DENTAL ANTHROPOLOGY - Scott & Turner 1988
DENTAL ANTHROPOLOGY - Scott & Turner 1988
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DENTAL ANTHROPOLOGY
G. Richard Scott
Department of Anthropology, University of Alaska, Fairbanks, Alaska 99775
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Christy G. Turner II
Department of Anthropology, Arizona State University, Tempe, Arizona 85287
INTRODUCTION
The term "dental anthropology" first appears in the title of an article published
in 1900 by George Buschan, although Klatsky & Fisher (97) are credited with
its formal introduction (44). The field is rooted in French, German, and
English encyclopedic mammalian odontological treatises of the past two
centuries. That teeth possess qualities valuable for anthropological study (i.e. ,
they are durable; evolutionarily conservative and yet adaptable; rich with
genetically determined traits; and reflective of behavior, ecology, and diet)
was recognized, if not fully explicated, by such 19th century natural histo
rians as L. Rousseau, G. Koch, 1. Henle, and R. Owen.
The breadth and depth of dental anthropology are evidenced by the contri
butions to past International Symposia on Dental Morphology (28, 45, 102,
1 48, 164a). These major congresses were preceded by a landmark symposium
on dental anthropology sponsored by the Society for the Study of Human
Biology (24). Other recent compendia are devoted to various facets of dental
anthropology and allied fields (46, 93, 157), while a recent monograph
entitled Teeth (84) provides both archaeologists and dental anthropologists a
useful overview of the dentition.
Research categories in dental anthropology include morphology, metrics,
health, evolution, growth, genetics, usage, forensics, and ethnographic treat
ment. Although primate and fossil hominid dental studies fall within the
purview of dental anthropology, this review is limited to research on post
Pleistocene human populations and major themes in the discipline.
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100 SCOTT & TURNER
studies on crown morphology (41, 42,83, 88, 89, 104) indicated the potential
of such traits to discriminate among the major geographic races. In particular,
the affinities between native Americans and Asians and the distinctiveness of
both groups from Europeans in crown morphology were consistent findings.
Works that included basic odontometric descriptions for a wide variety of
groups (30, 50, 59, 134, 139, 146, 182, 183) also demonstrated broad
variation in human tooth size. This variation is more difficult to interpret
historically than differences in crown morphology, suggesting that size is
more responsive to selection and/or environmental factors than is morpholo
gy.
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the development of tooth crown size (23, 191). For morphologic crown traits,
workers have acknowledged polygenic modes of inheritance (65, 80, 81,
1 77), although the action of major genes may be detected within the broader
context of a multifactorial system (98, 140). Environmental factors, in par
ticular maternal effects and nutrition, do play a role in the development of
tooth size and morphology (57, 1 00, 143).
Tooth size, number, and morphology have a sufficiently strong genetic
basis to make them useful variables for assessing biological relationships and
microevolutionary trends. When groups are compared for mean dimensions or
morphologic trait frequencies, measurements of between-group similarity
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yield phenetic rather than genetic distances; this is, of course, true for many
other variables studied by physical anthropologists.
Mongoloid (76) and Caucasoid Dental Complexes ( 1 21). While such general
characterizations provide useful guides in forensic assessment of ethnic
affiliation, they have no explanatory value by themselves. For assessing
population affinities using tooth morphology, distance statistics are used to
estimate relative degrees of similarity among groups that diverged from
commOn ancestors many hundreds or thousands of years in the past.
It is desirable to check population relationships determined dentally with
other types of biological, historic, linguistic, and cultural information. Brew
er-Carias et al (22) observed several crown traits in seven Yanomama villages
that had also been surveyed for genetic markers. A high level of dental
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single root trait (3-rooted lower first molars) to distinguish three primary
subgroupings within the Americas: Eskimo-Aleut, Na-Dene, and all other
Indians of North and South America (199). Subsequent research on more than
20 crown and root traits observed in native Americans from both North and
South America confmned this pattern (202, 204, 205). The inference thus
derived from dental morphologic variation is that the three primary subgroups
evident in the Americas correspond to three major and separate movements of
peoples into the New World. Evidence from linguistics, archaeology, and
serological genetics has been used to support this three-migration model (66,
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67).
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alized crown and root trait frequencies. The much simplified Asiatic Indian
and European dentitions may ultimately have arisen from an Upper Paleolithic
Sundadont population in southeast Asia; further work on early skeletal series
in those regions will test this hypothesis (207).
Dental morphology suggests that prehistoric Jomon peoples are ancestral to
the modem Ainu (200). Moreover,Ainu dental characteristics resemble those
of Micronesians and Polynesians more than those of Chinese and Japanese
groups (210). Despite the traditional depiction of the Ainu as "archaic Cauca
soids," Ainu teeth do not fit the European dental pattern but, along with the
Jomon, exhibit Sundadonty. Modem Japanese populations show the Sinodont
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pattern and are thus thought to have descended from mainland Asian groups
that invaded Japan beginning about 2200 years ago. As noted above, this
reconstruction is also indicated by patterns of tooth size variation (19).
In contrast to the morphologically complex dentitions of Asians and their
derivative populations, Europeans exhibit simple crown morphology. Eu
ropean groups are characterized by moderate frequencies of incisor shoveling
with few pronounced shovel forms (125); a high frequency of Carabelli's cusp
(178,215); simplified lower molars often lacking the hypoconulid (especially
on the second molar); and low frequencies of cusp 6, cusp 7,the protostylid,
and perhaps the deflecting wrinkle (121, 177). Although there have been no
detailed studies of European dental microdifferentiation, Kirveskari (96) has
shown that Skolt Lapp dental morphology indicates a primary European rather
than Asian origin for this enigmatic group. In this instance, dental morpholo
gy corroborates the findings of serological genetics. Asiatic Indians also
comprise a major grouping with presumed genetic ties to Europe. Although
living and skeletal samples from India and Pakistan have only recently been
profiled for dental morphologic variation, the samples studied indicate genetic
affinity to European rather than east and north Asian groups (113, 115).
MICROEVOLUTIONARY PERSPECTIVES
Selection
Most mammals need an intact and functional dentition to survive. When the
teeth fail to function through extreme wear or loss-for example, in in
dividual baboons (2l)-death soon follows. This was true for our hominid
ancestors of the Pliocene and Pleistocene; in most cases, fossil hominid jaws
have intact dentitions. At some point in human history, cultural mechanisms
started to buffer the equation of tooth loss and death. During the late Pleis
tocene and Holocene, perhaps as many as 10% of adults lived many years
after the loss of most or all of their teeth, Dietary changes and new food
preparation techniques allowed them to survive . Thus post-Pleistocene dental
changes must be viewed in a biocultural context.
One trend in the dentition of post-Pleistocene human populations is tooth
106 SCOTT & TURNER
gathering groups, a viable dentition was essential for survival , but during the
early Holocene, with the emergence of food production and the development
of new food preparation techniques, the advantage to large-toothed in
dividuals diminished .
Brace (13, 14) has long contended that structural reduction in the dentition
(and other systems) is a natural consequence of reduced selection pressure and
his "probable mutation effect." As long as a certain tooth mass is essential to
normal function and survival, selective pressures maintain (or increase) tooth
dimension. When selection pressure is reduced, in this case through cultural
buffering (e. g . the use of earth-ovens, pottery, eating utensils, etc) , recessive
mutations hitherto selected against accumulate in the gene pool . These muta
tions, acting in concert, affect development negatively and bring about the
reduction of a structure-in this case, elements of the dentition.
A common counterargument to the probable mutation effect is that selec
tion pressure is not actually reduced when a structure (tooth size) becomes
less essential to survival . Rather, selection then favors a decrease in tooth size
because the development of large teeth would use energy that might be better
channeled to other organ and tissue systems. This hypothetical process has
been referred to as "the somatic budget effect" (94), or "selection for total
organism efficiency" (68).
The definitive explanation for Holocene tooth-size reduction awaits further
work. It seems unlikely, for example, that natural selection underlies signifi
cant secular tooth-size increases from one generation to the next (55, 64, 106)
or an 8.5% increase in tooth size in Swedish males between 1810 and 1970
(51). Maternal effects (57), as well as nutritional and disease stresses in early
childhood, can impact crown size development. Environmental factors should
be examined closely before tooth-size trends are attributed to genetic changes
brought about by selection.
It is difficult to demonstrate that natural selection has brought about
changes in any organ or tissue system. However, taking a cue from Kurten's
(101) classic study of selection on dental characteristics of European cave
bears, Perzigian (149) attempted to demonstrate differential survival in an
DENTAL ANTHROPOLOGY 107
stressed at an early age would have a greater mortality risk during childhood
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and adolescence than those who had undergone normal development. Con
sidering the developmental flexibility of tooth size (57), it is plausible that
stress rather than genetic selection is the primary contributor to reported age
differences in tooth size.
The effect of natural selection on crown morphological variability is even
more hypothetical than its effect on tooth size. One proposal is that surficial
traits of the tooth crown enhance the mass and masticatory efficiency of teeth
subjected to implemental use and high levels of attrition. Shovel-shaped
incisors have been discussed more than other traits in this context,as shovel
ing commonly occurs in Neandertals and Eskimos-groups that subjected
their anterior teeth to pronounced stresses. In an attempt to delineate environ
mental and cultural factors associated with the distribution of shovel-shaped
incisors, Mizoguchi (125) computed correlations between frequency and
degree of shoveling in dozens of human groups with physical, meteorologi
cal, and cultural factors. His findings-which included significant negative
correlations between shoveling and mean annual temperature, history of
milking, and stature; and significant positive correlations with latitude and
annual rainfall-are suggestive but not conclusive. The worldwide distribu
tion of shoveling is relatively straightforward, ranging from groups in north
east Asia and the Americas, who show the highest frequencies and most
pronounced expressions, to Europeans and Asiatic Indians, who have mod
erate frequencies and slight expressions. Between these extremes occur most
other human groups,including Africans,southeast Asians,and populations of
the Pacific (176, 203, 204). If the latter intermediate groups exhibit the
ancestral condition of Homo sapiens sapiens, it is necessary to explain why
shoveling increased in northeast Asia and decreased in Europe. Beyond the
argument that selection favored individuals with pronounced shoveling in a
northern temperate-to-Arctic regime where meat eating and the implemental
use of teeth was stressed, there is only indirect evidence to suggest that
selection played a role in the current pattern of shoveling variation.
It has been suggested that another common dental morphologic trait,
Carabelli's cusp, increased in frequency and expression in Europeans because
108 SCOTI & TURNER
it added mass to the upper molar crown and thereby. impeded rapid rates' of
attrition (43a). While plausible, this remains an unproved hypothe$is� Such
inconclusiveness applies to all surficial morphologic crown or accessory root
traits that might increase the masticatory efficiency, tooth longevity, and
fitness of individuals who possess the traits.
Indeed, why do some groups, notably Europeans and Asiatic Indians, show
crown simplification and retention of earlier root numbers? The most popular
explanation, beyond reduced selection pressures on agricultural groups, is
that morphologically simpler crowns are less susceptible to caries because
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there are fewer pits and fissures to collect food debris (144). Greene (68, 69)
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Gene Flow
Estimating the degree and rate of gene flow using crown size or morphologic
variables is possible both diachronically and synchronically. Turner (196-
198), for example, compared skeletal and living gr�)Ups of Kodiak Islanders,
Aleuts, and Hopi Indians and found that European admixture was evident in
the crown morphology of the living populations. Moreover, rates of gene
flow, calculated using dental trait frequencies, were relatively close to es
timates based on gene frequencies, and consistent with known history. The
disadvantage of using morphologic traits, compared to systems with simple
modes of inheritance, is that trait frequencies are not reducible to gene
frequencies. The advantage provided by tooth crown variables is that one can
directly observe the ancestral populations of a particular living group. This is
impossible with serological genetic traits.
Genetic Drift
Genetic drift, including founder's effect, has likely played a major role in
human dental differentiation in post-Pleistocene times. Dental morphological
differentiation among groups that share recent common ancestry has been
well documented in Melanesia (80), Polynesia (208), the American South
west (179, 180), Venezuela (22), and the middle East (190). In all cases,
dental divergence parallels genetic differentiation which, because of the short
time spans involved, is viewed as a consequence of colonization events,
population structure, and the accumulation of random genetic changes reflect
ing small population size-i.e. drift and founder's effect. The fact that we can
estimate consistent times of divergence between groups separated by many
thousands of years (204, 206) also suggests that drift is the key ,mechanism
involved in generating patterns of crown and root trait variation.
DENTAL ANTHROPOLOGY 109
Mutation
The major morphological features observable in recent human populations
have long evolutionary histories, extending back to earlier hominids or even
Miocene apes. One trait may, however, illustrate a recent mutational event in
human history. This feature, termed the Uto-Aztecan premolar by Morris and
his colleagues (135), is a rare and distinctive attribute of the upper first
premolar that occurs only among Indians of the New World (204). It may
reflect a specific mutation that occurred after the peopling of the Americas.
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Eskimos show pronounced wear due to frozen or tough dietary items that
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(127, 165),changing dietary patterns through time (216), and the influence of
acculturation on native groups recently adopting the highly refined diet of
modem industrial societies (47). The modem diet has radically reduced crown
wear. In today' s industrialized societies, it is so unusual to see levels of wear
comparable to those exhibited by earlier populations that dentists once
thought excessive attrition was pathologicaL
Sex differences in crown wear vary between populations. As males are
larger than females, they require more food intake and also generate stronger
vertical occlusal forces in mastication, so one would expect males to show
more attrition, all other things being equaL In some studies males exhibit
more wear than females, but in others there is no significant difference (95,
112,118,127) or females show more wear (129,159). Even among Eskimos,
where females are commonly thought to show more wear than males because
of their sex-specific activities (e. g. chewing skins for leather, softening frozen
boots; 108),some studies have found no sex difference or even greater wear
in males (47,58,193,209). Clearly, every population must be assessed in its
own biocultural context to understand why there is or is not a male-female
difference in attrition .
Related to crown wear are instances where tooth substance is lost through
traumatic fractures of the crown. Enamel subjected to enough stress will
fracture or chip. The amount lost may range from a tiny fragment to an entire
cusp or even a whole tooth. Dental chipping is best known for Eskimos (108,
146), who traditionally subjected their teeth to a variety of heavy stresses,
both in terms of dietary constituents and implemental uses. A comparison
between Arctic and sub-Arctic groups showed that Eskimos had significantly
higher frequencies of dental chipping than either Aleuts or northern Indians
(209). Milner (122) noted, however, that chipping occurred frequently ill
Mississippian period Indians from Illinois.
the occlusal and interproximal surfaces of their anterior teeth were likely
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Many groups throughout the world intentionally modify their tooth crowns
in a wide variety of ways (most of which are painful to contemplate; 161,
185). Such alterations, commonly referred to as dental mutilations, may
involve complete removal of one or more teeth, removal of parts of specific
teeth through filing or chipping, or modifications of crown surfaces through
incising or drilling, sometimes followed by inlays of precious metals or
gemstones. Such mutilations involve the more visible anterior teeth, es
pecially the upper incisors. The stimulus for modification may relate to rites
of passage, status differentiation, simple cosmetics, or other cultural motiva
tions. In some cases, tooth-tool use results in patterns of wear that simulate
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intentional dental mutilation. Blakely & Beck (8) outline criteria to distin
guish incidental from intentional dental modifications.
Disease processes of the jaws and teeth that vary spatially and temporally and
that have been interpreted relative to dietary and cultural differences and
trends include caries, periodontal disease, periapical osteitis (abscessing),
ante-mortem tooth loss, and malocclusion, including tooth crowding.
From an anthropological standpoint, the most important dental disease is
caries, because of its association with the evolution of food production. A
carious lesion is characterized by demineralization of enamel, dentine, and/or
cementum by acidogenic microorganisms (e.g. Lactobacillus acidophilus,
Streptococcus mutans) that permanently inhabit the plaque on tooth crown
surfaces. Such lesions can develop on either the occlusal (crown) or cervical
(root) surfaces. As most humans have similar oral microflora, the extent of
dental caries is not determined primarily by the presence or absence of
bacteria, but by host resistance and diet (1).
Factors of host resistance include morphological crown complexity (144),
tooth size (3, 213), trace elements in food or water (1, 84), developmental
defects (35, 91), dental wear (155), and immunologic and nonimmunologic
characteristics of saliva (119). Dietary factors that affect caries etiology
include foods eaten, methods of food preparation, eating habits, and degree
and duration of mastication. Refined carbohydrates are the most cariogenic
dietary elements, especially sucrose, glucose, and fructose (99, 184).
Periodontal disease affects the gingiva (gums), periodontal membrane, and
bone (alveolus). Plaque and its associatcd microbiota can extend below the
gumline and lead to inflammation of the soft tissues and periodontal mem
brane surrounding tooth roots. It is commonly held that long-standing in
flammation stimulates the formation of periodontal pockets and alveolar bone
loss, although this viewpoint is being reassessed (32). Alveolar resorption
progresses either horizontally (occurs along the entire tooth row) or irregular·
114 SCOTT & TURNER
the alveolus; bone may also be destroyed on the lingual wall (2, 84). Per
iapical osteitic lesions are typically the result of pulp exposure, brought about
by rapid attrition, caries, trauma, or periodontal disease.
Tooth loss is another generalized indicator of oral disease readily studied in
human skeletal material. In modem populations, most missing teeth are
attributed to rampant caries. DMF (decayed-missing-filled) counts are used to
characterize groups for caries intensity. Even today, some tooth loss is due to
causes other than caries, especially periodontal disease; but in earlier pop
ulations, the etiology of tooth loss was more complicated. In groups with low
caries rates, teeth may be lost owing to excessive wear or trauma followed by
pulpal necrosis, periapical osteitis, bone resorption, and evulsion. Alveolar
resorption resulting from periodontal disease may also be a factor. The most
susceptible teeth are the upper and lower molars while the teeth least likely to
be lost are the canines; incisors and premolars fall between these extremes.
This tooth survivorship pattern appears to apply to hunter-gatherers, early
agriculturalists, and modem populations.
When the teeth are arranged in the jaws in a parabolic arcade and proper
intercuspal contact is maintained between upper and lower teeth, occlusion is
said to be normal. Two factors can produce abnormality: (a) insufficient space
in the jaws for the full complement of teeth, or (b) over- or underdevelopment
of one jaw relative to the other jaw. In the first instance, lack of harmony
between tooth size and jaw size results in crowding. Differential development
of the jaws leads to pronounced overbite, underbite, or other forms of
malocclusion. Problems of occlusion are developmental structural defects or a
secondary consequence of tooth loss.
a similar incidence of periapical osteitis for Indian Knoll (48%) and Zuni
(52%) but in the former, severe attrition caused 99% of the lesions while in
the agricultural Zuni, 70% of the abscesses were caused by caries. Hunter
gatherers, who exhibit moderate rates and degrees of wear and a low in
cidence of caries, show a correspondingly low incidence of periapical osteitis;
16% of Sioux skulls ( l 09) and 19% of Eskimos skulls ( l 08) exhibited
abscessed teeth. Among the Eskimo, 1 0% of all abscesses were attributed to
trauma.
Data on ante-mortem tooth loss in skeletal series are reported by percentage
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of total teeth lost per number of observable sockets (25), percentage of tooth
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loss by tooth type for a sample ( 1 08), or by sex and age category (38).
Because the factors associated with tooth loss are strongly correlated with
age, between-group comparisons can be skewed when samples have different
age profiles. In general , however, tooth-loss trends in recent human evolution
correspond to changes in other dental maladies. In hunting-gathering groups
with low caries rates , moderate wear, and limited periodontal disease, tooth
loss is relatively infrequent. In groups practicing an agricultural or mixed
economy, an increase in caries and periapical osteitis (due to caries and rapid
wear rates) elevates the percentage of tooth loss. The highest rates of tooth
loss are found in modem populations that utilize highly processed foods and
have limited access to dental care . The extremely high rates of caries in
modem groups would result in correspondingly high rates of tooth loss were it
not for the intervention of dental health practitioners.
Tooth crowding does occur in earlier human populations, including some
fossil hominids , but pronounced forms of malocclusion are a relatively recent
development (90, I l l ). The reduced need for vigorous mastication following
the adoption of agricultural foodstuffs and advances in cooking techniques
apparently leads to an underdevelopment of jaw size without a fully corre
sponding reduction in tooth size. In some individuals, differential growth of
the jaws leads to over- or underbite and impaired occlusion; and when teeth
are lost, the mesial drift of the remaining teeth can alter occlusal relationships .
Along with caries and periodontal disease, malocclusion i s a cost associated
with the benefits of civilization.
The teeth in the two sides of the jaw mirror each other structurally. Strong
developmental canalization also results in regular and predictable tooth de
velopment and eruption sequences. Significant deviations from such normal
development can help researchers to assess nutritional, morbid, and other
types of stress in both extinct and extant human populations .
DENTAL ANTHROPOLOGY 117
Fluctuating Asymmetry
Van Valen's (21 4) description of various asymmetries apparent in biological
systems encouraged anthropologists to look more closely at the human denti
tion. The teeth in the two sides of either jaw-e.g. left 12, right I2-are called
antimeres. Antimeres are almost identical in crown dimensions and morphol
ogy, but observable differences sometimes exist. As there is no tendency for
either to be larger or to express morphologic traits differentially, teeth exhibit
fluctuating asymmetry (214).
Twin studies have shown that fluctuating asymmetry in the dentition has no
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Developmental Defects
Developmental defects of the dentition are being used to measure stress and
the nutritional and health status of living and prehistoric groups. This area of
research has been reviewed extensively and found promising (26, 33, 163).
Tooth crowns and roots are formed systematically under genetic control.
Amelogenesis and odontogenesis, terms for the formation and deposition of
118 SCOTT & TURNER
Over the past two decades, the American Journal of Physical Anthropology
and other journals have published numerous articles on the dentition of living
primates, fossil hominids, and skeletal and living human populations. An
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