AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 135:6474 (2008)

Oral and Physiological Paleohealth in Cold Adapted

Peoples: Northeast Asia, Hokkaido
Marc F. Oxenham1* and Hirofumi Matsumura2
1
2

School of Archaeology and Anthropology, Faculty of Arts, Australian National University, Canberra, ACT, Australia
Department of Anatomy, Sapporo Medical University, Sapporo, Hokkaido, Japan
KEY WORDS

bioarchaeology; health; jomon; Okhotsk; Alaska

ABSTRACT
This paper examines variables useful in
reconstructing oral (caries, antemortem tooth loss, alveolar defects) and physiological (cribra orbitalia, linear
enamel hypoplasia) well-being in two bioarchaeological
assemblages from Hokkaido, Japan: Okhotsk (n 5 37
individuals) and Jomon (n 5 60). Findings are compared
and contrasted with each other, with published series
from Honshu Japan, and samples from climatically nearequivalent Alaska. It was found that more meaningful
comparisons of Hokkaido paleohealth could be made
with Alaskan material, rather than the more southerly
Jomon. Results were ambiguous with respect to physiological well-being. Low levels of LEH in the cold-adapted
samples suggest operating in arctic and subarctic environments with marine-based subsistence regimes is not
physiologically expensive. However, the relatively high
levels of cribra orbitalia in Hokkaido, relative to Alaska,

suggest the picture is not straightforward: the reasons

for elevated cribra orbitalia in Hokkaido are unclear.
The subarctic and arctic samples formed three broadly
similar groupings in terms of oral health profiles: (1)
Aleuts and Eskimo; (2) Ipiutak and Tigara; (3) Hokkaido
Jomon, Okhotsk, and Kodiak Island. Differences between
these groupings could be explained with a combination of
sample demographics and subsistence orientations. The
extremely high frequency of caries in one sample, caribou
hunting Ipiutak, may have been influenced by factors such
as low levels of dietary magnesium and potentially cariogenic foodstuffs, such as preparations of caribou stomach
contents. It was concluded that oral health profiles are
potentially sensitive to differences in subsistence strategies among cold-adapted huntergatherers, although
they lack predictive value. Am J Phys Anthropol 135:64
74, 2008. V2007 Wiley-Liss, Inc.

The island of Hokkaido is of interest in the study of

human adaptation and behaviour in marginal environments due to both its long occupation and subarctic environment. The earliest occupation of Hokkaido extends to
the Upper Palaeolithic at least 20,000 yr BP, while the
period of interest in this paper begins with the early
Jomon at approximately 6,000 yr BP and continues
through to the final Jomon in 2,400 yr BP (Habu, 2004).
In Hokkaido a further stage, not seen in the rest of
Japan, is identified as the Epi-Jomon (or Zoku Jomon)
which continues through to approximately 1,400 yr BP
(Crawford and Takamiya, 1990). This period is contemporaneous with the Yayoi and early Kofun periods in
other parts of Japan, which saw the introduction of agriculture amid mass movements of migrants from the
mainland (Imamura, 1996; Hudson, 2003).
The Jomon culture-complex is best described for the
central Japanese island Honshu where the earliest evidence for the Jomon arguably dates back to 16,500 yr
BP (Habu, 2004, 42). The occupation of Hokkaido by
Jomon peoples was a rather later event and the timing
was no doubt influenced by the need to develop a new
set of adaptive strategies to operate in a subarctic environment. The Jomon, following the Palaeolithic and lasting for over 10,000 years, is often characterized as a
complex huntergatherer society that used and produced
pottery (Habu, 2004). Despite this long stretch of time
and considerable regional diversity throughout the Japanese archipelago, the Jomon (as the use of the single
name also suggests) is still perceived as having a certain
degree of unity and homogeneity; in terms of material
culture this manifests as the virtually ubiquitous cord
marking commonly applied to the surface of pottery as

well as the high frequency of hunting tools (Aikens and

Higuchi, 1981; Imamura, 1996). A further reason for this
perception is perhaps the physical characteristics, or biological unity, of the Jomon people which likely share a
common ancestry with Neolithic Southeast Asian populations (Turner, 1976, 1989, 1990, 1992; Hanihara, 1990,
1992a,b, 1993; Matsumura, 1994, 1995, 2006; Matsumura and Hudson, 2005) and are cranially (Ogata, 1981;
Yamaguchi, 1982; Mouri, 1986; Kondo, 1994) and dentally (Matsumura, 1989, 2007) relatively homogenous on
the one hand and distinct from the Yayoi and later modern populations of Japan on the other.
Apart from the Jomon peoples, portions of Hokkaido
and outlying islands were occupied by the Okhotsk people between the 6th and 12th centuries AD (Maeda,
2002; Hudson, 2004). These relative latecomers were
morphologically Northeast Asian, physically very robust
and originally from the Amur River basin in Russia
(Yamaguchi, 1982; Ishida, 1988, 1996). The Okhotsk
were already well adapted to the sub-Arctic zone and

C 2007
V

WILEY-LISS, INC.

Grant sponsor: Sapporo Medical University, Hokkaido, Japan.

*Correspondence to: M.F. Oxenham, School of Archaeology and
Anthropology, AD Hope Building, College of Arts and Social Sciences, Australian National University, Canberra, ACT 0200, Australia.
E-mail: marc.oxenham@anu.edu.au
Received 15 January 2007; accepted 14 July 2007
DOI 10.1002/ajpa.20709
Published online 4 September 2007 in Wiley InterScience
(www.interscience.wiley.com).

65

PALEOHEALTH IN HOKKAIDO
brought technological and behavioral skills to the region
that allowed them very successfully to pursue the
hunting of large marine animals, such as whales and sea
lions, and also exploit the diverse range of other resources in the region (Hudson, 2004).
The overall nature and patterns of human health in
the prehistoric and protohistoric periods of Hokkaido is
still incompletely understood despite a number of bioarchaeological (e.g., Chisholm et al., 1992; Hanihara
et al., 1994, Minagawa, 2001; Oxenham et al., 2006a)
and biodistance (e.g., Yamaguchi, 1974, 1981; Ishida,
1988, 1996; Dodo and Kawakubo, 2002; Matsumura
et al., 2006) studies. Further, the biological costs associated with living in sub-Arctic conditions in the past,
whether in Northeast Asia, northern America or Europe,
require additional work. Nonetheless, So (1980) has suggested that despite marginal conditions such populations
were well adapted and were not as unhealthy as may
have been predicted, particularly given their high protein and low carbohydrate diets. In terms of diet,
at least, even extremes in the relative contribution of
carbohydrates and protein is not necessarily inconsistent
with a balance in required nutrients.
The aim of this paper is to reconstruct aspects of the
behavior and health of two culturally and temporally
discrete populations from Hokkaido and outlying islands,
namely the northern Jomon and Okhotsk, using curated
adult skeletal and dental samples. Specifically, summary
health profiles of these samples will be examined by focusing on cribra orbitalia, enamel hypoplasia, dental caries,
alveolar defects and antemortem tooth loss (AMTL). The
health profiles of these samples will then be compared and
contrasted with: (1) each other; (2) other, southern, adult
Japanese prehistoric samples; and (3) non-Japanese adult
samples from climatically and ecologically near-equivalent
contexts (a range of Alaskan samples).

MATERIALS, METHODS AND

BIOCULTURAL CONTEXT
Japan
Hokkaido, with a temperature that fluctuates between
a low of 2108C in the winter (ice flows occur in the Sea
of Okhotsk during this time) and a maximum of 258C in
the summer, is the largest northern most continuously
occupied island in the Japanese archipelago. The climate
in Hokkaido has fluctuated in a complex manner over
the course of the Holocene but can be summarized as
being relatively warmer during the initial through middle Jomon and colder toward the end of the Middle and
into the Late Jomon (Koizumi et al., 2003). Regarding
the assemblages, sample size varies depending on
whether dental disease and enamel hypoplasia is being
assessed (dental sample) or cribra orbitalia (cranial sample). The Hokkaido Jomon sample derives from 15 separate sites (Tables 1 and 2, Fig. 1) and is represented by
62 dental individuals (1,458 alveoli, 1,092 teeth) and
58 cranial individuals. Approximately half the sample
derives from three sites: Funadomari (on Rebun Island),
Kitakogane and Irie. The Okhotsk sample is represented
by 7 sites, approximately half come from Wakkanai
(Tables 1 and 2), of which there are 35 dental individuals (833 alveoli, 530 teeth) and 36 cranial individuals.
The relatively small sample sizes for the individual sites
necessitated grouping the assemblages into separate
composite Jomon and Okhotsk samples. Further, given
that plant husbandry seems to have [still] been periph-

eral to their way of life (Crawford and Takamiya, 1990,

p907) the Epi-Jomon samples are subsumed within the
general Jomon sample for Hokkaido. Individual sites
will not be referred to in the subsequent analysis and
discussion. Because of differences in the age-at-death
categorizations in the comparative material, samples are
characterized as being relatively old or young with reference to the proportion of each sample below or above 35
years of age. Both Hokkaido series have approximately
equal numbers of older ([35 years old) and younger
(\35 years old) individuals.
In terms of subsistence behaviors for the samples in
this study, the Jomon and Okhotsk communities from
Hokkaido can be characterized as sea mammal hunters
(whales, sea lions, seals, sea otters for the most part)
who were also involved in fishing and plant gathering to
a degree. The stable isotopic data indicate that 4346%
of dietary protein came from sea mammals in contrast to
only 35% from land mammals for Jomon communities
in Hokkaido (Minagawa, 2001). This is in stark contrast
to the situation in central Japan where the marine mammals only contributed 317%, while the land mammal
contribution was 2037% (Minagawa, 2001). Further,
archaeological evidence suggests plants and terrestrial
animals were more important in the diet of southern
Jomon (Honshu) communities compared to Hokkaido
(Habu, 2004) suggesting a diet high in proteins and
relatively low in carbohydrates in the north (see also
Minagawa, 2001). However, Crawford and Takamiya
(1990) note that some form of horticulture was practiced
by the Jomon in Hokkaido. For the Okhotsk, archaeological evidence suggests they shared similar dietary and
subsistence strategies to the Hokkaido Jomon (see Hudson, 2004), although stable isotopic data suggests an
even greater marine dependence for the Okhotsk (Chisholm et al., 1992). Okhotsk populations also appear to
have utilized and cultivated a range of plant foods
(Hudson, 2004).
Comparative dental data, for caries at least, are extensive for Honshu, and Fujitas (1995, see Table 1) data for
14 sites and 195 individuals are used here. Unfortunately, fewer comparative Jomon data are available for
linear enamel hypoplasia (LEH) and cribra orbitalia.
Hirata (1990) provided details on cribra orbitalia in 44
individuals from the late to final Jomon in central Honshu, while Yamamoto (1988) summarized the level of canine and incisor LEH for 31 individuals (119 teeth), the
majority being from the middle to late Jomon.

Alaska
These samples derive from both Arctic and sub-Arctic
contexts and range in age from just before pre-European
contact to 3,500 yr BP (Table 1). The arctic zone is characterized by temperatures ranging from 2498C in the
winter to 258C in the summer, whereas the sub-Arctic
zone has a similar temperature range to Hokkaido. The
Alaskan material includes: three Arctic samples from
Point Hope and Point Barrow; and two sub-Arctic samples from Chaluka, Umnak Island, in the Aleutian chain
and Jones Point, Kodiak Island (Fig. 2). In terms of subsistence, fish and seal played important dietary roles for
both Point Hope communities, although the Ipiutak have
been characterized as caribou hunters and the Tigara as
whale hunters (Costa, 1980). The Ipiutak and Tigara
samples have relatively young age profiles with more
younger (\35 years old) than older ([35 years) individu-

American Journal of Physical AnthropologyDOI 10.1002/ajpa

Location

American Journal of Physical AnthropologyDOI 10.1002/ajpa

Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita

(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)

This study
This study
This study
Keenleyside
(1998)
Costa (1980)
Costa (1980)
Costa (1980)
Keenleyside
(1998)

Reference
b,c

Early Jomonh
MiddleLate Jomon
MiddleLate Jomon
Late Jomon
MiddleLate Jomon
MiddleFinal Jomon
Late Jomon
LateFinal Jomon
Final Jomon
Early Jomon
Late Jomon
LateFinal Jomon
Early Jomon
Final Jomon

Pre-Euro. contact
1,600
400200
1,450100

5,1001,400
5,1002,400b,c
1,500600
3,000500

Date
(yrs BP)
(58)
(47)
(37)
(54)

7
9
14
19
31
14
9
12
26
11
18
14
5
6

79
46
200
128 (118)

60
49
35
65

Individuals
na

2,258
1,409
5,887
3,483/3,502g

1,458
1,152
833
1,566/1,558g

Tooth
positions

28.6
30.6
29.4
27.4

24.3
23.5
23.8
24.1

Positions/n

179
151
245
241
705
292
130
186
345
113
113
196
115
118

1,566
922
4,375
1,840

1,092
853
530
807

Preserved
teeth

Numbers in parentheses refer to sample sizes for cribra orbitalia assessment.

Initial Jomon starts "8,000 bp, but only three individuals derive from this period in the sample.
Uncalibrated radiocarbon dates.
d
Jomon excluding the epi-Jomon sample.
e
Age-at-death distribution based on a subsample of 17 individuals with assessable auricular areas.
f
Age-at-death distribtion based on a subsample of 27 individuals with assessable auricular areas.
g
Abscess demonimator/AM denominator.
h
Fujita does not provide absolute dates, although uncalibrated dates for the Jomon of central Japan are similar to those for Hokkaido (see Table 2).

Hokkaido
Hokkaido
Hokkaido
Aleutian Islands
(Chaluka)
Kodiak Island
Jones Point, Alaska
Ipiutak
Point Hope, Alaska
Tigara
Point Hope, Alaska
Eskimo
Pt. Hope/Pt. Barrow,
Alaska
Jomon samples (Honshu)
Uwasato
Northern Honshu
Nakazawahama
Northern Honshu
Kaitori
Northern Honshu
Sanganji
Northern Honshu
Ubayama
Central Honshu
Kasori
Central Honshu
Yoyama
Central Honshu
Ikawazu
Central Honshu
Hobi
Central Honshu
Hikozaki
Southern Honshu
Yosekura
Southern Honshu
Tsugumo
Southern Honshu
Kou I
Southern Honshu
Kou II
Southern Honshu

Jomon
Jomond,e
Okhotskf
Aleuts

Sub-Arctic
samples

TABLE 1. Adult dental summary: Japanese and sub-Arctic samples used in this study

25.6
16.8
17.5
12.7
22.7
20.9
14.4
15.5
13.3
6.3
6.3
14.0
23.0
19.7

19.7
20.0
21.9
14.4

18.2
17.4
15.1
12.4

Teeth/n

54.4
69.6
59.5
44.5

47.0
52.0
20.0

Young
% !35
years

45.6
30.4
40.5
55.5

53.0
48.0
80.0

Old
[35
years

66
M.F. OXENHAM AND H. MATSUMURA

67

PALEOHEALTH IN HOKKAIDO
TABLE 2. Sample description for Hokkaido Jomon and Okhotsk
Sample size
Sample name

Map no.

Location

Period

Midorimachi
Kitakogane
Kotan-Onsen
Funadomari
Takasago
Irie
Usujiri
Tenneru
Misawa
Onkoromanai
Rebunge
Minami-Usu
Bozuyama
Chatsu4
Total

1
2
3
4
5
6
7
8
9
10
11
12
13
14

Abashiri City
Date City
Yakumo Village
Rebun Island
Abuta Village
Abuta Village
Minamikayabe Village
Kushiro City
Chitose City
Wakkanai City
Toyoura Village
Date City
Ebetsu City
Tomari Village

Initial Jomon
Middle Jomon
Middle Jomon
Late Jomon
Late Jomon
Late Jomon
Late Jomon
Late Jomon
Final Jomon
Epi Jomon
Epi Jomon
Epi Jomon
Epi Jomon
Epi Jomon

Ohmisaki
Hamanaka2&1
Oshonai
Utoro-Jinjayama
Menashidomari
Pirikatai
Tomiiso
Total

15
16
16
17
18
15
19

Wakkanai City
Rebun Island
Rebun Island
Shari Village
Kitami-Esashi Village
Wakkanai City
Wakkanai City

Okhotsk
Okhotsk
Okhotsk
Okhotsk
Okhotsk
Okhotsk
Okhotsk

Datea (yrs BP)

"8,0005,100
5,1004,050
5,1004,050
4,0503,000
4,0503,000
4,0503,000
4,0503,000
4,0503,000
3,0002,400
2,4001,400c
2,4001,400c
2,4001,400c
2,4001,400c
2,4001,400c
5501200
5501200
5501200
5501200
5501200
5501200
5501200

ADd
ADd
ADd
ADd
ADd
ADd
ADd

Individualsb

Alveoli

Teeth

(1)
(9)
(4)
(15)
(6)
(9)
(2)
(0)
(1)
(2)
(2)
(4)
(2)
(1)
(58)

39
214
123
302
147
224
54
32
17
74
44
97
57
34
1,458

10
161
87
249
129
157
26
22
12
67
33
71
35
33
1,092

17 (19)
4 (4)
3(3)
6 (6)
1 (1)
2 (1)
2 (2)
35 (36)

411
111
53
165
7
58
28
833

255
74
22
116
6
52
5
530

3
9
4
14
6
9
2
1
1
3
2
4
2
2
62

Habu (2004, 42).

Dental individuals (cribra orbitalia individuals).
c
Crawford and Takamiya (1990)
d
Hudson (2004).
b

Fig. 1. Location of Jomon and Okhotsk samples from Hokkaido examined in this study: 1 Midorimachi; 2 Kitakogane; 3
Kotan-Onsen; 4 Funadomari; 5 Takasago; 6 Irie; 7 Usujiri; 8
Tenneru; 9 Misawa; 10 Onkoromanai; 11 Rebunge; 12 MinamiUsu; 13 Bozuyama; 14 Chatsu4; 15 Ohmisaki and Pirikatai; 16
Hamanaka2&1 and Oshonai; 17 Utoro-Jinjayama; 18 Menashidomari; 19 Tomiiso.

als. The diet of the Eskimo from Point Barrow has been
described as similar to that of the Point Hope communities although a preference for either caribou or whale is
not mentioned (Keenleyside, 1998). The Eskimo sample
is demographically similar to the Jomon samples,
although with a ratio of 45:55 favoring older individuals
may be better seen as demographically older.
Regarding the two sub-Arctic communities, the Aleut
diet also focused on a wide range of marine resources
including seals, otters, sea lions, whales, fish, shellfish
and birds (Keenleyside, 1998). Kodiak Islanders con-

Fig. 2. Location of arctic and subarctic north American comparatives samples referred to in this study: 1 Point Barrow
(Eskimo); 2 Point Barrow (Ipiutak); 3 Point Hope (Tigara); 4
Jones Point (Kodiak Island); 5 Chaluka (Aleutians).

sumed fish and seal as staples and whales to a lesser

degree when compared to the more northerly Tigarans
at least (Costa, 1980). The Aleut sample is unique in
having a much greater proportion of older individuals,
while the Kodiak sample is demographically similar to
the Eskimo and Jomon. Keenleyside (1998) provides
comparative Alaskan frequencies of cribra and enamel
hypoplasia for the Aleutians (54 individuals) and Eskimo
(118 individuals).
Differing reported age-at-death data from published
studies drawn on for this paper, and the need for a sin-

American Journal of Physical AnthropologyDOI 10.1002/ajpa

68

M.F. OXENHAM AND H. MATSUMURA

gle consistent age-at-death summary of each sample,

only allowed for a dichotomization of each sample into
young (\35 years) and old ([35 years). Both Keenleyside
(1998) and Costa (1980) provide age-at-death data that
allow for a division of the Aleust, Kodiak Island, Ipiutak,
Tigara, and Eskimo into old and young at a 35 years of
age break point. Keenleyside defines adults as #18 years
while Costas (1980) youngest adults are 16 years.
Because previous curation methods in Hokkaido involved
the dismantling of complete individuals and storage by
element type, age determination of dental material was
difficult. Dentally adult individuals were defined as having a complete adult dentition, including the eruption of
at least one M3, or where M3s had not erupted and
agenesis was suspected, M2 wear consistent with at
least several years of post eruption use. In order to estimate the proportion of old versus young in the sample
the auricular areas of separately stored os coxae were
assessed for age-at-death using Buckberry and Chamberlains (2002) method and assigned to old and young age
categories. An assumption was made that the age distribution of the pelvic remains was consistent with that of
the dental remains.
Methods and procedures for scoring and recording the
three dental conditions (caries, alveolar defects, or
abscesses and AMTL), and two markers of physiological
well-being (LEH and cribra orbitalia) examined in the
Japanese Hokkaido series follow those set out in Oxenham (2006) and Oxenham et al. (2006b). While this
study focuses on adult health, enamel hypoplasia of the
permanent dentition and cribra orbitalia (see StuartMacadam, 1985) form during subadult development. The
recording protocols used in this study are believed to be
broadly commensurate with those employed by authors
of the comparative data for Alaska and more southerly
Jomon Japan. In interpreting the results relating to the
teeth themselves (e.g. caries and LEH) potential under
representation of certain tooth classes need to be considered. While sample representation of teeth by type (incisors, canines, premolars, molars) were not available for
Costas (1980) data (Ipiutak, Tigara, Kodiak), the position/n figures in Table 1 indicate better tooth preservation than any of the other Arctic and sub-Arctic samples.
The observed percentage of anterior (% observed/
expected incisors and canines) to posterior (% observed/
expected premolars and molars) teeth for the Jomon
(50.7/60.6), Jomon excluding the epi-Jomon (49.5/57.3),
Okhotsk (42.4/50.3), Aleut (38.7/48.7), and Eskimo (27.8/
45.4) samples indicate relative under-representation of
the anterior teeth. While in low caries situations anterior teeth will be less likely to be affected by lesions (see
Hillson, 2001, 251252). The absence of these teeth will
reduce the denominator, potentially causing calculated
caries rates in these samples to be inflated.

RESULTS
Oral health profile
As not all of the comparative samples have data on
dental conditions by individual, oral conditions are only
reported by tooth count. In terms of the frequency of
caries in Japan, 11/14 southern samples (Honshu Jomon)
display statistically significantly (v2 range 10.9259.32,
P ! 0.001) higher rates than the northern (Hokkaido
Jomon and Okhotsk) series (Table 3). The three southern
Jomon samples with lower caries rates than Hokkaido
have sample sizes ranging from 5 (115 teeth) to 9 (151

teeth) individuals and the Uwasato and Kou I samples

did not display any sign of carious lesions. Kou II, on
the other hand, displayed the second highest caries rate,
after Sangangi, at 12.7% tooth count (all four individuals).
Figure 3 (see also Table 3) summarizes the oral health
comparison between the Hokkaido samples and Alaskan
material. The Aleut and Eskimo samples have similar
oral profiles with the highest frequencies of AMTL, both
at 18.3%, highest levels of alveolar defects (7.5% and
6.0% respectively) and the lowest frequency of carious
lesions (0% and 0.05% respectively). The Aleuts and
Eskimo also have similar diets in terms of their focus on
seal, fish, and whale and both samples have the oldest
age-at-death profiles.
The Ipiutak appear anomalous with a very high frequency of carious lesions (statistically significantly
higher, v2 range 59.59131.23 P ! 0.001, than any other
sample at 14.3%) and AMTL (15.1%). The frequency of
alveolar defects (1.6%) is depressed in comparison to all
samples with the exception of Kodiak Island. The Ipiutak are unique in being the only group where caribou
forms an important dietary staple. The Tigara share
similarities with the Ipiutak and the only statistically
significant difference (v2 131.23, P ! 0.001) between the
two is in terms of caries which is only 4.4% in the Tigara
assemblage. Further, these assemblages have the youngest age-at-death profiles sampled in this study. In terms
of subsistence, the Tigara are unique in targeting whale
to a greater extent than any other sample.
The Hokkaido samples, Jomon and Okhotsk, have similar oral health profiles and only AMTL is statistically
significantly higher in the Jomon (7.5% compared to
4.8%, v2 6.52, P ! 0.025). The Kodiak Island sample is
also arguably broadly similar to the Hokkaido Jomon
with no significant statistical difference between the
rates of caries and AMTL. While Kodiak Island AMTL is
low relative to any other Alaskan sample, it is significantly greater than the Okhotsk rate (6.7% and 4.8%
respectively, v2 3.89, P ! 0.05). The most pronounced difference between Kodiak Island and the northern Jomon
is the low rate of alveolar defects in the Kodiak Island
sample (0.4% compared to 3.4%, v2 48.27, P ! 0.001). All
three assemblages have broadly similar diets in terms of
animal protein and share similar age-at-death profiles.
While individual teeth were not analyzed for signs of
extramasticatory wear or task-wear-facets, two Jomon
period individuals with relatively complete dentitions
displayed wear patterns consistent with the use of the
teeth as tools. While the mandible of KO9 (adult male,
Kotan-Onsen) was edentulous with extensive alveolar
remodeling, the maxilla displayed extensive angled wear
to the labioocclusal aspect of all preserved teeth (Fig. 4).
Another adult male from the same site (Fig. 5) exhibited
asymmetrical maxillary and mandibular wear with
extensive crown loss on the left-hand side.

Physiological well-being
Linear enamel hypoplasia. Figure 6 displays the frequency of canine and incisor LEH, by tooth count, for
samples from southern (Honshu) and northern (Jomon
and Okhotsk) Japan, two Alaskan samples (Eskimo and
Aleut) and two early (5,0001,800 yr BP) samples from
Southeast Asia, illustrating the upper end of LEH
frequency range for Asian samples. The rate of combined
canine and incisor LEH is statistically greater in the
central Jomon (v2 17.2569.12, P ! 0.001) than any of

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69

PALEOHEALTH IN HOKKAIDO
TABLE 3. Adult oral pathology summary: Japanese, Arctic, and sub-Arctic samples
Arct & sub-Arctic
samples

Site

Jomon
Hokkaido
Jomonc
Hokkaido
Okhotsk
Hokkaido
Aleuts
Aleutian Islands (Chaluka)
Kodiak Island
Jones Point, Alaska
Ipiutak
Point Hope, Alaska
Tigara
Point Hope, Alaska
Eskimo
Pt. Hope/Pt. Barrow, Alaska
Other Japanese Samples
Uwasato
Northern Honshu
Nakazawahama
Northern Honshu
Kaitori
Northern Honshu
Sanganji
Northern Honshu
Ubayama
Central Honshu
Kasori
Central Honshu
Yoyama
Central Honshu
Ikawazu
Central Honshu
Hobi
Central Honshu
Hikozaki
Southern Honshu
Yosekura
Southern Honshu
Tsugumo
Southern Honshu
Kou I
Southern Honshu
Kou II
Southern Honshu
a
b
c

Antemortem
tooth lossa
7.5
8.0
4.8
18.3
6.7
15.1
5.1
18.3

(48.3)
(51.0)
(42.9)
(73.7)
(65.3)
(na)
(na)
(65.3)

Alveolar
Defectsa
3.4
2.9
5.3
7.5
0.4
1.6
3.0
6.0

(36.7)
(36.7)
(40.0)
(66.7)
(na)
(na)
(na)
(56.8)

Cariesb

Reference

2.6
2.6
1.9
0
3.5
14.3
4.4
0.05

(23.3)
(18.4)
(14.3)
(0.0%)
(na)
(na)
(na)
(0.9)

This study
This study
This study
Keenleyside (1998)
Costa (1980)
Costa (1980)
Costa (1980)
Keenleyside (1998)

0.0
4.6
9.0
14.1
8.4
6.5
10.8
10.8
10.4
6.6
8.8
8.2
0.0
12.7

(0.0)
(44.4)
(50.0)
(63.1)
(58.1)
(42.8)
(77.7)
(50.0)
(65.4)
(54.5)
(22.2)
(50.0)
(0.0)
(100)

Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita
Fujita

(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)
(1995)

% Affected teeth (% affected individuals).

% Carious teeth (% individuals with caries).
Jomon excluding the epi-Jomon sample.

Fig. 4. Probable extramasticatory induced dental wear pattern in an adult male (KO9) from Kotan-Onsen, middle Jomon.

Fig. 3. Arctic and subarctic adult oral pathology summary.

the sub-Arctic and Arctic samples, Hokkaido and Alaska.

Hokkaido Jomon, Okhotsk, and Eskimo levels of combined tooth LEH are similar and all are significantly
greater (v2 17.2527.38, P ! 0.001) than the Aleut. Central Jomon rates of LEH are commensurate with the levels of LEH seen in subtropical agricultural and hunter
gatherer samples, although higher than that seen in a
metal age sample from Ban Chiang, Thailand.
Cribra orbitalia. Figure 7 shows the frequency of cribra orbitalia using the same samples examined in Figure
6. While there is no statistically significant difference
between the Hokkaido Jomon and Okhotsk (v2 3.26,
P ! 0.1), cribra orbitalia in the Hokkaido samples is significantly greater than that seen in either of the north
American samples (minimum v2 23.7, P ! 0.001) and
also the Honshu Jomon sample (minimum v2 14.2,
P ! 0.001). As with the LEH results, the Southeast
Asian figures are presented to simply show the consider-

able variation in cribra orbitalia present in Asian bioarchaeological assemblages.

DISCUSSION
The Hokkaido and Alaskan assemblages represent
groups of people culturally adapted to operating within
challenging sub-Arctic and Arctic environments, although the social and technological manifestations of
this were quite different. While plant foods played a role
in the subsistence of all of these samples, animal protein, particularly of marine origin, played a major dietary role. Given that the past populations sampled from
Hokkaido and Alaska are believed to have lived in their
respective regions for long periods of time, both were
presumably physiologically and perhaps morphologically
adapted to cold environmental conditions (see a discussion
of this for North American populations in So [1980]). For
these reasons, we argue that meaningful comparisons can

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70

M.F. OXENHAM AND H. MATSUMURA

Fig. 5. A further example of probable extramasticatory

induced dental wear pattern in an adult male (KO16) from
Kotan-Onsen, middle Jomon.

Fig. 6. Comparison of rates of permanent tooth LEH (tooth

count). Note Neolithic northern Vietnam (n560 canines, 39 incisors; 50004000 BP) and Metal Age Vietnam (n575 canines, 71
incisors, 25001700 yr BP, Oxenham 2000); Ban Chiang, Thailand (n530 canines, 26 incisors, 4100 to 1800 yr BP; Pietrusewsky and Douglas 2002).

be made between Hokkaido and Alaskan oral health and

physiological well-being. Differences in health between
these populations, within and between the two regions,
may reasonably be interpreted in terms of the respective
nature, and perhaps effectiveness, of the adaptational
suites employed: including specific dietary emphasis, technological differences and culturally mediated behaviors
that may or may not have adaptive value.

Hokkaido and Honshu

The marked differences in oral health and physiological well-being between the Hokkaido Jomon (and also
the Okhotsk) and more southern Jomon samples reinforces the view espoused here that such a comparison has
limited value given the very different adaptive strategies
and environmental conditions experienced in each
region. Nonetheless, the very low level of dental caries
in Hokkaido relative to Honshu does demonstrate that
huntergather differences in oral health can be marked.
Perhaps the otherwise relatively subtle differences in
huntergatherer diets can be observed by way of oral
health profiles. The higher plant, and hence carbohy-

Fig. 7. Comparison of rates of adult cribra orbitalia (individual count). Note Neolithic northern Vietnam and Metal Age
Vietnam (n558 and 56 individuals respectively; Oxenham
2006); Ban Chiang, Thailand (n533; Pietrusewsky and Douglas
2002).

drate, content of Honshu diets may be a factor contributing to their greater level of dental caries (Fujita, 1995).
A factor suppressing the level of caries in Hokkaido is
perhaps the marine dietary orientation of these people.
Marine diets are arguably cariostatic in terms of
increased fluoride levels (Kelley et al., 1991; Larsen
et al., 1991), often associated with an increased degree of
tooth abrasion and the antibacterial effects of increased
oral cavity alkalinity (Powell, 1985; Sealy et al., 1992;
Littleton and Frohlich, 1993).
Signatures of physiological well-being among the
Jomon samples provide a mixed message. Hokkaido
(Jomon and Okhotsk) levels of canine LEH are depressed
relative to the one central Jomon sample available for
comparison. The complex etiology of LEH means specific
causes cannot be isolated. It might have been expected
that reduced dietary breadth and the biological costs of
living in a sub-Arctic region would have led to comparatively higher levels of LEH in the Hokkaido sample. Further, marine-oriented subsistence economies can lead to
the maintenance of high parasitic loads (Walker, 1986)
which may be reflected in elevated levels of physiological
stress markers, such as LEH or cribra orbitalia. The
level of LEH seen in Honshu Jomon is comparable to
that seen in huntergatherer and agricultural samples
from subtropical northern Vietnam, where high tropical
parasitic loads and haemoglobinopathies, such as thalassemia, are implicated (Oxenham et al., 2005). Yet, the
sample from prehistoric Ban Chiang, Thailand, demonstrates the variability in this stress marker even in tropical samples. In terms of LEH, at least, a sub-Arctic
environment and marine focused subsistence regime
does not seem to have been physiologically expensive.
The picture is quite different when physiological wellbeing is assessed by way of cribra orbitalia. The very
high rate of this condition seen in the Hokkaido Jomon
and Okhotsk samples compared to central Japan is at
odds with the findings from LEH. The etiology of cribra
orbitalia is also complex and is most often attributed to
anemia (Stuart-Macadam, 1985), itself a complex condition with multiple etiologies, or various inflammatory
diseases (Wapler et al., 2004) and also scurvy (Ortner
et al., 1999). Given the subsistence economy of the Hokkaido populations, it is unlikely that a lack of dietary
iron was an issue, although at least one study has found
depressed levels of folic acid in Canadian Eskimos
(Sayed et al., 1976). Folic acid deficiency can lead to

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PALEOHEALTH IN HOKKAIDO
megaloblastic anemia which is associated with suboptimal erythropoiesis and can presumably also lead to cribra orbitalia. While vitamin C is available through the
consumption of raw meat (So, 1980), Clow et al. (1975)
found ascorbic acid deficiencies in modern Canadian
Inuits and, while this could be due to changed modern
dietary conditions, the level of vitamin C in prehistoric
sub-Arctic and Arctic populations is unknown. As noted
above, scurvy, a condition associated with vitamin C
deficiency, can manifest skeletally as cribra orbitalia.
The high parasitic load of marine diets mentioned in reference to LEH may also be an important factor in the
elevated level of cribra orbitalia seen in the Hokkaido
samples.

Hokkaido and Alaska

The dental profile for these samples suggested three
groups identified by way of broadly similar patterns of
oral health: Aleuts and Eskimo; Ipiutak and Tigara; the
Hokkaido Jomon and Okhotsk with the Kodiak Island
sample. Examining the Aleuts and Eskimo first, the
enormous amount of AMTL and elevated levels of alveolar defects in both samples are consistent with their
older demographic age. It is intriguing that the oldest
samples (in terms of age-at-death composition) have the
lowest frequency of dental caries. The true caries level is
likely being masked by the elevated level of AMTL. This
study has not employed a caries/AMTL correction factor
and the published comparative data is not reported in a
way that would facilitate this. Moreover, such corrections make a number of untested assumptions that may
affect the reliability of corrected data (Hillson, 2001;
Oxenham et al., 2006b). The high levels of alveolar
defects are also consistent with the rates of AMTL and
are likely the product of advanced tooth wear and subsequent pulp chamber exposure. While the Eskimo sample
represents the most northerly Arctic community and the
Aleuts the most southerly Alaskan group, both are
reported to have had a broad based marine diet targeting fish, whale, and seal.
The Ipiutak and Tigara assemblages both derive from
Point Hope, in the arctic region and they are each demographically the youngest samples. Their chief similarity
is in terms of elevated AMTL and moderate levels of alveolar defects. They also both display the highest levels
of dental caries for Hokkaido and Alaska combined,
although Ipiutak caries are extremely high even compared to Tigara. That the samples with the youngest
age-at-death profiles have the highest frequency of caries
is unexpected. Presumably, many individuals were dying
before they lost teeth through caries induced infection
and/or wear. Madimenos (2005) has recorded greater levels of wear in the Tigara sample and suggested this as a
possible cause for the lower rates of caries compared to
the Ipiutak sample. A number of studies have suggested
high tooth wear environments inhibit caries formation
(e.g., Larsen, 1995; Powell, 1985). Notwithstanding this,
the rate of Ipiutak caries is so high (14.3% of Ipiutak
teeth compared to 4.4% Tigara teeth had carious lesions)
as to suggest either a cariogenic diet and/or a caries
inducing oral environment.
The chief difference in diet between these groups was
the preferential targeting of caribou by the Ipiutak and
whale by the Tigara. This marked dietary difference
may be implicated in the differences in dental caries.
The Ipiutak was the only community that preferentially

71

consumed terrestrial animals and thus presumably did

not reap the entire benefit of the cariostatic nature of a
marine dependent diet (see Discussion above). Moreover,
an important source of carbohydrates in modern caribou
hunters is the partially fermented stomach contents of
caribou. Presumably the potential cariogenicity varied
with the specific stomach contents seasonally and
Gubser (1965) notes variations in the color of Alaskan
caribou stomach contents with seasonal changes in diet.
Andersen (2005) reports there is 11.4 g carbohydrate per
100 g in the stomach contents of Greenland caribou,
although a summer through winter range in values is
not given. Samuel Hearne, an Arctic explorer in the late
18th century, describes several caribou (Hearne refers to
them as deer in his journal) stomach contents preparation methods used by indigenous people in the Hudson
Bay area. The stomach contents could be consumed immediately after a fresh kill, they could be mixed with
blood and water and boiled to make a sort of porridge, or
this mix with the addition of meat and fat could also be
smoked and hung within the stomach lining for several
days giving it an agreeable acid taste (Hearne [1795],
1958, p204). Such a sticky, carbohydrate rich food stuff
is worth considering as a possible etiological agent in
elevated Ipiutak caries rates.
Given the potential for vitamin and mineral deficiencies in traditional Arctic diets, it is worth considering
the possibility of magnesium deficiencies and rates of
oral disease. While the link between magnesium levels
and dental caries is complex and controversial (Luoma,
1988), animal model research suggests a clear link
between magnesium deficiency and increased rates of
dental caries (Kleber and Fehlinger, 1988). The concentration of magnesium (measured as mg of Mg per 100
mg) varies depending on the animal and body part consumed (the following values are based on Andersen,
2005, p3337). Concentrations for Canadian Arctic whales
vary from a low of 11.6 mg in raw blubber (Beluga) to
72 mg in dried muscle (Narwhal). Blubber, and mattak
(skin with blubber), has very low levels of magnesium in
all marine mammals. Concentration in seal muscle
varies from 24 to 30 mg in most seal species reviewed by
Andersen (2005), while intestines have similar concentrations. Salmon muscle, depending on species, ranges
from 21 to 74 mg, while smoked salmon is 143 mg. Shellfish have high values, clams at 122 mg, and blue
muscles 58 mg when eaten raw. Compared to the high
magnesium values seen in marine-based diets, caribou
ranges from 29 to 32 mg for raw and cooked muscle,
although in dried caribou increases to 72 mg. Other body
parts range in value from 1.8 mg (raw bone marrow) to
34 mg (boiled heart). Notwithstanding this, even if it is
assumed that caribou-hunting Ipiutak were consuming
less magnesium than the Tigarans, and other Arctic and
sub-Arctic huntergatherers for that matter, it is unclear
if they were necessarily deficient in magnesium. Other
reasons for greater cariogenicity being associated with a
terrestrial animal-based diet are not obvious. Food preparation, preservation, and consumption techniques and
behaviors may have differed between the two groups and
also contributed to the difference in caries rates.
The final broad clustering of samples includes both
series from Hokkaido and another island sample, Jones
Point on Kodiak Island. These assemblages share a similar, generalized, marine diet and have middle-aged demographic profiles. The main difference in their oral profiles is a very low rate of Kodiak Island alveolar defects,

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72

M.F. OXENHAM AND H. MATSUMURA

contrasting with a slightly higher rate of caries. The

only other two samples with higher rates of caries than
alveolar defects are from Point Hope. The oral health of
all three assemblages is quite good relative to the other
subarctic and arctic samples, and that of Kodiak Island
is arguably the best. It is perhaps fortuitous that these
samples derive from Island contexts. While dental task
wear facets or evidence for extramasticatory use of the
dentition was not purposefully examined, the Hokkaido
Jomon sample provided two clear examples of such
behaviors, but had a relatively low rate of AMTL. Recent
North American communities are known to have used
their dentitions as tools (e.g., Balikci, 1970) in addition
to tackling tough, preserved foodstuffs as, no doubt, did
their forebears: Madimenos (2005) found evidence for
extramasticatory use of the dentition in both the Tigara
and Ipiutak samples. The wear patterns seen in the
illustrated Jomon examples are consistent with the use
of the teeth in processing tough foods or skins and/or
using the mouth as a form of vice. Extramasticatory use
of the dentition may have played an important role in
the etiology of AMTL in all samples, although a direct
correlation between the rate of AMTL and level of extramasticatory behaviors cannot be assessed in this study.
The explanation for the relatively low levels of LEH
seen in the Hokkaido samples likely applies to the similarly low rate of LEH in the Eskimo series and very low
frequency in the Aleut sample. Keenleyside (1998)
suggested the effects of seasonal food shortages differentially affected the Aleuts in comparison to the Eskimo.
Given that Keenleyside (1998) reports that the Aleuts
had a higher rate of postcranial signatures of infectious
disease than the Eskimo, it seems counterintuitive that
they also experienced an extremely low frequency of
LEH and cribra orbitalia. Perhaps there was a child vs.
adult difference in physiological and infectious assault
among the Aleuts, although what could have caused this
is unclear. While the etiology of LEH in these regions
remains ambiguous, it would seem that climatically challenging environments (arctic/subarctic Alaska and subarctic Japan) are characterized by populations with reasonably good levels of physiological well-being. This is
not the case, for Hokkaido at least, when well-being is
measured using cribra orbitalia. While the rate of cribra
orbitalia in the two Alaskan series remains very low, it
is elevated in the Hokkaido samples in comparison to
central Jomon huntergatherers and is comparable,
especially the Okhotsk figures, to some prehistoric subtropical Vietnamese samples. The underlying etiology of
cribra orbitalia is perhaps quite different, but currently
unknowable, in prehistoric Alaska and Hokkaido.

CONCLUSIONS
The main objectives of this paper included an examination of oral and physiological well-being in adult skeletal assemblages from Jomon and Okhotsk Hokkaido
Japan. To contextualize these results the discussion
focused on comparing the Hokkaido material to hunter
gather data from central Japan and assemblages from
environmentally similar contexts in Alaska. Hokkaido
oral health and physiological well-being was generally
better, with the exception of cribra orbitalia, than that of
their southern Jomon counterparts. Perhaps more important is the finding that Hokkaido oral health is
more meaningfully comparable, not to other southern
Japanese samples, but to assemblages from similar envi-

ronmental zones that are biologically and culturally

adapted to sub-Arctic and Arctic conditions.
Despite operating within similar environmental zones,
the adaptive strategies of these various communities
varied and likely contributed to the complex picture of
sub-Arctic and Arctic human health seen here. Behavior
was not the only variable influencing the oral and physiological health profiles of these samples. With the exception of the Ipiutak and Tigara, the other assemblages
had broadly similar subsistence bases and formed two
groups defined by similar oral profiles. The differences
and similarities between the Eskimo and Aleuts on the
one hand, and Hokkaido samples and Kodiak Island on
the other hand, were influenced by differences in age-atdeath composition as well as heterogeneity in life-ways
and specific adaptive strategies. The Ipiutak and Tigara
were somewhat similar in oral demographic profile, but
were unique in targeting caribou and whale respectively.
Clearly there is a great deal of variability in cold climate
huntergatherer oral and physiological health profiles.
This variability suggests health, oral particularly, is sensitive to the subtle differences in the diet and life-ways
of such cold adapted communities, in a similar sense to
the way agricultural and huntergatherer communities
can often be differentiated by way of their patterns of
oral health. Of course, these profiles lack predictive
value and cannot be used to determine subsistence and
adaptive strategies in and of themselves, just as few
would attempt to identify a dental assemblage as representing an agricultural community based on the frequency of oral health traits alone. Finally, the comparative context for Jomon and Okhotsk paleohealth is not
prehistoric and protohistoric central and southern
Japan, but rather other cold adapted peoples.

ACKNOWLEDGMENTS
The authors thank the following people for generously
donating their time and commenting on previous versions of this paper: David Bulbeck, Colin Groves, Mark
Hudson, and Lorna Tilley. Many thanks to Brenton Hill
for drawing the accompanying maps.

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