Archaeology and the study of teeth
Simon Hillson
Archaeologists reconstruct the past from a variety of surviving articles recovered by excavation.
Among these the study of teeth is particularly rewarding both because their physical structure and
configuration throws much light on the nature and life history of the animals from which they came,
and because they have a high survival rate even under adverse conditions.
Archaeological excavations produce
many different finds. Not only are
there stone, pottery, and metal arte-
facts, but also the remains of plants
and animals brought in as food or raw
material, or living on the site when it
was occupied. The most abundant
animal remains are the bones and teeth
of mammals, greatly outnumbering
sherds of pottery on many sites. They
have been recognised as important
finds since the earliest days of archaeol-
ogy. Remains of extinct mammals
found alongside artefacts were impor-
tant evidence for man’s antiquity in the
19th century English cave excavations of
William Pengelly. During the present
century research on ancient bones and
teeth is an established part of archaeolo-
gy. The study of the remains of man and
his extinct relatives has become the
thriving discipline of palaeoanthropolo-
gy, with several international research
institutions and journals. Work on
ancient diseasehas led to the formation of
an international Palaeopathology Asso-
ciation whose membership includes
surgeons, physicians, pathologists, and
anthropologists. Study of non-human
mammal remains has also attracted
researchers from many different back-
grounds and there is now an International
Council for Archaeozoology which
maintains links and organises confer-
ences.
Teeth and jaws are a major part of
the biological evidence for mammals in
the past. The jaws typically form part
of the waste of the carcase in food
animals - discarded early on in the
butchery process - and are therefore
prominent in the debris found in
middens, pits, and ditches on
Simon Hillson, Ph.D.
Was born in 1953 and graduated in geology
and archaeology from the University of Birm-
ingham, in 1974. He carried out postgraduate
research at the Institute of Archaeology,
University of London, on the biology of ancient
Egyptian and Nubian human remains. Since
1978 he has been lecturer in archaeological
science at Lancaster Universitv. He now works
largely on dental remains fioi archaeological
sites and has written a textbook ‘Teeth’ (1986).
Endeavour. New Series, Volume 10. No. 3. 1986.
016GS327/86 $0.00 + so.
archaeological sites. They are robustly
constructed and survive well in a
variety of soils. The tissues of the teeth
are especially tough, particularly the
hard enamel which coats the tooth
crown. These tissues have to survive an
environment of powerful abrasion,
leaching, and attack by microorganisms
in the living mouth, so it is perhaps not
surprising that they survive well in the
similar conditions of the soil. Under
some soil conditions, the enamel coat
of the crown may be all that survives.
Another reason for the importance
of teeth is their highly complex biolo-
gy. The form of the dentition is closely
related to diet and there are detailed
differences in tooth morphology be-
tween most species of mammals. In
some groups of rodents the dentition is
the main basis for identification even in
living animals. It follows that, although
much of the skeleton has features
which allow the identification of small
fragments of bone, the teeth and jaws
are particularly useful. Teeth also vary
between the members of one species;
in the form of the roots, in the size of
the crown, and in small details of the
pattern of cusps, folds, points, and
blades that decorates the surface of the
crown. The overall dimensions of the
crown are routinely measured for a
variety of species, but detailed features
of crown morphology are normally
recorded in discrete categories, asses-
sed by eye. For this reason, they are
known as non-metrical variants and can
be recorded very rapidly, not only in
archaeological material, but also in the
living. Non-metrical variants seem to
vary consistently between population
groups in man so. for example, it
seems to be possible to distinguish the
native inhabitants of North America
from those of South America on this
basis; Eurasia from the Americas; and
Australasia from Eurasia. There have
been few archaeological studies, but
there is considerable potential for
reconstructing the biological rela-
tionships of ancient and living human
populations.
Teeth and jaws are the commonest
fossils in the record of human evolu-
tion. Our extinct relatives from the
Pliocene and early Pleistocene of Afri-
ca, the australopithecines, walked up-
right on two legs almost as we do, but
had a brain about half the size of ours.
They also differed in the size and form
of teeth, jaws, and their supporting
structures. Their incisor and canine
teeth were about the same size as our
own, but their cheek teeth (premolars
and molars) were considerably larger
and the most robustly built australo-
pithecines had vast third molars, up to
22mm across. The jaws were corres-
pondingly longer, more protruding,
and more heavily buttressed, and were
worked by larger muscles spreading
over a larger area of the smaller brain
case. Forms such as Homo habilis and
Homo erectus, thought to be intermedi-
ate between the australopithecines and
Homo sapiens, generally have teeth
which are intermediate in size, or fit
within the overall range of modern
man. The teeth of Homo sapiens may
have become smaller in some areas of
Europe and Asia since the start of the
late Pleistocene some 150 000 years
ago, particularly around 10 000 years
ago, although this is not entirely
proven [5]. In living man, the largest
teeth, on average, are found amongst
the Australian aborigine people, but
they do not hold the published world
record of two U.S. marines whose
teeth overlapped australopithecines’ in
size [6].
Tooth size may also be used to
establish the sex of a skeleton. Some of
the toothed whales show extreme
sexual dimorphism in tooth size - for
example in the great tusk of the male
narwhal. Many of the carnivores show
sexual dimorphism in the size of their
canine teeth and blade-like carnassial
teeth, and tooth measurements have
been used to distinguish between male
and female cave bears, cave lions, and
weasels. The distribution of tooth sizes
in the mouth also differs between men
and women. Multivariate statistical
studies of measurements of several
classes of teeth have been successfully
used to distinguish between the sexes
in archaeological.material. The method
works not only for permanent teeth,
but also for the deciduous, or milk
teeth. This is of considerable import-
ance for archaeology, because it is
difficult to distinguish between boys
and girls on their bony skeleton alone.
Reduction in size seems to have
accompanied the early stages of domes-
145

Figure 1 Section through a Neolithic
human incisor from Tunisia (scale bar is
Icm). The worn crown is at the top of the
picture and thick cement coats the root.
The dentine has been affected by a
tunneling mycelium, but an area of
transparency in the root dentine can still
be seen. The size of this area yields an
age-at-death estimate of 68 + 11 years.
tication in several mammals. This is
true, for example of dogs, sheep and
goats, pigs, and particularly cattle. In
some mammals this size reduction is
also shown in teeth. Wild cattle, or
aurochs, in Mesolithic Denmark had
substantially larger teeth than the
domestic cattle of the Neolithic. In
dogs and pigs, jaw and tooth reduction
seems to have occurred even before
overall body size reduction, and tooth
measurements are used as indices for
domestication. The current published
record for earliest domestication is the
reduceddentitionofal2 OOOyearolddog
jaw from Palaeolithic levels of Palegawra
Cave in Iraq [2]. The processesinvolved
are not understood but, since this first
reduction the teeth of dogs seem to have
remained roughly the same size. Dogs
with longer jaws simply have the teeth
spaced further apart and dogs with
146
shortened jaws have the teeth crowded
together. This is particularly marked in
the modern bulldog, whose cheek teeth
are stacked-up one beside another.
Determination of age-at-death is yet
another area where teeth are invalu-
able to archaeology. The teeth grow,
migrate through the bone of the jaws,
and erupt into the mouth in a reason-
ably consistent sequence for each
species. Different classes of teeth start
to form, are completed, and erupt at
different times. The stage of dental
development reached by young indi-
viduals can thus be used to assesstheir
relative maturity. Once the teeth are
functional and in wear due to the abra-
sive nature of the diet, then the extent to
which they are worn is proportional to
the length of time for which they have
been in use. This is dependent on the
diet being constant and on many other
factors, but seems to hold reasonably
true for many mammals. Numerous
recording schemes for tooth growth,
eruption, and wear have been devised;
for example in man, sheep and goats,
cattle, pigs, horses, various deer, lla-
ma, hippopotamus, elephant, dugong,
dog, cat, lion, fox, badger, weasel,
woodchuck, voles, and housemouse.
Not all are directly applicable in
archaeology, but they can be adapted
[5]. In many of these schemes, the
stage of dental development is associ-
ated with a range of ages, at which
each is attained in living animals. From
these, the age-at-death is suggested.
There is some danger in this, because
tooth formation and eruption are
known to vary with sex, ethnic group,
and economic background (in man)
and breed (in domestic animals). The
Figure 2 Canals produced by a tunneling mycelium in dentine from the tooth in
figure 1. Scanning electron micrograph of polished, etched surface (field width c.
140pm). The canals are lined with secondary minerals, deposited whilst the tooth
was in the ground.
rate of wear varies similarly. Work is
being carried out on modern domestic
breeds in order to assessthe constancy
and timing of the sequences. Despite
uncertainties, dental development is
much more clear cut and regular than
most of the age-related changes in the
skeleton and provides a more hopeful
basis for age determination,
In man, several age-related changes
inside the tooth are used for age
determination in forensic studies. Of
these, the most reliable seemsto be the
growth of a transparent zone of dentine
in the roots of the teeth. This is due to
changes at a microstructural level, but
can be monitored on a simple slice
through the tooth (figure 1) by measur-
ing the transparent zone. These
measurements are then converted into
rough estimates of age-at-death using
standard formulae [l]. The method has
not been widely used in archaeology,
but has particular potential because it
is applicable to older adult individuals
which are difficult to age by other
means.
The dental tissues - the enamel
coating the crown, the dentine forming
the main body of the tooth, and the
cement which coats the roots - have a
very intricate microscopic structure.
They are, however, mineral/organic
composites and are variably affected by
decomposition of their organic compo-
nent over time. This has little effect on
enamel, which is 99 per cent mineral,
but dentine and cement contain 65-75
per cent of mineral when fresh and
may be softened or made brittle by
decomposition. Fresh dentine contains
about 19 per cent protein by weight
and this can be reduced to as little as
0.1 per cent in badly affected
archaeological specimens. The extent
to which proteins and other parts of the
organic component are lost varies from
site to site. This is partly a factor of
time, but is also affected by the
chemistry of the soil in which the teeth
are buried. The calcium phosphate
minerals of the inorganic component
remain stable in a wide range of
environments, but parts of the organic
component also may be preserved. Cell
walls have been demonstrated, still in
place, in Bronze Age human dentine
from the U.S.S.R. and dried cells from
the pulp have been found adhering to
the side of the pulp chamber in teeth
from early Mediaeval Belgium. In both
cases, no other soft tissues were
preserved [5]. Very little work has
been done on this aspect and it may
well be an important area for future
research. Dentine and cement may be
adversely affected by the tunnelling of
mycelium-producing microorganisms
(figure 2). These spread out from the
pulp chamber deep inside the tooth
(figure 1) and also extend in from the
root surface, forming canals up to
20pm in diameter. They are often
lined by a skin of secondary minerals,
deposited from the ground waters
which percolate through the soil.
Enamel may be attacked by a microor-
ganism which produces all the histolo-
gical features of the disease dental
caries and care must be exercised in
palaeopathological studies. The mic-
roorganisms involved have not yet
been identified, but they seem to affect
the teeth early on in the process of
decomposition.
The microscopic structure of ancient
enamel may help in the identification
Figure 3 Prisms in the enamel of a Mediaeval human tooth from York, England.
Scanning electron micrograph of fractured, etched surface (field width c. 50Fm). The
alternate swellings and constrictions
shadows in figure 5.
Figure 4 Complex structure in the enamel of a porcupine incisor from a
l\leolithic site in Tunisia. Scanning electron micrograph of fractured, etched surface
(,‘ield width c. 2OO~m).
of species. Enamel is composed of
prisms; bundles of tiny crystals (figure
3) woven together into a complex
‘felted’ mass. The type of weave varies
between different groups of mammals.
It is, for example possible to disting-
uish the enamel of carnivores, of
pinnipeds (seals and sealions), pri-
mates, ungulates (hoofed mammals),
lagomorphs (hares, rabbits etc.), and
rodents. The highly complex enamel of
the rodents makes it possible to
distinguish between families and even
some genera (figure 4). This type of
evidence may be useful for small
fragments of teeth, or the incisors of
rodents, which are difficult to identify
represent cross striations, seen as small
from their external form alone.
Growth of the crown leaves its mark
in the microscopic structure of the
enamel. The prisms alternately bulge
out and constrict along their length
every 5pm or so (figures 3 and 5).
These structures are known as cross
striations and are thought to record a
circadian rhythm in enamel growth.
The progress of growth is also recorded
in the layering of enamel. Boundaries
between layers occur at spacings of 6 to
14 cross striations and are seen under
the polarising microscope as fuzzy
brown lines, the brown striae of
Retzius (figures 5 and 6). These
probably correspond to a break or
dislocation of the prisms. The cross
striation count between brown striae is
constant for each individual (7 or 8 is
the average in man) and the layering is
thought to represent another biological
rhythm. Cross striations and brown
striae are more prominent in some
mammals than others, and are especial-
ly marked amongst the primates and
the carnivores. Almost all archaeolo-
gical work on these enamel structures
has been carried out on human teeth.
The brown striae vary in ‘intensity’ in a
way that seems to be under systemic
control, because the sequence of brown
striae in one tooth crown may be
matched with other crowns which were
growing at the same time. One applica-
tion of this is that isolated teeth from
one individual may be matched up: this
may be useful in a burial where
remains from several skeletons are
mixed together. Another application is
based on the fact that crowns from the
different classes of teeth are formed at
different times during the growth
period. In permanent teeth this begins
147
 with the first molar, which starts to
form around birth and ends with the
third molar crown which is completed
around the mid-teenages. An overlap-
ping series of other tooth classes
connects the two and the zones of
overlap in the sequence may be
connected by matching the sequences
of brown striae. In this way, an
extended sequence of brown striae and
cross striations can be built up from
birth (which sometimes registers as a
particularly pronounced brown stria) to
the end of the crown growth period
around 15 to 17 years of age. If
someone was unfortunate enough to
die within this period, then their
age-at-death can be determined with
considerable precision by counting the
layers up to the point when enamel
formation stopped. It is also possible to
follow growth rate by determining the
age at which different stages of the
dentition were completed. More con-
troversial is the idea that the intensity
with which each brown stria is de-
veloped is related to some systemic
disturbance which is in turn related to
environmental stresses. In studies of
American and Nubian prehistoric
material [8, 91 the relative frequency of
occurrence of particularly pronounced
brown striae has been used to suggest
differences in diet and infectious dis-
ease between different groups.
All these features of enamel need to
be studied from sections of tooth
crowns. These are time-consuming to
prepare and destroy valuable material.
Another possibility is to study in-
cremental structures on the surface.
Brown striae of Retzius form concen-
tric, dome-shaped layers during the
first part of crown formation, but when
the enamel of the cusps is finished, the
rest of the crown is completed as
sleeve-like layers (figure 6). For each
sleeve, where the brown striae reach
the crown surface, a small trough is
formed. Several of these together give
the appearance of a series of waves,
.called perikymata (figure 7). Careful
counts of perikymata correspond very
closely to counts of brown striae in the
sleeve region, so a similar chronology
of crown formation can be built up.
Disturbances to enamel formation reg-
ister at the crown surface as defects
called enamel hypoplasia: these vary
considerably in form. At their simplest,
they are little more than especially
pronounced perikymata, producing a
groove-like defect around the tooth
(figure 8). Other forms appear as a
band of pitting around the crown and
the most serious defects greatly disrupt
the crown form. Enamel hypoplasia of
these kinds is caused by a variety of
growth disturbances, ranging from
neurological conditions to diabetes, but
the most common causes are thought
to be infections which produce fever,
148
and dietary deficiencies. Inherited
hypoplasia also occurs, but it is rare
and can be distinguished because it
affects all the crowns of all teeth,
whereas growth-disturbance hypoplasia
occurs as bands representing particular
periods of disturbance. These bands
can be matched up between teeth and
counts of the perikymata in between
make it possible to reconstruct a
chronology of growth and its disturb-
ances. Another method is to record the
position of the defect on the crown and
apply standard tables for crown forma-
tion timing to produce a similar, but
less detailed chronology. These techni-
ques are rapid and can be based on
simple silicone rubber casts of the
crown surface. Only modest magnifica-
tion is needed, so that low-power
microscopes can be used and there are
possibilities for recording the defects
automatically from replicas of the tooth
surface. Information on growth dis-
turbances during childhood is pre-
served in this way, even in the teeth of
adults. Such studies shed light on the
diet and health of children in ancient
communities [3, 41.
Similar diurnal and longer-period
growth rhythms can be studied by
examining layering in dentine. These
can be matched up between teeth in a
similar way to enamel and hypoplasia
also registers as a defect in dentine
mineralisation. In marine mammals
such as the toothed whales and some of
the pinnipeds, there may also be a
seasonal or annual layering in the
dentine. Counts of these layers are
used in zoology to determine age-at-
death [7] and the technique has been
established by studies of tagged anim-
als. Marine mammal teeth do occur on
coastal archaeological sites and the
method has potential as an age-
determining technique. The difficulty
for all this in archaeology is that
dentine is frequently poorly preserved.
Similar difficulties obtain for cement,
which also commonly has layers which
seem to occur in response to annual or
seasonal rhythms. This is particularly
true of some toothed whales and seals,
but also seems to be the case in large
herbivores on land, in hibernating
mammals , a range of carnivores, and in
man himself (figure 9). Tests of the
method on animals of known age have
proved less reliable in terrestrial than in
marine mammals, but there is consider-
able potential for determining age in
archaeological material.
Traces of dental pathology are fre-
quently found in material from
archaeological sites [5]. This includes
not only human material, but also teeth
and jaws from a variety of mammals.
Some conditions, such as dental caries
(decay) present themselves as in living
patients (figure 10). The hard, mineral-
ised deposits of calculus (figure 11)
even preserve the outlines of bacteria
from ancient dental plaque. Periodon-
tal disease, however, is primarily an
inflammatory condition of the gums
and their attachment to the tooth. The
10~sof bone which is the sign of the
disease in archaeological material
(figure 12) is only one part of the
process. The same is true of some
other conditions of the jaws, but by
and large there are fewer problems of
diagnosis than elsewhere in the
skeleton. Dental caries can be di-
agnosed by normal methods of ex-
amination and radiography, but care is
needed because conditions of preserva-
tion may mimic the stain, pitting, or
radiolucency which would normally
demonstrate the presence of a carious
lesion. Clear signs of the disease are,
however, present in australopithecine
jaws over one million years old, in
Homo erectus and in material of all
dates up to the present. In Europe, the
rate of caries attack seems to have
increased only slowly through Prehis-
toric and Mediaeval times, with the
main change to modern levels occur-
ring in the 19th century. The pattern of
caries seems also to have changed. In
modern times (although the position is
changing somewhat now) most caries
occurs in the fissures of the cheek teeth
and at the edges of the contact areas
between neighbouring teeth. The
mediaeval and pre-mediaeval pattern
was for caries to occur most commonly
around the neck of the tooth, where it
meets the gums. This change may well
reflect a major increase in importation
and consumption of refined sugars
during the 19th century. Caries has also
been found in mammals other than
man. It is quite common amongst
domestic horses and dogs, but ancient
examples also include the teeth of
mastodons, cave bears, and sabre-
toothed cats. It is not clear what role, if
any, sugar had in such cases.
Periodontal disease is found in a
range of animals; in the primates
(including man himself), rodents,
hoofed mammals, and carnivores. It is
more common in domestic animals
than in the wild state, especially in
dogs, cats, horse, cattle, and sheep. In
man, the disease affects particularly the
supporting tissues of the cheek teeth
and bone is lost around their roots, so
that eventually they fall out. In most
cases, the disease progresses slowly,
active phases alternating with quies-
cent, and the amount of bone lost
increases with advancing age. This
pattern can be seen quite clearly in
archaeological material, although there
are some problems with diagnosis.
Bone loss clearly related to periodontal
disease has been recognised in Homo
erectus and Neanderthaler fossils tens
and hundreds of thousands of years
old, and is very common in later
 Figure 5 Prism outlines, cross striations
and brown striae of Retzius in the
enamel of a human tooth from Nubia (c.
1700 BC). Polarising microscopy of
polished thin section (field width
2OOpm). Prism outlines run from lower
right to upper left. Cross striations are
seen as small shadows along the prism
boundaries. Brown striae of Retzius are
the fuzzy brown lines running
horizontally. 8 cross striations can be
counted between each brown stria.
Figure 6 Brown striae of Retzius in the
enamel of a human tooth from Nubia (c.
1700 BC). Polarising microscopy of
polished thin section (field width c.
5mm). The enamel layer is on the left of
the picture and to the right is the
dentine, with the pulp chamber of the
tooth in the lower right corner. The
brown striae run diagonally through the
enamel layer and vary in their
development.
Figure 8 Enamel hypoplasia in human
teeth from Nubia (c. 1700 BC). The
defects run like grooves around the
circumference of the tooth crowns and
represent at least three separate growth
disturbances through childhood.
Figure 10 Dental caries in a human jaw
from Nubia. One of the premolar tooth
crowns has been completely destroyed
Figure 7 Perikymata in the crown by the disease.
surface of a human tooth from Mediaeval
York. Scanning electron micrograph
(field width c. 2mm).
collections of human skeletons. There
is also evidence for bone loss, probably
related to periodontal disease, in
ancient sheep. It has been demons-
trated in Roman and Mediaeval mate-
rial from Britain, but is quite unlike the
pattern of periodontal disease in mod-
ern sheep. The most common dental
disease of today’s sheep involves rapid
loss of the front teeth, causing the Figure 11 Dental calculus deposits on
animal to die from starvation, without human teeth from Nubia.
greatly affecting the bone of the jaws.
By contrast, the archaeological condi-
tion mainly affects the cheek teeth and
the extensive changes suggest that the
animals must have lived with the
disease for some time.
Studying teeth represents good value
for archaeology. A great deal of
potential information is locked up in
small fragments of tissue and for a
given research effort, a disprop-
ortionately large amount of detail can
be added to the record of an archaeolo-
gical site. Ancient teeth also represent
an unusual and interesting area for Figure 12 Periodontal disease in a
Mediaeval human lower jaw from Britain,
applying zoology, histology, physical The cheek teeth have all been lost and
anthropology, pathology, dentistry,
the bone has healed over again.
and forensic science. There are still
many avenues of research waiting to be
exploited.
References
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[3] Goodman, A. H., Armelagos, G. J.
and Rose J. C. Human Biol., 52,
515, 1980.
[4] Hillson, S. W. World Archaeology, 11,
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[5] Hillson, S. W. ‘Teeth’, Cambridge,
1986.
[6] Keene, H. J.’ Amer. J. Physical
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[7] Morris, P. Mammal Rev. 2, 69, 1972.
[8] Rose, J. C., Armelagos, G. J. and
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Figure 9 Cement layers in the tooth [9] Rudney, J. C. J. Human Evol., 12,
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149