Ann. Rev. Anthropol. 1988.

17:99-126
Copyright 1988 by Annual Reviews Inc. All rights reserved
DENTAL ANTHROPOLOGY
G. Richard Scot
Department of Anthropology, University of Alaska, Fairbanks, Alaska 99775
Christ G. Turner I
Department of Anthropology, Arizona State University, Tempe, Arizona 85287
INTRODUCTION
The term "dental anthropology" frst 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 refective 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 Interational 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 pst
Pleistocene human populations and major themes in the discipline.
WRITING HISTORY WITH TEETH
The groundwork for the use of dental characteristics in studies of popula
tion relationships dates to the frst half of this century. Classic comparative
99
0084-6570/88/1015-0099$02.00
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Quick links to online content
Further
ANNUAL
REVIEWS
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 odontometic 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.
Tooth size generally refers to two measures: maximum crown length
(mesial-distal diameter), and breadth (buccal-lingual diameter) (63, 134).
From these two variables, workers can derive different crown indexes or
estimate total crown area. Further precision can be gained through measuring
intercuspal distances or calculating total crown area through the use of
photographs and planimeters.
In contrast to the continuous variation of tooth size, crown and root
morphology is typically discrete (present or absent) in expression. As
morphologic trait presence can range from slight to pronounced, this type of
variation is quasi-continuous (72) rather than discontinuous. Crown traits take
the form of accessory ridges, tubercles, styles, and/or cusps expressed on the
lingual, buccal , or occlusal surfaces. Root traits are manifest primarily as
deviations in root number from the norm (e.g. lower molars usually have two
roots but can exhibit, as variants, either one or three roots).
A biological trait can be useful in historical-evolutionary analyses only if a
significant component of its variation is genetic. Phenotypic differences
between groups through time and space can then be assumed to reflect
temporal gene frequency changes or underlying genetic differences, respec
tively. Aspects of the human dentition most likely to have a strong genetic
component-tooth size, morphology, and number-have been analyzed in
populations, families, and twins in an attempt to sort out the respective effects
of genes and environment on dental development (for reviews, see 6, 54, 116,
138).
Dental genetic studies over the past three decades have significantly altered
earlier views of crown size, agenesis, and morphology as largely invariant
reflections of the genotype. No dental trait, excluding certain rare abnormali
ties, has been shown to have a simple mode of inheritance. There are general
size factors related to crown dimensions (81a, 110, 123, 124, 153, 154, 195) ,
but it is still unknown whether these are attributable to major genes or to
numerous minor genes acting together. Hypodontia (baving fewer teeth than
normal) and byperodontia (having superumerary teeth) are, in general ,
controlled by the same system of multifactorial inheritance that is involved in
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DENTAL ANTHROPOLOGY 101
the development of tooth crown size (23, 191). For morhologic 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 materal 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 suficiently 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
yield phenetic rather than genetic distances; this is, of course, true for many
other variables studied by physical anthropologists.
Tooth Size: Population Histor and Relationships
The teeth of native Australians (the largest-toothed living populations) are
3035% larger in total crown area than those of small-toothed groups such as
the Bushmen, Lapps, Iranians, etc. To what extent is such odontometric
variation historically meaningful?
The general consensus is that absolute or net tooth size is not a good
indicator of population relationships (77, 78, 82, 96, 134, 151 ). By contrast,
the Penrose "shape" component, which provides some measure of among
tooth proportionality, is thought to yield patters of relationship more in
accord with known population histories. In a survey of world-wide odontome
tric variation, Perzigian (151 ) computed size and shape distances among 42
human samples (fossil, recent skeletal, and living). In his dendrogram based
on size distances, samples were discriminated at the highest level between
large-toothed groups (Homo erectus, Neandertals, modem Australian natives)
and medium to small-toothed groups. In some instances, very distantly related
groups such as American whites and prehistoric Peruvian Indians clustered
closely. The shape distances exhibited a much higher level of taxonomic
consistency. His 1 7 native American samples were nonsensically scattered
throughout the size-based dendrogram, but in the dendrogram based on shape
these Indians and four Asian samples clustered as expected. Thus factors such
as technocultural complexity and a diet based on agricultural products may
lead to tooth size reduction in different geographic areas and dimensional
convergence between distantly related populations. Relative between-tooth
proportions, as measured by the Penrose shape component, are not as greatly
afected by such factors, so this component can discriminate among pop
ulations.
While comparative odontometric studies discriminate among groups at high
levels of population diferentiation, the results are not as straightforward in
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102 SCOTT & TURNER
comparisons among closely related groups. Schneider and her colleagues
(167) analyzed tooth size variation in ten central US American Indian pop
ulations dating from 1000 Be to AD 1550. Of 45 pairwise distance values
(Mahalanobis D2), they found only six to be statistically significant. There
was some tendency for areal but not temporal clustering; the general finding
was one of among-group homogeneity. In contrast to this, O'Rourke &
Crawford (141) found a relatively high level of odontometric microdifer
entiation in four modem Tlaxcaltecan communities in Mexico. Most general
ized distance values were significant among the samples, with females show
ing more between-group differentiation than males. Premolar dimensions
were the most useful in discriminating among the Tlaxcaltecan samples, a
finding also reported in odontometric studies of Melanesians (53) and New
Guinea Highlanders (10).
Although most workers prefer distance statistics for intergroup com
parisons of tooth size, Brace and his co-workers (15-20) compare the
summed cross-sectional crown area of all permanent teeth. While recognizing
the limitations of the method, Brace (15) feels that a single value is more
meaningful than tooth by tooth comparisons. He also maintains that natural
selection would more likely operate on overall crown dimensions than on
individual tooth measurements. Although a detailed review of Brace's work is
not possible here (see comments in 16, 17), he and his colleagues have
developed the general historical hypotheses that: (a) tooth size reduction is
greatest in groups with the longest history of agricultural practice and associ
ated changes in cooking techniques (15); (b) tooth size variability among
native Australians is greater than generally recognized, including a north-to
south cline of increasing tooth size that suggests a "trickle" of biological
infuences from the north (16); (c) the Pacific was first peopled by large
toothed Pleistocene hunting and gathering groups, whose descendants now
include Australians and the peoples of large-island Oceania, and much later
by small-toothed, seafaring, agricultural migrants from Asia who ultimately
peopled Micronesia and Polynesia and mixed with Austral-Melanesian pop
ulations in boundary areas (17); and (d the small-toothed Ainu of Japan are
the likely descendants of aboriginal Jomon peoples while the larger-toothed
modem Japanese population is afiliated primarily with Asian mainland
groups who began migrating to Japan around 300 Be during the Yayoi
invasion (19).
Tooth Morphology: Population Histor and Relationships
Surficial morphologic traits of the tooth crown exhibit significant differences
in frequency among the major geographic races (41, 42, 83, 88, 217). For
some taits, including shovel-shaped incisors, Carabelli's trait, and cusp 6,
the between-group diferences are so pronounced that workers have defned
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DENT AL ANTHROPOLOGY 1 03
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
microdifferentiation correlated significantly with both genetic and geographic
distance. Harris (80) assessed crown morphologic variation among 14 villages
on Bougainville in the Solomon Islands, correlating phenetic distances based
on crown morphology with those based on other biologic variables. He found
significant correlations with odontometric and anthropometric distances but
not with serologic distance. Additional studies have contrasted patters of
relationships based on crown morphology with those derived from gene
frequencies and have found generally congruent results (180, 1 81, 189, 190).
Although dental morphologic distances do not mirror precisely those based on
genetic traits, agreement is suficiently close to suggest both are measuring
the same process-genetic differentiation.
Similarly, variations in dental morphology can be compared with those in
language. While language families are not closed biological entities, many
authors have noted parallels in linguistic and biological differentiation. Harris
(80) performed a cluster analysis among six Austronesian and four non
Austronesian language stocks in Melanesia using a battery of 44 crown traits
and found a close correspondence between relationships indicated by the
dentition and those indicated by linguistic affiliation. Among natives of the
American Southwest, workers have also reported a high correspondence
between classifications based on dental morphology and those based on
language families (179- 181).
While there is some controversy regarding the utility of dental morphology
in studies of human microdifferentiation (4, 142), most workers concur that
these dental traits are useful in discriminating among tribal groups and local
races. The majority of recent dental morphologic studies have focused on
groups at these levels of diferentiation (70, 74, 113, 115, 158, 160, 190,
196, 202, 204, 205, 207, 218). Much of this work is devoted to delineating
the dental variation of Asian and Asian-derived groups. Less attention has
been paid to historical questions in Europe, India, Africa, Australia, and
Oceania.
The distinctiveness of American Indian dental morphology was clearly
shown in the classic studies of Dahlberg (42, 43). In 1971, Turer used a
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104 SCOTT & TURNER
single root trait (3-rooted lower frst 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 rot taits observed in native Americans from both North and
South America confmned this patter (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,
67).
Turer (204, 206) also developed a new method to estimate times of
divergence between local races based on dental morphologic distances. This
method, dentochronology, assumes that dental differentiation occurs at a
relatively regular rate following the separation and isolation of populations.
Using the mean measure of divergence (MMD) to calculate between-group
distances and an estimated dental microevolution rate of 0. 01003 0. 004
MMD per 1000 years, times of divergence were calculated among a wide
variety of groups. Although the method is experimental, results suggest most
North and South American Indians diverged from north Asians between
13,000 and 16,000 years agoates supporting the hypothesis that the New
World was peopled during the terminal Pleistocene. The divergence of Es
kimos and Aleuts was estimated at 3888 years, closely approximating linguis
tic evidence, with the ancestral Eskimo-Aleut population diverging from
north Asia around II ,000 years ago. Overall, dentochronological estimates
agreed with independent lines of evidence of human origins and migrations,
including linguistics, sea-level fuctuations, and 14C dates from the archaeo
logical record.
Turer also (207) described a large number of mainland Asian and Pacifc
populations. He delineated not a single Asian or Mongoloid Dental Complex
but two basic patters within Asia: Sinodonty and Sundadonty (203). The
Sinodont patter is characterized by morphologic trait intensification (high
frequencies and pronounced expressions of the traits thought to characterize
Mongoloids) and is evident not only in modem Chinese, Japanese, and
Siberian groups but also in all New World populations. Turer infers that all
native Americans were derived from parts of this Sinodont complex in north
Asia. The Sundadont patter, which involves trait retention and simplifica
tion, is observed in southeast Asians, Polynesians, Micronesians, and the
Jomon in Japan. Australians and Melanesians are not viewed as part of the
Sundadont complex, although they may have been derived from early Sunda
dont groups in southeast Asia. Turer (207) proposes that the distinctive
features of the Austral-Melanesian dentition evolved locally in Australia and
Melanesia following the early peopling of these areas. Sinodonty is viewed as
a specialized derivative of Sundadonty because the latter exhibits more gener-
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DENTAL ANTHROPOLOGY 105
alized crown and root trait frequencies. The much simplifed 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 patter but, along with the
Jomon, exhibit Sundadonty. Modem Japanese populations show the Sinodont
patter 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 patters 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 microdiferentiation, 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
afinity to European rather than east and north Asian groups (113, 115).
MICRO EVOLUTIONARY 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
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106 SCOTT & TURNER
size reduction. Although a few studies have reported no change or even an
increase in tooth size through time in a particular region (12, 56, 171, 173),
trends in tooth size reduction have been noted in Europe (20, 107), China
(15), India (114), north Africa (29), and North America (87). The role of
natural selection in trends toward tooth size reduction has been viewed in
several ways: (a) With a reduction in jaw size, there has been positive
selection for smaller-toothed individuals to maintain harmony in jaw and
tooth size; (b) in the Neolithic, natural selection favored individuals with
smaller and morphologically simpler teeth, which were less susceptible to
dental caries and attendant pathologies; and (c) in Pleistocene hunting and
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, afect 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). Materal efects (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 difcult to demonstrate that natural selection has brought about
changes in any organ or tissue system. However, taking a cue from Kurten's
(10 1) classic study of selection on dental characteristics of European cave
bears, Perzigian (149) attempted to demonstrate differential survival in an
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DENTAL ANTHROPOLOGY 107
Arikara Indian skeletal series by comparing the dental dimensions of juveniles
and adults. After finding that adults had significantly larger teeth than ju
veniles, Perzigian maintained that this difference suggested differential sur
vival of large-toothed individuals in a high-attrition environment. However,
an alterative explanation was ofered by Guagliardo (73), who found the
same juvenile-adult tooth-size disparity in a recent Tennessee Indian sample.
This author proposed that individuals in a group suffered differential stress
during the period of dental development (pre- and postnatal), and this contrib
uted to variation in tooth size. Moreover, the small-toothed individuals
stressed at an early age would have a greater mortality risk during childhood
and adolescence than those who had undergone normal development. Con
sidering the developmental fexibility of tooth size (57), it is plausible that
stress rather than genetic selection is the primary contributor to reported age
diferences in tooth size.
The efect 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
norther 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 patter of shoveling variation.
It has been suggested that another common dental morphologic trait,
Carabelli's cusp, increased in frequency and expression in Europeans because
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108 SCOT & 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 surfcial morphologic crown or accessory root
traits that might increase the masticatory efciency, tooth longevity, and
ftness 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 morhologically simpler crowns are less susceptible to caries because
there are fewer pits and fssures to collect food debris (14). Greene (68, 69)
believes this hypothesis helps to explain trends toward dental morphological
simplification in Egypt and Nubia, assuming genetic isolation.
Gene Flow
Estimating the degree and rate of gene flow using crown size or morphologic
variables is possible both diachronically and synchronically. Turer (196
198), for example, compared skeletal and living grps 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
fow, 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 particula living group. This is
impossible with serological genetic traits.
Genetic Drit
Genetic drift, including founder's efect, 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 diferentiation 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 refect
ing small population size-i.e. drif and founder's efect. 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 patters of crown and root trait variation.
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Mutation
DENTAL ANTHROPOLOGY 109
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 atribute of the upper first
premolar that occurs only among Indians of the New World (204). It may
refect a specific mutation that occurred after the peopling of the Americas.
READING BEHAVIOR FROM TEETH
Dental anthropologists are also interested in environmental efects on teeth
that enable them to identify temporal and spatial differences in subsistence
strategies and culture-specific behaviors.
Dental Wear: Masticator
Normal tooth use wears away the chewing or occlusal surfaces; in exteme
cases, the entire crown wears off, leaving root stumps for chewing. This
process, dental wear, has two components: attrition (resulting from direct
tooth-on-tooth contact) and abrasion (produced by contact with foreign mate
rials). Wear involves the inherent abrasive properties of food objects, abra
sives added to food, the duration and strength of chewing movements and
associated tooth-on-tooth contacts, and even the pathological grinding of the
teeth during sleep (bruxism) or out of habit. Wear also occurs at the mesial
and distal surface contacts of adjacent teeth (interproximal or approximal
attrition) owing to slight movement when they are used.
Dental wear occurs throughout life, but the nature of its variation has made
it diffcult to record with precision. Broca's fve-grade scale was once com
monly used to characterize groups for degree of crown wear. More recently,
precise ranked scales, where categories are defined by patters and degrees of
dentine exposure, have been employed for dietary assessments (127, 136,
174). Attempts to estimate areas of dentine exposure even better through
photographs and planimeters (129, 159, 216) are promising. Measuring
crown height, standardized by total tooth size, is another way to quantify
degree of dental wear (193). Recent work has also focused on the angle of the
wear plane as a key to relating teeth to dietary behavior (75, 127, 186).
Because skeletal and living samples usually differ in their age profles,
comparing groups for absolute degrees of wear presents a challenge. To
circumvent this diffculty, inta-individual rates of attition rather than mean
degrees of wear are ofen compared. By taking advantage of the natural
eruption sequence of the permanent molars, where there is an approximate
6-year interval between the eruption ofMI, M2, and M3, it is possible, with
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110 SCOTT & TURNER
respectable confidence, to derive age-independent rates of wear. To compare
samples based on rates of wear, one can compute mean rank (or area)
differences between MI-M2, M2-M3, and MI-M3 (105, 118, 137), calcu
late correlations (188), or perform principal axis analyses (159, 175).
Studies of degree, rate, and angle of crown wear have focused on regional
differences, temporal trends, sex diferences, and relationships to subsistence
strategies. Hunter-gatherers and agriculturalists can both show pronounced
degrees and rapid rates of wear, but the wear results from diferent causes.
Although meat is nonabrasive compared to plant foods, the carivorous
Eskimos show pronounced wear due to frozen or tough dietary items that
require extensive chewing, powerful chewing muscles that exacerbate tooth
on-tooth attrition, gritty contaminants on the food, and the use of their teeth as
all-purpose tools. Agriculturalists exhibit degrees and rates of wear equalling
or exceeding those of Eskimos, but this is because plant foods have more
abrasives than meat; when seeds and nuts are processed by milling or grinding
stones, fine abrasives are added to the food that greatly accelerate wear rates;
and the environments inhabited by many agriculturalists have more dust in the
air, adding even more abrasives to the diet. For these reasons, agriculturalists
would probably exhibit uniformly more wear than hunter-gatherers, as Leigh
(109) noted, were it not for the common use of food preparation techniques
(e.g. baking in earth ovens, bag or pot stewing, boiling, etc) that reduce the
consistency of both plant and animal foods and lessen the need for prolonged
vigorous mastication.
Although degrees and rates of wear do not always distinguish groups with
different subsistence means, other components of wear may be more diagnos
tic. B. H. Smith (186) studied molar wear angle in five hunter-gatherer and
five agricultural groups from around the world. She found that agriculturalists
consistently showed a more oblique wear angle than hunter-gatherers. She
also found that diachronic Nubian samples, dating from 5000, 4000, and 2000
BP, showed more steeply angled wear through time, suggesting a trend toward
an increased reliance on agricultural products. Wear angle promises to be a
useful variable to further understanding of the origins and development of
agriculture in various regions.
Hinton (86) discovered in his study of prehistoric Tennessee Indian teeth
that interproximal wear can also distinguish subsistence modes. Interproximal
wear facets were significantly broader in Archaic intensive foragers (6000-
500 Be) than in Mississippian agriculturalists (AD 1300-1550), even when
adjustments were made for tooth size and occlusal wear. A Woodland sample
(AD 7501150), characterized by a mixed economy, fell between these ex
tremes. Hinton attributes this trend to increasing reliance on cultigens, accom
panied by food preparation techniques that reduced heavy chewing.
Hinton (85) also analyzed anterior tooth wear in two hunter-gatherer and
two agricultural groups, finding another distinction between wear characteris-
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DENTAL ANTHROPOLOGY III
tics of these subsistence categories. The hunter-gatherers exhibited more
absolute and relative wear on their anterior teeth than did the agriculturalists.
Moreover, the anterior teeth of hunter-gatherers more frequently exhibited
labially rounded wear, suggesting the use of these teeth as tools. Agricultural
ists showed more cupped wear on their incisors and canines, which may
refect higher rates of molar and premolar tooth loss (due primarily to
increased caries and periodontal disease) and an associated shift of chewing
function to the anterior teeth.
Other regional and micro geographic wear studies in various parts of the
world have revealed other differences attributable to subsistence strategies
(127, 165), changing dietary patters 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 diference 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 norther Indians
(209). Milner (122) noted, however, that chipping occurred frequently i
Mississippian period Indians from Illinois.
Dental Wear: Nonmasticator
Several nonmasticatory behaviors can be identified in distinctive types of
dental wear. Generally, these behaviors must be practiced over a long period
to leave marks on the teeth. In some cases, pattered wear can result from
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112 SCOTT & TURNER
using the teeth in a particular task, or it may be incidental to a certain cultural
practice (31, 128, 192).
Used as pliers, vises, or a third hand, the anterior teeth often show
pattered wear unrelated to chewing food. This wear may take the form of
grooves or notches on the occlusal surfaces; or the distal and mesial margins
of two adjacent teeth may be disproportionately wor, resulting in a deep V
or U-shaped depression below the level of crown wear induced by normal
chewing. For example, Northwest Coast, Califoria, and Great Basin Indians
manufactured nets, baskets, mats, and other plant-based objects. Grooves on
the occlusal and interproximal surfaces of their anterior teeth were likely
produced by pulling abrasive fibrous materials through and across them
(40, 103, 168). The manipulation of sinew (for bow strings, ropes, etc) can
result in similar anterior tooth grooving. Some implemental uses of the
anterior teeth, which might involve grasping objects evenly across all
these teeth, do not leave abraded grooves but are indicated instead by labial
rounding (85).
Most dental wear is limited to the occlusal and interroximal surfaces, but
certain dietary habits can wear the lingual surfaces of the teeth. One example
is lingual surface attrition of the maxillary anterior teeth (LSAMAT) reported
for prehistoric Brazilian (211) and Panamanian (92) skeletal samples. As the
wear is on a surface of the crown that does not participate in normal occlu
sion, and there is no corresponding wear on the labial surfaces of the
mandibular incisors and canines, LSAMAT is interpreted as the result of an
unusual oral practice: pulling an abrasive material, perhaps manioc root,
between the tongue and lingual surfaces of the upper anterior teeth. As this
unusual wear form is found in association with high rates of dental caries,
processing a carbohydrate-rich plant material was a more likely cause than
shellfish or meat consumption.
Cultural practices sometimes affect the dentition. The use of labrets among
Alaskan and Pacific Northwest natives is indicated by polished or wor labial
or buccal tooth surfaces (40, 147). Such faceting is readily distinguishable
from masticatory and implemental wear. Habitual pipe smoking may also
leave a distinctive wear patter. Pipes clenched between the teeth over long
periods lead to approximately equal abrasion on the upper and lower teeth,
often resulting in an ovate hole observable when the teeth are occluded. The
use of probes to remove debris and perhaps bring relief from dental pathology
is also thought to leave distinctive interproximal grooves. Typically, this wear
is at the cervico-enamel junction, so both crown enamel and the root are
affected (5,212). While only a few workers have concentrated their efforts on
habitual behaviors refected in the dentition, this is a promising avenue of
biocultural research for archaeologists and ethnographers, as well as dental
anthropologists.
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DENTAL ANTHROPOLOGY 113
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 diferentiation, simple cosmetics, or other cultural motiva
tions. In some cases, tooth-tool use results in patters of wear that simulate
intentional dental mutilation. Blakely & Beck (8) outline criteria to distin
guish incidental from intentional dental modifications.
ECONOMICS OF DENTAL PATHOLOGY
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 (14),
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 afect 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 sof tissues and periodontal mem
brane surrounding tooth roots. It is commonly held that long-standing in
fammation 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
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114 SCOTT & TURNER
ly (pockets develop around specific teeth). Some workers identify periodontal
disease in dry bone only when resorption is found in conjunction with
pathological alterations of the alveolar crest, including the loss of cortical
bone (2, 32, 39).
When the area surrounding the tip of a root is invaded by bacteria, a painful
disease process can set in. The white cell and phagocyte defense against this
invasion lea.s to fluid and dead cell accumulation in the periapical region,
and a well-defned spherical hole develops. Usually the pocket breaks through
the alveolar wall, producing an abscess. This invasive-destructive process,
periapical osteitis or abscessing, is usually manifest on the buccal surface of
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
resortion 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 extemes.
This tooth survivorship patter 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) insufcient 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 frst 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.
Changing Subsistence Strategies and Declining Oral Health
Some individuals in a population are caries free because of host-resistance,
dietary habits, or some combination of the two. At one time we believed that
certain so-called primitive populations who had no or very few caries might
possess an inherent resistance to this disease-.g. Eskimos, who were
almost caries-free prior to European contact (108, 145). This theory was
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DENTAL ANTHROPOLOGY 115
dispelled in the 20th century when caries rates in long-isolated Eskimo
populations increased dramatically in Alaska (34, 156, 164), Canada (120),
and Greenland (145) only a few decades afer these groups adopted many of
the refned carbohydrates of the European diet.
Although caries occurred in Paleolithic hunting and gathering groups (25),
this disease was minimal until the Holocene emergence of food production.
Turer (201) reviewed worldwide caries frequencies and found that hunters
and gatherers had less than a 2% caries rate, mixed economies about 5%, and
agricultural economies 10% or more. The practice of intensive agriculture and
increased use of sticky carbohydrates clearly exacerbates caries formation
(see also 109, 155, 166).
Multi-generational epidemiological studies employing skeletal populations
have discered significant caries-rate trends around the world (27, 36, 79,
117, 130--133, 194). In some instances, increased or decreased caries rates
associate with specific historic events (79). Brabant (11) noted several long
term trends since the Mesolithic: an increase in the number of individuals in a
sample who exhibit caries, an increase in affected teeth per individual, greater
involvement of the anterior teeth, earlier onset of caries, and an increased
frequency of deciduous tooth lesions. From the observations of Lunt (117)
and others, a trend is also evident in caries type. In prehistoric and medieval
groups, most lesions were approximal and cervical (i.e. root caries). Only in
recent human populations do occlusal caries, in pits and fissures and at
contact points between the teeth, predominate in frequency. This changing
caries patter is likely due to diminished crown wear; pits and fissures do not
wear away in modem humans who use highly refined foods.
Periodontal disease shows a strong relationship to age (2), but it can be
signifcantly influenced by general dietary practices. Leigh's (109) classic
work on North American Indians showed that hunter-gatherers (Sioux) and
intensive foragers (Indian Knoll) exhibited less periodontal disease than
agriculturalists (Arikara, Zuni). Moreover, groups that practiced a hunting
gathering or mixed economy generally exhibited alveolar resorption only in
the molar districts; among the agricultural Zuni, generalized resorption
affected all teeth.
Anthropologists may have traditionally overestimated the role of per
iodontal disease in horizontal alveolar bone loss (32). About 90% of the
individuals in a wide range of premoder groups were free of buccal alveolar
bone loss. In two modem samples, bone loss attributable to periodontal
disease was about three times greater than in premoder series. The work of
Clarke and his colleagues (32) suggests that the significance of periodontal
disease is a more recent phenomenon than the rise in caries frequencies.
Dental abscess may occur at about the same rate in groups practicing
different subsistence economies, though its causes vary. Leigh (109) reported
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116 SCOTT & TURNER
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
of total teeth lost per number of observable sockets (25), percentage of tooth
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, diferential 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 benefts of civilization.
THE IMPRINT OF STRESS ON TEETH
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.
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Fluctuating Asymmetr
DENTAL ANTHROPOLOGY 117
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-.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
fuctuating asymmetry (214).
Twin studies have shown that fuctuating asymmetry in the dentition has no
significant genetic component (152), so it is viewed primarily as a refection
of developmental noise induced by environmental factors. Because fuctuat
ing asymmetry might serve as a generalized indicator of stress, both
archaeological and living populations have been studied. Perzigian (150)
found significantly more crown size asymmetry in Archaic Indians from
Indian Knoll than in late prehistoric and protohistoric agricultural groups.
This greater asymmetry was associated with other indicators of stess, includ
ing Harris lines, enamel hypoplasia, small adult stature, and high infant
mortality. Less dental asymmetry has been found in modem human groups
than in archaeological samples (49), although exceptions have been noted (7).
Fluctuating asymmetry may reflect between-group differences in environ
mental stress, but recent studies have pointed out potential limitations of this
measure. Smith et al (187) found that dimensional asymmetry, which is
typically dwarfed by overall tooth size, varies closely with sample size. These
authors demonstrated that between-group differences in antimeric correlations
or root mean squares might be attributable to sampling error when sample size
is less than 100, and many archaeological series studied for dental asymmetry
fall below this number. Greene (71 ) pointed out that between-observer
measurement error may compound the problem of between-group com
parisons of asymmetry; real diferences may be obscured while significant
diferences may be claimed where none exists. Associations between specific
measures of prenatal stress and increased levels of asymmetry remain to be
identified (48, 187). Although fluctuating asymmetry may help us measure
environmental stress, we must overcome the problems of sampling and
measurement error and focus on the phenomenon's proximate causes (cf 172).
Developmental Defects
Developmental defects of the dentition are being used to measure stess 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
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118 SCOT & TURNER
enamel and dentine, respectively, can nonetheless be disrupted during de
velopment. Severe environmental insults can permanently afect the structure
of the enamel or dentine. Some such imprints are visible macroscopically
(e.g. hypoplasia, hypocalcification); others can be seen microscopically (e.g.
striae of Retzius, Wilson bands) .
The best-studied developmental defect, surface enamel hypoplasia, in
volves enamel thinning and is manifested as horizontal circumferential lines,
grooves, and/or pits on a tooth crown. One or more hypoplastic lines may be
evident on a single tooth, with expression concentrated on the middle third of
the crown, followed in frequency by the cervical and incisaVocclusal thirds
(60). Because crown formation occurs within well-defined temporal limits, it
is possible to determine the ages, usually by half-year intervals, at which
specific hypoplastic lines developed (37, 61, 62). Hypoplasia is usually
bilateral and can occur on any tooth; it is most common and pronounced on
the incisors and canines (52, 60). The wide variety of associated environmen
tal stressors, including malnutrition and disease, produce similar hypoplastic
lines (see references in 62, 163).
Studies of within- and between-group variation in the frequency, severity,
and age distribution of hypoplastic lines have yielded interesting results.
Students of prehistoric midwester American Indian series have found that the
shift from an intensive collecting or mixed economy to a focal agricultural
economy was accompanied by a significant increase in developmental enamel
defects (61, 162, 169, 170). This trend is usually attributed to multiple
stressors, including increased population density and sedentism, diminished
health status, and potential nutritional problems when maize becomes a staple
food. The chronological distribution of hypoplastic lines, which developed
primarily between ages two and four, is used to support an interpretation of
stress related to weaning. Goodman et al (62) also found differences in the
distribution of hypoplasias between pre-Mississippian and Mississippian sam
ples that may indicate an earlier and greater use of a low-protein weanling diet
(maize gruel?) in the latter group. Hypoplastic expression peaked somewhat
later (3- years) in a Barbados slave population dating from the 17th to 19th
centuries than in most Amerindian samples, perhaps owing to a delayed and
taumatic post-weaning period (37).
In studies of mortality and morbidity in skeletal samples, workers have
found a relationship between age of onset of enamel defects in the deciduous
teeth (hypocalcification and hypoplasia) and mortality risk. Individuals with
enamel defects that developed prenatally were more likely to die in early
infancy that those who developed such defects postnatally; nonaffected chil
dren lived longer than those with defects (9). Another study found that
prenatally formed defects occurred more frequently in intensive agricultural
ists than in earlier Archaic and Woodland samples ( 1 69) . Cook & Buikstra
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DENTAL ANTHROPOLOGY 119
(35) found additional relationships between deciduous crown defects, circular
caries in these defects, and higher early-childhood mortality rates. Despite
medical progress, developmental defects in the dentition are about as common
today under conditions of poverty as they were prehistorically (91, 126).
THE GROWTH OF DENTAL ANTHROPOLOGY
Over the past two decades, the American Journal ofPhysical Anthropology
and other jourals have published numerous articles on the dentition of living
primates, fossil hominids, and skeletal and living human populations. An
thropologists are exploiting dental information to resolve a myriad of histori
cal, evolutionary, behavioral, health, and dietary questions. With the rapidly
accumulating volume of information on dental variation, dental an
thropologists should be encouraged to synthesize and summarize results so
that they are more accessible to students and nonspecialists. Although physic
al anthropologists are ofen depicted as "bone and tooth" specialists, the
"tooth" side of this characterization could be more fully represented in
physical anthropology, human biology, and even general anthropology text
books.
In earlier years, dental anthropological issues far outnumbered dental
anthropologists. Today, students at many universities can take formal courses
in dental anthropology at either the undergraduate or graduate level. This
opportunity for increased training, accompanied by a growing number of
graduate students specializing in aspects of the dentition, helped stimulate the
formation of the Dental Anthropology Association (1986). Now, numerous
researchers are positioned to address the broad range of anthropological
questions that can be resolved through study of the human dentition.
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