences in diet between females and males (White, 1994).
The variable nature of diet in comparison of isotope and dental caries
suggests that these data sets may reflect somewhat different aspects of diet.
Dental caries is caused by acids produced by bacteria] metabolism of
Isotopic analysis
279
280
Isotopic ana/ysis
281
grass, was present in central Europe by the late fifth millennium BC (Murray
& Schoeninger, 1988). Like maize in la ter contexts, millet has been assumed
to ha ve been primarily used as food for farm animals. Analysis of collagen
from !ron Age skeletons from Magdalenska Gora, Slovenia, indicates less
negative b"C values, in sharp contrast with virtually ali other noncoastal
European samples from various time periods (cf. B. Kennedy, 1989).
Because marine foods were probably not part of the diets of this group,
consumption ofC4 plants or animals consuming C4 plants best explains
these values (Murray & Schoeninger, 1988). lt is unlikely that C4-consuming animals contributed appreciably to !ron Age diets in Slovenia, because
isotope values derived from animal remains are very negative, unlike the
values determined from human remains.
Millet was also an important cultigen during the Neolithic in northern
China, especially in the H uang He (Yellow River) basin (Schwarcz &
Schoeninger, 1991; van der Merwe, 1992). Analysis of 13 C values indicates
that millet contributed well over 50% of the carbon in the diets of these
groups. From the period of the fifth century BC to the second century AD,
isotope values became increasingly negative, indicating a shift from C4
(millet) to predominantly a Ci diet based on rice and wheat. Today, the
isotopic signature of dietary carbon has again shifted to less negative
values, this time relecting a C4, maize-based diet (van der Merwe, 1992).
Along with millet, sorghum - also a C4 domesticate - dominated the
archaeological record for much of Nubian prehistory (White & Schwarcz,
1994). Analysis of bone and other tissues.indicates that, although these
plants were present in substantial amounts, diets were based principally on
Ci cultigens (e.g., wheat and barley) (White & Schwarcz, 1994), but
analysis of hair shafts from mummies indicates seasonal shifts from Cito
C4 staples (White, 1993). Isotopic evidence indicates a shift towards
consumption of more C4 plants in the X-group (AD 350-550), a period
characterized by political instability, alterations in trade patterns, and
decreased water availabi!ity as the Nile River lowered. In the following
Christian period (AD 500-1400), more negative isotope values revea! a
decline in c. plant consumption. This shift in diet appears to have
accompanied an increase in elevation of the Nile and generally improved
economic conditions.
Marine diets and coastal environments
282
orientation are documented in the comparison of Mesolithic and postMesolithic populations (e.g., Lidn, 1995; Tauber, 1981). In coastal
Mesolithic Danish populations, for example, generally less negative values
(- 11%0 to - 15%0) indica te a reliance on marine foods (Tauber, 1981). The
values are similar to those of populations known to ha ve depended on sea
food (e.g., Greenland Eskimos). In la ter coastal groups (Neolithic, Bronze
Age, early !ron Age), 1i 13 C values are progressively more negative (- 18%0
to - 23%0), indicatinga shift to terrestrial CJ foods, such as domestic plants
and farm animals. Isotopic evidence indica tes that post-Mesolithic Danish
and Swedish populations consumed few marine foods, despite close
proximity to them (Lidn, 1995; Tauber, 1981). Similarly, late Neolithic
populations from Alepotrypa Cave, coastal southern Greece, have very
negative ne values, for both collagen (mean= - 19.9%0) and apatite
(mean= - 13.1 %0) (Papathanasiou et al., 1995). These findings indica te
that the diets ofthesc groups were largely focussed on terrestrial C3 plants
and animals. The narrow rangc of values (collagen: - 19.5%0 to - 20.2%0)
indicates a rcmarkably high degree of homogeneity in diets.
An abrupt shift from tnarine to terrestrial food resources in the
Mesolithic and the less negative values are Neolithic. The presence of less
variability in the Neolithic period suggests an increasing homogeneity in
diet during the period of increased consumption of plan! and animal
domestica tes (Lubell et al., 1994). Unlike the pattern of increased prevalence of dental caries in New World settings with the increasing reliance on
plant domesticates, there is a marked decline in dental caries prevalence,
nuinber of carious tooth surfaces, and premortem tooth loss in permanent
mandibular molars in late Mesolithic and Neolithic Portuguese in comparison with earlier populations from the region. The relatively high
prevalence of dental caries in the Mesolithic period is probably related to
the consumption of cariogenic plants such as nondomesticated figs (Lubell
et al., 1994; and sec Chapter 3).
8.2.3
Nitrogen has two stable isotopes, 14 N and 15N. Field and laboratory
feeding studies demonstrate that stable nitrogen isotope ratios in an
anirnal's tissues, including bone, reflect similar ratios in the diet (DeNiro &
Epstein, 1981; Hare et al., 1991; Wada, 1980). The ratios determined from
Jsotopic analysis
283
284
in sorne desert settings may be explained by these factors (e.g., Ambrose &
DeNiro, l 986a, l 986b; Aufderheide, Tieszen et al., 1988; Heaton et al., 1986;
Schoeninger, 1995b). In sorne settings, terrestrial animals living in arid
environments have higher ii 15N values than marine animals. Because of
climatic and/or other ecological variables, Late Stone Age coastal and
interior foragers in Southwestern Cape Province, South Africa, actually
have nitrogen isotope ratios inverse to those expected, but car bon isotope
values clearly identify marine vs. terrestrial differences in resources (Sealy,
1986; Sealy & van der Merwe, 1985, 1986; Sealy et al., 1987). In other
settings, stable nitrogen isotope ratios are useful far identifying consumers
ofterrestrial vs. marine foods (e.g., Schoeninger& DeNiro, 1984; Schoeninger et al., 1983). These various findings indica te that local factors can play an
importan! role in isotopic composition and diet (see Ambrose, 1993).
Isotopic analysis
285
18
16
&
z~ 14
"'
12
10
COof90
o
o
o
3
Age (years)
Figure 8.2. Stable nitrogen isotope values rom birth to eight years; Prospect
Weaning
similar to the mother's and there is a lag in the registration of the colla gen in
the nitrogen isotopic signature (Katzenberg, 1993b; Katzenberg et al.,
1996; Tuross & Fo gel, 1994). The age of weaning coincides with morbidity
indicators of physiological stress, such as hypoplasias and circular caries in
the few studies in which nitrogen isotope values ha ve been determined (e.g.,
Reed, 1994). Moreover, the pattern of change in J 15N values corresponds
with the known age of weaning in populations in which data are historically
documented, such as in Euroamericans (e.g., Katzenberg & Pfeiffer, 1995).
Osteoporosis and nitrogen isotope ratios
286
Isotopic analysis
1986b). These findings suggest that physiological disruption, and not diet,
may be responsible for the variation in nitrogen stable isotopes (and borre
loss) in this population (White & Armelagos, 1997). More generally, it
strongly suggests that stable nitrogen isotopes are susceptible to nondietary
- especially physiological - factors and may serve as an indicator of
osteoporosis in past human populations.
8.2.4
"
14 r-
..
L.
'
LL.t,
L,
oc
12
'
..
"'z
-..,
"
r-
10
..
..
L.
L, L,
In many coastal areas of the New World, where marine foods and maize
were simultaneously consumed by human populations, the overlapping
carbon isotope ratios for both foodsprecludes the discrimination of diets,
at leas! with regard to assessing the relative contribution of marine foods
vs. maize. In order to circumvent this complication, the use of bivariate
plots of 15N and 13 C values has been advocated (Cooke et al., 1996:
Schoeninger et al., 1983, 1990). In the Georgia Bight, archaeological
evidence indica tes that marine foods continued to be heavily used throughout prehistory and into the contact period (Larsen, 1982). Maize consumption is largely based on circumstantial evidence in this region, including
changes in settlement (increased population size and aggregation), increasing social complexity, and increasing morbidity (e.g., dental caries,
periosteal reactions; see Chapters 2 and 3). Owing to poor preservation of
plan! remains in late prehistoric archaeological sites, dietary reconstruction is inconclusive.
Analysis of collagen samples from prehistoric foragers and farmers and
early and late contact mission Indians alleviates the incomplete picture of
die!. Isotopic analysis reveals a distinctive temporal trend showing increasingly less negative ,jllC values and less positive 1'N values (Hutchinson et
al., 1996; Larsen, Schoeninger et al., 1992; Schoeninger et al., 1990) (Figure
8.3). This trend indicates an increasing focus on terrestrial plants (maize)
and animals coupled with perhaps a decreasing reliance on marine
resources. This shift commences during the twelfth century AD, reaching its
peak in Spanish mission native populations inhabiting St. Catherines
Island, Georgia, in the early to middle seventeenth century and in the later
descendant groups on Amelia Island, Florida.
Additional analysis of collagen from the late prehistoric Irene Mound
site on the north Georgia coast indicates that, although maize generally
increased in importance, there is a marked decrease in 13C values for !he
period immediately prior to European contact, suggesting that use of maize
temporarily decreased following its initial introduction in the region
(Larsen, Schoeninger et al., 1992). The decline in maize consumption may
287
E:
"...
. ..
.. .. '
L,
L.
.o
.e
E,B,
:~
,ce .e
,e.e
,e
8 -
B1'c%. (PDB)
Figure 8.3. Plot of Georgia Bight stable carbon and nitrogen isotope 'values;
E, early preagricultural; L, late preagricultural; A, agricultura!; C, contact era.
Note the increase in carbon values and decrease in nitrogen values. (From
Larsen, Schoeninger et al., 1992; reproduced with pennission of Wiley-Liss,
Inc., a division of John Wiley & Sons, Inc.)
288
numerous potential foods from these settings. Sorne reef fish ha ve higher
c)l3C values and lower b15 N values than other ocean fish. The analysis of
car bon and nitrogen isotope ratios in archaeological human samples from
Jsotopic analysis
289
This temporal shift instable carbon isotope ratios suggests that Caribbean
populations becan1e increasingly marine-oriented in later prehistory.
8.2.5
Strontium, an alkaline earth element, expresses isotope ratios (87 Sr/86 Sr)
that identify the relative contributions of marine and terrestrial food
Oxygen isotopes
290
skeletal and dental apatite is in equilibrium with body water, the phosphal!'
oxygen isotope composition of these hard tissues directly reflects t h!'
temperature and climate at the time the organism was alive (Fricke et al.,
1995). In orderto test the hypothesis that hard tissues from archaeological
settings can be used to track the history of regional clima tes; Fricke ami
coworkers (1995) determined the temporal change in 18 0 values from
archaeological dental enamel (Eskimos and Europeans) in coastal western
Greenland and Denmark for the periods preceding, during, and following
the so-called 'Little Ice Age' of the Medieval period. In a comparison o\'
la ter with earlier populations, there is a 3%" decrease in b18 0 values, which
is consistent with increased cooling docun1ented in historical records
describing colder climates in the North Atlantic region from ca. AD 1400 to
1700. These changes are in accord with other studies that indicatcd
increasing dietary and climatological stress and eventual abandonment of
Greenland by Vikings during the Medieval period (Buckland et al., 1996;
Scott et al., 1991).
8.3
Elemental analysis
8.3.1
Background
Various clements found in both the organic (collagen) and the inorganic
(mineral or apatite)components ofbone tissue ha ve dietary and nutritional
significance. Most of these elements are contained in the inorganic
componen! (see Sandford, 1992); more than 99% of strontium in vertebrales, for example, is found in the bone mineral (Schroeder et al., 1972).
Major or bulk elements (carbon, hydrogen, iron, nitrogen, calcium,
phosphorus, oxygen, potassium, sulfur, chlorine, sodium, and magnesium)
perform critical functions in structura] maintenance and are generally
low levels (e.g., lead, mercury, cadmium), although virtually ali trace
elements are toxic if taken excessively.
Elemental analysis
291
itially, this research was received with enthusiasm, especially beca use it was
assumed that trace element values represen! accurate and unaltered
signatures of past diets, which evaluation had not been accomplished by
any other means in archaeological settings. Accumulating evidence shows
that the interpretation of trace elements in archaeological remains is far
more complex than had originally been anticipated. The enthusiasm for
elemental analysis has been tempered by the realization that a number of
factors, such as food preparation techniques, cooking utensils, geochemical variation, synergism between elements, nter- and intra-bone variation,
8.3.2
292
trophic leve! has been observed in field studies, in both aquatic and
terrestrial ecosystems (Elias et al., 1982; Ophel, 1963; Schoeninger, 1985).
Various local factors influence strontium chemistry, including geology
and regional levels of alkaline earths, and there is a considerable amount of
variability in strontium (and Sr/Ca ratios) within trophic levels (plants,
herbivores, carnivores) (Runia, 1987a; Schoeninger, 1985; Sillen, 1992;
Sillen & Kavanagh, 1982). Even within the same environment, economically importan! cereals and roots may have either elevated or lowered
Sr/Ca ratios in comparison with other plants (Runia, I987a). Owing to
variation within trophic levels, knowledge oflocal foodwebs and predator-
Elemental analysis
293
200
E
o.
-9'
E 150
"
E
iii
100
strontium (e.g., Sealy & Sillen, 1988). In addition, because bone strontium
and Sr/Ca ratios can be strongly influenced by consumption of highcalcium foods (e.g., seafood) and use of mineral additives, the simple
measurement of strontium in archaeological bone is not necessarily
ological samples must be made with a great <leal of caution (Buikstra et al.,
1989; Burton, 1996; Ezzo, 1994a; Radosevich, 1993; Sandford, 1992,
l 993b; Sillen & Kavanagh, 1982; Sillen et al., 1989).
Workers have also become increasingly aware that diagenesis in elemental analysis is the most important impediment to paleodietary reconstruction (e.g., Ezzo, 1992; Grupe & Piepenbrink, 1988; Lamber! et al., 1983,
1984; Nelson & Sauer, 1984; Nelson et al., 1986; Pate & Hutton, 1988; Pate
et al., 1991; Price, 1989; Sillen, 1981, 1992; and others). In addition to the
aforementioned factors influencing strontium in bone, there are a number
50 0~----=-5o~oc---~1~00~0,---.,-15='0"0---~2000
Year (AD)
Figure 8.4. Plot of strontium values in prehistoric and historie Ontario native
populations, showing an increase in maize in diets after AD 1000. (Data frorn
Katzenberg, 1984.)
294
Ied for there is a decrease in values that reflects a shift from plants high in
stronti~m (leafy plants and nuts) to low-strontium grasses (e.g., maize)
(Figure 8.4). The results are consisten! with findings based on nitrogen and
carbon stable isotopes (see above), although the approaches-isotopic and
elemental- relate to different parts ofthe diet, those that contribute to the
organic and to the inorganic fractions of bone tissue.
Schoeninger (1979) determined strontium values in members of groups
of different status at Chalcatzingo, Mexico. Her study suggested lower
values in higher-ranked individuals, which is consisten! with the hypothesis
that the elite in this society probably had greater access to animal protein,
whereas Iower-ranked individuals consumed relatively more plants, such as
maize (and see Blakely & Beck, 1981; Hatch & Geidel, 1985; Jacobs, 199 5,
for alternative patterns).
Weaning age has also been estimated on the basis of the ratio of
strontium to calcium in human bone samples. Sillen & Smith (1984)
suggested that, because strontium is discriminated against in relation to
calcium in milk production in the mammary gland and in the placenta,
newborn and young infant Sr/Ca ratios should be low in comparison to
those of adults. In contras!, plan! foods su ch as wheat and barley, which are
primary weaning foods, should exhibit relatively high Sr/Ca ratios.
Consisten! with these observations, a prehistoric Middle Eastern skeletal
series exhibits low ratios in newborns; ratios increase thereafter, peaking
between 1.5 and 3.5 years (Siilen & Smith, 1984).
Barium
Barium has received little attention in bioarchaeological chemistry and
paleodietary studies. This underrepresentation is surprising, since a significan! body of evidence indicates that the element is a highly sensitive
indicator of past foodways (Burton & Price, l 990a, l 990b; Ezzo, 1992,
1993, 1994a; Francalacci & Borgognini Tarli, 1988). Like strontium,
barium has no known biochemical function, it is nontoxic, it is structurally
similar to calcium, it is not tightly regulated metabolically, and it undergoes
fractionation with increasing trophic level (Burton & Price, l 990a, l 990b;
Burton & Wright, 1995; Elias et al., 1982). Barium values and Ba/Ca ratios
are Iower in herbivores in relation to the plants they consume, and
carnivores have lower values and ratios than herbivores. Most barium in
the geological environment comes in the form ofbarite (BaSO,), a chemical
that is relatively less soluble than the carbonates that are associated with
strontium (see Ezzo, 1992). This suggests that the soil-to-plant discrimination should be greater for barium than for strontium. Consisten! with this
Elemental analysis
295
296
Elemental analysis
297
0.25~----------.--------------,
833
-e- -0.75
!(i
Terrestrial diet
Marine diet
ee.
-1.00
o
_,"'
-1.25
-1.50
~I1
,, ..
~
~
-1.75
1-
-2.00
T
2
I3 I4
I6
!5
'
'
'
'
'
'
'
'
'
83.4
equal in terrestrial settings, but barium and Ba/Sr ratios in marine and
terrestrial settings are highly distinctive (Figure 8.5). Barium values and
Ba/Sr ratios are considerably lower in sea water and marine organisms than
in terrestrial organisms. Analysis of human remains from a wide range of
New World archaeological sites from predominantly marine settings,
coastal sites with agricultura! (terrestrial) consumption, and inland siles
with no access to marine foods reveals that Ba/Sr ratios also clearly
distinguish between human populations utilizing marine vs. terrestrial
resources (Burton & Price, 1990a, 1990b). Analysis of archaeological
human bone samples from desert settings (e.g., Stillwater Marsh, Nevada).
are an exception to the terrestrial pattern, and appear to be more similar to
marine values. This unusual pattcrn may result from strontium enrichment
in desert soils which would immobilize barium but not strontium (Burton
& Price, 1990a, 1990b) . Therefore, in at leas! sorne desert settings, the
differentiation of marine/terrestrial diets based on Ba/Sr ratios does not
appear to be possible. Ali other previously tested contexts clearly separate
human populations that use predominantly marine resources from those
using terrestrial rcsourccs.
Multi-element analysis
Deficieney in iron in past populations is inferred va analysis ofpathological lesions (cribra orbitalia, porotic hyperostosis) in a range of settings
worldwide (see Chapter 3). Sorne suggest that elemental analysis of
archaeological bone samples provides additional understanding of iron
status where pathology data are available (Edward & Benfer, 1993;
Fornaciari et al., 1981; Sandford et al., 1988; Zaino, 1968). On the basis of
information from limited samples, it was inferred that the presence oflower
iron levels in individuals with eribra orbitalia in Punic Carthage (third
century ec) eompared to nonpathological individuals suggested a link
between iron status and pathological indieators of anemia (Fornaciari et
al., 1981; although see Richtsmeier & Sheridan, 1996).
Juveniles from later Carthage (seventh century AD) have significantly
higher levels of iron than adults, which may indicate that iron in
arehaeological bone represents a biogenic signa! (Sandford et al., 1988),
Comparison of iron levels in the third century ec and seventh eentury AD
Carthaginians shows higher values in the later period; This suggests that
298
either iron status improved or iron intakes increased in the later perio<l
(Sandford et al., 1988).
Ezzo (l994a) has questioned the biological significance ofiron measurement in archaeological human bone. Except in regard to marrow, iron has
no apparent function in bone tissue, nor <loes borre actas a reservoir for the
element (Ezzo, 1994a). The lack of physiological evidence suggests that
varying amounts of iron in archaeological borre probably do not correlate
with the prevalence of iron deficiency anemia. Additionally, the iron
content in soil is considerably higher than that in bone, indicating that
unless all dirt is removed - a virtual impossibility in archaeological remains
-the iron content in the skeletal tissue is highly exaggerated (Ezzo, l 994a).
Zinc
F ollowing Gilbert's ( 1977) lead, there has been a general acceptance of zinc
as an importan! discriminator of diet (e.g., amount of mea! vs. plan!
consumption) and inferences about nutritional health in past populations.
Unlike for strontium, calcium, or barium, a theoretical basis has not been
developed for zinc as a paleodietary indicator (Ezzo, l994a, l994b).
Although zinc may be sensitive to dietary history, the lack of theoretical
models limits its usefulness in bioarchaeological chemistry and dietary
inference. Underlying the use of zinc is the unsubstantiated assumption
that the element is present at higher levels in meat and shellfish than in
plants. Zinc acquired in the diet will inftuence the levels in skeletal tissues of
growing animals, so that diets defi.cient in zinc will result in low values in
bone and other tissues (Ezzo, l 994b). Zinc appears to play an essential role
in growth: severe deficiencies of the element result in markedly reduced
growth and dwarfism. Therefore, in contrast to strontium and barium, the
essential nature ofzinc and its rather tightly controlled metabolic regulation suggest that it has little use as a paleodietary indicator.
Lead
Elemental analysis
299
300
individuals. Analysis of lead stable isotope ratios (206 Pb/204 Pb) suggests
that lead was traded to the Omaha from present-day Missouri. Lead was
also used to make red facial pigment, which would have been readily
absorbed. Thus, facial paint is probably the primary source of lead for
many of these individuals. Sorne skeletal lead may also have originated
from activities taking place after death, since red paint was also applied to
deceased individuals prior to their interment.
8.4
The two primary components ofbone tissue, apatite (mineral) and collagen
(organic), provide useful dietary information. In comparison with isotopic
analysis of collagen, elemental and isotopic analysis of the apatite is more
problematical, because of thc stronger inftuence of diagenesis and the
myriad of other factors both in living and in post-burial environments.
Measurement of the degree of alteration of apatite remains unknown for
matter) in collagen samples that could potentially alter the biological signa!
(Ambrose, 1987, 1990; Ambrose & Norr, 1992; Stafford et al., 1988). New
technology and better understanding of the biochemistry of the mineral
componen! (apatite) ofbone for trace element ami stable isotope analysis
has greatly expanded the range of possibilities for paleodietary study (e.g.,
Lee-Thorp et al., 1994; Cooke et al., 1996).
8.5
The application of isotopic and elemental analyses facilitates an understanding of foodways in past human societies by providing a nearly direct
record of diet. This approach addresses severa! issues in bioarchaeology
and archaeology that have heretofore been essentially speculative. Es-
301
pecially noteworthy are the timing and spread of C4 cultigens (e.g., maize
agriculture in the New World) and the relative contribution of marine and
terrestrial foods in coastal settings globally. Other critica! issues in dietary
ecology are curren ti y under investigation, such as the consumption of meat
by eastern African hominids living sorne two to three million years ago
(Lee-Thorp et al., 1994; Sillen, 1986, 1992). Both stable isotopes and trace
elements are subject to sources ofvariation that potentially impede dietary
interpretation, including climate, habita!, physiology, and diagenesis.
The approaches to dietary reconstruction and ecological and behavioral
inference discussed in this chapter should be considered as par! of larger
issues dealing with subsistence, adaptation, and nutritional ecology in the
past. Although qualitative means of subsistence documentation (e.g.,
analysis offood refuse) are subject to a range ofbiases- as is also the case
for isotopic and elemental analyses - it is importan! to consider them in
order to gain a comprehensive perspective of the foodways of earlier
societies. Direct information on die! in particular and the impact of diet
and nutrition during the juvenile and adult years in general can be acquired
only through the investigation of human remains.
Introduction
9.1
lntroduction
multidimensional scaling.
Although different biodistance statistics are used for metric and nonmetric data, these procedures have underlying strategic commonalities. The
multivariate Mahalanobis' 0 2 has become the benchmark statistic for
analysis of nonmetric data. The e. A. B. Smith Mean Measure of
Divergence (MMD) is the most commonly used statistic for nonmetric
303
nta. Until recently, the standard biodistance statisticfor metric traits has
een the 0 2 generalized distance (see Brace et al., 1990). More recently,
mean e-seores, 0 2 distances based on individual e-seores, and Q-mode
nnalysis of individual e-seores have gained favor as representations of
overall biological distance and intergroup patterning among populations
(e.g., Brace et al., 1990; Hanihara, 1994; Howells, 1986, 1989; Pietrusewsky, 1994). Another powerful multivariate analysis uses principal
componen! seores derived from tooth measurements for identifying broadscale population relationships (Harris & Rathbun, 1991). This method
reveals how size is apportioned across tooth types, thereby detecting crown
shape differences between populations.
Biodistance analysis is problematic, owing to the multifactorial nature of
the traits being studied. Skeletal traits are inlluenced by intrinsic genetic as
well as both local and general epigenetic and environmental factors: It is
thus not surprising that skeletal and dental heritability is imprecisely
understood (for sorne estimates of heritability, see Goose, 1971; Potter et
al., 1976; Sj0vold, 1984, 1995; Townsend, 1992). Thus, the application ofa
biological population model - which assumes at leas\ sorne degree of
genetic control of traits and that the distances based on these traits are
directly proportional to those derived from gene frequencies - is difliclt to
verify in human populations drawn from archaeological settings (and see
Saunders, 1989).
Neither metric nor nonmetric traits bear a one-to-one correspondence
with an individual's genome. The humeral sepia! aperture, a frequently
used postcranial nonmetric variant in biodistance analysis, for example,
has a high degree of association with robusticity. Similarly, spondylolysis
has often been assumed to be a population genetic marker. However,
spondylolysis is strongly inlluenced by activity load on the lower back (see
ehapter 5): eraniofacial and postcranial metric traits, as well, frequently
reflect activity-induced remodeling (Heathcote, 1994; Larsen, 1987; and see
ehapters 5 and 6). eonsistency offindings with other lines of evidence (e.g.,
archaeological or historical), especially when care is taken to screen out
traits influenced by activity or representing biomechanical adaptations
(Brace et al., 1990; Heathcote, 1994), suggests that these traits provide
importan! insight into population structure and relatedness. Minimally,
they are importan! for testing nonrandomness in skeletal series (see
Saunders, 1989; Spence, 1994). Familia! studies of modern humans (e.g.,
Saunders & Popovich, 1978), laboratory mice (e.g., Grneberg, 1952), and
rhesus monkeys (eheverud & Buikstra, 1981a, 198lb, 1982) support this
assessment. Phenotypicanthropometric analysis seems to provide the same
results as quantitative genetic analysis (Konigsberg & Ousley, 1995). In
304
305
9.2
1!
1
1
306
307
Measurement
Measurement
Interorbital breadth
Frontal chord
Parietal chord
Occipital chord
Fora1nen magnum length
Foramen n1agnum breadth
Mastoid length
Chio hcight
Mandible body height
Mandible body brcadth
Bigonial width
Bicondylar breadth
Minimum ramus breadth
Maximu1n ramus breadth
Mandible length
Mandible angle
mu!fipll,smal
~.,---+-4-
two, distlnd
population structure and history (e.g., Brace & Hinton, 1981; Calcagno,
1989; Harris & Rathbun, 1991; Hemphill, 1991; Lukacs & Hemphill, 1993).
More than 200 cranial nonmetric traits have been identified in humans
(Hauser & De Stefano, 1989) and include four primary types - ossiclcs
(small bones) within cranial sutures (e.g., lambdoidal ossicles); hyperostotic traits or unusual skelctal proliferations such as bridges ofbone (e.g.,
atlas bridging); hypostotic traits involving ossification deficiencies (e.g.,
metopic suture); and foramen variation (e.g., single or double supraorbital
foramen) (Buikstra & Ubelaker, 1994; Ossenberg, 1970; Saunders, 1989)
(Figure 9.1).
Built on earlier work by Dahlberg (1956), there are sorne 30 standard
nonmetric dental traits used at present in biodistance analysis (Hillson,
1996; Scott & Turner, 1997; Turner et al., 1991) (Table 9.2). These traits
include morphological variants on tooth crowns (e.g., incisor shovelling,
Carabelli's trait) as well as root variation (e.g., lower molar root number)
(Figure 9.2). There is a strong genetic componen! to these traits implied by
studies ofliving populations (e.g., Biggerstaff, 1970; Harris & Bailit, 1980;
Lundstrom, 1963). This factor, coupled with the conservative nature of
their evolution, indicates that dental traits are important for biodistance
analysis (e.g., Dahlberg, 1951, 1963; Greene, 1982; Griffin, 1993; Hemphill
et al., 1995; Lukacs, 1983, 1989; Lukacs & Hemphill, 1991; Sciulli et al.,
------
6.Parietalloramen
reMnlon pafletal
---------------~""-'~e
~'~
7b. coronal osslcle pre&ent
"" J
---z_/
-~
;...
~------
r'" '
I
"" ''""'' . ..
\
1f. Lambdold ossieles p11senl
---"',-
Figure 9.1. Cranial nonmetric traits. (Frorri Buikstra & Ubelaker, 1994;
reproduced With permission of authors and Arkansas Archaeological Survey.)
308
309
Teeth
Winging
Shovelling
Labial convexity
Double-shovclling
(a)
(b)
310
9.3.J
The traditional model of population structure and history in the Nile River
valley is based on the assumption that a series of migrations of racially
distinct populations replaced one another at various times. The model
posits that in the past there were two racial types living in the Nile River
valley. Thcse include a 'Caucasoid' type in the Upper Valley and a
'Negroid' type in the Lower Valley. Strouhal (1971) views biological
changes in the region as resulting from the arrival and grcater influence of
one race over another at various times (and see discussion by Armelagos et
al., 1981). Thus, the traditional model invokcs population movement and
gene flow to account for biological change in this region. From this
perspective, the reduction in craniofacial robusticity faund in X-group
populations is interprcted as signalling the arrival of a more 'Negroid'
population (Batrawi, 1946; and see Chapter 7).
In arder to evaluate thc traditional model of population replacement in
Nile Valley history, Greene and colleagues (Greene, 1967, 1972, 1982;
Greenc & Armelagos, 1972; Greene et al., 1967) analyzed a battery of
dental nonmetric traits within samples from Wadi Halfa, Sudan. Univariate statistical comparisons of nonmetric dental traits revealed no significan! differences between Meroitic and X-group samples. On the basis of
!he.se results, they concluded that population replacement was unlikely in
Temporal perspectives
311
this setting, 'but instead represent(s) a continuum of (temporally) successive local Mendelian breeding populations' (Greene, 1982:76). Although
they are suggestive, univariate statistical analyses are problematic, especially because such analyses incorrectly assume that individual traits are
independent. In arder to provide a more convincing argument far demonstrating biological continuity, Greene (1982) employed the more powerful
multivariate MMD biological distance statistic to dental nonmetric traits
to test the hypothesis that interpopulation biological differences are not
significan!. This analysis revealed extremely low divergence values, and
thus indicates no significan! change in biological groupings, or the virtual
lack of infusion of peoples with different biological histories into the
region. The implications of this study are importan!, because they lend
strong support to the argument that biological change in Nubia was dueto
local circumstances and not to gene flow.
Turner and coworkers (Irish & Turner, 1990; Turner & Markowitz,
1990) reassessed Greene and coworkers' continuity model of Nubian
population history. Although lauding the studies ofGreene and others for
using a nonracial, nontypological approach to Nubian population history,
they argue that extra-regional sources of variation ha ve been insufficiently
considered, especially sources that may explain temporal shifts in craniofacial morphology in this region. lmplicit in the work by Greene and
others is the assumption that population continuity extends al least as far
back as the late Pleistocene (ca. 12 000 BP). Turner and coworkers contend
that continuity can be claimed only if non-Nubian populations are also
considered in statistical analyses of dental trait variation in this region.
Turner and coworkers include in their analysis additional Nubian
dentitions from the late Pleistocene 'Upper Stone Age', Meroitic, X-group,
Christian period, and historie era European samples. Computed MMD
values, modifled for small samples (Green & Suchey, 1976) and tested far
significance (Sj0vold, 1973), revea! few significan! differences between
Meroitic, X-group, and Christian periods, thus confirming Greene's earlier
conclusions regardingpopulation continuity; However, significan! differences between the Pleistocene and la ter groups were clearly identified (e.g., the
frequency of incisor shovelling and three-rooted mandibular molars).
Because oftheapparent temporal discontinuity between the Pleistocene and
laterpopulations, Turner & Markowitz(l 990) hypothesize that theancestry
of recent Nubians was not derived from local late Pleistocene populations,
and that a population replacement event occurred during the Holocene in
Nubia. The origin of these later populations is unclear, but, solely on the
basis of dental traits, they argue that populations north ofNubia containing
European and Near Eastern traits are the most likely sources.
312
tinuity model can be accepted. Moreover, the west African collection used
for identifying dental trait frequencies is largely undocumented. Therefare,
although the similarities between late Pleistocene and west African dental
traits are interesting, they are not compelling. From the preponderance of
Temporal perspectives
313
314
between the two groups. Overall, social behavior has enormous itnplifa
tions for patterns of geneflow and biological variation in South Africa (:111d
see M orris, 1992).
Lastly, temporal comparisons of crania show very little evidencc frn
change before and after the shift from foraging to pastoralism amonp,
prehistoric Khoi in the coastal region. In contras! to traditional modcls fo1
the origins of Khoi coastal pastoralists, this finding argues that populat ion
replacement did not occur in this rcgion. Rathcr, pastoralisrn was intro
Population movernent has been a hallmark of European history. Biodistance analyses undertaken by Schwidetzky and Rosing for the Neolithic/
Bronze Ages (Schwidetzky, 1967), [ron Age (Schwidetzky, 1972), Rom:111
period (Schwidetzky & Rosing, 1975), and Medieval and recent periods
(R6sing & Schwidetzky, 1977, 1979, 1984; Schwidetzky & Rosing, 1982)
reveal a high degree of constancy in geographical and temporal differenti~
ation. Cluster analysis indicates two primary morphometric complexes -
Western and Eastern ~ with size gradicnts between the two. In the Western
complex, cranial length is greater than in the Eastern cornplex; in thc
Eastern complex, cranial width and bizygornatic breadth are greater (sec
R6sing & Schwidetzky, 1979). During the Early Middle Ages, the distinctions between the groups becorne less clear cut. Schwidetzky and R6sing
suggest that such a reduction in distinctiveness is due to increased
between observers and definition of characters, srnall sarnple size for sorne
si tes, the nature ofvariables used in the study, and incorrect sex identification. Cranial lengths, breadths, and facial size are developmentally plastic
(see Chapter 7). Therefore, sorne of the variability documented in European populations may also be inftuenced by shifts in technology or dietary
practices. Long-term contacts (e.g., Gothonic-Slavic groups), admixture
(e.g., Turkic-Mongolian groups), and widespread migration (e.g., ltalian
groups) probably inftuenced the patterns of cranial variation seen in these
Temporal perspectives
315
samples. In surn, then, no clear groupings are evident in this very broad
region. The association of geography, language, and temporal period with
craniometric variation suggests the presence of sorne regional variability
(and see Lalueza Fox et al., 1996; Sokal & Uytterschaut, 1987).
South and Central Asia
Among the vexing questions that surround biological and cultural history
in south Asia is the origin, rise, and decline of the Harappa culture of the
Indus River valley of Pakistan. Harappa culture arase and peaked in the
lower Indns Valley during the third millenniurn ne, and is the first
large-scale society in the Indian subcontinent (Chakrabarti, 1980; Fairservis, 1971). Marshall (1931) argued that Harappa culture developed out of
earlier pre-Harappan cultures within the Indus Valley, whereas sorne later
scholars argued that it was the product of 'stimulus diffusion' or migration
frorn the West, especially Mesopotarnia (Gadd, 1932; Gordon, 1958;
Wheeler, 1968; and see discussions by Hemphill et al., 1991; Lukacs, 1989).
Similarities in ceramics, metallurgy, and other material culture in the
archaeological record appear to link the Indus Valley and Mesopotamia.
Other archaeological evidence appears to contradict this assessrnent
(reviewed by Hemphill et al., 1991).
Hemphill and coworkers (1991, 1997) employed multivariate statistical
analyses of craniometric, cranial nonmetric, and dental nonmetric traits for
Indus Valley samples that might resolve problems in understandirig
biocultural change and population origins in this exceedingly complex
region. Cluster and principal components analyses of Harappan-phase
dentitions and skulls from Cemetery R37 at the site of Harappa provide
two principal findings. First, the differences between Neolithic and the later
Chalcolithic dentitions from the Indus Valley suggest a lack of population
continuity between about 6000 and 4500 ne. Second, a general similarity in
traits between sites in the region after 4500 ne indicates the presence of
biological continuity during the rise, apogee, and decline of the Harappa
culture. !ron Age populations (ca. 200 ne) are markedly different in their
pattern of dental nonmetric variation, suggesting a discontinuity during
la ter prehistory after the demise of the Harappa culture. Analysis of cranial
nonrnetric traits indicates that the source ofthis discontinuity may be gene
ftow into the region from the Iranian Plateau in the Near East. In sum, the
results of this research indica te the lack of biological continuity from the
Neolithic to the period of Christianity. Rather, two discontinuities existed;
the first between 6000 and 4500 ne and the second between800 and 200 ne.
These findings suggest that, although the Harappa culture was an indigen-
316
ous development within the Indus Valley, the region nevertheless interne
ted on a widespread leve) at various times, especially with populations lo
the west.
Unlike in the Indus Valley, the biological and archaeological history of
central Asia is poorly documented, in par! because the region has long
been envisioned as culturally barren, serving primarily as a buffer betwecn
the Harappa culture to the east and the Mesopotamian complex societics
to the west (Wheeler, 1968). Archaeological research beginning in thc
1970s and l 980s revealed that ihe central Asian borderlands were inhabited by various complex societies, including the Oxus culture, associated with cases in Bactria and Morgiana, located. in Uzbekistan,
Turkmenistan, and Afghanistan (see Hemphill & Christensen, 1994;
Hemphill et al., 1995, 1997; and B. E. Hemphill et al., unpublished
manuscript). This region may have played an importan! role in the
diversification and spread of Indo-Iranian languages into the lranian
plateau and the Indian subcontinent (Hiebert & Lamberg-Karlovsky,
1992; and discussion by B. E. Hemphill et al., unpublished manuscript).
Based on archaeological evidence, and especially material culture, an
argument is put farward by sorne that the Bactrian culture is an eastward
expansion of a central Asian oasis society originating in western Turkmenistan and northwestern Iran. The presence of Harappan artifacts in
Bactrian sites indicates to others that the origins of Bactrian culture are
Temporal perspectives
317
.O
KUZ
SKH
HAR
MOL
318
Temporal perspectives
319
dental and cranial biodistance analyses. For example, Yayoi teeth are sorne
14% larger than preceding Jomon teeth and are much more similar in
overall size to Chinese Neolithic dentitions than to dentitions from other
populations (Brace & Nagai, 1982). In all likelihood, sorne admixture of
Jomon and indigenous Yayoi took place, but the reduction in tooth size
following this event can best be explained by processes associated with in
situ change in post-Yayoi populations (Brace & Nagai, 1982).
The study of dental traits and size is also consisten! with craniometric
and cranial discrete trait analyses completed by various workers (Dodo,
1986; Hanihara, 1979, 1985; Howells, 1966, 1986, 1989; Nakahashi, 1993;
Ossenberg, 1986; Yamaguchi, 1982). Multivariate analyses indicate closer
biological links between Jomon and living Ainu than between Jomon and
living Japanese. Collectively, these analyses support the alterna ti ve point of
view that the biocultural discontinuity during the late Jomon period
resulted from migration and gene flow. These studies ccmcur that after the
Yayoi period, tooth size reduction and brachycephalization represen! in
situ adaptive responses to cultural changes and subsistence shifts. These
analyses emphasize the fact that this discontinuity is not a precipitous
population replacement. The gradient ofbiological distances in nonmetric
cranial traits from Siberians-Kinki-Kanto-Ainu-Jomon suggests the
presence of gene flow and hybridization between mainland immigrants and
indigenous Japanese during the Yayoi and following periods (Ossenberg,
1986).
transition at about 300 BC were dueto immigration and gene flow from the
Based on the strong anatomical similarity between Native New World and
Asian populations, especially in dental characters (e.g., both share high
frequencies of incisor shovelling), the origin of Native Americans is
unequivocally traced to northeastern Asia (Siberia) with immigration
taking place across the now-submerged Bering land bridge to present-day
Alaska (Dahlberg, 1951, 1963; Harper& Laughlin, 1982; Hrdlicka, 1912;
Turner & Bird, 1981). Archaeological evidence suggests that the immigration ofld World peoples to the New World took place by 11 200yearsago
(Hoffecker et al., 1993; Meltzer, 1993, 1995; Patterson & Larsen, 1997;
Rogers et al., 1992; and others). Earlier dates are suggested, but they are
problematic with regard to either the dates themselves or the association
with human agency (Meltzer, 1995).
Analysis of dental traits byTurner (1985) indicates that Native Americans are par! ofthe Sinodont group. His extensive study- based on sorne
320
third population migration (Nadene). Turner's dentochronological analysis suggests that these migrations were separate and were completed by
9000 BP. Greenberg and coworkers (1986:485) argue that there is 'a
remarkably good lit' between Turner's tripartite dental groupings, linguistic divisions of Native American languages, and genetic evidcncc of living
populations far inferred divergence and migration dates. The apparent
concordance of the three lines of evidence- dental, linguistic, and genetic-
has been met with strong criticism. For example, Turner (1985) assigns
prehistoric dentitions in the Aleut region to the Aleut category, which
assumes that the past and present population distributions are the same.
This assurnption is tenuous, especia11y in consideration of the re1narkable
time depth and diversity ofsamples studied (see also comments by Mcltzer,
1993, 1995; Steele & Powell, 1992). The minimal overlap in Nadene
Ianguages and dental groupings, along with other disjunctions in the
dental, language, and genetic evidence, led Szathmry to state that Turner
'interprets his analytic results in the light of a preexisting hypothesis that he
simply assumes to be true' (1986:490).
Analysis of mtDNA evidence in living Macro-lndians, Nadene groups,
and Asians provides an independent means for testing Turner's migration
Temporal perspectives
321
322
The prehistoric and early historie North American Great Plains was the
scene of a number of population movements that have implications for
Temporal perspectives
323
The call for the identification of biological and cultural links between living
na ti ve groups and poten ti al archaeological ancestors as a result of new laws
on the repatriation of Na ti ve American remains in the U nited States has
created additional incentive for biodistance analysis. Biodistance provides
a powerful means of linking present and past populations in North
America or other regions where repatriation ofhuman remains is a point of
discussion. On the other hand, it is difficult to identify relationships
between living and ancient groups, owing to the fluidity of ethnic groups
after contact.
Alaska is a highly visible focal point of repatriation discussions, in large
part owing to the presence of a highly vibran! native population in the
region. In order to identify links between living and past groups, a massive
amount of dental data ha ve been collected from both contexts on Kodiak
Island. Prehistoric and living populations have been studied by biological
anthropologists seeking information about ancestral and descendant
relationships (see Scott, 1994). Scott (1994) analyzed dental morphological
variables from prehistoric archaeological dentitions from the Uyak site,
which includes a population succession from the Kachemak (or preKoniag [1500 BC-AD 1100]) and Koniag (AD 100-1763) periods. Univariate
statistical treatment of crown trait frequencies reveals only two (of 14)
significan! differences between the pre-Koniag and Koniag archaeological
populations. Analysis of living Koniag populations reveals no significan!
intervillage variation. Trait comparisons indicate only three statistically
significan! differences between pre-Koniag and modern Koniag and two
differences between prehistoric and modern Koniag. This findings suggest
324
9.4
9.4.J
Spatial perspectives
325
RY
H-t-t-HJz
YO
Figure 9.4. Map generated from Womble analysis of cranial discrete traits in
west-central Illinois populations. The analysis identifies a series of tiles_
northern from a southern group (LD, Ledders: KL, Pete Klunk; GI, Gibson;
SH, Schild; KO, Koster Mounds; HN, Helton; HR, Hacker South Mound 2).
(From Konigsberg & Buikstra, 1995; reproduced with permission of authors
and Plenurn Press.)
326
9.4.2
Spatial perspectives
327
Stefano, 1989). Unlike the Wise series, precise familia! relationships among
individual skeletons could not be determined, owing to the lack of
association between grave markers and human remains. Historical records
indicate that individuals interred in the cemetery included primarily
members of a nuclear family and their el ose kin (Larsen, Craig et al., 1995).
Thus, an extraordinarily high frequency of metopism is consisten! with
archaeological and historical documentation that the cemetery is
dominated by the presence of closely related individuals.
There is a striking degree of homogeneity of traits in skeletal samples
from family cemeteries. When observed in large skeletal samples, such
homogeneity suggests the presence of endogamous mating patterns.
Biodistance analysis of dental nonmetric traits of sorne 300 individuals
from the Early Bronze Age site, Babedh-Dhra', in southern Jordan reveals
an unusually high degree ofhomogeneity- about 80% ofthe sample shares
specific traits (Bentley, 1991). There is also a high degree of similarity
between adult females and males. Analysis of trait distribution reveals a
clustering of rare traits within particular tombs. For example, only 8%
(13/158) ofthe Bab edh-Dhra' population possess mandibular molars with
six cusps; ali individuals with the trait are clustered in six of25 tombs. Third
molar agenesis is present in only five tombs. In summary, the high overall
degree ofhomogeneity suggests group endogamy, but with the clustering of
traits indicating burial of related individuals in the sme tomb.
The conclusion that trait homogeneity represents group endogamy is
based on the assumption that traits become homogeneous through time in
biological lineages (Kennedy, 1981; Konigsberg, 1987; Konigsberg &
Buikstra, 1995). Hypothetically, then, in endogamous populations, adult
males and females should exhibit the same degree of intrasex variability. In
contras!, exogamous populations should exhibit a relatively high degree of
variability as well as significan! sex differences. This suggests, therefore,
that the natal group or nonmigratory componen! of the population should
be relatively more homogeneous in trait expression than the migratory
componen t.
Beginning with the influential studies of Lane and Sublett (1972) and
Spence (1974a, 1974b), inferences have been drawn about postmarital
residence patterns by various researchers, especially in relation to intergroup and intragroup variance in nonmetric traits (see also Birkby, 1982;
Buikstra, 1980; Corruccini, 1972; Droessler, 1981; Kennedy, 1981). Spence
(l 974a, l 974b), for example, emphasized within-group variation based on
his modification ofthe simple matching coefficient. In his model, a high
degree of similarity in traits within a group represents low variability. Thus,
a higher mean similarity coefficient (i:e., decreased variability) in adult
328
males than in females (and see discussion by Spence, 1994). The greater
similarity among males than females in these traits suggest that the
corporate social group may have becn organized around male kin who
maintained their residence over the period of the occupation of the
beca use they do not deal with population variation within the framework
of formal quantitative genetics. In arder to allay this shortcoming,
Konigsberg proposes a population genetic basis far inferring residence by
applying a modification ofWright's ( 1951) island model and the migration
matrix method (e.g., Wood, 1986). Unlike the earlier approaches, application of quantitative models provides separate estimates offemale and male
genetic variance in a formalized tnanner.
Focussing on the relationship bctween migration rate and within-group
Spatial perspectives
329
330
open interclass communication, these results suggest that the Monte Albn
class structure may not have been strongly endogamous.
Other social distinctions aside from rank have also been explored va
biodistance analysis. Multivariate craniometric analysis of historie era
Arikara from the Leavenworth site, South Dakota, reveals a very high
degree of biological variability, far exceeding what would be experted in a
population practicing village endogamy (Byrd & Jantz, l 994; Key & Jantz,
l 990). Comparisons of this series with populations that were known to
have been distinctive in earlier Arikara history (e.g., Larson site) revea! an
unusually high degree ofheterogeneity in craniometric variation. Consisten! with archaeological evidence for the use of specific burial areas by
groups attempting to maintain corporate identity (see O'Shea, l 984) and
linguistic evidence for the presence of two dialects spoken by Leavenworth
inhabitants (Byrd & Jantz, 1994), there is a significan! hurial area effeet in
the analysis, specifically identifying two distinctive groups from two
respective areas of the site. This bipartite grouping pattern probably
reflects social subdivisions such as bands or attempts at maintaining earlier
village identities. During the late eighteenth century, different bands with
their own social, historical, and linguistic identities amalgamated into
Island in the eastern Pacifie (Owsley et al., 1994) and the northern Great
Plains (Jantz & Owsley, 1994b).
European contact with Easter Islanders initially occurred in the early
eighteenth century. From the outset of this contact, Europeans perceived
the native people as the primary resource for exploitation. In addition to
serving asan important labor source, the island was known to sailors 'for
the charms of its women' (Owsley et al., 1994:163). Sexual relations
between Europeans and native islanders are well documented in historical
accounts. Multivariate (discriminan! function) analysis of cranial measure-
331
ments from Easter Island populations reveals strong evidence for the
presence of Europeans among the native individuals (Owsley et al., l 994).
Visual inspection ofthree individuals shows distinctive morphology that is
characteristic of Europeans. F or example, a young adult male cranium
from the Hekii site has features - large and straight nasal bones, parabolic
palate, well developed nasal spine, and wide cranial base - that contras!
with those of native Easter Islanders. Additionally, this and two other
individuals possess anatomical features- such as forward projection of the
midfacial region - suggesting admixture of Europeans with Easter Islanders. Owsley and coworkers suggest that admixture may ha ve involved
a European father and native mother, a conclusion well supported from the
historie accounts of native and nonnative contacts.
In the Great Plains, European exploration and settlement increased the
opportunity for admixture between Europeans and native groups. There
are a number ofhistorical accounts of Euroamerican residen! traders who
married native women and raised offspring (Jantz & Owsley, l994b).
Multivariate analysis ofhuman remains from Swan Creek, South Dakota,
indicates the presence of a morphologically unique cranium - an old addt
male from a multiple grave including five individuals. The cranium
possesses a number ofEuropean or Euroamerican traits, including reduced
facial prognathism, triangular parabolic palate, prominent nasal bones,
and narrow nasal aperture. Three craniofacial indices that discriminate
between population groups suggest that the outlier male has Euroamerican
ancestry (Gil! & Gilbert, 1990). On the basis of the archaeological and
mortuary context, the individual was probably a residen! trader or a
captive. The 'foreign' origin of the individual is confirmed by his unique
element composition of dental enamel (see Schneider & Blakeslee, 1990),
suggesting that the enamel had formed while the individual was living in'
another geographical setting. The similarity of tooth wear pattern with
other individuals in the Swan Creek series suggests that the individual lived
a predominantly native lifestyle, at least with respect to the use of the
dentition in masticatory functions. The teeth are heavily worn, a pattern
typical ofnative but not Euroamerican dentitions in this setting.
9.5
332
10
10.1
Introduction
1
1
334
Sample representa/ion
10.2.1
10.2
Sample representation
335
Growth patterns
336
Sample representation
337
The essay by Wood and coworkers (1992) reminds us that the task of
interpreting skeletal characteristics is neither easy nor straightforward;
Especially problematical is the interpretation of age,at-death and popula'
tion structure from archaeological human remains. It would seem that an
increase or decrease in average age-at-death in temporal series refiects a
respective improvement or decline in health. Should not average age-at-
338
Sample representation
339
Late Woodland
Pike County sites
(and see Cohen, 1977, 1989). Contrary to intuition about age structure,
average age-at-death in a skeletal series <loes not reflect mortality or life
expectancy. Rather, it has been shown mathematically that average
age-at-death is a measure of birth rate and fertility if the population is
closed and not subject to great fluctuations in size (Buikstra et al., 1986;
Johansson, 1994; Johansson & Horowitz, 1986; Milner et al., 1989; Paine,
1989; Sattenspiel & Harpending, 1983). As applied to archaeological death
samples, this indicates that the presence of a relatively high number of
young individuals (low average age-at-death) in a skeletal series represents
more individuals entering the population through higher fertility. Conversely, relatively large numbers of older individuals (high mean age-atdeath) indica tes an abatement of births.
If skeletal series can be relied upon for population structure, then it
should be possible to examinepatterns offertility and population growth in
the past. Buikstra and coworkers (1986) present a case study from
west-central lllinois that demonstrates the importance of skeletal samples
in the complex link between dietary transitions and population structure especially with regard to theadoption and intensification of agriculture and
regional population increase. Mortuary and other archaeological data
identify an increase in population size after AD 1000 in the successive Late
Woodland and Mississippian periods in this region. On the basis of
Sattenspiel & Harpending's ( 1983) calculation that birth rate is about equal
to the in verse of mean age-at-death, they use a proportion of number of
deaths in composite age categories: number of deaths over age 30 (Dio+)
divided by number of deaths over age five (Ds+). By removing the
individuals who are less than five years, the problem of infant undernumeration - commonplace in archaeological samples - is avoided. Calculation
ofthe proportions far a temporal succession of eight skeletal series from the
Middle Woodland through Mississippian periods shows a decrease,
indicating an increase in birth rate in association with an increase in
Site(s)
DJo+/Ds+
Middle Woodland
Pike County sites
Gibson/Klunk
86
404
0.7791
0.6757
185
131
187
125
205
0.5676
0.5344
0.5134
0.4800
0.4585
232
0.4138
Helton
Koster
Ledders
Schild Mounds
Mississippian
340
10.4
341
and data used when the collection is no longer available for study.
10.3
342
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General index
General index
433
Angle, E.H. 242
Anglo-Saxon 11, 80, .145, 152, 254
animal protein 73, 76, 80, 127, 294
animal skins, processing of 262
ankylosis 166
anterior teeth 46, 77, 82, 95, 21 O, 234,
247, 256-258, 260, 262
anterior tooth use 236
anteroposterior bending forces 213, 216,
222
anteroposterior breadth 199
anteroposterior plane 201, 216, 217
anthropogenic factors 299
anthropologists !, 5, 6, 19, 65, 84, 119,
154, 159, 163, 166, 172, 197, 210, 233,
236, 271, 323, 334, 340, 342
anthropometric history 13
anvil fracturing 134
apatite 272, 282, 289, 290, 300
apatite signatures 272
archaeological context 3, 18, 51, 64, 79,
106, 123, 126, 243, 267, 270, 312, 318,
339
archaeological refuse deposits 38
archaeozoology 3
archaic and modern humans, differences in
bone remodeling 219
archaic hominids 169, 212
archaic Homosapiens 51, 113, 115, 168,
185, 219
Archaic period 1O, 27, 72, 117, 177,
179-180, 213, 221, 232, 240, 243,
253-254
archers, professional 221
Acc!ic 41-42, 104, 120, 131-133, 156, 158,
159, 189, 237, 320, 324
Argentina 63
Arikara 10, 58, 61, 124, 126, 175, 187,
322-323, 330
Arizona 92, 133, 146, 174, 275, 289
Arkansas I, 42, 96, 172, 250, 273
Armelagos, G.J. 333
Arnhem Land (Northern Territory) 141
articular joint dislocations 121
articular joint pathology 173
articular joints
amphiarthrodial 162
knee, interphalangeal, fibrous capsule
162, 166
diarthrodial (synovial) 162
interphalangeal 162
articular modifications 185, 186, 187, 193;
metatarso-phalangeal joint 185
articular size 219, 225
ascorbic acid 29
ascribed social rank 131, 178
General index
434
Ashanti 312
Asia
14, 17, 21, 42, 52, 56, 70, 80, 96, 104,
176, 179, 228, 256, 265, 266, 315, 316,
317, 318, 319, 321
east 317, 318
western 113, 115, 265
Asian
319, 321
Australian Aborigines
257, 334
435
General index
178, 198-199, 203, 217, 226-228, 235,
247, 249, 250, 252, 254, 256, 262, 268,
270, 284, 299, 314, 334-335, 340
behavioral change 51, 181, 211, 213
behavioral inference 5, 220, 223, 267, 301
behavioral reconstruction 168
beheading 126, 145, 149
bejel; see endemic treponematosis
Belemnitella 271
Belize 53, 76, 276, 287
Bell Beaker period 289
bending 196, 198, 199, 201, 204, 216, 217,
218, 223, 234
bcnding forces 199, 216
bcnding strength 201
bending stresses 217, 234
Bennike, P. 142
Berget, K.A. 206
Bering land bridge 319, 320
biceps brachii 189
bilateral asyn1n1etry 173, 188, 210, 212
structural, 210
bilateral symmetry 174
bioarchaeological data: in history,
economics, and nutrition science 4,
339, 340
bioarchaeological method and theory 340
bioarchaeological research 59, 291, 305,
332, 340
bioavailability 16, 29
biochemistry 290, 300, 340
biocultural I, 4, 64, 91, 102, 103, 233, 288,
315, 319
biocultural discontinuity 319
biodistance 302-306, 316, 319, 321-326,
328-332
statistics 302
biogenic signal 291, 297
biohistorical 304, 305
biological anthropologists !, 84, 172, 197,
210, 233, 236, 323, 334, 340
biologica\ anthropology 2, 3
biological discontinuities 324
biological diversity 329
biological population model 303
biological relationships: inter- and
intrapopulation 302, 304, 326
biological sclectivity 334, 336-337
biological variability 3, 330
biomechanical adaptation 2, 219, 303
biomechanical approaches: temporal
trends 213
biomechanica\ function 223
biomechanical geometry 221
biomechanical variation 203
biomechanics 197, 226
ne
271
263, 211, 212, 213, 214, 215,
276, 277, 278, 279, 280, 281, 282, 286,
288
211, 212, 214, 275, 219, 280, 281, 282
use of 280
Ci (Calvin-Benson) 271
Ci plants 271, 272, 279, 280, 282
C 3-C diet 280
c. 271, 272, 274, 275, 276, 280, 281, 301
G (Hatch-Slack) 271
plants 271, 272, 274, 275, 281
CA 58, 60, 199-200, 203, 206-210, 214,
216-217, 223, 225
C.A.B. Smith Mean Measure of Divergence
(MMD) 302
Caddo region 42, 219
Caddoan speakers (Pawnee and Arikara)
322
Cadien, J.D. 267
cadmium 290
Calcagno, J. M. 25
calcaneus 185, 189
calcium 57, 59, 61, 285, 290--294, 298
calcium-oxylate 67
calcium/phosphorus ratio 58
11c
e,
c4
436
General index
culinary 135
ritualizcd 135
Cantogrande 23
Canzo, Governor 182
carbohydrates 56, 65, 68, 72, 74-76,
80--82, 184, 252. 272, 279
carbon 183, 208, 271-272, 275, 276-277,
279-281, 284, 286-290, 294, 300
carbon isotopc ratios 271, 275, 276, 279,
281, 286-288
carhon isotope values, dictary signatures
of 277
carbon isotopic signaturcs 208, 272
carbon stable isotopes 294
see a/so stable isotopes
Caribbean 288
caries; see dental caries
caries sicca 95
cariogenesis 55, 66, 67, 72, 73, 78
cariogenic plant carbohydrates 184
carious lesion development, factors
intlucncing 65
carious lcsions 66
size, \ocation 70
Carlson, D.S. 230, 233-234
carnivores 121, 284, 291 295
Carthaginians 297
cartilagc 40, 165
case studies 3, 50, 96, 97
Catholic missions 53, 87, 263
catt\e 239, 256, 313
Caucasoid 229, 310
cause ofdeath 100, 134
Cave, AJ.E. 168
cavitation 65
cementoenamel junction 43, 48, 49, 77
central Canadian Arctic 42
central Europe: Ba\kan region 231
central lllinois River valley 10, 122, 325
Central Plains 323
Central Va\ley 18, 41
ceramics 299, 315
cervical vertebrae 132, 137, 145, 175, 176,
177
cervical vertebral osteoarthritis 176
Chaco region 28
437
General index
Cha!colithic 256, 266, 315
Chamay, A, 195
Charles, D.K. 185
Chaupicruz phase 277
chemical signatures 270
chenopod 70, 179
child abuse 65, 157
childhood 9, 11, 13-14, 20, 21, 32, 39,
51--53, 56, 58-59, 61, 106, 114, 157,
191, 284, 313
childhood anemia 39
childhood growth 14, 21, 313
children 8-10, 12-13, 17, 20, 23, 28, 32,
35, 37, 42-43, 50, 63, 68, 94,-95, 125,
127, 130, 133--134, 157, 210, 243, 260,
265, 339
Chile 18, 36, 93, 96
coastal 96
northern 18, 93
Chilean mummies 99, 100
China 46, 57. 281, 318-320
northern 281
chipped stone tools 267
ch\orine 290
chronic conditions 12
chronic diarrhea 29. 34
chronic helminth disease 38
chronic pathogen loads 62
chronic systemic disease 50
Churchill, S.E. 206
circular caries 55, 56, 285
circularity of femur: temporal increase 222
circumpolar groups 235. 237
city-states 143
Civil War military 188
C!arke, N.B. 77
C!assic period Maya 56, 76, 276, 328
clcft palate 24
climate 14, 163, 237, 273, 283, 289, 301,
342
- and body breadth, afld body surface
area 14
climatic patterns 289
cloacae 84
cluster analysis 302
cnemic index 21
coastal environments 281
coastal sites 296
Coatsworth, J.H. 4
cognition 63
cognitive development 6
cold adaptation 235, 236
cold stress 235, 237
Cole, M.S. & Cole, T.M. 235
collagen 271, 272, 281-286, 300
collagen samp\es 271, 286, 300
11
Colonal period 14
Colonial-era North America 299
Colorado 1, 11, 133, 336
compensatory hypothesis 208
compensatory remodeling 234
competition 122, 126, 130, 131
compression 19, 171, 173, 196, 200
computed tomography (Cn 202
computer analysis 202
computer-simulation sampting 28
computing 333
conftict 109, 112, 118-120, 122, 124, 126,
128, 130, 136, 139-140, 143, 146, 148,
153-157, 160
congenital defects 24
congenital syphilis 24
Constandse-Westennann, T.S. 212
contact period 27, 33, 36, 38, 52, 68, 87,
89, 125, 128, 138, 146, 148, 181, 211,
216, 263, 275-276, 286, 322, 339
conta1ninated water 37, 38
continuity 311, 312, 315, 321, 322, 324
continuity model 311, 312
Cook, D.C. 10, 43
cooking utensils 291
Coon, C.S. 235
coprolites 2, 37, 67, 270
co-residence 328
Corruccini, R.S. 242
cortical area (CA) 199
cortical bon e 12-13, 56, 77, 82, 90, 117,
199, 204, 207-208, 220, 223
thickness of (deficiencies) 12-13, 58,
201, 223
remodeling 208, 219
cortisol 11
Costa Rica 276, 287
covariance matrices 328
Coville, F. V. 259
cranial base flattening 20
cranial base height 19
cranial depressed fractures 119, 124, 129,
130, 142, 154, 157
cranial discrete trait analyses 319
cranial form, temporal change in 229
cranial length 314
cranial morphology 226, 230
cranial nonmetric traits 306, 307, 315.
324, 326, 328
divided supraorbital foramen 326
metopic sutures 326
open foramen spinosum 326
patent mendosal suture 326
temporal squamous foramen 326
cranial robusticity 226
cranial shape 228, 305
303
,!
:;.
438
General index
General index
dental comple:idty 312
dental crowding; see tooth crowding
dental damage 267
dental defects; see enamel defects
dental development 23, 26
eruption timing 24
dental dimensions 243
dental disease 76, 154
dental impaction 246
dental plaque 65
dental tissues 3, 5, 23, 43
dental trauma 156, 268, 269
dental variation 268, 305, 321
trait variation 311, 318
dental wear; see tooth wear
dentin 248, 252
dentitions ;24-28, 48, 51, 53-54, 61, 67, 70,
81-82, 156, 169, 226, 230, 235-237,
242. 250, 253-254, 257-263, 265-266,
268-269, 284, 305, 311, 315, 317, 319,
320, 323, 331
deciduous 23, 25, 47, 51, 55, 56
dentochronology 320
Denver 11, 336
depopulation 330
depressed fractures 118, 121, 123, 128,
130, 139-142, 145, 150, 153
see a/so cranial depressed fracture
deprivation 14, 22, 29, 145, 156
desert environments 36, 91, 171, 203,
283-285, 296
developing nations 8, 9, 62
development 6, 8, 23, 45, 46, 60, 65, 68,
70, 89, 113, 185, 191, 202, 233-234,
241, 266, 312, 316, 321-322, 334, 341
developmental disturbances 25, 44
developmental geneticists 27
developmental instability 28, 29
developmental plasticity 224, 228
DeVore, l. 161
diabetes 27
diagenesis 292, 293, 297, 300, 301
diaphyseal morphology 212, 222
remodeling 209, 214
shape 22, 222
strength 219
structure, 208, 218, 219, 224
diaphyses, 21. 98-99, 195-197, 199, 200,
202, 203, 209, 217, 222, 224--225
diaphysis length: femur; tibia 10
diarrheal infections 38
diarthrodial joints; see joints
Dickel, D.N. 31
diet 2, 10, 27, 29, 33-37, 56, 58-59, 65, 68,
70--76, 82, 87, 91, 136, 174, 179, 207,
228, 233, 244, 247. 250--251, 254, 258,
439
disease vectors 86
dislocation 193
dismemberment 1~9, 121, 124-125,
132-133, 138, 144, 147, 150--151
distal phalanges 188
disuse hypothesis 243
DNA, ancient 332
DNA synthesis 62
dolichocephalic 228
domesticated plants 70, 71, 80, 250
domesticates 16, 70, 73, 280, 282
double-shovelling 318
Droessler, J. 322
drought 17, 91, 131
dura mater 153
Duray, S.M. 55, 56
dwarfism 298
Dynastic-era Egyptians 76
early adulthood 8, 22
early hominids 51, 210
early Homo 218, 233
early lron Age 282
Early Middle Ages 314
Early Middle period 54, 112, 130
Early Mississippian pre-maize populations
263
early modern Europeans 225
early Neolithic 251
early nineteenth century 58, 173, 175
early prehistoric 18, 31, 100, 117, 174,
184, 192, 232, 295
Easter Island (Rapa Nui} 120, 139, 140,
158, 330--331
eastern Medterranean 19, 20, 33, 244
Eastern Woodlands of North America; see
American Eastern Woodlands
eburnation 165, 166, 168, 171-172, 184
definition 165
economic crises 9
economic historians 14
Ecuador 17, 18, 31, 37, 69-70, 88, 113;
137, 186-187, 277
Efe 73
Egypt 229
Egyptian dynastic tomb' art 187
Egyptians 76, 226
elbow osteoarthritis; see osteoarthritis
elemental (major and trace) constituents
270
elemental analysis 270, 291, 292, 297
elemental concentrations 291
elevated morbidity 334
elevated mortality 334
elevated systemic infection 336
elite individuals 93, 299
440
General index
dentoenamel junction 47
gross
55
epidemiological studies 50
in nonhuman primates 46
prevalences 46, 54, 56
susceptibility 46
enamel deficiency 46
enamel deposition 49
enamel elemental composition 65
enamel hypoplasias 45, 45, 50; see a/so
hypoplasias
enamel opacities 65
endemic treponematosis 91, 96, 97, 99
endemic warfare
124
Endo, B. 234
endogamous mating patterns 327, 329
endosteal bone loss 59, 208
endosteal contraction 209
endosteal or medullary area (MA) 199
endosteal section contours 203
endosteal surfaces 200, 203, 210, 221
endurance kayaking 189
engineering 197
England 11, 20, 51, 92, 106, 112, 152, 159,
176, 287
enslavement 14, 48, 139
enteric bacteria 36
enthesopathies 188, 189
high prevalences in postcrania 189
enthesophytes; see enthesopathies
environment 3, 5, 6, 9, 13-14, 20, 23-28,
33-34, 40, 43-44, 51, 54, 57-58, 62,
64-65, 90, 94, 108-109, 127, 131, 135,
137, 139, 155-156, 171-172, 178, 191,
195, 197, 203, 206, 215, 217, 219, 224,
226-227, 239, 240, 242, 245, 247, 249,
252, 268, 270, 272, 275, 287-288, 292,
294-295, 299, 303-304, 335-337
environmental constraints 6, 23
environmental degradation 131, 139, 295
environmental disturbance 13, 28, 275,
287
epidemics 64, 136
epidemiological studies 56, 64, 118, 163,
164
epigenetic 303, 305
epiphyseal fusion 8
episodic diarrheal disease 8
Equus caballus 115
Erlandson, J.M. 71
erosion 151, 165, 24 7
General index
eruption
Escherichia co/i 84
Eskaleut (Eskimos and Aleut of the
Arctic) 320
Eskimo craniofacial morphology 235, 236
Eskimos 61, 82, 131, 169, 170, 173-174,
178, 187, 235-237, 239, 256, 258, 260,
267, 282, 290, 320
estrogen 57
ethnobioarchaeological 334
ethnographers 334
ethnographic documentation 76
ethnographic evidence 33, 236
ethnohistorical accounts 136
Etruscan period 18
Euclidian map analysis 28
Euphrates River valley 265
Euroamericans 14, 28, 58, 113, 117, 126,
149, 172, 206, 220, 242, 279-280, 285,
326, 331
Europe 14, 17, 21, 70, 76, 80, 88, 95, 96,
101, 106-107, 142-143, 149, 155,
157-158, 179, 212,.228, 229, 231, 240,
281, 314
central 229, 231, 281
northern 88, 101, 120, 142, 143, 153, 158
western 17, 142, 143, 185, 212
European contact 96, 100, 104, 175, 211,
280, 286, 330
European Neandertals 185
European-introduced diseases 36, 53, 339
Europeans 9-10, 16, 37, 53, 70-71, 87, 89,
96, 99, 100, 104, 113, 115, 120, 126,
128, 139, 142, 146, 148, 152, 153, 156,
175, 181, 185, 211, 220, 225, 227, 237,
252, 254, 274, 279-280, 286-287, 290,
311, 313, 314, 330-331
evolution 9, 218, 243, 245, 247, 306, 313
evolutionary (genetic) change 245
evolutionary history 304-305
excessive masticatory loading 77
excessive mechanical loading 173
ex:ercise 57, 164, 195, 196, 337
exfoliation 23, 78, 267
ex:ogamous mating patterns 313, 327
exogenous materials added to foods 250
ex:tension 82, 115, 162, 173, 175, 185, 186
cxtramasticatory activities 77, 82, 236,
257-258, 260, 267-268
unintentional changes on tee.th arising
from 258
Ezzo, J.A. 275, 298
face and jaws, dietary and nondietary uses
of 5, 227, 233
face height 240, 318
441
ex:cessive intake 56
folklore 140, 145
food acquisition 173; 179
food availability 9
food collection 122, 252, 256
food preparation 39, 72, 175, 179, 186,
211, 239, 244--245, 250-252, 254,257,
266, 291
manner of 251
techniques 39, 239, 245, 291
technology 244, 250-252, 254, 266
food processing 35, 68, 173, 213, 253, 334
food production 4, 213, 229-230, 251,
252, 256
food shortages 9, 51, 126, 136
food taboos 39
foodways 68, 247, 270, 276, 280, 294,
300-301
foot osteoarthritis; see osteoarthritis
foragers 5, 10, 16-17, 27, 35-38, 41--43,
51, 55, 66-1;8, 70-74, 77, 78, 80, 84-88,
91, 117-118, 130, 140, 159, 161, 176,
178-179, 181-183, 189, 194, 203-204,
206, 213, 221, 224, 230, 232, 234,
236-237, 244--245, 250-252, 254-257,
260, 265-266, 284, 286, 293, 295, 312
foraging 10, 17, 25, 36, 51, 54, 61, 68, 71,
72, 80-81, 85, 88, 90, 141, 169, 179,
180, 183, 211, 214, 221, 229, 230, 241,
251, 252, 254, 258, 265, 266, 297, 313,
314, 337, 338
see a/so gathering
foramen magnum enlargement 138
foramen variation 306
forearm 94, 111, 112, 116, 117, 118, 189
forearm trauma 117
forensic sciences 109
fortifications 119, 122, 148, 173
forwardly placed zygomas 236
fossil hominids 202, 218, 266
fractionation factor 271
fractures 40, 57, 84, 92, 110, 112'--118,
121-124, 128, 132, 134, 137, 139, 141,
144, 150, 152-156, 158, 166, 171, 173,
190, 191, 235, 258, 267
age-specificity and prevalence 115
Cotles's 11 J
cranial depressed fractures 119. 124,
130, 142, 154, 157
crushing fractures 126
diaphyseal fractures 111
forearm fractures 112, 118
parry fractures 111, 112, 118
postcranial fractures 118
prevalences 115-118
rib fractures 112, 137, 154
l
442
General index
443
General index
Frayer, D.W. 76
French and Indian War 149
Fricke, H.C. 290
rontals 121, 124, 126, 139
FueguianjPatagonians 236
runctional hypothesis 235
functional studies 219
functional-mechanical paradigm
234
189
gangrene 246
Garn, Stanley M. 5, 12, 43
gathering 54, 72, 81, 87, 183, 192, 221,
231, 263; see also foraging
gender-based masticatory and
extramasticatory activities, 257
gender-specific behaviors 257
gene flow 233, 304, 310-315, 318-319
genealogies 4
with 288
geology 289, 292
geometric analysis 202-203, 216, 220, 221
Georgia 16, 25, 38, 49, 70, 72, 86--87, 91,
24, 26
retardation 10, 11, 24, 33
velocity 8, 9, 334
growth and development 19, 22, 54, _6162, 203, 209, 223, 270, 289, 329
growth arrest Iines 40; see a/so Harris
lines
growth plate 40
growth recovery 12, 22, 40
growth status 9, 13, 50, 62
growth suppression 1O, 11, 13
growth velocity 8, 9, 334
Guagliardo, M.F. 241
Guatemala 8, 50
link to status 92
Hearne, Samuel 132
height
285
historical accounts
recent 9
Hume, l.N.
humeral septal apertufe 303
humerus 8, 10, 110, 114, 125, 146, 151,
196, 221
hypocalcifications 44, 54, 55
hypoplasias 44, 45, 46, 47, 48, 50, 51, 52,
444
General index
immobilization
152, 196
297-298
bioavailability of 29, 34
iron absorption 16, 29, 38
Iron Age 21, 80, 142, 254, 281-282, 314,
Industrial Revolution 21
industrial urban living 16
malnutrition 88
infection prevalence 85, 88-93
infection: systemic 246
infectious disease 9-11, 17, 34, 42, 52, 61,
445
General index
315
33-40, 62, 158, 298
288-289, 300
Israel 240, 293, 341
ltalan groups 314
J (polar second moment of area)
20 f, 203,
317, 318
Japanese 9, 14, 76, 228, 244, 258, 318, 319
Jequetepeque River valley 137
joint morphology 218
joint size 218, 219
Jomon period 244, 251, 317, 318, 319
Jomon-Yayoi transition 318
Jones, Joseph 95
Jurmain, R.D. 163, 170
juvenile growth, pattern of 9, 22
juveniles 8-11, 22-23, 25-26, 32, 34, 43,
Kom 326
Koniag 133, 323
Konigsberg, L.W. 324, 328-329
Kricun, M.E. 112
Kyushu 265
La Florida 146
laboratory animals 25, 27, 46, 198, 227
mice 303
sheep 50
lactation 39, 57, 292
lactic acid 29
Lactobacillus acidophilus 65
Lake Ontario 149
Lallo, J.W.
10, 85, 86
lambdoidal ossicles
Lambert, P.M.
306
52, 54, 58, 66, 70, 72, 88, 90, 91, 93,
97, 107, 117-118, 120, 122-123,
125-126, 128, 132-134, 136, 148, 155,
158, 174, 178, 181, 184, 186, 192,
207-208, 218, 223, 229, 232, 250, 252,
254, 274-275, 277, 280, 286, 287, 289,
322
Late Woodland period 10, 13, 42, 52, 55,
68, 85, 102, 128, 183, 187' 240, 268,
321-322, 328, 338
lateral tibia! condyle 185
Latin America 52, 136, 138
latte stones 193
Jaw of bone remodeling 195
Lawson, John 98
Layrisse, M. 29
105
history of 106
incubation period 104
Medieval leper cemeieries 104
Mycobacterium /!prae 104
primary stage of 104
leucine-isoleucine ratio 16
life expectancy 34, 338
lifestyle 2, 3, 39, 57-58, 93, 110, 113, 117,
23, 336
morphology 172, 21 O, 226
structural analysis 183
Louisiana I, 42, 96
Lovejoy, C.O. 11, 118, 340
LSAMAT: as a primarily New World
phenomenon 260, 262
Lubell, D. 70, 256
193
Lunt, O.A.
lysine 16
25
446
General index
General index
mastication 227, 231, 234-235, 241, 246,
247, 252, 257
masticatory adaptation 246
masticatory and nonmasticatory functions
269, 305
masticatory behavior 226-228, 235, 247,
249-250, 252, 256, 262-263,267, 268
masticatory complex 107
masticatory function 226, 230, ~35, 239,
267, 331
masticatory-functional hypothesis 230, 232
masticatory loading 77, 227, 235-237, 239,
243, 253
masticatory muscles 227, 230, 232, 233,
234, 236
masticatory stress 233, 235, 237, 243, 244,
249
skeletal indicators of 249
material culture 183, 315, 316, 324
maternal health 12, 24, 25, 245
iron status 34
matrilocality 329
maxillae 258
maxillary 26, 98, 105, 106, 235, 237, 242,
257, 260, 262, 266, 268, 316
dentition 242
sinusitis 106
teeth 26, 262
Maya 8, 53, 56, 76, 276, 277, 278
Mayan Lowlands 279
Mbuti 73
mean age-at-death 52, 63, 86, 338
mean e-seores 303
measles 10
measuren1ent 12, 28, 61, 154, 199, 223,
270, 292, 298, 302, 314, 340
measurement error 28
meat 29, 42, 61, 72-74, 77, 174, 236-237,
239, 257, 266, 275, 292, 295, 298, 301
mechanical adaptation 234
mechanical demand 77, 82, 164, 170-171,
177-178, 181-182, 185, 192-193,
196-197, 199, 201, 214, 217, 220, 225,
235-239, 244, 249, 258
mechanical environment 172, 178, 191,
195, 197, 215, 217, 219, 224, 249, 268
mechanical integrity 59, 208, 209
mechanical loading 59, 173-175, 177, 179,
192, 194, 204, 206, 210-213, 21~-221,
224, 225, 227, 230, 233, 243
onjoint size 219
mechanical stress 163, 165, 166, 171, 175,
179, 184, 187-189, 191, 198, 214, 239
argument 163
model 191
medial epicondyle 188
447
Middle Ages 142-143, 146, 153, 158,
314
Middle East 70, 101, 228, 293, 294
Middle Mississippian period 42, 52, 85,
183, 263
Middle Woodland period 10, 13, 20, 55,
68, 70, 102, 178, 263, 268, 321, 322,
324, 328, 338
midshaft 214, 216, 217, 221, 222, 225
midshaft '~ 216
mid-South 70
Midwest 16, 35, 39, 96, 102-103, 118, 120,
122, 127, 151-152, 154, 158, 172, 222,
279, 321, 323, 339
migration 310, 314, 315, 319, 320, 321,
322, 328, 329
military campaigns 120, 148
military preparatory schools 14
military records 14
millet 33, 58, 280, 281
Milner, G.R. 258
mineral 16, 58, 197, 290, 292, 300
mineralization 40, 45, 60, 283
missile weapons 189
mission context 88
mission lndians 53, 217, 286
mission period 38, 54, 68, 72, 86, 87, 146,
148, 181, 182
.
missionization 68
Mississippi 69, 93, 96, 250, 263, 265, 274
Mssssippi River valley 93, 250, 263, 265
middle and lower 69
Mississippian 10, 36, 39, 42, 52, 58, 85-87,
93, l 02, ll 8, 122, 126, 128, 173, 177.
179-180, 182-183, 213, 221, 232, 240,
244, 253, 254, 263-264, 274, 287,
321-322, 328, 338
Mississippian Acculturated Late
Woodland 42, 52, 85
Missouri 10, 27, 123, 124, 126, 235, 273,
300, 322, 323
mixed foraging and farming 10, 61
Mizoguchi, Y. 237
MMD biological distance statistic 302,
3ll, 316, 317, 317, 320
mobility 1O, 85, 90, t 62, t 66, 194, 205,
206, 216, 289
patterns of 205, 206, 289
skeletal indicator of 216
molars 47-48, 67-68, 80, 95, 248, 251,
253-257, 263, 265-267, 282, 289, 31 l,
318, 327
Moldavia 34
M0ller-Christensen, Vilhelm 104-106
Molleson, T. 112, 187, 265
monozygous 28
448
General index
history 44
Morgiana 316
morphometric 233, 305, 314
mortality 56, 64, 98, 102, 109, 155,
334-339
mortuary studies 4
mtDNA sequence data 320
291, 297
N
N
282, 283
282, 283, 284, 285, 286, 288
General index
neutral axis 198, 198, 199, 200, 201, 214
Nevada: west-central 258; western, 171
Newell, R.R. 212
New England 287
Newman, M.T. 244
New Mexico 1, 92, 133, 136, 146, 170,
208, 217
New World 16, 19, 35, 52, 70, 95-96,
99-101, 104, 107, 120, 146, 155, 158,
175-176, 178, 228, 245, 260, 262, 279,
282, 286, 288, 296, 301, 319, 321, 330,
339
New York 115, 149, 155, 172
Ngarulurutja 72
niacin 16
Niagara River 150
Nile Valley 33, 34, 81, 89, 90, 229, 310
Nile Valley of Sudanese Nubia; see
Sudanese Nubia
nitrates 283
nitrogen 271, 282-285, 288, 289, 290, 294,
300
nitrogen isotope ratios 282, 284, 285, 288
nitrogen isotopic signature 285
nitrogen stable isotopes 286; see also
stable isotopes
nitrogen-fixers 283
nomadic pastoralism 313
nondomesticated plants 71, 179, 217, 250,
292
nonheme 29
noninvasive imaging 202
nonmetric traits 303, 305, 306, 310, 315,
324, 326--329, 340
nonspecific infection 38, 42, 88, 91-92, 99
periosteal 11, 18
prevalence: sex differences 91
nonspecific periosteal reaction 85, 90, 99
masticatory-functional hypothesis 230, 232
Norr, L. 272
North America 5, 9, 14, 20, 35, 51, 57,
68-70, 72, 80, 85, 88, 90, 96, 99, 117,
120, 128, 131, 133, 148-149, 151,.
154-155, 158, 169, 175, 179, 183-184,
203-205, 229, 232-233, 237, 250, 258,
272-273, 275-276, 279, 299, 320,
322-323, 326, 332
North A tlantic region 290
North Carolina 96, 98, 164
North Dakota 124, 234, 322
northeastern Australia 37
juvenile crania 37
northern Africa 33
Northwest Coast 36, 96, 102
Northwest Territories, Canada 131
northwestern South America 38
Norway 238
Nubia 13, 17, 20, 24, 33, 43, 57, 71, 89,
106, 112, 115, 116, 209, 220, 229, 230,
234, 252, 254, 255, 285, 310, 311, 312
see also Sudanese Nubia
Nubian 25, 34, 58, 60, 61, 152, 230, 254,
281, 285, 311
agriculturalists 254
craniofacial anatomy 230, 231
females 58
prehistory 281
nutrition 2, 4, 8-10, 14-:18, 20, 23-24, 26,
46, 52, 56, 57-59, 61-63, 65, 78, 86, 88,
163, 217, 222-223, 245, 270, 276, 301,
304, 335
nutritional deficiencies 10, 13, 19, 22, 44,
51, 56
nutritional deprivation 4, 18-19, 24-:25,
46, 219
nutritional ecology 2, 270, 301
nutritional immunity 62
nutritional insult 335
nuts 29, 73, 292, 294
observation of microwear 249
occipital condyle size 321
occipital precondylar tubercles 326
occipital tori 236
occlusal abnormalities 242-245, 247, 269
increasing prevalence of in
agriculturalists 244
occlusal enamel 248
occlusal margins, notching of 260
occlusal n1icrowear 265
occlusal surface 49, 65, 66, 228, 247, 248,
250, 252, 254, 256-258, 260-263,
266-267, 305
complexity 305
grooves in anterior teeth 259, 260
wear 66, 77, 80-81, 250, 254, 257-258;
see also tooth wear
Odense 106
odontometric fluctuating asymmetry 27
offertory items 138
Ohio 42, 70, 256, 273, 274
Oklahoma I, 42, 123, 250, 275
southeastern 123
Otd World 53, 70. 88. 95, 96, 100, 104,
144, 156, 158, 175, 179, 221, 234, 245,
250, 265, 279, 280, 319
older adults 23, 52, 80, 117, 118, 130, 139,
171, 187, 208, 214, 220, 238, 240, 242,
248
Ornaba and Ponca, historie era 175
omnivores 291
Oneota 120, 122
449
Ontario 11, 18, 58, 80, 89, 91, 113, 149,
188, 250, 254, 272, 273, 280, 293, 299,
326
southern 58, 254, 280
ontogenetic age pattems, 209
ontogenetic development 241
opportunistic infections 102, 107
oral hygiene 78, 81
oral medicines 152
oral tradition t 32
orbital roof thickening 30
orofacial adaptation 244
orofacial morphology 247
Ortner, D.J. 65, 173
osseous exostoses 188
osseous tissues 64
osteitis 82
osteoarthritis 144, 162-164, 166-184, 187,
189, 192, 194, 206, 213, 239
age variation in 178
and equestrian lifeway 175
appendicular 177, 180
diachronic trends 170
differences by sex in cervical spine 176
in elbow 169, 173-174, 179, 184
etiology and pathogenesis 162
in foot 194
gender differences in 176
highest prevalence in lumbar spine 176
in hip 164, 168
in knee 163, 164, 168, 177, 179
in acromioclavicular joint 169
in ballet dancers 164
in contact era populations 181
in cotton mili workers 164
in foundry workers 164
in industrial laborers 163
in manual laborers, farmers 164
in miners 163
and grinding implements 174
patterns in appendicular skeleton 170
and pneumatic tools 164
population-specific patterns of 167
prevalence of, between adult females and
males 176
right-side dominance in 170, 174
sexual dimorphism in 176
spinal 165, 166, 192
in temporomandibular joint 239
unilateral and bilateral 174
osteoarthritis prevalence 163, 170, 171,
176, 178-179, 181, 182, 184, 194, 206,
213
in prehistoric adult males 171
osteoarthrosis; see osteoarthritis
osteoblasts 83
"'1
450
General index
osteoclasts 59
osteoid 59, 60
osteological analysis: guides to
methodology and case studies 3
osteology 2. 333
osteomyelitis 82-84, 84, 86, 95, 246
osteon area 219
osteon population density 219
osteonal densities 220
osteons 59, 60, 61, 219-220
osteophytes 165, 166, 176, 188, 193
osteoporosis 23, 57, 58, 286, 340
premature 58
Type I 57, 61
Type Il 57, 61
osteoporotic fracture 57
ovarian function 161
overbite 242. 243
overpopulation 139
Owsley, D.W. \, 58, 125, 139, 331
Oxus culture 316
oxygen 29, 290
P.L. (Public Law) 101-601, 341; see a/so
NAGPRA
Pacific 36, 96, 120, 128, 131, 139, 183, 330
Pakistan 315
pa\atine tori 235, 237, 238
paleodemography 2, 338
paleodietary indicator 298
paleodietary reconstruction 292
paleodietary study 300
Paleo-lndian 322
paleopathology 2, 3, 64, 142
Panama 261, 262, 276, 287
Panamint women 259
Panicum mileaceum 280
parabolic palate 331
Paraguay 28
parasitism 30, 34, 36--39
Ascaris lun1bricoides 37
pinworm infection 37
Enterobius vermicularis 37
Moniliformis clarki 37
Strongyloides stercoralis 37
Trichuris trichuris 37
parathyroid hormone, overproduction of,
link with maize-based diet 59
parity status 39
pars interarticularis vascular foramina 191
particu!ate plant matter 300
parturition 46
Pastor, R. 256, 262
pastoralists 237, 266, 313, 314
Patagonia 236
patellar insertions 188
pathogen load model 62
General index
pathogenic agents, exposure to 5, 91
pathogens 62, 64, 86, 88, 90, 91-92, 94,
107, 108
pathophysiology 164
patient interviews 164
patrilinealclan lines 313
patriloca\ residence 328, 329
Patterson, D.K. 268
Pawnee 322
Pecos Pueblo Native Americans 170
Pecos region lndians 67
pectoralis majar 188
Pecdee geological formation, South
Carolina (PDB) 271
pelvic inlet shape 61
pelvic morphology 20
Pennsylvania: western 250
per cent cortical area ('YoCA) 58, 208, 209,
223
per cent osteonal bone 219
pericarp (bran) 16
perimortem fracture 110, 133, 157
perimortem skelctal modification 132, 136
perimortem trauma 134, 137, 149, 151
periodontal disease 65, 77, 78, 79-81, 246
temporal trends 78
periosteal expansion 208, 209, 223
periosteal reactions 54, 82, 84-85, 87, 89,
90, 92, 94, 99, 286, 336, 340
periosteum 82, 86, 94, 203, 223
periostitis 83, 84-86, 93, 95, 96, 99, 103,
106
permanent teeth 26, 28, 265, 268
Persians 226
Peru 18, 23, 27, 36, 37, 59, 75, 136, 139,
153
coastal 136
Peruvian Indians 173
Perzigian, AJ. 27
pewter 299
phenylalanine 16
Philadelphia 21, 172
phosphate oxygen isotope composition
290
phosphorus 59, 285, 290
photon absorptiometry 202
physical demands 164, 182, 216
physiologica! disruption 44. 50. 286
see a/so stress
physiological disturbance 16, 44, 49, 55,
61, 148
physiologica\ stress 2, 5, 10, 18, 22, 25, 40,
45, 51, 53, 60, 61, 158, 172, 245, 285
physiology 285, 290, 301
phytates 29, 33
phytoliths 67, 248, 267
308
Polynesia 56, 139, 207, 268
population aggregation 38, 39, 54, 61, 88,
295
population boundaries 304, 324
population crowding 37, 38, 88
population density 42, 85, 89-91, 96,
102-103, 139, 143, 155, 219
population health 334
population history 5, 226, 228, 304,
310-313, 317-318, 321-322, 332
population mobility t O
population replacement 18, 230, 310, 311,
314, 319
population size 18, 86--88, 90, 106, 107,
127, 266, 286, 338, 339
increase in 18, 86, 88, 90, 338
population structure 303, 304, 306, 310,
337, 338, 339
population-oriented approaches, 120
porion-basion distance 19
porotic hyperostosis 10, 30, 31, 32-39, 63,
136, 297
active 32
etiology 31
in juveniles 32
prevalences 33~34, 36, 39
Portugal 67, 71, 81, 251, 282
Portuguese 67, 256, 282
post-agricultura! 70
post-burial environments 300
postcranial nonmetric variants 303
postdepositional alterations 109
451
cracking 109, 268
small carnivore 109
root stains 109
weathering 109, 124, 137
postdepositional factors 335
posterior cruciate ligament 185
posterior teeth 25, SI, 80, 247, 256, 264
see also molars
postmarital residence patterns 304,
326-328, 332
postmenopausal women 117
postmortem alterations 291
postmortem breakage 11 O
post-Pleistocene 251
potassium 290
Pott's Disease (Sir Percivall Pott) 100
see a/so tuberculosis
pottery 266
Powel\, M.L. 93, 98-99
precontact 36, 53, 87, 96--97, 100, 112,
126, 133, 136, 138, 153, 155, 182,
210-211, 214, 254, 263
precontact Australia 254
precontact Eskimo 254
'precontact Latin America 136, 138
precontact Mexico 138
pregnancy 75, 78,292
pregnancy and lactation 292
prehistoric Australian populations 118
prehistoric Georgia coastal foragers 16
prehistoric Georgia lndians 233
prehistoric hunter-gatherers 192, 212, 219,
222, 256
prehistoric Jomonese 318
prehistoric lower 11\inois River valley 10
prehistoric maize agriculturalists 58, 70,
86, 117
prehistoric manioc agriculturalists 260
prehistoric Michigan 157
prehistoric Mississippian localities 39
prehistoric North America 9, 175, 183,
229, 258
prehistoric northwest Alabama 177
prehistoric Ontario 80, 91, 272, 293
preindustrial warfare 120, 143
pre-Koniag 323
pre-Medieval Euro pe 107
pre-modern reccnt humans 213
premolars 67, 68, 253, 257, 266, 318
pre-pottery Neolithic 265, 266
preservation status 336
preservation-related biases 270
pre-Yayoi 318
Price, T.D. 295
primary flexor 189
primates, nonhuman 46, 202
452
General index
prison inmates 14
projectile injuries 121, 128, 130, 135, 142,
154
projectiles 123, 143, 150, 157, 158
proliferative exophytic growths 165
proliferative Jipping
171
74
30
453
General index
rib fractures 112, 137, 154
rice, 29, 73, 280, 281, 318
Richards, L.C. 257
rickets 20, 21, 33, 40
right hand dominance 188
right radial tuberosity 189
right-dominant bilateral asymmetry 210
rigidity 199
ritual sacrifice 138
ritualized violence 138, 156
Roanoke River valley 250
Roberts, C. 106, 153
robusticity 76, 168, 185, 206, 209, 218,
219-222, 224-227, 229, 230, 232-236,
241, 244, 253, 275, 303, 310, 312, 335
rodeo cowboys 196
Roman period 106, 152, 158, 314
Roman Britain 158
Romano-British 34, 112, 145, 176, 299
root resorption 235
Rose, J.C. 70, 81, 263
ROsing, F.W. 314, 326
rubella 24
Ruff, C.B. 208-209, 216, 222-223, 240
Rufus Valley 36, 91
rural 46, 50, 94, 112, 164, 172, 242, 326
Russell, M.O. 233, 234
sacrifice 119, 137, 138, 145, 160
sacrificial death 136
Sadlermiut Eskimos 169, 268
sagittal keeling 236
Sagne, S. 26
saline soils 283
Salmonelfa typhi 84
sample size 28, 93, 278, 314, 336
San 35, 36, 148, 257, 313
sanitation 9, 11, 37, 39, 50, 86, 94, 107
Santa Barbara Channel 17, 36, 38, 41, 54,
71, 73, 90, 91, 92, 112, 120, 128, 131,
156, 158, 183, 184, 252, 288
Santley, R.S. 329
Sattenspiel, L. 338
Saunders, S.R. 11-12
scalping 119, 121, 121-126, 150, 154, 158
Scandinavia 71, 106, 142, 281
scanning electron microscopy (SEM)
analysis 258
Schistosoma 30
schistosomiasis 34
Schmorl's nodes 172
Schoeninger, M.J. 273, 293-294
Schurr, M.R. 274
Schwarcz, H.P. 277
Schwidetzky, l. 314
Scott, G.R. 239, 324
Sinodonts 317-321
Sioux 126
site characteristics 119
skeletal deformation 21
skeletal development; see growth
skeletal infections; see infection
skeletal modifications 133, 138, 185, 189,
227
skeletal morphology 169, 185, 197, 204,
333
1
1
454
General index
southeastern Australia 36
Southern United States 1
Southhampton Island, Northwest
Territories
169
southwestern Asia 42
Southwestern Cape Province 284
Spanish 25, 38, 69, 92, 128, 146, 148, 181,
216-217, 286
191
spondylolisthesis 193
spondylolysis 190, 190-194, 303
predisposing factors 191
spondylolytic fracture 193
squatting 185
Sri Lanka 189
455
General index
Streptococcus mutans 6,s
streptomycetes 90
stress 2, 5, 6, 8, 9-11, 13, 17, 18, 21, 22,
23, 24, 25, 26, 27, 28, 29, 31, 32, 34,
38, 39, 40, 42, 43, 44, 45, 46, 47, 48,
49, 50, 51, 52, 53, 54, 55, 56, 59, 60,
61, 62, 63, 78, 84, 90, 91, 126, 127,
131, 135, 148, 155, 158, 163, 165, 166,
171, 172, 179, 184, 188, 189, 191, 198,
208, 214, 217, 234, 235, 237, 239, 243,
244, 249,270, 275, 284, 285, 290, 295,
304, 313, 335, 336, 337
chronic mechanical 188
climatologica\ 290
early childhood 21, 51
environmental 27, 34, 155, 295
functional ,eonsequences 2, 5, 6, 9, 10,
11, 13, 17, 18, 22, 23, 25, 26, 27, 28,
29, 31, 32, 34, 38, 39, 40, 42, 43, 44,
45, 46, 47, 48, 49, 50, 51, 52, 53, 54,
55, 56, 59, 60, 61, 62, 63, 84, 90, 91,
126, 127, 131, 148, 158, 163, 165, 166,
171, 172, 179, 184, 188, 189, 191, 198,
214, 217, 234, 235, 237, 239, 243, 244,
249, 270, 275, 284, 285, 290, 304, 313,
335, 336, 337
loads 10, 184
mechanical 163, 165, 166, 171, 175, 179,
184, 187, 189, 191, 198, 214, 239
nutritional 18, 22, 52, 60, 61, 135
psychological 8, 78
stress history 22, 40, 43, 48, 54, 59, 61
stress indicators 6, 1O, 24, 42, 51, 55, 61,
63, 126, 158, 172, 337
identification of 55
stress model 7, 191, 237
stressors 6, 10, 17, 23, 24, 39, 42, 45, 53,
339
striae of Retzius 46
strontium 289, 290, 291, 292, 293, 293,
294, 295, 297, 298
strontium isotope ratios 288, 289
strontium/calcium ratio 292, 293, 294
Strouhal, E. 31 O
structural analysi~ 183, 198, 217, 220, 333
structural morphology 172, 215
subchondral bone 165
Sublett, A.J. 327
subperiosteal area (TA) 199
subperiosteal expansion 59, 224
subperiosteal surface 56
subperiosteum 83
subsistence 5, 10-11, 16, 58, 60, 68, 71-73,
85, 87, 116, 158, 169, 170-171, 173,
176, 179, 180, 183, 194, 203-206, 211,
214, 229, 230, 233, 235, 239, 242, 244,
199, 200,
taxonomy 333
technology 68, 71, 117, 146, 158, 160, 173,
183, 214, 220, 229,'235, 242, 244, 250,
251, 252, 254, 266, 268, 300, 314
teeth, I, 6, 23-28, 30, 46, 47, 51, 55, 56, 61,
65-68, 70, 72, 75-78, 80, 81-82, 95,
106, 110, 151, 154, 210, 226, 228,
234-236, 239, 242-245, 247, 248, 252,
256-260, 262-268, 289, 305, 318, 319,
331, 341
Tehuacan Va\ley 250, 276
Temascalcingo 50
temporal lines 235
temporal patterns 11 O, 130
temporal succession 181, 229, 253, 302,
338
temporal trends 14, 53, 67, 86, 116, 180,
228, 232, 279
temporalis 227, 230, 233, 236
temporomandibular joint 168~169, 233,
239, 253
size 230
Tennessee 25, 39, 70, 95, 96, 103, 112,
117-118, 154, 174, 177, 207, 221, 232,
233, 244, 250, 253, 260, 273
tennis players 195. 196, 209, 212
tension 196, 200
Tenninal Classic period, Maya 276, 279
tenninal height 13, 14, 18, 335
terrestrial plants 271-272, 283, 286, 288
terrestrial vertebrales 283
Terry Collection Blacks and Whites 187
tetracyclines 90
Texas 1, 42, 67, 71, 72, 117, 172, 259, 261,
262, 275
central 71, 72, 259
thalassemia 30, 33, 40
thennoregulatory model 236
third-world; see developing nations
thoracic vertebrae 142, 168, 175, 190,
193
three-rooted mandibular molars {M 1) 311,
318
Thule Eskimo 189
libia 8, 1O, 21, 40, 85, 86, 94, 11 O, 125,
144, 185, 188, 222, 225
lengths 10
tibial midshaft diaphyses 222
tibia! plateau 185
tibia! transverse lines 42
Tierra del Fuego 236
Tokyo 34
Tombigbee River valley 128, 155
tombs 18, 178, 327
tangue excisions 124
tooth antimeres 28
'
456
General index
268
tooth crowding 34, 37, 38, 88, 117, 242
temporal increases in
243
234
anterior incisa] loading 234
patterns of 234
tooth loss 23, 65, 67, 75--82, 105, 240, 246,
258, 282
prevalence in past human populations
79
tooth size
267, 334
tooth wear
257
systematic differences in human
populations 254
tooth wear pattern 254, 331
T orroni, A. 320
torsion 196, 198, 199, 201, 204, 205, 207,
218
24, 209, 211
457
General index
torus frequency 238
Townsend, G.C. 28
toxicity 297, 298
trabecular bone ll5, 219
trace elements 290, 301
trade, 58, 71, 73, 86, 184, 275, 281
transverse lines 40, 41, 42, 43
trauma 40, 82, 85, 109,)10, 112-118, 116,
124
mutilation of teeth 11 O
percussion 134
weathering 109, 124, 137
trauma-induced subperiosteal lesions 157
traumaticinjury 47,83, 112-113, 117, 119,
318-319, 320
behavioral interpretations 11 O
decapitation 119, l 24, 125, 132, 137,
213, 225
Upper
Upper
Upper
Upper
316
120
interpersonal 91, 119
mutilation 110, 119, 121, 124, 125, 132,
153, 154
violent death 120, 123, 125, 126, 132, 137
Virgin lslands 260, 261, 262
Virginia 48, 250, 299
virilocal 328
vitamin D 20, 21
vitamin D deficiency 20
Vogel, J.C. 272
voter registration 14
Waldemar, King of Denmark 143
Waldron, H.A. (T.) 164
Walker, P.L. 36, 71, 73, 77, 90, 128, 156,
63
vertebral osteoarthritis 169, 177, 183; see
a/so osteoarthritis
vertebral wedging 23
Viking period 142
village endogamy 330
violence 5, 109, 118-120, 122-126, 128,
I 19
War of 1812
warfare
292, 294
age or 285
weaning hypothesis 48
weaponry 119, 139, 143, 151, 158, 160,
458
General index
Woo, S.L.Y.
195
Site index
Bennett 268
Black Mesa 35, 92
Bogebakken 142
Cabet;:o de Arruda 67
Cahokia 39, 72, 93, 274
Campbell 27
Canyon de Chelly 35
Carlston Annis (Bt-5) JO
Carrier Mills 295
Carthage 34, 297
Punic 297
Cases Grandes 35
Chaco Canyon 35
Chalcatzingo 294
Chaluka 296
Chavez Pass 174
Chimu 137
Chin-Tafidet 188
Cholula 321
Chucalissa 118
Cinco Cerros 153
Cirencester 112, 113, 176
Clifts Plantation 299
Copn 56, 76, 277, 278
Corond 260
Crag Point 133
Cross 279, 326
Crow Creek 120, 123-126, 150-151, 157-158,
275
Cuicuilco 321
Curbridge 145
Dallas 207
Dickson Mounds 22-23, 42, 52, 54,
56. 85--87, 91, 118, 177, 183. 279,
297
Djarkutan 316, 317
Dolores 35
Edwards Plateau 72
Elephantine 326
Elizabeth 324, 325
Feurt 274
First African Baptist Church 172
Fort Ancient 274
Fort Erie 150. 172
Fort \Villiam Henry 149
Fourche-Maline 66
Gartner 274
Gibson 13. 20, 59, 179, 325
Glen Williams 80
Gran Quivira 35
Grasshopper Pueblo 275, 295
Gujarat 189
Hacker South Mound 2, 325
Hadar 168
Harappa 81. 266, 315, 316, 317
Cemetery R 37, 315
Harvie 280
Hawikku 35, 107, 187
Hayonim Cave 293
Hekii 331
Helton 325
Hertfordshire 145
Huang He (Yellow River) basin 281
Humboldt Lake Basin 259
lnamgaon 317
lndian Knoll 27. 117, 177, 183, 240
lnscription House 35
Irene Mound 97, 98, 103, 286
Jones Point 82
Kiavak 296
King 66
Kleinburg Ossuary 268
Koger's lsland 120, 123, 126, 127, 128, 158,
244
Koster 321, 325
Krapina 51, 80
Kulubnarti 34, 43, 90, 115, 116, 209, 239
Kuzali 317
La Chapelle-aux-Saints 80, 168, 218
La Florida (site) 277
La Plata Valley 35, 92, 136
La Ventilla B 328
Lake Shelby 275
Lamanai 53, 76, 276, 277, 278
Langhnaj 71
Larson Village 27, 120, 124, 125, 126, 150,
151, 330
Leavenworth 175, 330
Ledders 13, 59, 178, 325
Lepenski Vir 231
Lerna 222
LeVesconte Mound 268
Libben 10, 11, 55, 118, 152, 218, 256, 336
Little Egypt 101
Magdalenska Gora 281
Mahadaha 266, 317
Mahujhari 21
Mali Hassi el Abiod 188
Mancos Canyon 134, 135, 135, 136
Mary Rose 221
Matucana 153
Mauer 79
McCutchan-McLaughlin 120, 123
Mehrgarh 256, 262, 317
Melides 251
Mesa Verde 35
Mobridge 187
Moche 137
Moita 67, 251
Molali 317
Monte Albn 296, 321, 329-330
Morgiana 316
Moundville 39, 91, 92, 98, 99. 103, 117, 128,
287
Nadene 320
Nrestved 104, 105, 106
Nanjemoy Ossuary 187
Navajo Reservoir 35
Nasca 137
461.
Site index
Site index
460
326
145
Radley 145
Rajasthan 189
Raunds 11, 12
Ray 59, 324, 325
Rio Viejo 296.
Riviere aux Vase 120, 122
Rolling Bay 296
Sabastiao 67
Safety Harbor 97
San Cristobal 35, 148
San Luis de Talimali 148
Santa Barbara Channel Islands 38, 41, 54,
90, 92, 112, 129, 184, 252
Santa Catalina de Guale 25-26, 26, 38,
215-216
Santa Catalina de Guale de Santa Maria 32,
38, 79
Santa Catalina Ossuary 83
Santa Rosa lsland 71
Sapalli Tepe 316, 317
Sarai Khola 317
Saunaktuk 120, 131, 132, 133, 134, 156, 158
Schild 102, 280, 321, 325
Schild Knolls 329
Shanidar 113, 114, 114, 115, 168, 218
Snake Hill 149. 158, 172, 188, 280, 299
Spitalfields 21, 113, 164, 176, 191
St. Catherines lsland 25, 38, 215, 263, 286
St. Lawrence Island 324
St. J0rgensgard 104, 105, 106
Stanton Harcourt 145
Stillwater Marsh 49, 167, 171, 203, 206, 296,
296
Uyak Bay
133, 323
310
Weeki Wachee 97
Westerhus 48, 55, 76, 177
Wickliff Mounds 274
Windover 59
Wisby 143, 144, 146, 152. 153, 157, 158
Wise 326, 327
Yokem
325
Zalavar 76
Zebree 263
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