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Encyclopedia of Global Archaeology

Claire Smith
Editor

Encyclopedia of
Global Archaeology

With 2619 Figures and 106 Tables

Editor
Claire Smith
Department of Archaeology
Flinders University
Adelaide, SA
Australia

ISBN 978-1-4419-0426-3
ISBN 978-1-4419-0465-2 (eBook)
ISBN 978-1-4419-0466-9 (print and electronic bundle)
DOI 10.1007/978-1-4419-0465-2
Springer New York Heidelberg Dordrecht London
Library of Congress Control Number: 2013953915
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Further Reading
COLLINS, M., M. BUCKLEY, H.H. GRUNDY, J. THOMASOATES, J. WILSON & N. VAN DOORN. 2010. ZooMS:
the collagen barcode and fingerprints. Spectroscopy
Europe 22: 6-10.
EL-ANEED, A., A. COHEN & J. BANOUB. 2009. Mass spectrometry, review of the basics: electrospray, MALDI,
and commonly used mass analyzers. Applied Spectroscopy Reviews 44: 210-230.
ROBINSON, N. E. & A. B. ROBINSON. 2004. Molecular
clocks: deamidation of asparaginyl and glutaminyl
residues in peptides and proteins. Cave Junction:
Althouse Press.

Zooarchaeology: Methods of
Collecting Age and Sex Data
Deborah Ruscillo
Department of Anthropology, Washington
University in St. Louis, St. Louis, MO, USA

Introduction and Definition


Zooarchaeology has taken great strides since the
early 1900s when a zoologist or veterinarian
might occasionally have been consulted to examine animal bones recovered from archaeological
excavations, if the context in question was
believed to be especially important. Faunal
reports from that era were often simply lists of
species present: specific elements may or may not
have been listed, and the age and sex of individuals represented were certainly not included.
Having consulted the zoologist at all was a big
enough step in archaeological scientific inquiry at
the time. Most excavations before 1960 did not
even bother collecting animal remains, and it was
not until the 1970s when the processualist movement took hold in archaeology that excavators
began consistently to consider bones a valuable
source of environmental and anthropological
information. However, in order to obtain that
information, more data was needed from the
animal remains themselves.
Understanding past lifeways through faunal
remains is contingent on the amount of

Zooarchaeology: Methods of Collecting Age and Sex Data

information one can extract from those remains.


Determining the age and/or sex of animals
slaughtered for daily food requirements or ritual
purposes can aid in the reconstruction of herding
strategies or ceremonial procedures, respectively.
For example, identifying a predominantly older
female population among sheep remains can
suggest a secondary product strategy. In this
instance, ageing and sexing information from
animal remains is crucial to the interpretation of
culturally deposited bone.
The evolution of the study of animal bones
was facilitated by adopting methodologies
pioneered originally for the study of human
bones. Thus, systematic studies of dental eruption
and attrition, and epiphyseal fusion of postcranial
bones using modern specimens enabled the
definition of stages of growth and development
which could then be applied to archaeological
specimens. These techniques were proven to be
effective for the study of archaeological human
remains and were then applied to common
domesticates. As more techniques developed in
the study of humans, so too did the application of
similar methods to animal bones. Metrical and
morphological studies proliferated, especially for
dental analyses. More recently, technological
innovations in morphometric and microscopic
analyses have been used to age and sex animal
bones, as have genetic and histological methods.
These techniques are outlined below in simple
terms, from the most basic to the most advanced
available today. This entry is not meant to present
a summary of every study produced on ageing
and sexing such a feat would require a volume
unto itself. Rather, it describes many of the more
common methods available for distinguishing
age and sex.
To date, there are three major volumes that
compile methods for ageing and sexing of archaeological animal bones. These are Ruscillo (2006a),
Pike-Tay (2001), and Wilson, Grigson, and Payne
(1982). Countless articles in journals and edited
volumes have also been written, each presenting
its own unique application of known methods of
ageing and/or sexing or presenting new methods
for applicability to animal bones.

Zooarchaeology: Methods of Collecting Age and Sex Data

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Key Issues and Current Debates


Ageing Methods
Cortical Bone Texture
Perhaps the most basic method of determining age
is to inspect the texture of cortical bone. Fetal and
juvenile specimens exhibit rough developing cortex which becomes smoother with age. This texture difference is due to extracellular
modifications of collagen as well as other developmental processes involving the mineralization
of bone. Histological studies show microscopic
differences in the structure of juvenile and adult
bone tissue, but more useful to the bone analyst are
the visual and tactile differences of the bone cortex
which will immediately distinguish younger from
older individuals. Although not always useful for
suggesting precise ages, this method can be used in
a bone assemblage to establish the presence of
juvenile and immature specimens. Figure 1
shows the cortical differences between juvenile,
immature, and adult long bones.
The juvenile bone (Fig. 1a) has a rough cortex, similar in texture to fine sandpaper. The
younger the juvenile individual is, the finer the
granularity. Fetal bones are smaller and less
developed and have a very fine granular texture.
Fetal specimens are often difficult to distinguish
from neonatal individuals and can be described
as perinatal i.e., having died close to the date
of birth, either before (prenatal) or after
(postnatal).
The immature specimen (Fig. 1b) has residual
rough cortex around the epiphyseal joint, especially towards the ends of the bone diaphysis
(shaft). This roughness decreases with age as
the cartilage between the epiphysis and diaphysis
ossifies. By maturity (Fig. 1c), the cortex is
smooth with well-defined muscle attachments.
A good synopsis for identifying fetal bones in
archaeological contexts is Prummel (1987). For
a useful reference work on the ageing of fetal
bones, see Habermahl (1975).
Epiphyseal Fusion

Analysis of epiphyseal fusion is the most common age determination technique used for

Zooarchaeology: Methods of Collecting Age and Sex


Data, Fig. 1 Differences in cortical bone texture in (a)
juvenile bone, (b) immature bone, and (c) adult bone
(Image by the author, with thanks to Lucretia Kelly for
the specimens)

postcranial bones. The fusion of bone epiphyses


to the adjoining diaphysis occurs in regular stages
during development and is fairly predictable with
age, thus representing a good age indicator.
Silvers reference work The Ageing of Domestic Animals (1969) greatly aided faunal analysts
when zooarchaeology was still in its infancy. Not
only did Silver gather information from older
veterinary and zoological studies on tooth
eruption from nine major domesticates (horse,
ass, cattle, sheep, goat, pig, dog, cat, and
camel), but he also recorded the ages of epiphyseal fusion for almost all postcranial elements.
Despite some problems, this work is still
consulted for good estimates of age based on the
state of epiphyseal fusion of specific bone ends
and also for dental eruption timings (see below).
Many studies following Silver have attempted
further to define ages from epiphyseal states
while also considering influences such as castration, breeding, and environmental factors (e.g.,
see Davis 2000 for the delayed epiphyseal fusion
of bones due to castration in a population of
Shetland sheep). Problems with this method
arise when trying to determine ages over four
years for ungulates and over a year for smaller
mammals such as canids and felids. After the
complete fusion of long bones, ageing individuals is impossible without incorporating some

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Zooarchaeology: Methods of Collecting Age and Sex


Data, Fig. 2 Epiphyseal fusion drop-off curve for sheep
at Halaf period Fstkl Hoyuk, Turkey. Data points represent the percentage of specimens fused or fusing in each of
five sequential fusion groups (as defined by Zeder 2006).
Numbers in brackets are the sample sizes for each group
(Image from David Orton, used with permission)

other method, such as dental attrition which continues throughout life.


Since the fusion states of individual specimens
typically only provide age estimates of the form
younger than x or older than y, moving from
this to an estimate of population structure or kill-off
pattern is not straightforward. The usual approach
is to combine elements into a number of groups,
from the earliest-fusing to the latest-fusing (e.g.,
Zeder 2006), and to calculate the percent of specimens that are fused within each of these. One
naturally expects to see a drop-off in percentage
fused as one moves towards the later-fusing
groups, but the rate and position of this drop-off
can indicate kill-off structure (Fig. 2).
Tooth Eruption

The most commonly used age determination


methods involve some form of dental analysis,
partly due to the high preservation rate of
teeth and mandibles. As with humans, other
mammals have fairly predictable ages of tooth
eruption and development. Mandible and maxilla
fragments with teeth still intact can thus often
offer fairly accurate estimates of age-at-death,

Zooarchaeology: Methods of Collecting Age and Sex Data

especially for complete mandibles or maxillae


and for individuals under 48 months of age,
the time when most medium-large mammals
have completed eruption of all teeth.
If the permanent dentition is fully erupted,
other methods of ageing teeth can be adopted,
such as measurements of dental attrition or
cementum annuli counts (see below). Studies
such as Deniz and Payne (1982) on goats and
Jones (2006) on sheep have developed methods
of ageing which consider both eruption stages
and tooth wear to produce a complete mortality
profile of a population. Zeder (2006) considered
both tooth eruption and wear along with long
bone fusion to further refine age determination
accuracy in caprids. Similar work has been
performed with wild and domesticated pigs by
Bull and Payne (1982).
Tooth Wear and Dental Attrition

Much work has been performed on recording


the rate at which dentition wears for different
species. Theoretically, dental attrition progresses
at a predictable rate so that wear can be measured
within an accuracy of months to determine the
age at death of an individual. Factors which affect
the rate at which attrition occurs include feed,
captivity, and environmental conditions for grazing. Sex and side of dentition do not generally
seem to significantly affect attrition.
Methods usually involve observing and
recording the wear on the occlusal surfaces of
a row of teeth. Diagrams using standardized
coding, such as those shown in Fig. 3, can be
drawn to show patterns by indicating the level
of wear of the occlusal enamel, cementum, and
dentine of a specimen (after Grant 1982; see also
Payne 1973). Theoretically, individual permanent teeth can be aged in this manner if given
a dataset from known age/wear individuals, but
usually this method is performed on complete or
partially complete mandibles.
Tooth Crown Heights

Examining tooth crown heights is another


method for gauging attrition. Typically, the
distance between the occlusal surface and the

Zooarchaeology: Methods of Collecting Age and Sex Data

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Zooarchaeology: Methods of Collecting Age and Sex


Data, Fig. 3 Dental attrition on a cattle mandible from
Vindolanda (V07 69A 18174) with corresponding diagram of tooth wear patterns (Images by Deb Bennett,
used with permission of the Vindolanda Trust)

Zooarchaeology: Methods of Collecting Age and Sex


Data, Fig. 4 Cementum annuli as seen under
a microscope with dye for contrast (Image copyright
Gary Matson/Matsons Lab, used with permission)

line separating the enamel of the crown from the


dentine of the root is measured for each relevant
tooth on the buccal side (outlined in Klein &
Cruz-Uribe 1984: 46-47). These measurements
can then be analyzed in various ways, including
Kleins and Cruz-Uribes own Quadratic Crown
Height Method (QCHM) which predicts age
based on eruption age, unworn height, and an
estimate of the age at which the crown would be
completely worn down. Alternatively, linear and
quadratic regressions have been performed using
measurements on specimens of known age. This
technique has been applied to cervids, bovids,
and equids.

seasonal climates may exhibit less characteristic


seasonal growth patterns in dental cementum
because there are less distinct differential feeding
and growth periods.
Not only can this method be used to age animals at death, but the season in which the animal
died can also be postulated.

Cementum Annuli Counts

Tooth cementum is laid down continuously, but


seasonal differences in deposition rate result in
growth banding analogous to tree rings. When
a tooth is thin-sectioned and stained for greater
visibility, the rings (annuli) can be counted under
a microscope and used to estimate age. Figure 4
shows dark annuli that have been deposited in the
winter months, while the larger lighter-stained
cementum represents the growth period in the
spring and summer. Most wild animals can be
aged this way, especially those from northerly
latitudes where seasonal temperatures are more
distinct. Domesticated animals and species in less

Sexing Methods
The number of diagnostic features in a skeleton
that indicate the sex of an individual differs from
species to species. The predominant difference
between male and female individuals is size
dimorphism; males tend to be larger than females.
Some species like pigs and deer are highly dimorphic, while many smaller species, such as hare, are
less so. Thus, osteometric data can be used to
estimate sex ratios for many species if the sample
size is large enough. More reliable for identifying
males from females, however, is the recognition of
primary, secondary, and tertiary sexual characteristics. These are outlined below.
Primary Sexual Characteristics

Primary sexual characteristics are, of course, the


reproductive organs. These features are useful for
sexing individuals while they are alive, but
after death reproductive organs generally do
not survive in the archaeological record.

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The baculum or penis bone, however, can preserve in certain conditions. Carnivores such as
procyonids, mustelids, canids, felids, ursids, and
pinnipeds have bacula. Occasionally, some of
these species may have a female equivalent
bone in their clitoris, the os clitoridis or
baubellum, but this element is often absent or
cartilaginous and cannot be predictably found in
female individuals archaeologically.
The presence of a baculum in an archaeological sample or in a burial of a complete skeleton
signifies a male individual, but not finding
a baculum does not necessarily mean that males
are absent. Some bacula are small and fragile and
do not preserve well. Alternately, finding
a baculum does not mean that females do not
exist in a sample. The presence of a baculum is
most useful for sexing when excavating whole
skeletons. Canids, in particular, are sexed in this
manner because many cultures bury companion
animals purposefully.

Zooarchaeology: Methods of Collecting Age and Sex Data

winter, and these may be collected by humans.


Both sexes of caribou produce antlers. The males,
like most cervids, will shed their antlers after the
rut, while females will shed theirs during the
winter months unless they are pregnant, in
which case they will retain their antlers until the
spring.
Horns differ from antlers in that the horncores
actually form part of the cranium; they are made
of bone and are not shed. Many species exhibit
horns on male individuals only, but some bovids
have produced breeds where both males and
females have horns. Work on distinguishing
between the sexes by identifying horncore morphology has been performed so that the horns
themselves can be sex indicators (Armitage &
Clutton-Brock 1976, although see Sykes &
Symmons 2007). Studies have shown that castrating male individuals early in life will affect
the development of the horn cores. Thus, horncore
morphology may also be useful in identifying
castrates in a bone sample (Fig. 5).

Secondary Sexual Characteristics

Secondary sexual characteristics include features


that males possess which females do not. These
commonly include horns, antlers, tusks, large
canines, and spurs. The identification of sex
based on these secondary features is not
a relative value like ageing data. Instead, it is
based on a present or absent classification as
with bacula. For example, if large canines from
a wild boar are present, the identification of the
individual as male is certain. A secondary trait
that is relative is body size; mature males are
larger than females in most mammals.
A comparison of size data within a faunal sample
can often distinguish size dimorphism.
Headgear Horns and especially antlers are good
indicators of male individuals. If a frontal bone is
found with pedicles from antlers, identifying the
individual as male is certain (with the exception
of caribou/reindeer). Caution must be used
when interpreting sex based on pieces of antler
finding antler is not always evidence that male
deer was hunted. Male deer molt their antlers
after the annual rut in the late autumn or early

Canine
Morphology Another
potentially
diagnostic difference between male and female
animals is canine size. This is true of many
species, including suids, equids, moschids,
hippopotamids,
and
elephantids,
among
others. With the exception of equids and
moschids, male tusks grow perpetually throughout
life; an old adult wild boar can have tusks of
a considerable size. In equids and moschids,
canine teeth are much more prominent in the
male individuals. Only 25 % of female horses
produce canines, and these are usually small
in size. For felids and canids, canines seem to be
proportionate with size. Females, being diminutive in stature, will naturally have smaller canines.
It is difficult, however, with domesticated dogs
and cats to use canines as sex indicators because
manipulated breeding has produced great size
variety which plays a large role in determining
the overall stature of male and female individuals
and their sexually dimorphic characters.
Size Dimorphism or Trimorphism Size
dimorphism is the most inclusive of all the

Zooarchaeology: Methods of Collecting Age and Sex Data

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Zooarchaeology:
Methods of Collecting
Age and Sex Data,
Fig. 5 Horn-core
differences in male, female,
and wether sheep
individuals as determined
from radiographs (After
Clutton-Brock et al. 1990)

sexually dimorphic differences in animals.


Almost all mammals with the exception of the
spotted hyena and the blue whale exhibit larger
male stature and/or female gracility. This size
dimorphism is measurable using standardized
bone measurements, the collection and manipulation of which is known as biometry or more
specifically osteometrics.
If the bone sample size is large enough, one
can plot bone sizes (e.g., length versus width of
any complete bone) to reveal size dimorphism in
a population. Usually, there will be great overlap
between the sexes, especially if there are wild
and domestic populations or multiple domestic
breeds present at the site but also because adult
females and immature males are often similar
in size. The challenge when interpreting
osteometric data is that male animals are often
culled from the population before maturity is
reached, while females are often kept until they
are too old to produce offspring or secondary
products like milk. Males that are kept for meat,
wool, or labor will often be castrated, further
complicating the interpretation of the data.
Castrated mature males are still larger than
females, but their economic contribution to the
herd has been altered: males can be used as studs
or for sport (e.g., bullfighting), while castrates are
kept either for meat (as in pigs and bovids), for
wool (e.g., sheep), or for labor (as in cattle or
equids). One cannot usually tell if the larger
individuals in a site population are male or
castrated individuals from size alone without
also examining morphological differences using
morphometrics, discussed below.
Bones that preserve well in the archaeological
record and for which osteometrics have proven

particularly useful for distinguishing between


males/castrates and females include the
metapodials, especially among bovids.
Birds Birds also exhibit secondary sexual
characteristics in the form of spurs (males) and
calcium deposits in the medullary cavity
(females). Plumage is also a secondary trait,
males often displaying more flamboyant colors
and longer tail feathers, but these features rarely
survive in the archaeological record.
Medullary bone is deposited commonly in
the leg bones of female birds (femur, tibiotarsus,
and/or tarsometatarsus) during egg production.
Figure 6a shows a layer of mineral deposits in
a tarsometatarsus of a hen. The medullary bone
acts as a repository for the storage of calcium
required for eggshell development. The medullary cavity can have a significant amount of
mineral deposition, or the mineral stores can be
represented by a fine bony web within the cavity.
The extent to which the medullary bone is laid
down is dependent on the species of bird, its age,
its egg productivity, the mineral content in its
diet, and other environmental factors. In order
to examine this trait, however, one would need
either to break the bones open or to use radiographs (OConnor & OConnor 2005). Medullary
bone can also be considered a tertiary sexual
character, for it arises after the sexual maturity
of the animal and not during puberty.
Male spurs are more obvious on the tarsometatarsus on most adult galliform birds (Fig. 6b).
This family includes turkeys, chickens, quails,
pheasants, and peacocks. These spurs are used
in mate and territory competitions and are useful
indicators of sex.

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Zooarchaeology: Methods of Collecting Age and Sex Data

Zooarchaeology:
Methods of Collecting
Age and Sex Data,
Fig. 6 Sexual dimorphic
characters of Gallus, (a)
medullary bone from a hen,
and (b) a spur on
a tarsometatarsus of a cock
(Image copyright Lena
Strid, used with
permission)

Tertiary Sexual Characteristics

Tertiary sexual characters differ from secondary


traits in that they arise during maturity and not as
a direct result of sex hormones during puberty.
Tertiary traits develop as a result of sex-specific
behaviors (i.e., pelvic changes due to pregnancy
and birth) or in support of secondary traits (i.e.,
atlas changes due to muscle development for
headgear support). These affected bones are
found in different places in the body but mainly
involve the neck, the pelvis, and the lower
extremities.
Pelvic Morphology The easiest and most
common method for identifying sex through skeletal morphology is by examining the shape of the
pubis. For ruminants, if the pubis is oval or round
in section, it is a male individual. If the pubis is
thin and slightly twisted, it is female (Fig. 7).
Caution should be taken when examining female
pubes a male castrated within the first few days
of life will develop a thinner pubis as well; the
pubis of the castrated individual will resemble
that of a female who has not yet reproduced.
The absence of androgens during the early development of males will default the female
morphology.
The shape of the obturator foramen, the space
between the acetabulum, ischium, and ilium, can
also be diagnostic of sex for ruminants. The female
foramen is symmetrical and when traced resembles the shape of an avocado pit. The male foramen
when traced is less symmetrical, bulging on one
side compared to the other (see images in Ruscillo

Zooarchaeology: Methods of Collecting Age and Sex


Data, Fig. 7 Bovid pubes of a male and female

2003). The foramen of a castrated individual interestingly resembles that of a male, probably due to
its equitable size. Therefore, if one finds
a complete innominate of a ruminant that has the
pubis resembling that of a female and an obturator
foramen of a male, then the identification of
a castrated individual is likely.
A method for distinguishing sex in suids specifically is by measuring the height of the medial
border of the acetabulum. Males have a higher
medial wall than females the female pubis is
thin and the medial wall by association is also
gracile (Greenfield 2006).
Mean Shape Method Morphometrics is
a branch of mathematics which examines the
metrical and statistical properties of shapes and
shape changes of objects. Using techniques such
as elliptic Fourier analysis (EFA) or eigenshape

Zooarchaeology: Methods of Collecting Age and Sex Data

8007

Zooarchaeology:
Methods of Collecting
Age and Sex Data,
Fig. 8 Mean shapes of
sheep atlases for (a) male,
(b) female, and (c) wether
(From Ruscillo 2003)

analysis, for example, bone morphology can be


compared between the sexes in quantifiable
terms. These techniques are particularly useful
for examining tertiary sexual characteristics
which are not immediately visible to the eye.
Ruscillo (2003) explores tertiary sexual
characteristics using what she has called the
Mean Shape Method. By digitally outlining profiles of selected bone elements using a statistically
sound number of specimens of known sex,
eigenshape analysis can produce an average
shape for each element by sex. Profiles were
obtained by the use of profile gauges. These
mean shapes can then be compared to the shapes
of the archaeological specimens to determine the
sex of the individual. The method was fruitful for
cervids and bovids and indicated several bones that
were sexually dimorphic. The transverse processes
of the atlas were significantly dimorphic for both
families of animals. Figure 8 shows the mean
shapes of sheep atlases produced after scanning
over 30 individuals of each sex including males,
females, and wethers (castrated males).
Male and female bovids and cervids showed
significant differences between shapes of the
glenoid process (the articular part of the scapula)
and the astragalus. For sheep, the transverse processes of the atlas, the astragalus, and the obturator foramen each showed a third shape
corresponding to wethers. In general, the mean
wether bone shapes are characterized as having
the morphology of females with the size of males.

Presumably, similar shapes would be expected of


castrated goats, cattle, and cervids.
Table Test Sexing canid remains can be difficult
unless a whole skeleton is present with a baculum.
Isolated bones of canids can be problematical to
sex because millennia of breeding the species have
produced highly variable sizes and even bone
shapes. However, a complete mature canid
humerus can help suggest the sex of the animal
with 85 % accuracy. The Table Test is
described by Ruscillo (2006b) and is fairly simple.
The complete humerus is placed on its
anteroventral side on a level surface. If the
humerus falls over to one side, then it is likely
a male individual; if it rests stably on its
anteroventral side, then it is more likely to be
a female specimen, though several male humeri
were stable as well in individuals which were lap
animals and did not exercise normal male
behaviors. These individuals may have been castrated as well. The morphological difference being
measured by this simple test is the size of the
deltoid tuberosity which is significantly more
developed in most male individuals because of
the more robust deltoid muscle.

Future Directions
Much work has been done in the past 60 years to
identify useful methods of ageing and sexing

8008

animal bones in the field and in the laboratory.


There is still much to do, however. With advancements in technology, future work in ageing and
sexing will no doubt involve more microscopic,
chemical, and molecular analytical methods, as
these become more accessible and affordable.
Osteohistology
Osteohistology is defined as the study of the
microscopic structure, chemical composition,
and function of bone. The stage of structural
development of bone can be observed and
recorded by viewing cross sections of bone
under the microscope. The patterns of lamellar
bone, osteons, resorption lacunae, and Haversian
canals correspond to different ages. There are six
primary stages of long bone structure in cross
section (shown in Dammers 2006: 23; Fig. 8).
With a detailed study of the bone structure of
each taxon, age profiles can be recorded and
used to age archaeological specimens. Many of
these studies have yet to be done for specific taxa,
but an overview of how this method can be
applied to different species is explained in
Dammers (2006). The development stages and
patterns of bone structure are also distinctive of
sex. Future work in this area is likely to prove
invaluable.
Imaging
Medical imaging and 3D modeling technologies
may be used to refine morphometric analysis for
distinguishing between the sexes. Magnetic
resonance imaging (MRI) technologies have, as
yet, seen limited use with archaeological animal
bone. Although radiography is currently
preferred for inquiries related to bone, the image
produced by MRI is superior for reproducing any
object accurately in three dimensions, recording
morphological and even histological information.
With further refinement, minute histological
details of bone structure may be more readily
identified and accurate ageing assessed from
bone samples.
There are many programs which have enabled
the 3D imaging of objects. These programs replicate the specimen under examination by

Zooarchaeology: Methods of Collecting Age and Sex Data

plotting coordinates on three planes. By comparing the location of a specific point or group of
points plotted on the surfaces of an object, the
principle components of shape can be measured
and contrasted. Eigenshape analysis has already
been used in comparing the eigensurface on
paleontological species (MacLeod & Krieger
2007). This kind of technology will be especially
useful for comparing sexual dimorphic differences in any species.
DNA and Molecular Studies
Every year strides in genetic technology are
made, and improvements in cost and reliability
are starting to make molecular sexing a viable
option in archaeology. Genetic studies based
on polymerase chain reaction (PCR) are
already being implemented in projects involving
various aspects of the origins of domesticated
species.
In a study by Telldahl et al. (2012), for example, sex classifications based on measurements of
cattle metapodia are corroborated by molecular
studies. A sample of archaeological individuals
was sexed using both conventional morphometric
criteria and genetic analysis, and the results compared to establish the accuracy of the former. No
doubt as DNA analysis becomes more mainstream and affordable, sex determination from
bone remains will become easier and more
accurate.
The future of ageing and sexing methods in
zooarchaeology will likely be imaging and biomolecule based. New techniques will give rise to
more accurate and more frequent identifications
of age and sex, permitting creation of mortality
profiles for animal populations from communities in the past. This should in turn give rise to
a clearer picture of economics, trade, ritual, and
daily lifeways of ancient peoples.

Cross-References
Biometry in Zooarchaeology
Geometric Morphometrics and Environmental
Archaeology

Zooarchaeology: Methods of Collecting Age and Sex Data

Secondary Products and the Secondary


Products Revolution
Zooarchaeology

References
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bones from archaeological sites (BAR British series
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CLUTTON-BROCK, J., A. VALBONESI, D. CASTILLO-FERNANDEZ &
E. DAHL. 1990. The osteology of soay sheep. London:
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mandibular dentition as a guide to ageing Turkish
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Oxford: Archaeopress.
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of domestic ungulates, in B. Wilson, C. Grigson & S.
Payne (ed.) Ageing and sexing animal bones from
archaeological sites (BAR British series 109):
91-108. Oxford: Archaeopress.
GREENFIELD, H.J. 2006. Sexing fragmentary ungulate
acetabulae, in D. Ruscillo (ed.) Recent advances in
ageing and sexing animal bones: 68-86. Oxford:
Oxbow Books.
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(ed.) Recent advances in ageing and sexing animal
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Further Reading
CRABTREE, P.J. 1990. Zooarchaeology and complex
societies: some uses of faunal analysis for the study
of trade, social status, and ethnicity, in M.B. Schiffer
(ed.) Archaeological method and theory, Volume
2: 155-205. Tucson: University of Arizona Press.
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season of capture, age and sex of archaeofaunas.


Oxford: Oxbow Books.
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University Press.
MCGRORY, S., E. SVENSSON, A. GOTHERSTROM, J. MULVILLE,
A.J. POWELL, M.J. COLLINS, & T.P. OCONNOR. 2012.
A novel method for integrated age and sex
determination from archaeological cattle mandibles,
Journal of Archaeological Science. doi:10.1016/j.
jas.2012.05.021
OCONNOR, T.P. 2006. Vertebrate demography by
numbers: age, sex, and zooarchaeological practice, in
D. Ruscillo (ed.) Recent advances in ageing and sexing
animal bones: 1-8. Oxford: Oxbow Books.
UERPMANN, H.-P. 1973. Animal bone finds and economic
archaeology: a critical study of osteoarchaeological
method. World Archaeology 4(3): 307-322.

Zvelebil, Marek
Peter Rowley-Conwy
Department of Archaeology, Durham University,
Durham, UK

Basic Biographical Information


Marek Zvelebil (19522011) was born in Prague,
capital of the then-communist country of Czechoslovakia. He relocated to the west in his teens
with the rest of his family and studied at Sheffield
and Cambridge. He spent 30 years as part of the
faculty in Sheffield until his death in 2011.
Mareks interests were highly diverse. His
mastery of eastern European languages enabled
him to study shamanism among Siberian
hunter-gatherers; he collaborated with his
father, the eminent linguist Professor Kamil
Zvelebil of Leiden University, in disputing the
Early Neolithic origins of the Indo-European
language group; he collaborated in fieldwork
projects in Ireland and the Outer Hebrides; and
after the fall of communism, he started to work
in the Czech Republic. But Marek is mainly
known for his work on the hunter-gatherers of
the Baltic region and their part in the transition
to agriculture. His Ph.D. was on this topic. His
study visits to Finland were always via ferry and

Zvelebil, Marek

car not by air for fear that a flight


approaching a fog-bound Helsinki might be
diverted to nearby Leningrad (now St. Petersburg), potentially resulting in his enforced
return to Communist Czechoslovakia.

Major Accomplishments
Marek was part of the small group of scholars
who argued that hunter-gatherer lives did not
have to be nasty, brutish, and short, and that
in favorable areas like the Baltic coasts, huntergatherer populations might be as dense and society as hierarchical as those of early farmers. The
availability model for the origins of agriculture
in the Baltic made the hunter-gatherers active
players in the transition to agriculture by defining
three phases in the process (Zvelebil & RowleyConwy 1984):
The availability phase: hunter-gatherers are
aware of agriculture practiced by neighboring
farmers but adopt little or none of it; agriculture produces <5 % of food.
The substitution phase: farming is being
adopted and the two ways of life are in competition due to scheduling clashes between
farming and foraging; agriculture produces
550 % of food.
The consolidation phase: farming matures and
spreads into suboptimal parts of the region;
agriculture provides >50 % of food.
Marek further developed this model 2 years
later, in a paper (Zvelebil 1986) from which
Fig. 1 is taken. This model was in direct opposition to immigration or wave of advance models
of the spread of agriculture into Europe, in which
the hunter-gatherer population did not even feature. In immigration models, the first appearance
of any domesticated species constituted the
Neolithic Revolution, but in the availability
model, revolutionary change occurred only
towards the end of the substitution phase and
the transition to the consolidation phase.
The existence of lengthy availability phases of
a millennium or more in various parts of the

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