International Journal of Osteoarchaeology

Int. J. Osteoarchaeol. 18: 262279 (2008)

Published online 31 August 2007 in Wiley InterScience
(www.interscience.wiley.com) DOI: 10.1002/oa.934

Dental Pathology and Diet at

Apollonia, a Greek Colony on
the Black Sea
A. KEENLEYSIDE*
Department of Anthropology, Trent University, Peterborough, Ontario, K9J 7B8, Canada

ABSTRACT

Dental pathology has the potential to provide insight into the composition of the diet and to
reveal dietary differences based on age, sex and social status. Human skeletal remains from
the Greek colonial site of Apollonia (5th to 2nd centuries BC) on the Black Sea coast of Bulgaria
were analysed for various forms of dental pathology in order to: assess the prevalence of
dental disease in the population; compare the dental pathology data from Apollonia with
dietary data derived from ancient literary texts and from previous stable isotopic analysis of the
colonists remains; explore variations in dental disease with respect to age and sex; and
compare the prevalence of dental pathology in the Apollonians with that of other Greek
populations. The composition of the diet, as indicated by the dental pathology data, is
consistent with the stable isotopic evidence from Apollonia and with the ancient literary texts,
both of which indicate the consumption of a relatively soft, high carbohydrate diet. The higher
frequency of dental caries, abscesses, calculus, and antemortem tooth loss in older adults
compared with younger ones reflects the age-progressive nature of these conditions. The lack
of significant sex differences in caries, abscesses, calculus and tooth loss corresponds with
the stable carbon and nitrogen isotopic data derived from bone collagen, which indicate no
significant sex differences in the consumption of dietary protein. In contrast, these findings
conflict with the ancient literary texts, which refer to distinct dietary differences between males
and females, and with the stable carbon isotopic values derived from bone carbonate, which
indicate sex differences with respect to the overall diet. Despite the lack of marked sex
differences in dental pathology, overall trends point to subtle dietary differences between
males and females. A greater degree of tooth wear in males also hints at possible sex
differences in the use of the teeth as tools. Copyright 2007 John Wiley & Sons, Ltd.
Key words: dental pathology; diet; Greek colonial populations; Black Sea

Introduction
Dental pathology is commonly recorded in archaeological samples, and its use in palaeodietary
reconstructions is based on the recognised link
between dental pathology and diet (Nelson et al.,
1999: 333). Studies of dental caries, antemortem
tooth loss, and other forms of dental disease
have explored the relationship between these
* Correspondence to: Department of Anthropology, Trent University, Peterborough, Ontario, K9J 7B8, Canada.
e-mail: akeenleyside@trentu.ca

Copyright # 2007 John Wiley & Sons, Ltd.

conditions and modes of subsistence (Littleton &

Frohlich, 1993; Tayles et al., 2000), investigated
dietary changes over time (Walker & Erlandson,
1986; Lillie, 1996), and examined age-, sex-, and
status-based differences in diet (Frayer, 1984;
Lukacs, 1996; Cucina & Tiesler, 2003). As an
indirect source of information on diet, dental
pathology has also been combined with direct
evidence of diet derived from stable isotopic
analysis of bone collagen and bone carbonate
(Sealy & van der Merwe, 1988; Lubell et al., 1994;
White, 1994; Lillie & Richards, 2000; Prowse,
2001).
Received 10 July 2006
Revised 30 April 2007
Accepted 11 May 2007

Dental Pathology in a Greek Colony

The earliest examination of dental pathology
in Greek skeletal samples was conducted by
Angel (1944), who compared dental pathology in
samples of early and modern Greek teeth with the
goal of assessing temporal changes in dental
health. Subsequent studies have examined dental
morphology and disease in approximately 1500
Middle Minoan (17501550 BC) teeth from
Knossos (Carr, 1960), explored the relationship
between dental caries and selenium and fluoride
levels in archaeological and modern Greek teeth
(Hadjimarkos & Bonhorst, 1962), examined the
impact of changing subsistence patterns on dental
wear in Mesolithic and Neolithic remains from
Franchthi Cave (Smith & Cook, 1991), and
compared the skeletal and dental evidence for
social status in a Late Bronze Age skeletal sample
from the Athenian Agora (Smith, 2000).
Dental pathology data have been recorded in a
small number of Greek colonial samples, most of
them from southern Italy (Becker & Donadio,
1992; Becker, 1995, 1996; Henneberg, 1998;
Henneberg & Henneberg, 1998, 2003). In
contrast, little is known about the dental health
of Greek colonists on the Black Sea. Few studies
of ancient Greek populations have explored sex
differences in dental health, and none have
compared data on dental pathology with that
derived from literary sources and stable isotopic
analyses to test the utility of each as sources
of information on diet (for an excellent example
of this approach applied to Roman remains see
Prowse, 2001). The ancient literary texts refer to
distinct sex differences in diet among the ancient
Greeks. At Apollonia, stable carbon isotope
values derived from bone carbonate indicate
sex differences with respect to the overall diet,
while stable carbon and nitrogen isotope values
measured in bone collagen revealed no sex
differences in the consumption of dietary protein
(Keenleyside et al., 2006).
The aim of this study is to assess the prevalence
of dental pathology in a skeletal sample from the
Greek colonial site of Apollonia on the Black Sea
coast of Bulgaria, explore variations in dental
disease with respect to age and sex, compare the
dental pathology data from Apollonia with
dietary data derived from ancient literary texts
and from previous stable isotopic analysis of the
colonists remains, and compare the prevalence of
Copyright # 2007 John Wiley & Sons, Ltd.

263
dental pathology in the Apollonians with that of
other Greek populations.

Archaeological and biocultural context of the

sample
According to literary sources, the Greek colony
of Apollonia was founded in 610 BC by the city of
Miletus in Asia Minor, on the site of the
present-day town of Sozopol on the Black Sea
coast of Bulgaria (Figure 1). Thought to have been
occupied by Thracians at the time of the colonys
foundation, the site had a number of desirable
characteristics, including an excellent harbour,
good fishing, and an easily defensible location
(Nedev & Panayotova, 2003). Its strategic
location contributed to its establishment as a
major trading centre, and Apollonia became a
prosperous urban centre during the 5th and 4th
centuries BC, minting its own coins, producing its
own ceramics, erecting monumental public
buildings, and establishing secondary settlements
in the surrounding territory (Nedev & Panayotova, 2003). Archaeological evidence indicates a
trading relationship between the colonists and
the local Thracian population, and close connections between the two groups are further
indicated by the presence of Thracian names on

Figure 1. Map of Apollonia (Sozopol) showing the

location of the necropolis.

Int. J. Osteoarchaeol. 18: 262279 (2008)

DOI: 10.1002/oa

264
several Greek tombstones excavated from the
necropolis (Nedev & Panayotova, 2003). Like
other Greek colonies on the Black Sea coast, the
inhabitants of Apollonia attempted to recreate
the same way of life they had enjoyed in their
homeland (Tsetskhladze, 1997). While they
brought the religious and political institutions
of their mother city with them, and maintained
commercial links with Miletus and other cities in
Asia Minor, they remained independent and
self-sufficient (Nedev & Panayotova, 2003).
In 1938, the chance discovery of a grave in an
area known as Kalfata, located on the shore of the
Black Sea approximately 2.5 km south of Sozopol
(Figure 1), prompted scholars to direct their
attention to this area, and from 1946 to 1949,
large-scale excavations uncovered 801 graves
consisting of simple unlined pits, stone cists, tile
graves, wooden coffins and urn burials (Venedikov, 1963). Extending to a depth of five metres in
some places, most of the burials were typical
Greek inhumation burials containing single
individuals laid out on their backs in an extended
position, with their heads oriented primarily to
the east. The remains dated from the second half
of the 5th century to the beginning of the 2nd
century BC, based on associated grave goods
(Venedikov, 1963). A small number of cremation
and flexed burials, dating from the mid to late 4th
century BC, have been tentatively identified as
Thracian burials (Venedikov, 1963). Since 1992,
excavations led by Dr Kristina Panayotova of the
Institute of Archaeology in Sofia, of the Kalfata
necropolis and several adjacent sites have
uncovered an additional 400 burials (Panayotova,
1998; Nedev & Panayotova, 2003). Palaeopathological analysis of some of these remains has
revealed evidence of iron deficiency anaemia,
trauma and non-specific infections among the
colonists (Keenleyside & Panayotova, 2005,
2006). The sample utilised in this study is derived
from this most recent excavation.

Dental pathology and diet

Dental caries, the most common form of dental
pathology, is an age-progressive disease process
characterized by the focal demineralization of
dental hard tissues by organic acids produced by
Copyright # 2007 John Wiley & Sons, Ltd.

A. Keenleyside
the bacterial fermentation of dietary carbohydrates, especially sugars (Larsen, 1997: 65).
Of the factors underlying the development of
caries, diet has been identified as the most
important (Powell, 1988). Diets containing
refined sugars are more cariogenic than those
consisting of unrefined carbohydrates, while diets
high in protein and fat and low in carbohydrates
are known to impede caries formation (Powell,
1988). The texture of food also plays a role in the
development of caries, with soft, sticky foods
being more cariogenic than hard, abrasive foods
(Larsen, 1997). Other factors influencing tooth
decay include tooth crown morphology, attrition,
enamel hypoplasia, oral hygiene, fluoride consumption, oral bacteria and the composition of
saliva (Dawes, 1970; Lillie, 1996; Hillson, 2000).
Numerous studies of skeletal samples worldwide have revealed higher rates of caries in
agricultural populations compared with huntergatherers (Turner, 1979; Larsen, 1995; Hillson,
1996), reflecting their greater consumption of
carbohydrates. Agricultural diets have, in fact,
been inferred from high caries rates in the
absence of other types of dietary evidence
(Cohen, 1989: 107). In the western Mediterranean region, for example, the transition to
agriculture was accompanied by an increase in
the rate of caries (Meiklejohn & Zvelebil, 1991).
Hunting and gathering populations, in contrast,
typically show an absence of caries, reflecting
their non-cariogenic, high protein diet; also,
populations heavily reliant on marine resources
typically show low rates of caries (Macchiarelli,
1989; Nelson et al., 1999).
One of the possible consequences of dental
caries is antemortem tooth loss, indicated by the
resorption of alveolar bone in the tooth socket.
Other causes of tooth loss include attrition,
trauma and periodontal disease, the latter being
one of the main causes of antemortem tooth loss
in modern populations (Hillson, 2005: 306).
Dental caries can also lead to periapical
abscessing, which occurs when bacteria invade
the exposed pulp cavity and proceed down the
root canal into the alveolar bone, causing an
infection of the surrounding tissue and the
formation of an abscess (Hillson, 2005). Pulp
exposure resulting from trauma or attrition can
also lead to abscesses.
Int. J. Osteoarchaeol. 18: 262279 (2008)
DOI: 10.1002/oa

Dental Pathology in a Greek Colony

Dental calculus, or mineralised plaque, is
common in archaeological remains. Occurring
both above (i.e. supragingival) and below (i.e.
subgingival) the gums, it is most frequently found
on teeth closest to the salivary glands (Roberts &
Manchester, 1995). The relationship between
diet and the formation of dental calculus is not
straightforward (Hillson, 2001: 265), and high
calculus rates have been associated with both
high protein and high carbohydrate diets
(Meiklejohn & Zvelebil, 1991; Lieverse, 1999;
Lillie & Richards, 2000). When combined with
dental caries, however, it can be used to
determine the relative contribution of proteins
versus carbohydrates to the diet. High calculus
rates and low caries rates have been found to
characterise populations subsisting on a high
protein, low carbohydrate diet (Lillie, 1996: 140),
while heavy calculus deposits and high caries
rates have been observed in populations that
consumed a high carbohydrate diet (White,
1994: 283). The formation of calculus is also
influenced by a number of non-dietary factors,
including oral hygiene, the mineral content of
drinking water, the rate of salivary flow, cultural
practices such as chewing betel nuts and coca
leaves, and the use of the teeth as tools (Lieverse,
1999).
Dental wear can provide valuable information
on the types of foods consumed, their consistency and texture, and their methods of preparation. Known to vary widely among populations, high levels of tooth wear recorded in
agricultural populations have been attributed to
the consumption of coarse cereal grains and the
use of grinding stones which can add a significant
amount of grit to the diet (Larsen, 1997).
Differential wear between the anterior and
posterior dentition, seen in some archaeological
populations, has been interpreted as reflecting
different causes of tooth wear (e.g. diet vs. tool
use) (Littleton & Frohlich, 1993).
Enamel hypoplasia refers to a deficiency in
enamel thickness caused by a disruption in the
secretory phase of enamel matrix formation
(Goodman et al., 1984). Formed during tooth
crown development, hypoplastic defects are not
remodelled in later life, and therefore provide
permanent markers of stress episodes during
infancy and childhood (Goodman et al., 1984).
Copyright # 2007 John Wiley & Sons, Ltd.

265
More than 100 different factors have been linked
to the development of enamel hypoplasia, with
particular attention being paid in the literature to
nutritional stress and childhood infections. Since
it is rarely possible to link enamel defects with
their underlying aetiology, they are considered to
be non-specific indicators of physiological stress
(Goodman & Rose, 1990). In some archaeological
samples, a higher prevalence of hypoplastic
defects in younger individuals compared with
older adults has been interpreted as reflecting an
increased likelihood of individuals who experienced stress during childhood dying at a younger
age (Duray, 1996).

The Ancient Greek diet

Information on the ancient Greek diet derives
from a variety of sources, including literary texts,
most notably Athenaeus Deipnosophistae (Professors
of the Dinner-Table) and Galens On the Properties of
Foodstuffs, archaeological evidence in the form of
plant remains, animal bones, food preparation
utensils, storage vessels, ritual food offerings
associated with graves, and stable isotopic
analysis of human skeletal remains. These sources
indicate that the ancient Greeks relied on a staple
diet of cereals, legumes, olive oil and wine for
their subsistence. Olive oil was the main source of
dietary fat (Martin, 1996), and wine mixed with
water was the preferred beverage. Cereal crops,
most notably wheat and barley, comprised the
primary source of protein and carbohydrates
(Garnsey & Whittaker, 1983: 7; Rathbone, 1983:
46; Garnsey, 1999: 15). Dry legumes such as
lentils, broad beans, peas and chickpeas were also
an important source of protein. Several varieties
of millet were cultivated but were considered an
emergency resource for small farmers (Garnsey,
1988: 52). Similarly, oats were grown mainly as a
fodder crop (Brothwell & Brothwell, 1969: 101). A
variety of fresh fruits, vegetables and nuts were
also consumed, and honey, dates and figs were
the main sweeteners (Brothwell & Brothwell,
1969; Grmek, 1989; Dalby, 1996).
Of the domesticated animals, sheep, pigs and
goats were the primary sources of meat (Dalby,
1996), although meat was relatively scarce and
generally made up only a small component of the
Int. J. Osteoarchaeol. 18: 262279 (2008)
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A. Keenleyside

266
diet (Jameson, 1983: 10; Garnsey, 1999: 16, 123).
Goat and sheep milk was consumed in diluted
form (Brothwell & Brothwell, 1969: 50), and
cheese made from the milk of both animals was
also eaten (Brothwell & Brothwell, 1969: 52). Fish,
including shellfish, were a highly valued part of
the diet of many Greeks (Dalby, 1996) and
garum, or fish sauce, was a popular ingredient in
many dishes.
According to literary sources, sex differences in
diet existed among the ancient Greeks. Males
were given preferential access to foods, while
women were denied meat and other nourishing
food items (Garnsey, 1999). Consequently,
women were at higher risk of nutritional
deficiency and disease, and may have experienced
a higher incidence of malnutrition than men
(Garnsey, 1999). Given this evidence for sexbased variation in diet, one should therefore
expect to find sex differences in dental pathology
in ancient Greek skeletal samples. Such differences might, for example, take the form of lower
caries rates in males resulting from a greater
consumption of meat and fish. Fluoride and
strontium, elements present in high quantities in
marine foods (Siebert & Trautner, 1985; Malde
et al., 1997), are known to impede the formation
of carious lesions.

Archaeological and stable isotopic evidence of

diet at Apollonia
The archaeological evidence recovered from
Apollonia suggests that during the 5th to 2nd
centuries BC, the population subsisted on a varied
diet of grains, nuts, fish, meat and shellfish
(Panayotova, 1998). Ritual fireplaces associated

with some of the graves contained food remains

interpreted as evidence of funerary feasts
(Panayotova, 1998). These included animal
bones, grape seeds, acorns, almonds and hazelnuts, and ceramic fish plates and grills (Panayotova, 1998: 105; Nedev & Panayotova, 2003:
1389). Astragali from sheep/goats, found in the
graves of both children and adults (Panayotova,
1998), suggest the consumption of one or both
species.
Stable isotopic analysis of bone collagen and
carbonate samples taken from the remains of 54
of the adults examined for dental pathology in
this study revealed that the colonists relied on a
mixed diet of terrestrial and marine resources
(Keenleyside et al., 2006). Statistical comparisons
by age and sex revealed no significant differences
in d13C or d15N values for bone collagen,
suggesting similarities in the consumption of
dietary protein. In contrast, males had significantly higher d13C values for bone carbonate,
suggesting sex differences with respect to the
overall diet (Keenleyside et al., 2006). As such, it
is hypothesised that sex differences should be
seen in the prevalence of certain forms of dental
pathology at Apollonia.

Materials and methods

The dentitions of 162 adult skeletons were
examined in this study (Table 1). Sex determination of the skeletons was done using standard
morphological and metric criteria of the crania
and pelvic bones (Bass, 1987; Ubelaker, 1989).
Age estimation was based on pubic symphysis
morphology (Brooks & Suchey, 1990), auricular
surface morphology (Lovejoy et al., 1985) and
ectocranial suture closure (Meindl & Lovejoy,

Table 1. Age and sex distribution of the sample

Age (years)
Males
1835
3650
50
Total

Number of teeth

Number of individuals

22
37


14
73

Females
40
23
20
83

Indeterminate

3
3
0
6

Males

Females

418/511
762/908


171/388
1351/1807

851/1031
424/594
175/436
1450/2061

Indeterminate

65/72
73/73
0/0
138/145

Number of observable teeth/number of observable teeth plus teeth lost antemortem plus teeth lost post-mortem.
Statistically significant differences between males and females, P < 0.05.

Copyright # 2007 John Wiley & Sons, Ltd.

Int. J. Osteoarchaeol. 18: 262279 (2008)

DOI: 10.1002/oa

Dental Pathology in a Greek Colony

1985). Individuals were assigned to one of three
age categories: 18 to 35 years, 36 to 50 years, and
50 years.
All permanent dentitions were examined macroscopically for evidence of pathology. Dental
caries, antemortem tooth loss, abscesses and
calculus were recorded by individual and by
tooth/tooth socket following the methodology
outlined in Buikstra & Ubelaker (1994). Carious
lesions were recorded as present only if they had
penetrated the tooth enamel. The number and
location of lesions per tooth were also noted.
Teeth were scored as being lost antemortem if
their sockets exhibited evidence of resorption.
Dental abscesses were identified by the presence
of a drainage channel through the alveolar bone
at the apex of the tooth root, and their location
(buccal vs. lingual) was also recorded. The
location and degree of dental calculus were
recorded for each affected tooth, following the
criteria outlined in Buikstra & Ubelaker (1994).
The degree of dental wear was recorded for each
tooth using Smiths (1984) 8-stage system for the
incisors, canines and premolars, and Scotts
(1979) 10-stage system for the molars; average
scores were calculated for each individual and
each tooth type (after Prowse, 2001). Finally,
enamel hypoplasia was classified according to the
Developmental Defects of Enamel (DDE) Index
(Commission on Oral Health, 1982). Hypoplastic defects were identified by visual examination
and were scored as present or absent for each of
the six anterior maxillary and mandibular teeth.
All observed defects were classified as either
linear arrangements of pits, horizontal grooves, or
both, and the total number of defects was
recorded for each affected tooth. Those teeth
that exhibited severe attrition resulting in the loss
of more than a third of the tooth crown, heavy
calculus deposits, severe caries or trauma were
excluded from the analysis.
Since the age and sex distribution of a sample
can affect interpretations of dental pathology
data, statistical comparisons were made by age
and sex to determine if there were any significant
differences in the number of individuals and the
number of observable teeth/tooth sockets. These
comparisons revealed that the sample contained
significantly more females than males in the
young adult age category (1835 years) with
Copyright # 2007 John Wiley & Sons, Ltd.

267
respect to the number of individuals (x2 5.29,
P 0.025) and the number of observable sockets
(x2 189.92, P 0.001) and preserved teeth
(x2 217.33, P 0.001). In contrast, there were
significantly more males than females in the
middle-aged category (3650 years) when comparisons were made by individual (x2 8.66,
P 0.01), by socket (x2 186.12, P 0.001),
and by tooth (x2 211.34, P 0.001). Frequencies were calculated for each type of dental
pathology recorded, and statistical comparisons
by age and sex were conducted using the
chi-square test and the Mann-Whitney U test.
All statistical analyses were performed using
SPSS, Version 14.

Results
When recorded by individual, dental caries was
the second most common dental disease observed
in the sample, affecting 53.8% (85/158) of
individuals (Table 2). In contrast, only 7.7%
(227/2939) of teeth had carious lesions (Table 3).
Of the four tooth types, the molars were the most
frequently affected (13.9%) (Table 4), reflecting
their more complex morphology. The majority of
lesions (54.2%) were located on the interproximal surfaces (Table 5), while only 2.9% were
located on the buccal or lingual surfaces. While
both males and females had the greatest number
of carious lesions on the interproximal surfaces of
their teeth, males had considerably more interproximal lesions than females (64.5% compared
with 44.5%), and females, in contrast, had more
cervical lesions than males (19.5% compared with
6.4%) (Table 5). When statistical comparisons
were made by the number of individuals affected,
there was no significant difference in the prevalence of caries between males and females for
any of the three age groups compared (Table 2).
When comparisons were made by the number of
affected teeth, young adult and middle-aged
females had a slightly higher frequency of carious
teeth than males (Table 3), while males over the
age of 50 had a slightly higher frequency than
females. There were no statistically significant sex
differences in caries rates for any of the age
groups examined (Mann-Whitney U test), and
for both sexes, the rates increased with age.
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A. Keenleyside

268
Table 2. Frequency of affected individuals by age and sex
Age (yrs)

Males

Dental caries
1835
11/22
3650
25/37
50
9/14
Total
45/73
AMTL
1835
6/22
3650
16/37
50
13/14
Total
35/73
Abscesses
1835
2/18
3650
11/36
50
6/14
Total
19/68
Calculus
1835
19/22
3650
31/37
50
11/13
Total
61/72
Enamel hypoplasia
1835
6/18
3650
9/32
50
1/7
Total
16/57

Females

Indeterminate

Total

50.0
67.6
64.3
61.6

16/39
14/23
10/18
40/80

41.0
60.9
55.5
50.0

0/3
0/2
0/0
0/5

0
0
0
0

27/64
39/62
19/32
85/158

42.2
62.9
59.4
53.8

27.3
43.2
92.9
47.9

6/40
14/23
19/20
39/83

15.0
60.9
95.0
47.0

0/3
0/3
0/0
0/6

0
0
0
0

12/65
30/63
32/34
74/162

18.5
47.6
94.1
45.7

11.1
30.5
42.9
27.9

4/40
4/22
4/19
12/81

10.0
18.2
21.0
14.8

0/2
1/2
0/0
1/4

0
50.0
0
25.0

6/60
16/60
10/33
32/153

10.0
26.7
30.0
20.9

86.4
83.8
84.6
84.7

28/39
18/23
13/18
59/80

71.8
78.3
72.2
73.7

2/3
3/3
0/0
5/6

66.7
100.0
0
83.3

49/64
52/63
24/31
125/158

76.6
82.5
77.4
79.1

33.3
28.1
14.3
28.1

9/35
4/16
1/8
14/59

25.7
25.0
12.5
23.7

1/3
0/3
0/0
1/6

33.3
0
0
16.7

16/56
13/51
2/15
31/122

28.6
25.5
13.3
25.4

Statistically significant, P < 0.05.

Table 3. Frequency of affected teeth by age and sex

Age (yrs)

Males

Dental caries
1835
19/418
3650
59/762
50
28/171
Total
106/1351
AMTL
1835
24/511
3650
40/908
50
142/388
Total
206/1807
Abscesses
1835
2/358
3650
15/721
50
13/309
Total
30/1388
Calculus
1835
176/414
3650
405/753
50
87/161
Total
668/1328
Enamel hypoplasia
1835
25/140
3650
33/254
50
2/43
Total
60/437

Females

Indeterminate

Total

4.5
7.7
16.4
7.8

43/851
50/424
26/175
119/1450

5.0
11.8
14.9
8.2

0/65
2/73
0/0
2/138

0
2.7
0
1.4

62/1334
111/1259
54/346
227/2939

4.6
8.8
15.6
7.7

4.7
4.4
36.6
11.4

7/1031
44/594
158/436
209/2061

0.7
7.4
36.2
10.1

0/72
0/73
0/0
0/145

0
0
0
0

31/1614
84/1575
300/824
415/4013

1.9
5.3
36.4
10.3

0.6
2.1
4.2
2.2

6/954
7/521
8/393
21/1868

0.6
1.3
2.0
1.1

0/42
2/49
0/0
2/91

0
4.1
0
2.2

8/1354
24/1291
21/702
53/3347

0.6
1.9
3.0
1.6

42.5
53.8
54.0
50.3

285/840
188/418
104/163
577/1421

33.9
45.0
63.8
40.6

13/65
26/72
0/0
39/137

20.0
36.1
0
28.5

474/1319
619/1243
191/324
1284/2886

35.9
49.8
58.9
44.5

17.9
13.0
4.6
13.7

43/307
3/124
3/67
49/498

14.0
2.4
4.5
9.8

2/22
0/26
0/0
2/48

9.1
0
0
4.2

70/469
36/404
5/110
111/983

14.9
8.9
4.5
11.3

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269

(7.7)
(1.6)
(10.3)
(44.5)
(11.3)

It is well-known that antemortem tooth loss

(AMTL) can have an effect on the calculation of
caries rates. To compensate for the antemortem
loss of teeth, some of which may have been
carious, Lukacs (1995) developed a caries correction factor which involves estimating the number
of teeth lost antemortem due to caries by examining the proportion of teeth with pulp exposure
due to caries. The caries correction factor was not
employed in this study, however, as data on pulp
exposure due to caries was not recorded.
Another calculation that relates AMTL to
caries is the Diseased Missing Index (DMI: Kelley
et al., 1991: 205), which involves adding together
the total number of carious teeth and teeth lost
antemortem and expressing this number as a
percentage of the total number of teeth and sockets observed. Application of this method to the
caries data from Apollonia yields a frequency of
17.3% (312/1807) for males and 15.9% (328/
2061) for females, values that are close to double
the standard caries rates (Table 6). However,
statistical comparisons again revealed no significant sex differences in the DMI (Mann-Whitney
U test).
Concern has also been raised about the
possible effect of post-mortem tooth loss on
caries rates. To compensate for the post-mortem
loss of teeth that may have been carious, Erdal &
Duyar (1999) developed a proportional correction factor designed for use on samples with a
high rate of post-mortem loss of the anterior
teeth. This method involves calculating the caries
rate of the anterior and posterior teeth and
multiplying it by a ratio representing the proportion of these teeth in one quadrant of the mouth
(e.g. 3 anterior teeth to 5 posterior teeth, or 0.6).
This correction factor was not applied to the
Apollonia data, however, as the ratio of anterior
to posterior teeth in the sample (0.6) matches the
expected ratio.
AMTL was also common in the sample, affecting 45.7% (74/162) of individuals (Table 2) and
10.3% (415/4013) of teeth (Table 3). Among
tooth types, the molars were most frequently
involved (16.1%). There were no significant sex
differences by individual for any of the age groups
compared (Table 2). Similarly, when comparisons were made by the number of affected teeth
(Table 3), males and females had a similar overall

(6.9)
(1.1)
(10.5)
(50.6)
(12.7)
111/1601
20/1858
226/2150
799/1578

67/527
(8.7)
(2.2)
(10.1)
(37.1)
(9.6)
116/1338
33/1489
189/1863
485/1308
44/456
(2.2)
(0.8)
(7.0)
(52.0)
(7.5)
15/676
7/878
72/1023
347/667
46/615
(3.2)
(0.9)
(4.7)
(48.4)
(17.7)
13/404
4/450
25/531
193/399
65/368

Statistically significant, P < 0.05.

55/821 (6.7)
13/873 (1.5)
90/1040 (8.6)
311/795 (39.1)

(13.9)
(2.5)
(16.1)
(42.2)
144/1038
29/1146
228/1419
433/1025

Caries
Abscesses
AMTL
Calculus
Enamel hypoplasia

Premolars
Molars

Tooth type
Dental pathology

Table 4. Dental pathology by tooth type and dental arcade

Canines

Incisors

Maxilla

Jaw

Mandible

227/2939
53/3347
415/4013
1284/2886
111/983

Total

Dental Pathology in a Greek Colony

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A. Keenleyside

270
Table 5. Frequency of carious lesions by location
Location

Males

Occlusal
Interproximal
Smooth
Cervical
Root
Large

Females

Indeterminate

8/110
71/110
2/110
7/110
11/110
11/110

7.3
64.5
1.8
6.4
10.0
10.0

9/128
57/128
5/128
25/128
11/128
21/128

7.0
44.5
3.9
19.5
8.6
16.4

0/2
2/2
0/2
0/2
0/2
0/2

0
100.0
0
0
0
0

Total

17/240
130/240
7/240
32/240
22/240
32/240

7.1
54.2
2.9
13.3
9.2
13.3

Surface of origin cannot be determined.

prevalence of AMTL. The prevalence of tooth

loss in both sexes increased with age.
Abscesses were the least common type of
dental pathology observed in the sample, affecting 20.9% (32/153) of individuals (Table 2) and
only 1.6% (53/3347) of teeth (Table 3). They
were found more frequently on the buccal side of
the alveolus than the lingual side, and affected the
molars more frequently than the other tooth
types (Table 4). On an individual basis, significantly more males had abscesses than females
overall (x2 3.87, P  0.05) (Table 2). When
comparisons were made by the number of
affected teeth (Table 3), however, males had
only a slightly higher prevalence of abscesses
than females (2.2% compared with 1.1%). For
both sexes, the prevalence of abscesses increased
with age.
Dental calculus was the most common condition observed in the sample, affecting 79.1%
Table 6. Comparison of caries index and DMI by age and
sex
Males
Caries index

1835
4.5
3650
7.7
50
16.4
Total
7.8
DMI


1835
8.4
3650
10.9
50
43.8
Total
17.3

Females

Indeterminate

Total

5.0
11.8
14.9
8.2

0.0
2.7
0.0
1.4

4.6
8.8
15.6
7.7

4.8
15.8
42.2
15.9

0.0
2.7
0.0
1.4

5.8
12.4
43.0
16.0

Total number of carious teeth  100/total number of
observed teeth.



Total number of carious teeth and teeth lost antemortem  100/total number of teeth and sockets
observed.

Copyright # 2007 John Wiley & Sons, Ltd.

(125/158) of individuals (Table 2) and 44.5%

(1284/2886) of teeth (Table 3). The incisors were
the most commonly affected teeth, followed by
the canines, molars and premolars (Table 4). Also,
significantly more mandibular teeth were affected
than maxillary teeth (x2 53.20, P  0.001)
(Table 4). The majority of affected teeth in both
males and females had only slight calculus
deposits. There were no significant sex differences when comparisons were made by the
number of affected individuals (Table 2). When
comparisons were made by the number of
affected teeth (Table 3), males had a higher
prevalence than females overall (50.3% compared
with 40.6%), and young adult and middle-aged
males had a higher rate than females of the same
age groups. None of these differences, however,
was statistically significant (Mann-Whitney U
test). For both sexes, the frequency of calculus
increased with age.
The majority of teeth in the sample showed
slight to moderate occlusal surface wear, as
indicated by the average wear scores calculated
for each tooth type (Table 7). The incisors,
canines and premolars had average scores ranging
from 3.3 to 4.1 out of 8 (using the Smith, 1984
system), and the molars had average scores
ranging from 12.7 to 20 out of 40 (using the Scott,
1979 system). While the use of two different
scoring systems makes it difficult to draw a direct
comparison of the degree of wear between the
anterior vs. the posterior teeth, both sets of teeth
show low to moderate levels of wear. Males in all
three age categories had higher average tooth
wear scores than females for each tooth type
examined (Table 8), but the differences were
not statistically significant for any of the age
groups (Mann-Whitney U test). When statistical
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Dental Pathology in a Greek Colony

271

Table 7. Average tooth wear scores for each tooth type

Tooth type

No. of teeth
observed

Average tooth
wear score

147
165
187
190
189
178
164
105

4.1
3.3
4.0
3.6
3.6
19.7
15.2
12.7

169
198
215
228
197
202
213
147

3.9
3.7
3.7
3.2
3.3
20.0
16.0
14.1

Maxilla
I1
I2
C
P1
P2
M1
M2
M3
Mandible
I1
I2
C
P1
P2
M1
M2
M3

comparisons of the posterior teeth (molars) vs.

anterior teeth (premolars, canines and incisors)
were conducted, however, males had a significantly greater degree of wear in the anterior teeth
than females overall (Mann-Whitney U test,
P 0.004), and young adult males had a significantly greater level of wear on the anterior teeth
than females of the same age group (Mann-Whitney U test, P 0.041). For both sexes, the
degree of tooth wear increased with age for all
tooth types.
Enamel hypoplasia involving the canines and/
or incisors was recorded in 25.4% (31/122) of
Table 8. Average tooth wear scores by age and sex
Category
Sexes combined
1835 years
3650 years
50 years
Males
1835
3650
50
Total
Females
1835
3650
50
Total

Molars

Premolars

Canines

Incisors

13.1
18.7
26.4

2.3
3.9
5.6

2.7
4.4
5.5

2.7
4.4
5.5

13.5
19.4
28.7
18.2

2.5
4.0
5.8
3.9

3.2
4.5
5.9
4.4

3.1
4.5
6.3
4.2

12.8
17.7
24.3
15.3

2.2
3.7
5.4
3.0

2.5
4.1
5.1
3.3

2.5
4.4
4.9
3.3

Copyright # 2007 John Wiley & Sons, Ltd.

individuals (Table 2) and 11.3% (111/983) of

teeth (Table 3). Statistical comparisons by the
number of affected individuals revealed no significant sex differences (Table 2). In contrast,
comparisons of the number of affected teeth
revealed that males had a slightly higher
frequency of defects than females (13.7% compared with 9.8%) (Table 3). For both sexes,
young adults had a higher prevalence of defects
than older adults.

Discussion

Dental caries
In contrast to other studies of caries which have
found significant differences in caries rates between the upper and lower dentition (e.g.
Thylstrup & Fejerskov, 1994; Lingstrom &
Borrman, 1999), there was no significant
difference in caries rates between the maxillary
and mandibular teeth in the Apollonia sample.
The significantly higher frequency of carious
lesions, by number of affected teeth, in older
individuals compared with younger ones is,
however, consistent with previous studies of
caries in archaeological populations, and reflects
the age-progressive nature of this disease
(Hillson, 2000, 2001).
Sex differences in caries rates have been observed in both archaeological and modern populations, with females typically showing higher
rates than males (Walker & Hewlett, 1990;
Hillson, 2001: 253). This has been attributed
to a higher carbohydrate intake by females, earlier
eruption of teeth, and behavioural differences in
subsistence pursuits (Larsen, 1997; Walker &
Hewlett, 1990). Changes in salivary composition
during pregnancy and lactation have also been
implicated in the development of caries in women
(Laine, 2002). In the Greek colonial sample from
rural Metaponto, a significantly higher number of
females (64%) were found to have carious lesions
than males (46%) (Henneberg & Henneberg,
2003: 34). Sex differences in caries rates in the
Apollonia sample are much less pronounced.
While young adult and middle-aged females had
a slightly higher frequency of carious teeth
than males, statistical comparisons revealed no
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A. Keenleyside

272
significant sex differences with respect to either
the caries rate or the DMI, suggesting that males
and females were consuming foods with similar
cariogenic properties.
Comparison of the prevalence of caries in the
Apollonia sample with that of other archaeological samples is hindered by the fact that different
methodologies may have been used to diagnose
and record carious lesions (Hillson, 2001: 249).
Further limiting such comparisons is the varying
age and sex composition of archaeological
samples, the failure to distinguish between tooth
types affected and the different forms of carious
lesions observed, and differing patterns of antemortem and post-mortem tooth loss. Despite
these limitations, it is useful to compare the
prevalence of caries (uncorrected) at Apollonia
with that of other archaeological samples, including those with varying subsistence strategies. As
expected, there is considerable variability in the
prevalence of caries between populations with
similar modes of subsistence (Turner, 1979;
Larsen, 1995: 188). This is not surprising given
methodological differences in the recording and
reporting of caries rates. At 7.7% of teeth, the
prevalence of caries at Apollonia falls at the lower
end of the range reported by Turner (1979) for
agricultural populations (2.126.9%, average
10.43%), and within the range reported for
populations with a mixed economy (1.010.3%,
average 4.84%). It also falls within the range
reported for other Greek skeletal samples
(3.236.9%) (Table 9).

It is interesting to compare the dental caries

data from Apollonia with those derived from
other coastal sites, particularly those for which
stable isotopic data have also been obtained. The
Neolithic site of Alepotrypa Cave and the Greek
colonial site of Metaponto in southern Italy are
particularly noteworthy in this regard. As indicated in Table 9, the rate of dental caries (10.5%)
for the rural cemetery from Metaponto is slightly
higher than that recorded at Apollonia (7.7%);
however, the average d13C and d15N values
measured in the Metaponto sample (19.6% and
10.5% respectively: Henneberg & Henneberg,
2003) approximate those measured in the
Apollonians (18.5% and 10.1% respectively)
(Keenleyside et al., 2006), indicating that both
populations relied heavily on terrestrial resources.
For Alepotrypa the caries rate was lower (3.2%),
but this calculation included both permanent and
deciduous teeth. The average d13C and
d15N values measured in the remains from
Alepotrypa (19.95% and 7.17% respectively)
also point to a primary reliance on a terrestrial C3
diet (Papathanasiou, 2005: 385).

Antemortem tooth loss

The rate of AMTL in the Apollonia sample is
similar to that seen in other agricultural populations and probably reflects the higher rate of
caries typically seen in such populations (Larsen,
1995). A significantly higher number of older

Table 9. Prevalence of dental caries in Greek skeletal samples

Sample/time period

Frequency by tooth

Neolithic (3rd millennium BC)

Neolithic (Alepotrypa Cave)

Middle Bronze Age (20001400 BC)
Mycenean (14001150 BC)
Lefkandi (Iron Age)

Early Iron Age (1150650 BC)
Knossos (7th or 8th c. AD)
Ailias (Middle Minoan)
Kato Zakro (Middle Minoan)

Khania (Late Minoan)

Armenoi (Late Minoan)
Classical (650150 BC)
Metaponto (7th2nd c. BC)

32/267
14/436
34/601
22/356
11/113
40/724
14/90
135/1498
11/152
76/206
367/2069
36/724
192/1828

(12.0%)
(3.2%)


(5.7%)
(6.2%)
(9.7%)
(5.5%)
(15.5%)
(9.0%)
(7.2%)
(36.9%)
(17.7%)
(5.0%)
(10.5%)

Reference
Angel (1944)
Papathanasiou (2005)
Angel (1944)
Angel (1944)
Musgrave & Popham (1991: 289)
Angel (1944)
Musgrave (1976: 46)
Carr (1960: 11922)
Becker (1975: 275)
McGeorge (1992: 39)
McGeorge (1992: 39)
Angel (1944)
Henneberg & Henneberg (1998: 535)

Coastal sites.
Permanent and deciduous teeth combined.

Copyright # 2007 John Wiley & Sons, Ltd.

Int. J. Osteoarchaeol. 18: 262279 (2008)

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Dental Pathology in a Greek Colony

individuals had lost teeth prior to their death than
younger individuals, a finding consistent with the
age-related increase in AMTL seen in modern
populations. Causes of AMTL include caries,
attrition, periodontal disease, trauma and abscesses (Hillson, 2000). Metabolic diseases such as
scurvy can also lead to tooth loss (Wols & Baker,
2004: 68). Determining the cause(s) of AMTL,
particularly in archaeological samples, may be
difficult. The fact that dental caries and AMTL
most frequently affected the molar teeth in the
sample suggests that caries may have been a major
cause of tooth loss in this population. Similar
rates of tooth loss among males and females
correspond with the caries data. Periodontal
disease was not scored in this study, but may also
have contributed to tooth loss among the colonists. Low to moderate levels of tooth wear in the
sample and the small number of cases of pulp
exposure resulting from wear suggest that tooth
wear was not a major contributor to AMTL
among the colonists.

Dental abscesses
The prevalence of abscesses in the sample is quite
low. While it is consistent with the low rate of
caries and the low to moderate rate of tooth wear
recorded in the sample, it may also reflect the
extraction of teeth prior to pulp exposure and
infection, the loss of abscessed teeth and the
subsequent remodelling of the alveolar bone long
before death, and/or the inability to observe
abscesses that have not yet penetrated the
alveolar bone (Hillson, 2000; Wols & Baker,
2004). The significantly higher frequency of
abscesses by tooth socket in older individuals
compared with younger ones is mirrored in other
archaeological samples, and reflects the higher
rate of caries and tooth wear typically seen in this
age group. Similar rates of abscesses in males and
females mirror the data on caries and AMTL.

Dental calculus
The Apollonian diet does not appear to have
promoted the formation of large amounts of
calculus on the teeth, as most of the affected teeth
Copyright # 2007 John Wiley & Sons, Ltd.

273
had only slight deposits. Dental calculus, like
caries, abscesses and AMTL, was significantly
more common in older individuals compared
with younger ones. This phenomenon has been
observed in other populations and reflects the
progressive build-up of calculus with age (Beiswanger et al., 1989). Few studies of Greek skeletal
remains report the prevalence of dental calculus;
however, the high rate of calculus in the
Apollonia sample is consistent with that of other
agricultural populations, and points to a reliance
on a high carbohydrate diet and/or poor oral
hygiene.
As noted earlier, the relationship between
calculus and diet is not straightforward (Hillson,
2001), and high calculus rates have been
associated with both high protein and high
carbohydrate diets (Meiklejohn & Zvelebil, 1991;
Lieverse, 1999; Lillie & Richards, 2000).
Significant sex differences, reported for some
populations, have been interpreted as reflecting
inequality in access to dietary protein (Lillie &
Richards, 2000). In the case of Apollonia, males
had a higher prevalence of calculus than females
overall (50.3% compared with 40.6%), and
young adult and middle-aged males had a higher
rate than females of the same age groups. None of
these differences, however, was statistically
significant (Mann-Whitney U test), suggesting
that males and females consumed a similar diet.

Tooth wear
Occlusal surface wear is an indicator of the type
of foods consumed and methods of food
preparation. Among ancient Greek populations,
high levels of tooth wear have been attributed to a
diet of uncooked and unrefined food items such as
coarse cereal grains, the consumption of poorly
washed foods, and the use of grinding stones
(Becker, 1975: 275; McGeorge, 1992: 3940). In
contrast, low levels of wear are associated with
the consumption of soft foods. While comparisons of tooth wear in the Apollonia sample with
those of other archaeological samples are
hindered by the use of different methodologies
to record tooth wear, as well as the failure of
publications to report the methods used, the low
to moderate levels of wear recorded in the
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274
Apollonia sample are consistent with those
recorded in other agricultural populations,
including Greek colonial samples from Pithekoussai (Becker, 1995: 279) and rural Metaponto
(Henneberg & Henneberg, 1998: 536), and
suggest the consumption of a relatively soft diet.
The association between tooth wear and age is
reflected in greater levels of tooth wear in older
individuals compared with younger ones. The
finding that males had a significantly greater
degree of wear on the anterior teeth than females
raises the possibility that non-dietary factors,
perhaps activities associated with fishing, may
have contributed to the wearing down of their
front teeth.

Enamel hypoplasia
The prevalence of enamel hypoplasia in the
sample is relatively low. The lack of significant
sex differences in the frequency of hypoplastic
defects among the colonists suggests that males
and females both suffered from physiological
stresses during early childhood. The higher
frequency of the condition in younger adults
compared with older individuals is consistent
with studies of other archaeological populations
(Larsen, 1997), and supports the hypothesis that
individuals who experience stress during childhood are more likely to die at a younger age
(Duray, 1996).
Comparison of the prevalence of enamel
hypoplasia in the Apollonia sample with that
of other samples is hindered by methodological
differences in the scoring of this condition, most
notably the lack of a minimum defect size
considered to be a hypoplastic defect (Goodman
et al., 1980: 527; Goodman & Rose, 1990: 92;
Hillson, 2000). Nevertheless, the frequency of
individuals with defects in the Apollonia sample
(25.4%) is not significantly higher than that
reported by Bisel & Angel (1985) for Hellenistic
period Greeks (18.8%) from other archaeological
sites. It is lower than that reported for Classical
period Greeks (37.9%) (Bisel & Angel, 1985), and
significantly lower than that reported for Metaponto, where 78% (88/113) of individuals in the
rural sample and 96% of individuals in the urban
Copyright # 2007 John Wiley & Sons, Ltd.

A. Keenleyside
sample had enamel hypoplasia (Henneberg &
Henneberg, 2001).
Hypoplastic defects in Greek skeletal remains
have been interpreted as evidence of nutritional
stress, childhood infections and/or weaning
stress. Angel (1984: 64), for example, attributed
the significant increase in the prevalence of
enamel hypoplasia in the Middle Bronze Age to
childhood epidemics resulting from increased
population size and density. Henneberg and
colleagues have suggested that the high frequency of defects observed in the rural sample
from Metaponto most likely reflects the presence
of some form of chronic, endemic infection,
possibly treponemal disease, which appears to
have been common in the population (Henneberg & Henneberg, 1998). With respect to the
urban sample from Metaponto, Henneberg &
Henneberg (2001) suggested that the high
prevalence of hypoplastic defects may reflect
infectious diseases spread by crowded living
conditions.
The factors underlying the development of
enamel hypoplasia in the Apollonian colonists
probably included both nutritional stress and
exposure to infectious diseases. As indicated by
the stable isotopic data, the colonists relied
heavily on a terrestrial diet of C3 plants, suggesting that nutritional deficiencies may have been
common in the population. Palaeopathological
analysis of 184 skeletons excavated from the site
revealed the presence of cribra orbitalia in 28% of
individuals, and iron deficiency anaemia has been
proposed as the most likely cause of these lesions
(Keenleyside & Panayotova, 2006). Periodic food
shortages, common in antiquity (Garnsey, 1999),
may also have contributed to the formation of
these defects. Ancient literary sources indicate
that Greek colonies on the Black Sea coast of
Romania and Ukraine suffered periodically from
food shortages during the Hellenistic period
(Garnsey, 1988: 13, 163, 274), and it is likely that
Apollonia experienced similar food shortages on
occasion.
The possibility that the hypoplastic defects
seen in the Apollonian sample resulted from
infectious diseases must also be considered.
While the skeletons analysed to date show no
evidence of chronic infection such as that seen at
Metaponto (Henneberg & Henneberg, 1998),
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Dental Pathology in a Greek Colony

this does not preclude the possibility of infectious
diseases having been a health problem among the
colonists. The population of Apollonia is estimated to have numbered approximately 3000
citizens during the 5th to 4th centuries BC
(Danov, 1948), and excavations have revealed
that during this time period, the centre of the
settlement was characterised by high-density
occupation (Nedev & Panayotova, 2003). It is
therefore conceivable that the population may
have been exposed to pathogenic microorganisms associated with crowded living conditions.
Other factors that have been linked to the
development of enamel hypoplasia include the
stresses associated with weaning (Corruccini et al.,
1985; White, 1994). Infant feeding vessels
recovered from child graves in Greece suggest
that weaning may have begun relatively early
(Fildes, 1986: 25), and literary sources indicate
that cereals were the basic weaning food among
the ancient Greeks (Garnsey, 1999). A recent
stable isotopic study of infant feeding practices at
Apollonia has revealed that infants were weaned
onto a diet of cereals and dairy products, foods
that would have predisposed them to nutritional
deficiencies (Kwok, 2007).

Comparison of the dental pathology data

with the stable isotopic evidence and ancient
literary sources
The composition of the diet, as indicated by the
dental pathology data, is consistent with the
stable isotopic evidence from Apollonia and with
ancient literary texts, both of which indicate a
heavy reliance on carbohydrates. The lack of
significant sex differences in dental pathology is
also consistent with the stable carbon and
nitrogen isotopic data derived from bone
collagen, which indicate no significant sex
differences in the consumption of dietary protein
(Keenleyside et al., 2006). In contrast, the dental
pathology data conflict with the ancient literary
texts, which highlight distinct dietary differences
between males and females, and with the stable
carbon isotopic values derived from bone
carbonate, which indicate sex differences with
respect to the overall diet.
Copyright # 2007 John Wiley & Sons, Ltd.

275
The lack of marked sex differences in dental
pathology is somewhat surprising given the
textual and isotopic evidence for dietary differences between males and females. It is useful,
however, to look at overall trends in the dental
pathology data. Higher rates of caries in females
and higher rates of calculus in males do hint at
subtle dietary differences in the form of greater
consumption of carbohydrates by females and
greater consumption of protein by males. This
corresponds with the stable carbon isotopic
values derived from bone carbonate, in particular
the isotopic spacing between bone collagen and
apatite (carbonate), which can be used as an
indicator of trophic level (Krueger & Sullivan,
1984). Large differences between d13Capatite and
d13Ccollagen reflect a more herbivorous diet, while
small differences reflect a more carnivorous diet.
In the Apollonia sample, the apatite-collagen
spacing is higher for females than males (8.8%
compared with 7.9%), suggesting greater carbohydrate consumption by females and greater
protein consumption by males.
When interpreting dental pathology data,
consideration must also be given to the nondietary use of the teeth. While stable isotopic
data and ancient literary texts may provide useful
information on diet, they tell us nothing about
non-dietary uses of teeth that might be reflected
in certain forms of dental pathology. As noted
earlier, greater levels of tooth wear in males
compared with females, particularly in the
anterior teeth, raises the possibility that nondietary factors, perhaps activities associated with
fishing, may have contributed to greater tooth
wear among males.

Conclusions
The prevalence of dental pathology observed at
Apollonia is typical of that seen in other
agricultural populations, and indicates the consumption of a relatively soft, high carbohydrate
diet. It is consistent with the stable isotopic
evidence for diet at Apollonia and with the
ancient literary texts, both of which indicate a
heavy reliance on carbohydrates. The prevalence
of caries among the colonists is similar to that
observed in the Greek colony of Metaponto, for
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A. Keenleyside

276
which isotopic data also indicates a primary
reliance on a terrestrial C3 diet.
The higher frequency of dental caries,
abscesses, calculus and AMTL in older adults
compared with younger ones reflects the ageprogressive nature of these conditions. Sex
differences in dental pathology are not pronounced, and, as such, are consistent with the
stable carbon and nitrogen isotopic data derived
from bone collagen, which indicate no significant
sex differences in the consumption of dietary
protein (Keenleyside et al., 2006). In contrast,
these findings conflict with the ancient literary
texts, which attest to distinct dietary differences
between males and females, and with the stable
carbon isotopic values derived from bone
carbonate, which indicate sex differences with
respect to the overall diet.
Despite the lack of strong evidence for sex
differences in dental pathology, overall trends in
the data, particularly when combined with the
isotopic data, do hint at subtle dietary differences
between males and females, with males consuming more protein and females consuming more
carbohydrates. As such, the integration of dental
pathology data with other types of evidence
provides greater insight into the diet of past
populations than reliance on one source of
information alone. A greater degree of tooth
wear in males also hints at sex differences in the
non-dietary use of the teeth, an observation not
recorded in the ancient texts.

Acknowledgements
I would like to thank Kristina Panayotova, Institute of Archaeology in Sofia, Bulgaria, and Dimitar Nedev, Director of the Archaeological
Museum in Sozopol, Bulgaria, for providing
access to the skeletal sample examined in this
study. I am grateful to Tracy Prowse, whose
research inspired this paper, and to Peter Garnsey,
Richard Lazenby, and the reviewers of an earlier
draft of this manuscript for their helpful comments. Fieldwork in Bulgaria was supported by a
Leverhulme Research Fellowship and a grant from
the Social Sciences and Humanities Research
Council of Canada (#410-2005-0452).
Copyright # 2007 John Wiley & Sons, Ltd.

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