Journal of Archaeological Science (1997) 24, 677–700
Dental Wear in Immature Late Pleistocene European
Hominines
Mark Skinner
Department of Archaeology, Simon Fraser University, Burnaby, B.C., V5A 1S6 Canada
(Received 28 June 1995, revised manuscript accepted 7 May 1996)
The relationship between dental attrition (nine stage scale) and specimen age, or functional age of teeth, is compared
between immature Middle Paleolithic (Neanderthal specimen count=28, tooth count=165) and Upper Paleolithic
(anatomically modern specimen count=54, tooth count=338) samples. All specimens are from Western Europe.
Maturation rate is treated as the same for both samples for analytical purposes and evaluated in terms of the results.
Analysis of reduction in crown volume with age indicates that the wear rate for the posterior permanent (P4–M3) and
primary teeth (dm2) does not differ between the samples; however the anterior primary teeth (di1–dm1) are more worn
in the UP sample with age held constant. By contrast, anterior permanent teeth (I1–P3) wear more rapidly in the
Neanderthal sample. Similar results are obtained in a comparison of relative attrition.
The MP sample of anterior primary teeth lags about a year behind the UP sample in the degree of accumulated wear
up to about age 5 years. Thereafter differences disappear. Analysis on the basis of individual specimens (ANOVA)
confirms these results with the MP sample of anterior primary teeth significantly older (P=0·0498) at successive wear
stages. While the rate of wear of the anterior primary teeth does not differ between the samples, the age of attrition
onset differs by about 1 year; age 2 years in the UP sample and 3 in the MP sample. It is concluded that dietary
supplementation of infants commences 1 year earlier on average in the UP. Data are insufficient to determine whether
the Late and Early Upper Paleolithic samples differ significantly in the study variables. Apart from the finding of earlier
onset of dental wear in the UP, the lack of difference in the relationship between age and tooth wear supports a model
of similar maturation rates and dietary abrasiveness in the Middle and Upper Paleolithic. Significantly earlier dietary
supplementation of UP infants may have reduced birth spacing through diminished hormonal inhibition of ovulation.
These results are compatible with demographic increase in the UP.
? 1997 Academic Press Limited
Keywords: LATE PLEISTOCENE, NEANDERTHAL, UPPER PALEOLITHIC, DENTAL ATTRITION,
AGE AT DEATH, DIET.
Introduction
Beyond descriptions of single specimens, there are no
integrative studies of dental attrition among immature
fossil hominines from Europe.
Wear on the teeth is a function primarily of the
interaction of the physical consistency of the diet and
the time during which a tooth has been functioning
(industrial tasks are ignored for the moment) (Molnar,
1971). With each variable alternately held constant, it
is possible to compare developmental rates and dietary
consistency between populations, such as those from
the Middle and Upper Paleolithic, in which neither
variable is well understood. For example, a study of
dental attrition in adults by Smith (1977a) from the
region and time span reported here found that increasingly recent hominines with smaller teeth and faces
showed elevated levels of dental attrition; leading to
the conclusion that facial reduction was not linked
to reduced functional demand on the face. A possible
weakness with this conclusion is our inability to be sure
of equivalent age compositions of samples composed
of adults of indeterminate age.
Furthermore, there are suggestions in the literature
that Neanderthals matured more quickly or, at least,
T
he study of tooth wear as a guide to behaviour
and life history has a long tradition in physical
anthropology (e.g. Molnar, 1972; Patterson,
1984 and contained references; Powell, 1985 and contained references; Dreier, 1994). However, fossil hominine of the Late Pleistocene have been little studied for
dental attrition. The Gibraltar Neanderthal child was
reported by Carbonell (1965) to show more wear than
observed in modern children. A study of microscopic
enamel striations on the same specimen (Fox & PerezPerez, 1993) led the authors to suggest a high meatcontent diet like that of eskimos. A similar result is
reported for the Middle Pleistocene individuals from
Arago (Tautavel) (Puech, 1982) who wore their teeth
much like those of meat-consuming aborigines of the
last century; at a lesser rate than observed among pure
vegetarians. Wallace discussed incisor rounding in La
Ferrassie 1 as an indication of industrial task wear
related to clamping with the jaws (1975); while Smith
(1977a,b) studied relative attrition and consequent oral
pathology in specimens from Riss to Late Würm.
677
0305–4403/97/080677+24 $25.00/0/as960151
? 1997 Academic Press Limited
678 M. Skinner
erupted some teeth earlier than did anatomically
modern humans (e.g. Legoux, 1966; Wolpoff, 1979;
Stringer et al., 1990; Zollikofer et al., 1995). While it is
inappropriate to attempt a complete discussion of the
topic here (see discussion in Skinner (submitted), it can
be pointed out that much of this viewpoint stems from
studies of the Gibraltar child whose perikymata count
is low and whose brain case is large (Dean et al., 1986;
Zollikofer et al., 1995). Minugh-Purvis (1988) has
shown that the Gibraltar child is unusual among
Neanderthals for its size while the utility of perikymata
counting for age at death estimation has been strongly
called into question by the results of Huda & Bowman
(1994). Recently, Tompkins (1991), Trinkaus &
Tompkins (1990) and Trinkaus (1995) have suggested
that Neanderthal patterns of dental formation, eruption and developmental dental histology fall within
modern human ranges of variation, although possibly
at the faster end, implying similar life history segments.
Nevertheless, the suggestion of Neanderthal precocity
can be evaluated by assuming dietary equivalence and
comparing dental wear. Alternatively, maturation rates
can be assumed to be the same so as to compare dietary
abrasiveness. Some have maintained that the advent of
stone boiling and preparation of soup occurred in the
Upper Paleolithic (Hadingham, 1979; Pfeiffer, 1986).
Arguably this would soften the diet and reduce dental
wear.
Several dental variables could potentially affect rates
of dental attrition; these include size, which is larger
on average in Neanderthals (Brace et al., 1987) and
enamel thickness. Two studies suggest that enamel
was thinner in Neanderthals (Zilberman et al., 1992;
Molnar et al., 1993). The latter study found that
modern permanent molar enamel was approximately
1·3 times thicker than in Krapina Neanderthals which
the authors tentatively attributed to systemic stress or
enlarged pulp chambers among the latter. Further
consideration of this problem (Smith & Zilberman,
1994) confirmed that Neanderthals had both thinner
enamel and enlarged pulp chambers.
All else being equal the following predictions can be
made for MP populations:
(a) given accelerated dental (and presumably somatic) maturation, dental wear will have an
earlier onset but accrue at a lesser rate (or alternatively, at equivalent wear stages, age will be
greater);
(b) given the absence of stone boiling, wear will be
greater;
(c) given larger teeth, wear will be less;
(d) given thinner enamel, wear will be greater.
The variables of tooth size, enamel thickness, tooth
formation and wear are considered in this study.
Attained tooth formation is used as a proxy for age at
death using modern normative standards. While it is
not possible to control all the variables at once, analysis of primary and permanent waves and evaluation of
the magnitude of observed differences will enhance the
investigator’s ability to judge the contribution that
each of the variables may be making. For example,
if the pattern of differences between the Middle
Paleolithic and the Upper Paleolithic changes from
early to late ontogeny (primary versus permanent
teeth) it can be argued that innate features of differences in developmental rate, tooth size and enamel
thickness are not the explanation (unless these change
their relative states during ontogeny).
Immature skeletal remains in which teeth are preserved permit the investigator to determine individual
age at death with more precision and accuracy than is
normally possible with adult remains (Ubelaker, 1978;
Buikstra & Meikle, 1985). Physiological and behavioural phenomena which are time dependent such as
introduction of solid foods in infancy, tooth wear,
and developmental rates can be calibrated from tooth
formation to investigate potential differences between
populations. The research reported here compares
wear on the teeth between immature remains from the
Middle and Upper Paleolithic of Europe.
Materials
Data were collected between May and July 1982. Some
additional data had been collected in the summer of
1973 and some information is taken from the literature
(Malegni & Ronchitelli, 1989). Eighty-two specimens
are reported here. A brief description of each specimen
with its provenance and repository is available from
the author.
Delimitation of samples
The Middle Paleolithic (MP) of Europe occurred in
Oxygen Isotope Stage 3, while the Upper Paleolithic
(UP) continues into Stage 2 (Straus, 1995). Although
the European Late Pleistocene artefactual record indicates fundamentally the hunting of large mammals
throughout, with a largely undocumented but presumably critical contribution from gathered resources,
there is general, but not total, agreement that the UP
with an emphasis on blade technology heralded a
profound cultural change (Mellars, 1973). This was
marked, in addition, by bone working, concentration
on usually a single species of mammal, especially
reindeer, increased social group size, increased population density, personal adornment, trade contacts, and
artistic accomplishment, including cave art (White,
1985). For this reason, it was felt acceptable to retain
the classic demarcation between the Middle and Upper
Paleolithic as the basis for comparison of dental attrition. The temporal delimitation of samples was based
on the assumption that the major, but certainly not
sole, factor in comparative dental attrition would likely
be changing subsistence strategies either in food base
or means of food processing. This is not meant to reify
the commonly held view of rapid cultural change
Dental Wear in Immature Late Pleistocene European Hominine 679
across the Middle/Upper Paleolithic boundary (see
discussion in Straus, 1995). Probably neither time
period was behaviourally and culturally static.
The upper time limit excludes Mesolithic individuals
who are felt to have been adapted to a lifestyle based
more on exploitation of smaller, more solitary forest
game and a wide variety of faunal resources (Frayer,
1978). The lower time limit is deliberately set at the
penultimate glaciation. One reason for this is the
pragmatic wish to include in a rather small Middle
Paleolithic sample, important pre-Neanderthal
immature specimens like La Chaise. The other is in
recognition of the apparent cultural continuity between
sites from this earlier time period and Early Würm
sites:
It is in fact difficult to think of a single well-characterized
tool-form found in middle Paleolithic industries which
does not have an ancestry extending back at least to the
penultimate (‘Riss’) glaciation . . . (Mellars, 1973, cited in
White, 1985).
Spatial delimitation of the samples is a function
primarily of non-theoretical considerations. The
majority of accessible specimens are to be found
in western Europe; in terms of the immature Late
Pleistocene fossil record, a ‘‘francocentric’’ emphasis is
unavoidable (cf. Straus, 1995). This and a 3-month
field season curtailed collection somewhat. Quite a
number of specimens examined were excluded from
consideration for reasons referred to above in the
discussion of suitability of specimens or because they
were deemed too ancient or recent (e.g. Badger’s Hole
and Mother Grundy’s Parlour). I have excluded seven
immature specimens from the MP sample because they
are either not Neanderthals, or are from North Africa
or the Middle East and, arguably, form a separate MP
clade of hominines (Stringer, 1994). These include
Skhul 1, Qafzeh 4, Qafzeh 11, Ihroud 3, Ksar ’Akil,
Rabat 1 and Haua Fteah*. The comparisons drawn
here are between two samples which differ biologically,
culturally and temporally.
The total sample consists of 82 individuals with
503 teeth ranging in individual age from about 1 to
18·6 years old. The earlier sample spans at least
150,000 years, including 28 individuals with 165 teeth
(mode=one tooth, six specimens), and the later sample
spans only about 25,000 years, including 54 individuals
with 338 teeth (mode=two teeth, 17 specimens). Since,
for the majority of specimens, it was not possible to
know the sex of the individual, it was judged better to
use mid-sex age determinations for each individual
despite acknowledged sex differences in dental maturation (Haavikko, 1970). The provisional nature of any
results must be appreciated in light of these facts. Not
all teeth were able to be included in all analyses.
In summary, each sample contains individuals differing in largely uncontrolled ways in temporal and
*Details of dental formation and attrition for these specimens may
be obtained from the author.
spatial origin (some groupings of individuals are from
single sites, however), sex, dietary behaviour, facial
configuration (at least in adults), and, possibly,
maturation rate. Furthermore, the two samples contain
individuals considered to differ at the subspecies or at
least ‘‘racial’’ level. The two samples are, however,
demographically similar in that the mean ages at death
of the two samples do not differ significantly:
MP=7·74 years, UP=7·02 years; t=0·142, P=0·5229
(Figure 1). Interpretation of the significance of the
results must consider the role that each of these
variables may play in having created any perceived
differences in attrition among these Late Pleistocene
immature hominines.
Methods
The basic relationship studied in this research is
between occlusal tooth wear and functional age of the
tooth. For a tooth to be included in the study it has to
possess the following attributes:
(a) the tooth had to be sufficiently erupted that it was
probably gingivally erupted;
(b) occlusal attrition, or the lack of it, had to be
reliably scoreable, i.e. post-mortem enamel loss
could not obscure attritional status;
(c) functional age of the tooth had to be ascertainable
by reference to a normative eruption age for that
tooth, i.e. age at death of the individual had to
be able to be determined, from which age the
eruption age of the tooth was subtracted.
Age at death was found from tooth formation, or from
some other age indicator such as root resorption where
normative standards for such existed, or from the
literature. Put another way, the tooth had to be from a
non-adult. For the purposes of this analysis, subadulthood is defined dentally as a tooth (and individual) in
which root formation (or that of another tooth from
the individual) was not complete. Root formation is
defined as full closure of the apical canal (Fanning,
1961). The age of socio-sexual maturation is not
indicated in this definition.
Calculation of individual age at death
A Fisher Portable X-ray machine (65–90 kVp,
20–10 mA) with collimator was transported to Europe
where dental film and powder developing chemicals
were purchased. A total of 258 radiographs were taken
on approximately 59 specimens to supplement those
earlier reported (Skinner & Sperber, 1982). All useful
radiographs were photographed and tooth formation
was determined from the photoradiographs and/or
original observations as detailed below. Age at death
was derived by converting the degree of tooth formation as observed in radiographs to an equivalent
dental age based on a combination of normative
standards. Nolla (1960) pioneered the use of dental
680 M. Skinner
7
6
4
3
2
1
0
Haavikko (1970)
Crown initiation (Ci)
Cusp coalescence (Cco)
Crown 1/2 to 3/4
Crown complete (Crc)
Root 1/4 (R1/4)
Root 1/2 to 3/4
Root complete, apex open (Rc)
Apex closed (Ac)
MP, N = 28 individuals
Mean age = 7.74 years
5
Count
formation for age determination but her standards
have been superseded by subsequent workers noted
below. That tooth formation is a preferable means of
determining age at death in immature remains is widely
accepted (Lewis & Garn, 1960; Demirjian, Goldstein &
Tanner, 1973; Stewart, 1979; Buikstra & Mielke, 1985;
Smith, 1991).
Scoring tooth formation was based on criteria
supplied by Demirjian, Goldstein & Tanner (1973),
modified for fossil hominine materials (Skinner, 1978).
This is a nine-stage scale, including ‘‘0’’, which was
converted into the scale provided by Haavikko (1970)
as follows:
2
4
6
8
10
12
14
16
18 20 years
This study
Crypt formation (1)
Initial crown formation (2)
Crown 1/2 (3)
Crown complete (4)
Root initiation (5)
Root advanced (6)
Root complete,
apical canal open (7)
Apical canal closed (8)
1
2
3
4
5
Haavikko’s normative standards are superior because
they include both upper and lower jaws and the third
molars. However, since they do not include the primary teeth, nor some of the beginning formation stages
for the early maturing teeth, recourse was made to the
following additional sources:
(a) Fanning (1961): primary canine, primary first
molar, primary second molar;
(b) Fanning & Brown (1971): female primary
canine—stage 8, male and female primary first
molar and primary second molar—stage 8, lower
permanent canine—stage 1, lower first (=third)
premolar—stages 1 and 2, lower second (=fourth)
premolar—stage 1, lower first permanent
molar—stages 1, 2 and 3;
(c) Aprile & Figun (1956, cited in Legoux, 1966):
isolated upper primary teeth and lower primary
first and second incisors;
(d) Moorrees, Fanning & Hunt (1963): lower permanent canine—stages 2 and 3.
These studies provide age equivalents for stages of
dental formation for each stage of each tooth (with
exceptions due to lack of detailed study) separated by
sex. Consequently, it is possible to derive two age
estimates for a fossil hominine: as though it were male;
and as though it were female. The ages at death
indicated by all teeth were combined and averaged and
a mid-sex age calculated for analytical purposes.
Calculation of a tooth’s functional age
Ideally, in an analysis of this sort one would have a
large series of dentitions with no missing teeth which
6
UP, N = 54 individuals
Mean age = 7.02 years
7
Figure 1. Age at death distribution compared between the Middle
Paleolithic (MP) and Upper Paleolithic (UP). Mean age at death is
almost identical in both samples.
would permit comparisons of dental attrition between
individuals of equivalent dental age. With fragmentary
fossil material this is not possible. To maximize sample
size, the tooth becomes the basis for comparison.
However, since teeth erupt and start to wear at characteristically different times, one has to calculate a
functional age for a tooth. This allows teeth with
equivalent functional ages, whether of different tooth
types or from different individuals, to be combined
for analysis. There is no question that different populations erupt their teeth at different times for a
combination of genetic and environmental reasons
(Haavikko, 1969). Since it is difficult to judge which
normative standards would be most suitable for
European fossil hominids, I elected to continue to use
Finnish data (which, after all, is northern European)
because Haavikko provides gingival eruption schedules
for the permanent teeth of the children whose tooth
formation was largely utilized in this study to age the
fossil hominine children. Also, Nystrom (1977) has
published gingival eruption standards for an additional
series of Helsinki children. Utilization of a single
geographical, and relatively genetically homogeneous,
reference group for age determination and for calculation of functional age of the teeth should provide a
high degree of confidence in the uniformity of results.
Dental Wear in Immature Late Pleistocene European Hominine 681
The Finnish data appear to accord with dental maturation studies from elsewhere (Moorrees, Fanning &
Hunt, 1963; Demirjian, Goldstein & Tanner, 1973).
The actual calculation of functional age for a tooth
is found by subtracting gingival eruption age for that
tooth from the previously calculated individual age at
death (see above). The latter figure is expressed as
mid-sex age since for the majority of specimens sex was
unknown. This procedure provides an estimate of age
in years during which a tooth has been gingivally
erupted and exposed to attritional forces, ie. attrition
can be compared to how long a tooth has been
wearing.
Attrition
Mammalian tooth size is assumed, generally, to be
appropriate to diet (van Valen, 1972). Thus, regardless
of variables such as differences in life history, tooth
size, enamel thickness, and dietary consistency, relative
attrition should not differ greatly between species.
Natural selection would militate against any variations
in tooth size that were needlessly large or so small as
to reduce reproductive success. Human choice could
presumably override this force at least in the short
term such that relative attrition could differ between
populations.
Attrition is defined as the progressive loss of occlusal
and interproximal contact tooth substance due to
masticatory behaviour. It embraces crown reduction
from tooth–tooth contact and tooth–bolus contact;
similarly it does not exclude tooth reduction due to
occasional or habitual non-dietary occlusion as in the
performance of industrial tasks or clamping of objects
between the jaws to aid in manipulation. The latter
cause of tooth substance reduction, while not felt to be
a significant factor in dentally immature individuals,
was specifically countenanced in this study by treating
the anterior and posterior teeth as separate analytical
units (see below). This study does not address potential
differences in enamel quality. Thorough reviews of
both ordinal and interval means of describing attrition
are provided by Patterson (1984) and Powell (1985).
The study reported here is designed to accomplish
both, and has built-in checks on replicability through
recourse to sketches, photographs and tracings. Furthermore, it permits the amalgamation of different
tooth types, teeth of different size, and both primary
and permanent teeth.
Attrition was recorded in four ways:
(a) by photographs of dentin exposure on occlusal
surfaces (see below);
(b) by sketches of dentin exposure on occlusal surfaces. The function of these was to aid in tracing
dentin exposure and crown outline onto acetate
sheets from which to calibrate dentin exposure on
a ratio scale (see below);
(c) by measurement of crown dimensions as follows:
height of cusp or crown from cemento-enamel
junction at cervix, vertically to the worn or unworn occlusal surface. This was taken labially on
the anterior teeth and buccally on the premolars.
Crown height of molariform teeth was taken at
the following locations: mesio-buccal cusp (5–4,
6–4); disto-buccal cusp (5–5, 1–6, 1–7, 1–8, and
their antimeres); mesio-lingual cusp height (7–4,
7–5, 3–6, 3–7, 3–8, and their antimeres). In some
analyses, crown heights were converted to z-scores
for statistical analysis in which single observations
are expressed as standardized deviations from the
mean for a tooth group (e.g. MP anterior primary
teeth), so as to allow teeth of different innate
heights to be compared for analysis of height
reduction as age increases. Cube root of crown
volume:
height#mesio-distal length#
labio(bucco)-lingual breadth
was calculated for linear regression against age;
(d) by assignment of an attrition score, subjectively
assessed at the time of examination. These scores
were reviewed later in light of the photographs
and sketches when all data were at hand for
evaluation. Only rarely was it felt that an initial
score was in error. It should be noted that the
numbers are not meant to imply equal intervals
between stages although they do reflect the
continuous nature of the variable. That is, the
attrition scale is an ordinal description of wear
on the teeth. Its advantage is that it expresses
early stages of enamel reduction (see below)
which is necessary in immature remains but which
are usually ignored in attrition studies based
on adults (Molnar et al., 1983). While more
detailed schemes for scoring attrition exist (e.g.
26 stages developed by Dreier, 1994), they cannot
be applied easily across tooth types as is necessary when working with quite fragmentary
remains.
The rating scale for attrition is described below.
Stage 0: No attrition. Teeth which have not pierced the
gum or have only recently done so will be at this stage.
Isolated (ageable) teeth with no attrition were excluded
from the study unless there was sufficient root formation to be sure it was not still in its crypt at the time
of death. A tooth had to show substantial projection
above the alveolar margin to qualify for inclusion in
the study when it was at this attrition stage.
Stage 1: Enamel faceting (trace). On individual cusps
of the primary molars and permanent molars this
earliest stage of attrition is visible as tiny planes or
facets which reflect light from their flat surfaces. On
canine teeth, faceting can be seen on the side or tip of
682 M. Skinner
the crown while incisors show initial reduction of
mammelons. Otherwise there is no reduction in crown
height at this attrition stage.
Stage 2: Enamel rounding (mid). Cusp tips are slightly
smoothed and rounded with loss of angulated faceting.
Main fissures and crenulations are largely pristine. On
incisors, the mammelons are worn away while the
incisal edge is still narrow and unflattened. There is
only minimal loss of crown height. (There was one
individual with primary molars best scored at this stage
but with a trace of dentin exposure—La Chaise 14,
7–4, 8–5.)
Stage 3: Enamel flattening (advanced). There is appreciable reduction in crown height resulting in broad,
flattish, low occlusal elevation. On cheek teeth the
majority of the occlusal surface is involved although
deeper fissures may be little affected. Cusp tips are
obviously rounded. There is trace, or typically no,
dentin exposure but dentin may be discernible through
a thin enamel layer. Canines may tend to show relatively more dentin than the other teeth at this stage.
Incisors exhibit a broad, flat incisal edge with crown
reduction and often darkly staining dentin visible
through a thin enamel veneer.
Stage 4: Slight dentin exposure. On molariform teeth,
this stage is differentiated from the next by the fact that
attrition tends to be angled such that dentin is exposed
first on one side of the tooth and only later on the other
as well. At this stage, one or two (rarely more) islands
of dentin are exposed on one side of the tooth (buccal
in lowers, lingual in uppers). On premolars one cusp, as
opposed to both, shows slight dentin exposure. For
canines, only a small spot of dentin is exposed—about
the size of a pencil dot. Incisors show a thin strip of
dentin, tapering mesially and distally—about the width
of a thin pencil line.
Stage 5: Dentin advanced. Dentin islands show on both
sides of molariform teeth of a size exceeding that of the
previous stage. There is no coalescence of dentin
islands. Larger spots and strips of dentin may be seen
on canines and incisors, respectively.
Stage 6: Strong dentin exposure. On molariform teeth
there is coalescence of two or more islands of dentin
even to the point where enamel remnants may only
remain on the central occlusal surface. There is
marked crown height reduction. Canine teeth show
large dentin exposure with crowns about half original
height. Incisors now resemble canine teeth with an
expanding circle of dentin within the dentin strip,
due to encroachment of attrition on the deep pulp
chamber.
Stage 7: Enamel ring. All occlusal enamel is worn away
on molariform teeth leaving only an enamel ring of
fairly uniform width circumferentially. There may be
darkly stained islands of secondary dentin. Canines are
judged to be at this stage by having circumferential
enamel of a width similar to that of posterior teeth.
Incisors show very strong height reduction with loss
of interproximal contact and round or oval dentin
exposure to a marked degree.
Stage 8: Root involvement. While self-explanatory, this
occurs on the labial and buccal side of mandibular
teeth and on the lingual side of maxillary teeth. This
stage was not observed in the current study.
Calculation of proportion of dentin exposure
It has proved difficult to quantify attrition on enamel,
which is why studies of attrition usually employ an
ordinal scale. However, it is feasible to measure the
degree of dentin exposure in a fairly replicable fashion
(see discussion in Powell, 1985). The procedure
adopted in this study is to trace the outline of exposed
dentin within the crown outline similarly traced from
enlarged photographs of occlusal surfaces. The amount
of dentin exposure is expressed as a proportion (%) of
crown outline. This method does not include enamel
attrition at all; a not insignificant factor when dealing
with infants. This, however, is accomplished through
the ordinal scale approach described above.
Given the variable appearance of worn occlusal
surfaces and the inherent difficulties of consistently
scoring dental attrition when travelling from collection
to collection, all occlusal surfaces were photographed
and sketched to record the extent of dentin exposure.
This was in addition to a system of scoring relative
dental attrition assessed visually at the time of examination. Close-up photography was done with a
200 mm f/5·6 Medical-Nikkor lens equipped with a
ring-light and a series of screw-on supplementary
lenses which permit magnifications ranging from
1/15# to 3#, i.e. the size of the image created on the
negative. More than 700 photographs were reproduced
based on about 82 specimens. These form the basis for
the study described below of proportion of dentin
exposure, plus they served as a check on the attrition
scoring. Ilford Pan F ASA 50 black-and-white film was
used with the camera set on ASA 25 followed by
prolonged development for extra high quality detail.
Statistical analysis
Similar analyses were made of ‘‘teeth in combination’’
(regardless of source) and ‘‘individuals’’. The former
were undertaken to search only for patterns, in recognition of the possibility of making a Type I error due to
statistical redundancy among teeth from single jaws.
The latter were made to test for statistically significant
differences. Most specimens are very incomplete and
may not contain the same teeth. Hence, the systems
for scoring dental attrition (classes) and dental height
Dental Wear in Immature Late Pleistocene European Hominine 683
Table 1. Comparison of enamel thickness between the MP and UP samples based on teeth in combination with minimal or no attrition
Second primary molar
Culture
MP
UP
Count
Mean enamel
thickness (mm)
4
11
0·92
1·19
First permanent molar
t-value
P-value
Count
Mean enamel
thickness (mm)
"0·270
0·0172
1
4
1·12
1·61
t-value
P-value
—
—
Table 2. Comparison of interaction of dental attrition and functional age between the Middle and Upper Paleolithic samples based on teeth in
combination (ANOVA—see Figure 3(a)–(d))
Tooth group
Anterior primary
Anterior primary (sans Att. Stage 6,7)
Posterior primary
Anterior permanent
Posterior permanent
Number
Mean age
difference
MP
UP
0·82
1·40
"0·73
"0·61
"0·55
69
61
33
31
32
113
91
62
82
71
(z-scores) were created so as to be comparable across
tooth types. By this means the probability of making a
Type II error was minimized. Specifically, individuals
are represented only by single teeth. These were found
by random selection of one tooth per individual.
In those analyses of teeth in combination, regardless
of the individuals from which they derived, analysis
was performed separately by location and wave. These
analytical categories were created to allow for the
possibility of differential (possibly non-dietary) wear
on the anterior teeth and the possibility of markedly
different attrition environments in infancy and later
childhood (as well as the difference in tooth size
between baby and adult teeth). The permanent teeth
were divided into an anterior and posterior moiety of
four teeth each composed of the I1 to P3, and P4 to
M3, respectively. Similarly, the primary teeth were
divided into an anterior group (i1, i2, c, m1) and a
posterior tooth (m2). The latter grouping divided the
primary wave into an early erupting group (c. 6–18
months) and a later erupting group (c. 27 months).
All statistical procedures were carried out using
StatView 4.01.
Results
The role of enamel thickness
While others have shown that Neanderthals have
significantly thinner enamel (Zilberman et al., 1992;
Molnar et al., 1993), further analysis is undertaken so
as to compare this attribute directly between the two
fossil samples under consideration here. Measurements
of enamel thickness (and mesio-distal crown length)
were taken directly on radiophotographs and scaled by
reference to crown length measures taken on original
specimens. Measures were taken at the visibly thickest
Culture+functional age
F-value
2·364
16·278
0·471
4·901
<0·0001
P-value
0·1261
<0·0001
0·4944
0·0291
0·9927
Culture+attr. class
F-value
P-value
3·432
2·617
3·422
3·486
1·683
0·0019
0·0270
0·0073
0·0060
0·1446
enamel. Analysis was undertaken only on the most
commonly preserved tooth types (mandibular primary
second molar and permanent first molar) and only on
those specimens with attrition stage 2 (enamel rounding) or less. The results are shown in Table 1. The
thicknesses shown are (scaled) absolute values; the MP
primary tooth enamel is 0·27 mm thinner (c. 22%)
while the permanent molar enamel is approximately
0·49 mm thinner (c. 30%). A larger sample reported by
Zilberman et al. (1992) showed the thin permanent
molar enamel of Neanderthals. The value obtained
for the UP permanent enamel sample (1·6 mm) is the
same as that reported for modern humans by Puech
(1982).
Absolute wear
Crown volume was calculated by multiplying the three
linear dimensions and finding the cube root so as to
generate a linear variable for regression against functional age. The results are shown in Figure 2(a)–(d).
Not surprisingly, given the contributing variables, the
correlation values are very low. Consequently, the
apparent differences among tooth groups and between
the MP and UP samples should be interpreted with
caution. Nevertheless, the figure is a good guide to
differences detected in subsequent analyses.
It seems that, apart from the obviously larger teeth
on average of the MP sample, there are few differences.
Apparently, the anterior primary and posterior permanent teeth wear at the same absolute rate while the
second primary molar may wear faster in the UP
sample while the anterior permanent teeth wear more
slowly in the UP sample. These are merely indications
whose validity and significance have to be determined
from more exacting analyses.
684 M. Skinner
12
11
12
(a)
11
10
10
9
9
8
8
7
7
6
6
5
5
4
4
3
–3 –2 –1 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14
2
CubRootVol. = 6.823 – 0.074 * FunctAge; R = 0.026 (MP)
2
CubRootVol. = 6.146 – 0.055 * FunctAge; R = 0.015 (UP)
12
Crown volume1/3
11
(b)
3
–3 –2 –1 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14
2
CubRootVol. = 8.277 + 5.062E-3 * FunctAge; R = 5.289E-4 (MP)
2
CubRootVol. = 8.133 – 0.083 * FunctAge; R = 0.35 (UP)
12
(c)
11
(d)
10
10
9
9
8
8
7
7
6
6
5
5
4
4
3
–3 –2 –1 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14
Functional age (years)
3
–3 –2 –1 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14
2
CubRootVol. = 9.132 – 0.199 * FunctAge; R = 0.178 (MP)
2
CubRootVol. = 7.815 – 0.039 * FunctAge; R = 7.793E-3 (UP)
2
CubRootVol. = 9.547 – 0.058 * FunctAge; R = 0.055 (MP)
2
CubRootVol. = 9.247 – 0.069 * FunctAge; R = 0.092 (UP)
Figure 2. Regression of cube root of crown volume against functional age of teeth in combination: (a) anterior primary (di1–dm1); (b)
posterior primary (dm2); (c) anterior permanent (I1–P3); (d) posterior permanent (P4–M3) teeth. In this analysis, the anterior primary and
posterior permanent teeth wear at the same rate in both samples, whereas the posterior primary teeth wear more slowly and the anterior
permanent teeth wear more rapidly in the MP sample. There is considerable overlap between the two distributions. ,: Middle Paleolithic;
+: Upper Paleolithic.
Relative wear
Figure 3(a)–(d) shows the relationship between attrition wear stage and functional age of a tooth for each
tooth group. This type of analysis does not assess
attrition rate per se, but rather compares the difference
in mean functional age at each successive wear stage
for the two samples (ANOVA) (Table 2). All such
analyses based on ‘‘teeth in combination’’ may show
Type I error due to statistical redundancy. Results
conform fairly well to the previous analysis. Because
the anterior primary teeth show the most consistent
and statistically significant difference in attrition
between the MP and UP samples, this tooth group will
be dealt with last, and in more detail.
Posterior primary tooth
Unlike the previous analysis, there is no indication
from ANOVA that the two samples differ consistently
in the functional ages at successive wear stages of the
posterior primary (dm2) tooth. While the rate of crown
volume reduction (Figure 2(b)) is faster in the UP
sample, a comparison (ANOVA) of mean functional
ages over successive attrition stages (Figure 3(b)) does
not differ between the two samples. Indeed, it is clear
from Figure 4 that this tooth does not differ in the
developmental age required to reach slight dentin
exposure (good large sample) nearly as much as do the
rest of the primary teeth. It may be concluded that in
this part of the mouth at the developmental ages
involved (¢3 years or so), what was a large difference
between the samples (UP showing more worn teeth
than the MP sample at similar ages) is starting to
disappear. It is distinctly possible that the thinner
primary enamel of Neanderthals is causing their teeth
to wear faster once it is breached.
Anterior permanent teeth
Neanderthals show (Figure 3(c)) a significantly
younger mean age (ANOVA) at successive wear
stages for these teeth (P=0·0291). In other words,
Dental Wear in Immature Late Pleistocene European Hominine 685
DentStrng
5 19
59
1 5
EnamAdv
10 12
EnamMid
8 17
DentAdv
1 7
DentAdv
24 12
DentSlght
4 13
DentSlght
11 9
EnamAdv
510
EnamMid
33
8 11
EnamTrace
10 21
DentAdv
5 18
DentSlght
4 15
EnamAdv
3 15
EnamMid
6 10
13
12
11
10
9
8
7
6
5
4
3
2
1
0
–1
–2
–3
–4
(b) ANOVA: NS
EnamTrace
DentStrng
(c) ANOVA: P = 0.0291
13
12
11
10
9
8
7
6
5
4
3
2
1
0
–1
–2
–3
–4
(d) ANOVA: NS
9
3
None
EnamAdv
EnamRing
8 21 28 25 12 16 7 18 1 4
DentAdv
57
DentSlght
313
EnamMid
None
5 9
EnamTrace
Error bars: +/– 2 S.E.
EnamTrace
13
12
11
10
9
8
7
6
5
4
3
2
1
0
–1
–2
–3
–4
(a) ANOVA: P = 0.1261 (all wear stages)
P = 0.0001 (wear ≤ DentAdv)
None
Functional age (years)
13
12
11
10
9
8
7
6
5
4
3
2
1
0
–1
–2
–3
–4
Attrition wear stage
Figure 3. ANOVA of relationship between functional age and attrition class for teeth in combination: (a) anterior primary (di1–dm1); (b)
posterior primary (dm2); (c) anterior permanent (I1–P3); (d) posterior permanent (P4–M3). Statistically significant differences are shown in the
figure; viz. MP is significantly older than the UP sample of the earlier stages of wear in the anterior primary teeth in combination, as well as
for anterior permanent teeth in combination. There are no differences between the two samples for the posterior primary and posterior
permanent teeth in combination. .: Middle Paleolithic; /: Upper Paleolithic.
Neanderthal anterior permanent teeth are wearing
faster or commence wear earlier. This could be
explained if Neanderthals erupted these teeth earlier;
but this would not explain their faster rate of wear
(Figure 2(c)). Thinner enamel could be a factor here
but, if so, would also have produced the same effect
among the posterior permanent teeth, for which there
is no evidence.
The same kind of analysis, avoiding type I error,
allows only one anterior permanent tooth to represent
the individual, but suffers from insufficient sample sizes
to show significant differences. Given the relatively
small sample for this tooth group and the possibility of
making a Type I error using teeth in combination, one
can only very tentatively conclude that anterior tooth
usage in late childhood/early adolescence may have
differed between the two samples such that Neanderthals wore their front teeth down more quickly at this
stage of life.
Posterior permanent teeth
The posterior permanent teeth wear at virtually the
same rate in both samples whether one is considering
the absolute reduction in crown substance (Figure
2(d)) or relative wear (Figure 3(d)). The latter is to be
expected because dental attributes of size, shape,
enamel hardness and thickness should be appropriate
for the growth rates and dietary abrasiveness of any
particular taxon. Virtually all studies of dental attrition
in traditional or prehistoric peoples concentrate on
adults (e.g. Richards, 1984; Dreier, 1994). An exception is the study by Molnar et al. (1983) of wear among
contemporary Australian aborigines eating a transitional diet. The first molar is reported to lose 0·5 mm
of enamel between the ages of 6 and 18 years. In the
results reported here for the same tooth, both MP and
UP samples lose 1·1 mm over the same period of time
(more than twice the rate).
686 M. Skinner
9
6
5
4
3
2
Error bars: +/– 2 S.E.
5
4
3
2
#2
4
2
#3
12 7
4 13
#4
#5
Primary tooth number
Figure 4. ANOVA of relationship between developmental age and
primary tooth position at a specific stage of dental attrition (slight
dentin exposure) chosen to maximize sample size. The MP sample of
primary teeth is significantly older at this attrition stage by an
average of 1·06 years. .: Middle Paleolithic; /: Upper Paleolithic.
Anterior primary teeth
The anterior primary teeth show the most obvious and
consistent difference across several analyses between
the two samples. Figure 2(a), showing reduction in
crown volume with increasing age, shows that the
anterior primary teeth appear to wear at the same rate
in both samples. There is no statistically significant
difference in age at death at successive wear stages
(ANOVA) between the MP and UP samples, considering all wear stages (Figure 3(a)). However, it is clear
that until the most severe wear stages (¢strong dentin
exposure) are attained, the mean functional age of the
MP sample is consistently older than the UP sample
(P=0·0001). It seems the MP sample lags consistently
behind the UP sample in wear of these teeth up to the
last wear stages. In other words, the MP children start
to wear their anterior primary teeth later, but at much
the same rate. This would suggest a delay in the onset
of wear in the MP sample (or, equally, earlier onset of
wear in the UP sample).
In order to test for real differences, with the effect of
statistical redundancy removed, the analysis shown in
Figure 5 compares the mean functional ages across
attrition classes (ANOVA); in this analysis each individual contributes only a single, randomly chosen,
tooth (increasing chances of a Type II error). Nevertheless, the ANOVA shows a statistically significant
difference (P=0·0498) such that the UP sample is
consistently younger at equivalent wear stages up to
‘‘dentin advanced’’ stage of wear.
The following section is designed to examine the
meaning of the difference in the timing of anterior
primary tooth wear between the MP and UP samples.
A potential criticism of previous analyses (e.g. Figure
3(a)) is that the wear stages are so broad (crude) that
0
2 1
1 2
3
4
57
74
DentAdv
9
DentSlght
#1
6
EnamAdv
7
EnamMid
n=6
EnamTrace
1
1
0
ANOVA: P = 0.0498
None
7
6
ANOVA: P < 0.0001
Functional age (years)
Developmental age (years)
8
7
Error bars: +/– 2 S.E.
Attrition class
Figure 5. ANOVA of the relationship between functional age and
successive attrition stages for MP and UP individuals for the anterior
primary teeth (di1–dm1). Overall, the MP sample is significantly
older throughout wear. .: Middle Paleolithic; /: Upper Paleolithic.
each stage could embrace statistically significant mean
functional age differences with no real difference between the two samples. This can be tested by using a
more precise estimate of attrition, viz. tooth height and
dentin exposure.
A regression of functional age against anterior
primary crown height (expressed as standardized deviations (z-scores) so as to remove the effects of innate
differences in crown height) is similar between the two
samples except that the intercept difference (value of Y
(age) when X (z-score)=0) is 0·9 years later in the MP
sample (Figure 6(a)). In other words, about halfway
through the process of reduction in crown height due
to attrition, the MP sample lags just under a year
behind. Restriction of wear to those teeth scored at
stages less than or equal to dentin advanced (for
comparability to Figure 3(a)) shows a similar relationship except that the intercept difference has increased
to 1·4 years (Figure 6(b)).
Figure 7(a) illustrates the regression of dentin
exposure against functional age. Unlike previous
analyses, this one, perforce, ignores earlier stages of
dental attrition. At equivalent ages, the MP sample
shows much less worn teeth. A more informative, if
less conventional, analysis is to regress functional age
against dentin exposure (Figure 7(b)). (This is not so
backward as it sounds since neither independent variable in these cases (7(a), (b)) is measured without error,
as strictly speaking it should be for linear regression.)
The value of this analysis is that it shows clearly that
the rate of wear is the same in both samples, but that
the MP sample lags behind the UP sample by about 1·1
years (difference in intercept values in Figure 7(b)). In
other words, the onset of attrition of the anterior
Dental Wear in Immature Late Pleistocene European Hominine 687
(a)
(a)
Percentage dentin exposure
–1
Functional age (years)
0
1
2
3
4
5
6
7
8
9
–2.5 –2 –1.5 –1 –0.5 0 0.5 1 1.5 2 2.5
Decreasing tooth height due to attrition (z-scores)
2
NegFunctAge = –3.24 – 1.005 * z-score; R = 0.308 (MP)
NegFunctAge = –2.343 – 1.299 * z-score; R2 = 0.405 (UP)
0
1
2
3
4
5
6
7
Functional age (years)
8
9
%Dent = –0.411 + 2.742 * FunctAge; R2 = 0.367 (MP)
%Dent = 1.974 + 3.985 * FunctAge; R2 = 0.48 (UP)
(b)
(b)
–1
9
0
8
1
2
3
4
5
6
7
Functional age (years)
Functional age (years)
60
55
50
45
40
35
30
25
20
15
10
5
7
6
5
4
3
2
8
1
9
–2.5 –2 –1.5 –1 –0.5 0 0.5 1 1.5 2 2.5
Decreasing tooth height due to attrition (z-scores)
0
5 10 15 20 25 30 35 40 45 50 55 60
Percentage dentin exposure
2
NegFunctAge = –3.028 – 0.827 * z-score; R = 0.245 (MP)
2
NegFunctAge = –1.65 – 0.396 * z-score; R = 0.105 (UP)
FunctAge = 2.365 + 0.134 * %Dent; R2 = 0.367 (MP)
FunctAge = 1.258 + 0.12 * %Dent; R2 = 0.48 (UP)
Figure 6. Linear regression of functional age of anterior primary
teeth (di1–dm1) against tooth height (expressed as a z-score). With
innate tooth height differences removed, the MP sample clearly lags
behind the UP sample by 1 year or more (difference in intercept value
of Y when X=0): (a) all wear stages; (b) wear stages up to and
including advanced dentin exposure (cf. Fig. 3(a)). ,: Middle
Paleolithic; +: Upper Paleolithic.
Figure 7. Linear regression using all attrition stages of (a) proportion (%) of crown surface showing dentin exposure against
functional age of anterior primary teeth in combination; (b) the
reverse. The UP sample has markedly more worn teeth; with the MP
lagging behind by about 1 year (cf. intercept values in (b)). These
differences probably reflect early onset of attrition in the UP sample.
,: Middle Paleolithic; +: Upper Paleolithic.
primary teeth appears to be delayed by about 1 year
on average in the MP sample, but they wear at indistinguishable rates thereafter.
All three means of assessing attrition (loss of crown
substance volume (Figure 2(a)) and height (Figure 6),
wear stage (Figure 3(a)), and dentin exposure (Figure
7), indicate close to a 1-year lag in the MP sample.
Thus the intercept values of Figures 6(a) & 7, which
provide the difference in Y (functional age) when X
(attrition)=0, are 0·9 and 1·1 years respectively. Similarly, the average difference in functional age with
attrition held constant (Figure 3(a)) is 0·9 years.
An alternative analysis, which includes the primary
second molar, is shown in Figure 4. Here, the relationship between individual age and primary tooth number
at just one attrition stage (slight dentin exposure for
which a reasonably large sample of teeth can be
amassed) is examined using ANOVA. The mean difference in functional age at slight dentin exposure
averaged across all five primary teeth is 1·06 years
(P<0·0001).
Another analysis based on teeth in combination is
designed to disclose the developmental age at which the
MP sample is demonstrably starting to lag behind
(Figure 8). This analysis requires that developmental
age be treated as a factor variable (age classes as shown
in the figure) with an ANOVA of percentage dentin
exposure at successive age classes compared between
the samples. The delay in dentin exposure in the
MP sample, while statistically significant (P=0·0097),
688 M. Skinner
Percentage dentin exposure
30
ANOVA: P = 0.0097
25
20
15
10
5
0
n = 9 16
< 2.4
13
23
2.5–3.4
29 24
> 3.5
Age class (years)
Figure 8. ANOVA of relationship between proportion (%) of dentin
exposure and individual age class in anterior primary teeth. By age
class ‘‘2·5–3·4 years’’ the MP sample has not increased dentin
exposure while the UP sample has; this probably reflects the earlier
onset of weaning in the latter sample. .: Middle Paleolithic; /:
Upper Paleolithic.
is only mildly expressed in infants aged less than
2·5 years, but is present by age 2·5–3·4 years and
maintained into older childhood. It would seem that
by age 2·5 years the UP sample is starting to wear its
anterior teeth quite significantly (already into dentine
exposure) whereas the MP sample is still no more worn
by age 2·5–3·4 years than it was in the previous age
class. A similar analysis compares average individual
age at the beginning of enamel attrition (trace wear).
Although samples are rather small (MP=three teeth,
UP=13 teeth), the difference in mean age is statistically
different: 2·0 years in the UP and 3·3 years in the
MP (t=2·787, P=0·0145). An obvious interpretation
to be placed on these results is that the introduction of
solid foods occurred 1 year earlier on average in the
UP sample, commencing by age 2 years.
Discussion
The essential findings of this study are that, overall,
statistically significant differences in attrition are few
and those that do exist are found in the anterior
dentition, most particularly the primary teeth which
are markedly more worn in the UP sample with age
held constant. The working assumption of this research
was that developmental rates did not differ between the
MP and UP hominines. Tompkins (1991) and
Trinkaus & Tompkins (1990) have reasoned that since
modern variation in eruption sequences and relative
tooth formation embraces those patterns seen in the
MP, there is no need to assume Neanderthals matured
at significantly different rates from anatomic moderns.
This view is not universal. Wolpoff (1979) has
suggested that third molar eruption in Krapina
Neanderthals occurred relatively early, at age 15 years
(assuming the rest of the dentition matured at modern
rates) while Stringer et al. (1990) reason that the
Gibraltar child died at age 3·9 years (as opposed to
c. 5·1 years reconstructed by Skinner & Sperber, 1982).
The Stringer study suggests that MP populations could
have matured in approximately 80% of the time
required for more anatomically modern samples. Substituting this maturation rate steepens the wear slope
for the MP sample, but has very little effect on the
previously disclosed patterns for all tooth groups,
except the anterior permanent teeth which wear even
faster in the MP sample.
Given the lack of significant differences between the
MP and UP tooth groups (except for the 1 year delay
in onset of wear on the anterior primary tooth group in
the MP sample), the simplest interpretation is that
neither the diet nor the overall growth rate differed
between the two samples during ontogeny. If one
assumed the validity of a faster growth rate for
Neanderthals, say 80%, then their diet must have
been proportionately more wearing (ie. 1·25#). If a
faster growth rate included the anterior primary teeth,
the difference is not enough to remove the effect of
the large lag in wear of these teeth in Neanderthals
(P changes from 0·0001 to 0·0274). A more economical
scenario is that the growth rates were the same in both
samples, and so was the abrasiveness of the diet.
The effect of thin enamel in the MP sample needs to
be considered. Clearly, this is not the explanation for
the relatively reduced wear of their primary dentition.
At all but the last two stages of wear for the anterior
primary dentition, the MP sample shows higher functional ages. In other words, it is taking the MP primary
tooth sample longer to attain these wear stages. Thin
enamel in the MP sample would tend to reduce the
observed difference, or, put another way, the UP
sample shows markedly more wear on the primary
dentition. As for wear of the permanent teeth, Figures
2(d) and 3(d) show no difference between the two
samples. Consideration of the thin enamel in the MP
sample indicates that the UP diet must have been as
hard as, if not harder than, that of the MP. The effect
of relative enamel thickness is less than that due to
other causes such as delay in onset of wear (anterior
primary teeth) or tooth usage (anterior permanent
teeth).
MP samples had larger primary and permanent
teeth on average. All else being equal, this would
tend to reduce their rate of wear. One implication of
this study is that ultimately the MP sample will run
out of anterior permanent tooth substance earlier;
thus root wear would commence a bit earlier in the
MP sample, which could conceivably lead to compromised health and/or productivity in older adults and,
in a demographic study, to their being over-aged
slightly. The difference seems mild. However, perhaps
the thinner enamel of MP teeth would exacerbate the
difference.
Dental Wear in Immature Late Pleistocene European Hominine 689
Table 3. Comparison between MP and UP samples of variance ratios for specimen age at successive attrition classes
¡6·2 years
Attrition
None
Enamel trace
Enamel mid
Enamel advanced
Dentin slight
Dentin advanced
Dentin strong
¢6·3 years
F-value
(MPn:UPn)
P-value
F-value
(MPn:UPn)
P-value
0·107
0·156
0·632
1·531
0·719
0·753
0·000
(5:19)
(11:25)
(10:17)
(18:29)
(25:34)
(8:18)
(2:6)
0·0218
0·0022
NS
NS
NS
NS
—
1·121
1·601
2·052
1·489
2·803
0·655
0·069
(9:16)
(11:31)
(14:27)
(11:38)
(22:34)
(12:18)
(6:19)
NS
NS
NS
NS
0·0117
NS
0·0039
Table 4. Comparison of interaction of dental attrition and functional age between the Early (EUP) and Late Upper Paleolithic (LUP) samples
based on teeth in combination (ANOVA)
Tooth group
Anterior primary
Posterior primary
Anterior permanent
Posterior permanent
Number
Culture+functional age
Culture+attr. class
Mean age
difference
EUP
LUP
F-value
P-value
F-value
P-value
1·94
0·83
"0·72
"1·25
11
17
26
23
53
34
53
43
10·534
4·012
0·0067
0·027
0·0019
0·0518
0·9349
0·8707
0·225
0·491
1·087
1·952
0·7990
0·7424
0·3701
0·1145
Variation
In order to examine Hayden’s proposition (1993) that
in the UP there was an increase in social heterogeneity,
the variance ratio for mean individual age with attrition class held constant is compared in Table 3. The
unit of analysis is the tooth, increasing the chance of
Type I error. At younger ages (c.<6·3 years old) there
are two instances out of seven in which the F-value
differs significantly between the MP and UP. In both
cases the UP sample is more variable. In the older age
class (>6·2 years), in one instance (out of seven), the
UP sample is more variable, and in one the MP sample
is more variable. The results provide mild support for
Hayden’s model that variation in the individual’s
experience with dietary attrition is increased in the UP.
Changes within the Upper Paleolithic
Justification was provided earlier for an analytical
distinction between the Middle and Upper Paleolithic.
Frayer (1984) demonstrated that during the Late
Pleistocene of Europe there is progressive decline in
dental areas with the Early Upper Paleolithic (EUP)
sample intermediate between the MP and Late Upper
Paleolithic (LUP). Consequently, it is appropriate to
compare the timing of dental attrition within the
Upper Paleolithic. The EUP sample consists of 77 teeth
from 15 individuals (i.e. about one-quarter of the total
UP sample) which is really too small for satisfactory
statistical analysis. Nevertheless, examination of the
result (Table 3) shows that the EUP sample, like the
MP sample, shows higher mean ages at successive wear
stages for the anterior and posterior primary dentition
(F=10·534, P=0·0019, and F=4·012, P=0·0518) and
no significant differences for the permanent dentition.
This analysis contains a heightened chance of Type I
error. It may be concluded that the apparent differences in dental attrition that exist between the MP and
UP arose, possibly, in the LUP.
Summary and Conclusions
A comparison of the interaction of tooth age and tooth
wear between immature cohorts of Middle and Upper
Paleolithic specimens shows that, apart from earlier
onset of wear of the anterior primary teeth in the UP
sample, and faster rate of wear for the anterior
permanent teeth of the MP sample, the other two tooth
groups do not differ significantly. With the possible
exception, then, of the anterior permanent teeth the
rates for the other tooth groups are indistinguishable.
Therefore, if one can assume dietary equivalence during the growing period, there is no evidence for their
maturing at different rates (contra Stringer et al., 1990).
If, on the other hand, the MP sample is deemed to
mature more quickly, then the UP diet must have been
less abrasive to an equivalent degree to maintain
similar wear rates. While soups may be an Upper
Paleolithic invention (see descriptions in Hadingham,
1979 and Pfeiffer, 1986), the advent of storage
(Soffer, 1985; Hayden, 1993) may have included the
smoking and drying of meats, rendering them more
abrasive. Consequently, it remains problematic
whether one can characterize the UP diet as generally
softer or harder than that of the MP.
Further discussion is limited to the anterior primary
teeth which commence wear significantly earlier in the
UP sample. Different interpretations can be placed on
690 M. Skinner
the significance of this observation. Supplementation
may occur as early as the first few days after birth, as
among the Apsaroke Eskimo who give the baby a piece
of partially dried blubber to suck until the mother’s
flow of milk is adequate (Gidley, 1976); and can
continue for years as among the !Kung who introduce
adult foods at 6 months or earlier but suckle very
frequently for 3–4 years (Gaulin & Konner, 1977).
Suckling frequency can override the potential effects of
dietary supplementation on ovulation (Gaulin &
Konner, 1977; Konner & Worthman, 1980). Thus, the
advent of dental attrition does not necessarily signal
weaning. For example, plants and pieces of meat are
given to !Kung infants as pacifiers and objects to
mouth (ibid.). Similarly, on the Pacific Northwest some
nursing infants were also given fat meat on a string
tied to their big toe (to prevent choking)
(McKechnie, 1972). Bullington (1987) found that age
at onset of dental microwear did not differ among three
prehistoric groups thought likely to wean at different
ages due to differences in group diet. Clearly, the
advent of dental attrition very likely signals the
introduction of solid foods and initiation of a weaning
process which could be abrupt or gradual.
A major determinant of fertility in noncontracepting mothers is suckling frequency
(McNeilly, 1977; Jain & Bongaarts, 1981). Human
reproductive ecology is a complex phenomenon of
which suckling frequency is only one of many important variables that influence ovarian function (Ellison,
1990). The contraceptive effect of breast-feeding is
debatable. Clearly, when breast-feeding continues
for more than a few months menses (and ovulation)
returns. Indeed weaning is often precipitated by a new
pregnancy (Santow, 1987). Nevertheless, dietary
supplementation of infants has been shown to have
important effects on suckling frequency, prolactin
levels and return of ovulation. Prolactin is secreted
within minutes of nipple mechanical stimulation; this
hormone in turn inhibits gonadal hormones which
control ovulation (Konner & Worthman, 1980; Wood,
1994). Use of a pacifier or the introduction of water or
solid foods can diminish prolactin levels (Jeliffe &
Jeliffe, 1979; Zeller, 1987). In other words, while supplementation need not herald weaning, it will probably
hasten the process.
This study echoes earlier work undertaken by
Brennan (1991) which concluded that UP infants may
have been weaned relatively early. We can ask how
likely is this conclusion. A recent study of enamel
hypoplasia among the same specimens reported here
found that the UP sample showed significantly more
developmental stress in infancy (Skinner, 1996). This
was tentatively attributed to combined effects of social
crowding, vitamin A malnutrition and infection. Early
dietary supplementation could contribute to this
problem through the oral introduction of pathogens.
This begs the question, however, of why UP mothers
would have felt able to, or had to, wean a year earlier
than was the case in the MP. Conceivably, infant
supplementation led to renewed pregnancies which in
turn would favour accelerated weaning. There is no
particular advantage in traditional societies to introduce early weaning. In fact the disadvantages are well
known; for example, loss of passive immunity, loss of
lactational amennorrhoea, loss of maternal–infant
bonding, and dietary stress for the weaned infant.
Rather, it seems likely that under traditional conditions the ideal would be prolonged lactation and
that a departure from this should be viewed as
unintentional and possibly deleterious.
Large mammal hunting, accompanied by an unknown contribution from plant foods, prevailed
throughout the Late Pleistocene (Gamble, 1986).
Climatic conditions probably limited the growing
season to about 3 months. Notably, climatic conditions
deteriorated during Oxygen Isotope Stage 2; the glacial
maximum occurred in the middle Upper Paleolithic
(Straus, 1995). If weaning at these times was tied to
seasonality, then inter-population differences at age of
weaning would be separated by yearly intervals, as
suggested by this study. Evidence of a delay in weaning
among Neanderthals, in the form of a 1-year lag in
dental attrition, may signal longer average birth
spacing in comparison to the UP sample. Birth spacing
is the most important variable affecting fertility
(Scott & Johnston, 1985). Profound demographic
and social changes ensue from reduced birth spacing
(MacCormack,
1982;
Constandse-Westermann,
Newell & Meiklejohn, 1984). Thus, the finding of
relatively early onset dental attrition provides a plausible mechanism for population increase in the UP.
Acknowledgements
The research for this article was completed during my
tenure as Senior Visiting Research Fellow, Department
of Archaeology and Prehistory, University of Sheffield.
I would like to express gratitude for assistance towards
completion of this project to the persons and institutions listed below: Chris Stringer (British Museum
Natural History), Alan Bilsborough, Bernard Denston
(Duckworth Laboratory, University of Cambridge),
M. Leguebe (Institut Royal des Sciences Naturelles de
Belgique), G. Ubaghs (Laboratoire de Paléontologie
Animale, Université de Liège), Anne-Marie Tiller,
Jean-Jaques Hublin, B. Vandermeersch (Laboratoire
de Paléontologie des Vertébrés et de Paléontologie
Humaine, Université Pierre et Marie Curie), Yves
Coppens (Laboratoire d’Anthropologie, Musée de
l’Homme), Henri Delporte (Musée des Antiquités
Nationales, Chateau de Saint Germain-en-Laye), Prof
Piveteau (Institut de Paléontologie), Jean-Louis Heim
(Institute de Paléontologie Humaine), Prof de Bonis
(Laboratoire de Paléontologie des Vertébrés, Université de Poitiers), A. Roussot, G. Chauvin (Musée
d’Aquitaine), Mme Prudhomme (Musée d’Histoire
Dental Wear in Immature Late Pleistocene European Hominine 691
Naturelle, Bordeaux), Jean Guichard, Andre
Morala (Musée National de Préhistoire, Les Eyzies),
Mme Edmée Lauder (Musée d’Histoire Naturelle,
Montauban), Michel Philippe (Museum d’Histoire
Naturelle, Lyon), Jean-Francois Bussiere (Musée
d’Anthropologie Préhistorique, Monaco), Jakov
Radovcic (Hrvatski Prirodoslovni Muzej, Zagreb),
Mirko Malez (Zavod za Paleotologiju i Geologiju
Kvartara).
Particularly I am grateful to the often unnamed
assistant curators who helped me cope with installing
the radiographic apparatus and tolerated my broken
French. Social Sciences and Humanities Research
Council of Canada provided funding. Student assistants were Anja Streich, Arne Carlson and Brian
Strongman. Jack Nance, Richard Lazenby and
Andrew Chamberlain gave me valuable advice. Jean
McKendry, my wife, was my very able research assistant and companion throughout the data collection and
analytical phases. Thank you.
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Dental Wear in Immature Late Pleistocene European Hominine 693
Appendix
List of specimens with attributes of age, height, and attrition
Group
SpecName
Spec
IndAge
Tooth*
FunctAge
EUP*
LUP*
Patau 26242
Ranis
1
2
1·0
1·0
uldi1
lldm1
lrdm1
lrdm1
lrdm1
lrdm2
lldi1
lldm2
lldm1
urdm2
urdm1
uldm1
urdc
uldi1
lrdi1
lrdi2
lrdc
urdi2
lldc
uldc
lldi2
urdi1
uldi2
lldi2
uldm1
uldm2
lldc
lrdi1
lldi1
lrdi2
lldm2
lldm1
lrdc
urdm1
lrdm1
lldm1
lldm2
uldm1
uldm2
lldm2
lrdm1
lrdm2
lldm2
urdm1
uldm1
uldm2
lrdm1
lrdm2
urdm2
lldm1
uldi2
urdm2
lrdm1
uldm1
urdm1
lrdm2
uldm2
lldm2
uldc
urdi2
uldi1
urdi1
0·20
"0·30
"0·30
0·10
0·10
"0·80
0·80
"0·80
0·10
"0·90
0·20
0·20
"0·10
0·60
0·80
0·40
"0·20
0·60
"0·20
"0·10
0·40
0·60
0·60
0·60
0·40
"0·70
0·00
1·00
1·00
0·60
"0·60
0·30
0·00
0·40
0·30
0·50
"0·40
0·80
"0·30
"0·20
1·10
0·20
0·60
1·60
1·60
0·50
1·50
0·60
0·50
1·50
2·00
0·50
1·50
1·60
1·60
0·60
0·50
0·60
1·30
2·00
2·00
2·00
LUP
LUP
LUP
Bruniqel545
EnfantsYoun
Figuier
3
4
5
1·4
1·4
1·6
LUP
Fées
6
1·8
LUP
Laugerie-5
7
2·0
LUP
LUP
LaugeriTooth
Isturitz-7
8
9
2·0
2·4
LUP
Brun540/541
10
2·8
LUP
Rochereil
11
2·8
TthHt
AttScor
AttClass
%Dent
6·30
4
1
1
0
1
0
5
0
0
0
1
1
0
4
4
4
0
1
0
0
3
4
1
4
4
0
3
5
5
4
1
4
3
4
4
2
1
1
0
2
3
1
2
3
3
2
3
2
2
3
2
0
2
0
0
1
0
1
0
2
5
4
DentSlght
EnamTrace
EnamTrace
None
EnamTrace
None
DentAdv
None
None
None
EnamTrace
EnamTrace
None
DentSlght
DentSlght
DentSlght
None
EnamTrace
None
None
EnamAdv
DentSlght
EnamTrace
DentSlght
DentSlght
None
EnamAdv
DentAdv
DentAdv
DentSlght
EnamTrace
DentSlght
EnamAdv
DentSlght
DentSlght
EnamMid
EnamTrace
EnamTrace
None
EnamMid
EnamAdv
EnamTrace
EnamMid
EnamAdv
EnamAdv
EnamMid
EnamAdv
EnamMid
EnamMid
EnamAdv
EnamMid
None
EnamMid
None
None
EnamTrace
None
EnamTrace
None
EnamMid
DentAdv
DentSlght
4·30
-
5·15
-
4·90
5·60
5·25
-
5·65
5·95
7·55
5·95
5·00
6·60
7·65
6·15
7·75
7·85
6·00
6·20
5·85
6·30
4·95
5·90
7·60
5·10
4·55
6·25
5·95
5·25
7·55
5·00
5·10
4·85
5·50
4·95
6·20
5·00
5·20
5·80
5·60
5·45
4·95
-
4·70
5·65
6·50
4·85
6·10
6·95
6·05
6·65
6·55
5·95
-
6·45
7·75
6·00
5·65
6·55
-
14·53
-
3·39
14·23
6·91
-
2·22
3·78
-
8·00
2·27
-
15·38
18·31
7·33
-
1·66
-
1·47
1·11
-
1·79
-
11·46
7·42
694 M. Skinner
Appendix continued
Group
LUP
LUP
SpecName
Spec
IndAge
FDI
Bruniquel 25
12
2·8
lldc
urdi1
urdi2
uldi1
uldi2
lrdm2
lrdm1
lldm2
lldm1
urdm2
urdm1
lrdm2
lldm2
uldm2
uldi1
uldm1
lldm1
lrdm1
lrdc
lrdi2
lrdi1
lldc
urdi1
uldi2
uldc
urdc
lrdm1
lrdm2
urdm1
urdm1
uldm1
uldm2
lldm1
lldm2
lrdm1
lrdm2
lrdi2
lrdi1
lldc
lldi2
lldi1
uldc
uldi2
uldi1
urdc
urdi2
urdi1
lrdm2
lrdm1
lrdm1
lldm2
uldm2
lldm1
lrdm2
uldm1
lrdm1
lrdm2
urdm1
urdm2
lldm1
lldm2
lrdm2
lldm2
uldm2
urM1
ulM1
urdm1
urdm2
llM1
lldc
uldm1
lldm2
lldm1
EnfantsOldr
13
3·0
EUP
Baoussé
14
3·2
LUP
LUP
Plac61401a
Madéleine
15
18
3·3
3·4
EUP
LUP
Rouset
Laugr2/4/6
19
20
3·6
3·8
EUP
Magrite2878
21
4·2
EUP
Patau26236
23
4·7
LUP
Plac61401b
24
4·8
LUP
Isturitz-6
25
5·0
LUP
St. Germ5–7
26
5·3
FunctAge
1·20
2·00
2·00
2·00
2·00
0·60
1·50
0·60
1·50
0·70
1·80
0·80
0·80
0·70
2·20
1·80
1·70
1·70
1·40
2·00
2·40
1·40
2·20
2·20
1·50
1·50
1·90
1·00
2·10
2·20
2·20
1·10
2·10
1·20
2·10
1·20
2·40
2·80
1·80
2·40
2·80
1·90
2·60
2·60
1·90
2·60
2·60
1·40
2·30
2·50
1·60
1·50
2·50
1·60
2·60
2·90
2·00
3·50
2·40
3·50
2·60
2·80
2·80
3·00
"1·10
"1·10
4·10
3·00
"1·00
3·70
4·10
3·10
4·00
TthHt
AttScor
AttClass
7·25
5·35
5·75
5·15
6·40
1
4
3
4
3
1
1
1
2
1
2
1
1
1
5
1
1
1
3
4
5
3
5
4
4
4
3
2
2
3
3
2
3
2
3
2
4
5
3
4
5
3
4
6
3
4
6
3
4
5
3
3
5
3
5
4
4
4
3
6
4
4
4
4
0
0
5
4
1
5
5
4
5
EnamTrace
DentSlght
EnamAdv
DentSlght
EnamAdv
EnamTrace
EnamTrace
EnamTrace
EnamMid
EnamTrace
EnamMid
EnamTrace
EnamTrace
EnamTrace
DentAdv
EnamTrace
EnamTrace
EnamTrace
EnamAdv
DentSlght
DentAdv
EnamAdv
DentAdv
DentSlght
DentSlght
DentSlght
EnamAdv
EnamMid
EnamMid
EnamAdv
EnamAdv
EnamMid
EnamAdv
EnamMid
EnamAdv
EnamMid
DentSlght
DentAdv
EnamAdv
DentSlght
DentAdv
EnamAdv
DentSlght
DentStrng
EnamAdv
DentSlght
DentStrng
EnamAdv
DentSlght
DentAdv
EnamAdv
EnamAdv
DentAdv
EnamAdv
DentAdv
DentSlght
DentSlght
DentSlght
EnamAdv
DentStrng
DentSlght
DentSlght
DentSlght
DentSlght
None
None
DentAdv
DentSlght
EnamTrace
DentAdv
DentAdv
DentSlght
DentAdv
-
5·35
5·70
5·55
5·55
5·45
6·05
5·35
5·80
5·30
4·90
7·35
6·00
4·35
7·40
5·35
5·55
7·40
7·00
5·35
6·55
5·45
6·15
6·15
6·60
5·45
6·15
4·35
-
6·20
4·60
7·65
5·80
4·30
6·90
6·05
5·70
7·30
6·50
5·65
-
4·45
5·50
5·90
4·45
6·80
5·15
4·75
5·60
5·55
6·65
4·20
5·80
5·05
5·15
5·40
7·25
7·65
5·15
5·75
6·70
6·75
4·95
5·10
-
%Dent
-
9·47
-
15·32
-
16·63
-
4·90
16·67
-
15·25
8·04
2·04
2·32
-
0·81
-
0·77
-
4·60
28·29
-
7·90
23·92
2·37
5·27
24·15
1·75
4·82
22·75
-
10·93
0·43
-
12·75
0·67
7·11
3·02
2·07
1·04
0·27
12·75
1·10
0·86
0·70
0·75
-
10·85
1·56
-
13·08
13·09
1·13
4·61
Dental Wear in Immature Late Pleistocene European Hominine 695
Appendix continued
Group
SpecName
Spec
IndAge
FDI
EUP
Solutré
27
5·4
LUP
EUP
LUP
Mas D’Azil
Fontéchvade
Laugerie-1
28
29
30
5·8
5·8
5·8
LUP
Laugerie-3
31
5·8
EUP
Roches
32
6·0
lldm1
lrdm2
lldm2
uldm2
lrdm2
lldm1
lldm2
lrdm1
lrdm2
lrM1
lrdm2
lrdm2
lrdm1
lldm2
llM1
lldm2
lrI2
lrI1
lldc
lrdc
lrdm2
llI1
llI2
lldm1
llM1
lrM1
lldm2
lrdm1
lrdm2
lrdm1
uldm1
uldm2
urdm2
urM1
ulM1
lrdm1
lrdm2
lrM1
lldm2
llM1
lldm1
lrdm2
lrM1
lrdc
lrI1
llI2
lldc
lrI2
lldm1
lldm2
lrdm1
lrdm2
lrM1
llM1
llI1
lrI1
lrI2
llI2
urI2
urdm2
urM1
lrM1
urdm1
lrdm1
lrdm2
uldm2
urdm2
lrdm2
lrdm1
lrI2
urdm1
urM1
LUP
Isturitz-5
33
6·2
EUP
LaQuina25
34
6·3
LUP
LUP
Bruniqel538
StGerm-3
35
36
7·1
7·2
LUP
Plac56029
37
7·5
EUP
Magrite2426
38
7·6
EUP
Vindija
39
7·6
EUP
Miesslingtal
40
8·0
LUP
LUP
LUP
LUP
LUP
StGerm-4
Laugri15107
Brun536/537
Laugerie-31
Plac61401c
41
42
43
44
45
8·3
8·9
9·5
9·8
10·0
FunctAge
4·10
3·20
3·20
3·50
3·60
4·50
3·60
4·50
3·60
"0·30
3·80
4·00
4·90
4·10
0·00
4·90
0·20
1·00
5·60
5·60
5·00
1·00
0·20
5·90
0·90
0·90
5·00
5·90
5·30
6·20
6·40
5·30
5·30
1·20
1·20
6·30
5·40
1·30
5·80
1·70
6·70
5·80
1·70
6·40
1·80
1·00
6·40
1·00
7·00
6·10
7·00
6·10
2·00
2·00
2·10
2·10
1·30
1·30
0·90
7·20
3·10
3·20
8·30
8·20
7·30
7·50
7·70
7·80
8·70
3·00
8·80
3·60
TthHt
AttScor
AttClass
4·80
5·25
5·70
5·60
5·90
3·10
4·55
4·20
5·25
3
3
3
4
3
6
5
6
5
0
2
4
6
3
1
5
1
2
6
6
5
2
1
6
3
2
5
6
5
6
6
4
5
2
2
6
5
1
4
2
6
4
2
6
1
0
6
0
7
6
6
5
3
3
4
4
3
3
2
5
1
1
6
7
5
4
6
7
7
4
7
3
EnamAdv
EnamAdv
EnamAdv
DentSlght
EnamAdv
DentRing
DentAdv
DentStrng
DentAdv
None
EnamMid
DentSlght
DentStrng
EnamAdv
EnamTrace
DentAdv
EnamTrace
EnamMid
DentStrng
DentStrng
DentAdv
EnamMid
EnamTrace
DentStrng
EnamAdv
EnamMid
DentAdv
DentStrng
DentAdv
DentStrng
DentStrng
DentSlght
DentAdv
EnamMid
EnamMid
DentStrng
DentAdv
EnamTrace
DentSlght
EnamMid
DentStrng
DentSlght
EnamMid
DentStrng
EnamTrace
None
DentStrng
None
EnamRing
DentStrng
DentStrng
DentAdv
EnamAdv
EnamAdv
DentSlght
DentSlght
EnamAdv
EnamAdv
EnamMid
DentAdv
EnamTrace
EnamTrace
DentStrng
EnamRing
DentAdv
DentSlght
DentStrng
EnamRing
EnamRing
DentSlght
EnamRing
EnamAdv
-
4·90
4·20
5·85
7·70
-
10·35
10·25
4·40
3·60
4·40
10·05
10·35
4·00
6·85
6·70
5·25
4·00
4·55
3·25
4·90
5·65
5·85
8·15
7·85
4·10
4·90
7·00
4·90
6·00
3·45
4·75
6·75
3·45
7·60
-
3·40
-
3·50
4·55
4·15
4·55
5·95
5·95
8·05
8·30
9·20
8·65
9·95
-
5·55
4·35
4·85
4·15
9·15
4·80
6·45
%Dent
-
0·06
0·67
0·96
0·22
35·56
9·15
16·02
13·27
-
1·68
15·90
2·42
-
30·40
46·49
19·43
-
22·54
-
10·56
32·30
17·02
37·51
22·94
4·39
5·14
-
14·40
14·44
-
54·51
32·02
19·53
16·51
-
0·21
8·68
4·54
2·33
0·96
-
8·94
22·08
75·61
49·01
3·00
15·70
-
696 M. Skinner
Appendix continued
Group
EUP
LUP
LUP
LUP
EUP
EUP
LUP
SpecName
Spec
IndAge
FDI
Rois
46
11·1
lrC
llP1
llM1
lldm2
lrdm2
llC
lrM1
lrP1
urI1
urI2
llM1
ulM1
llP1
llI2
llC
ulM2
lrM1
ulP1
lrM2
lrP1
llM2
llI1
lldm2
uldm2
ulI1
lrdm2
lrI2
lrC
ulI2
lrI1
lrdm2
lrM1
ulC
ulC
urI1
ulP1
ulP2
llM2
lrM2
llI1
lrI2
urC
urI1
urI2
lrC
lrI1
uldm2
llC
urP1
llI2
ulP1
urM2
lrP2
llP2
ulM2
ulM1
urM1
llP1
lrM1
ulI1
urP2
ulC
ulI2
llM1
lrM2
lrP1
llM2
llP1
lrdm2
llP2
lrP1
lrM1
LaugeriAdol
Morin
Farincourt
Parpallo
GrEnfants-6
RocDeSers
47
48
49
50
51
52
11·2
11·6
11·8
13·3
13·8
14·4
FunctAge
TthHt
AttScor
AttClass
1·30
1·10
4·80
8·90
8·90
1·30
4·80
1·10
4·40
3·20
4·90
4·80
1·20
4·20
1·40
"1·40
4·90
1·30
"0·60
1·20
"0·60
5·00
9·00
8·90
4·40
9·00
4·20
1·40
3·20
5·00
9·40
5·30
0·40
1·80
6·40
3·30
2·40
1·40
1·40
7·60
6·80
2·40
7·00
5·80
4·00
7·60
11·50
4·00
3·90
6·80
3·90
1·20
3·20
3·20
1·20
7·40
7·40
3·80
7·50
7·00
3·00
2·40
5·80
7·50
2·00
3·80
2·00
4·40
12·20
3·80
4·40
8·10
12·65
5·15
2
1
3
6
6
2
3
0
4
2
4
4
0
3
0
0
4
2
1
0
1
4
7
6
3
7
3
0
2
4
5
2
0
2
4
2
2
1
1
4
4
2
4
3
2
4
6
1
1
4
1
0
1
0
0
4
3
0
3
4
0
0
3
4
1
1
2
2
6
1
2
4
EnamMid
EnamTrace
EnamAdv
DentStrng
DentStrng
EnamMid
EnamAdv
None
DentSlght
EnamMid
DentSlght
DentSlght
None
EnamAdv
None
None
DentSlght
EnamMid
EnamTrace
None
EnamTrace
DentSlght
EnamRing
DentStrng
EnamAdv
EnamRing
EnamAdv
None
EnamMid
DentSlght
DentAdv
EnamMid
None
EnamMid
DentSlght
EnamMid
EnamMid
EnamTrace
EnamTrace
DentSlght
DentSlght
EnamMid
DentSlght
EnamAdv
EnamMid
DentSlght
DentStrng
EnamTrace
EnamTrace
DentSlght
EnamTrace
None
EnamTrace
None
None
DentSlght
EnamAdv
None
EnamAdv
DentSlght
None
None
EnamAdv
DentSlght
EnamTrace
EnamTrace
EnamMid
EnamMid
DentStrng
EnamTrace
EnamMid
DentSlght
-
3·70
3·95
13·20
7·80
-
11·90
9·85
6·50
7·20
3·70
9·65
12·85
8·00
-
8·30
7·05
-
7·25
9·60
4·45
4·25
11·80
3·75
10·20
12·95
10·00
9·40
-
11·65
-
8·35
9·85
11·80
11·25
-
8·80
4·70
12·65
9·20
9·20
-
6·10
-
6·90
6·60
7·05
-
11·05
-
11·40
-
6·55
4·20
5·05
5·30
4·50
7·05
%Dent
-
54·51
-
0·28
-
1·77
-
1·57
1·05
-
2·17
-
1·36
-
2·72
70·61
45·44
1·41
73·63
1·77
-
3·62
-
14·08
5·10
-
7·05
-
11·81
42·83
-
7·57
-
1·37
0·39
-
1·34
9·44
-
3·44
-
37·74
-
1·51
Dental Wear in Immature Late Pleistocene European Hominine 697
Appendix continued
Group
LUP
LUP
LUP
LUP
LUP
SpecName
Cap Blanc
Lachaud-5
Plac61397
LaugeriNoNo
Lachaud-3
Spec
53
54
55
56
57
IndAge
15·6
15·8
18·0
18·0
18·6
MP*
Chatoneuf1
60
2·0
MP
PechdeL’Azé
61
2·4
FDI
lrM2
lrC
llC
lrI1
llM1
llI1
lrI2
ulI2
ulI1
lrI1
lrC
lrI2
ulI1
urI2
urC
llP1
ulM2
llC
urM2
lrM2
llP2
llM2
ulP2
ulP1
lrP2
lrM1
llM1
llI2
ulM1
ulC
llI1
lrP1
urP1
urP2
urM1
llP1
lrI2
lrI1
llC
llP2
lrP1
lrM1
llM1
llM2
llI1
lrC
llI2
lrM2
lrP2
llM2
llM1
llP2
llC
lrM1
lrM3
lrM2
lrM1
llM2
llM1
lrdc
lrdm1
urdi1
urdm2
urdm1
urdc
lrdm1
lrdc
lldi1
lrdi2
uldi1
uldc
uldm2
FunctAge
2·60
4·60
4·60
8·20
8·10
8·20
7·40
7·60
8·80
9·40
5·80
8·60
8·80
7·60
4·20
5·60
3·00
5·80
3·00
3·80
5·00
3·80
4·80
5·70
5·00
9·30
9·30
8·60
9·20
4·20
9·40
5·60
5·70
4·80
9·20
5·80
8·80
9·60
6·00
5·20
5·80
9·50
9·50
4·00
9·60
6·00
8·80
4·00
5·20
6·20
11·70
7·40
8·20
11·70
"0·40
6·80
12·30
6·80
12·30
0·40
0·70
1·60
0·10
1·20
0·90
1·10
0·80
1·80
1·40
1·60
0·90
0·10
TthHt
-
12·65
12·65
9·05
6·10
9·00
10·40
-
4·60
7·60
6·95
10·15
6·55
4·65
6·30
5·30
6·90
7·00
9·65
8·75
6·60
5·95
-
6·20
4·70
11·05
6·35
10·70
6·15
5·55
6·05
5·65
8·00
6·35
6·00
6·15
6·05
7·95
5·00
6·50
5·20
6·95
6·30
7·60
6·05
AttScor
AttClass
2
3
2
4
4
3
4
3
4
4
1
1
4
3
4
1
2
1
2
1
1
1
3
3
1
4
4
1
3
1
4
1
3
3
3
3
5
5
3
3
3
5
5
3
5
3
4
3
3
1
3
2
3
5
0
3
5
3
5
0
3
4
1
4
0
3
2
5
3
4
0
2
EnamMid
EnamAdv
EnamMid
DentSlght
DentSlght
EnamAdv
DentSlght
EnamAdv
DentSlght
DentSlght
EnamTrace
EnamTrace
DentSlght
EnamAdv
DentSlght
EnamTrace
EnamMid
EnamTrace
EnamMid
EnamTrace
EnamTrace
EnamTrace
EnamAdv
EnamAdv
EnamTrace
DentSlght
DentSlght
EnamTrace
EnamAdv
EnamTrace
DentSlght
EnamTrace
EnamAdv
EnamAdv
EnamAdv
EnamAdv
DentAdv
DentAdv
EnamAdv
EnamAdv
EnamAdv
DentAdv
DentAdv
EnamAdv
DentAdv
EnamAdv
DentSlght
EnamAdv
EnamAdv
EnamTrace
EnamAdv
EnamMid
EnamAdv
DentAdv
None
EnamAdv
DentAdv
EnamAdv
DentAdv
None
EnamAdv
DentSlght
EnamTrace
DentSlght
None
EnamAdv
EnamMid
DentAdv
EnamAdv
DentSlght
None
EnamMid
%Dent
-
8·42
1·46
2·11
3·69
-
7·91
6·44
-
2·52
3·79
-
5·50
1·67
13·11
-
0·42
-
1·58
5·67
-
2·84
-
3·21
-
1·41
10·58
-
1·61
-
0·74
-
11·11
1·88
4·95
-
698 M. Skinner
Appendix continued
Group
MP
MP
MP
SpecName
Krapina 68
Archi
RocDeMarsal
Spec
62
63
64
IndAge
2·7
3·0
3·2
MP
La Chaise 13
65
3·2
MP
Molare
94
3·6
MP
Engis
66
3·6
MP
La Chaise 14
67
4·3
MP
Gibraltar 2
68
5·0
MP
MP
Krapina 67
KrapMand A
69
70
5·3
5·4
MP
Chatoneuf 2
71
5·7
MP
MP
Krapina 11
KrapMax B
72
73
6·3
6·5
FDI
FunctAge
TthHt
AttScor
AttClass
%Dent
lrdi1
lrdm2
uldm1
lrdm2
lrdm2
lrdm1
lldm1
lldm2
lldc
lldi1
uldc
uldi2
uldi1
urdc
urdi2
urdi1
lrdc
urdm2
uldm1
uldm2
lldm1
lldm2
urdm1
lrdm2
lrdm1
lrdi2
lldi2
lldc
lrdi1
lrdm2
lrdm1
lldm2
lldm1
lldm2
lrdm2
lrdm1
lrdm1
urdm1
urdm2
lrdm2
lldm1
lrdm2
lrdi2
lrdi1
lldm1
urdm2
lldm2
urdm1
lldm1
lldm1
lldc
uldi1
uldi2
urdi2
urdi1
uldc
lrdi1
lldm2
uldm1
uldm2
lldm1
lrdm1
urdm1
urdc
lrdm2
urdm2
lrdi2
urdi1
urdm1
urdi2
urdc
urdm2
1·80
0·20
1·20
0·50
0·80
1·70
1·70
0·80
1·40
2·60
1·70
2·40
2·40
1·70
2·40
2·40
1·60
0·90
2·00
0·90
1·90
1·00
2·00
1·00
1·90
2·20
2·20
1·60
2·60
1·00
1·90
1·00
1·90
1·40
1·40
2·30
2·30
2·40
1·30
1·40
3·00
2·10
3·30
3·70
3·70
2·70
2·80
3·80
4·00
4·10
3·80
4·90
4·90
4·90
4·90
4·20
5·10
3·50
4·50
3·40
4·40
4·40
4·50
4·20
3·50
3·40
5·30
5·70
5·30
5·70
5·00
4·20
5·05
5·65
6·45
5
2
3
1
1
2
2
1
0
5
2
4
4
1
3
4
1
1
4
1
4
2
3
2
3
4
4
0
5
3
4
3
5
1
1
1
4
4
3
3
2
2
4
5
4
3
3
4
5
6
6
4
4
4
4
4
5
3
4
3
4
3
4
4
3
3
5
6
4
6
4
3
DentAdv
EnamMid
EnamAdv
EnamTrace
EnamTrace
EnamMid
EnamMid
EnamTrace
None
DentAdv
EnamMid
DentSlght
DentSlght
EnamTrace
EnamAdv
DentSlght
EnamTrace
EnamTrace
DentSlght
EnamTrace
DentSlght
EnamMid
EnamAdv
EnamMid
EnamAdv
DentSlght
DentSlght
None
DentAdv
EnamAdv
DentSlght
EnamAdv
DentAdv
EnamTrace
EnamTrace
EnamTrace
DentSlght
DentSlght
EnamAdv
EnamAdv
EnamMid
EnamMid
DentSlght
DentAdv
DentSlght
EnamAdv
EnamAdv
DentSlght
DentAdv
DentStrng
DentStrng
DentSlght
DentSlght
DentSlght
DentSlght
DentSlght
DentAdv
EnamAdv
DentSlght
EnamAdv
DentSlght
EnamAdv
DentSlght
DentSlght
EnamAdv
EnamAdv
DentAdv
DentStrng
DentSlght
DentStrng
DentSlght
EnamAdv
10·58
0·06
1·09
-
5·40
7·55
5·85
6·15
7·40
6·30
6·85
7·85
6·45
6·55
6·15
5·60
5·80
6·25
5·60
5·50
6·50
6·60
7·80
4·95
5·75
5·75
5·95
5·40
6·40
6·60
6·20
5·20
5·90
5·75
6·15
5·20
6·15
6·50
4·35
5·15
5·80
6·00
5·75
4·95
-
5·70
5·15
5·40
5·45
5·35
6·30
4·85
5·45
5·40
5·50
4·50
4·90
5·65
6·80
5·50
5·35
5·70
4·70
-
4·65
6·45
-
-
13·26
-
8·21
5·79
-
1·75
2·55
-
1·62
-
1·38
-
1·18
-
0·46
6·46
6·04
-
11·99
0·23
1·51
0·27
-
1·23
8·38
27·78
9·45
2·76
2·57
13·32
-
16·85
15·36
14·60
11·22
12·92
11·61
7·38
18·80
1·27
9·06
1·26
7·09
1·22
5·06
3·62
1·57
1·64
-
15·55
3·81
23·25
3·68
0·29
Dental Wear in Immature Late Pleistocene European Hominine 699
Appendix continued
Group
MP
MP
MP
MP
MP
MP
MP
MP
MP
MP
SpecName
La Quina 18
Krapina 63
KrMdBMxA
Le Fate 2
René Simard
KrMdCMxC
Ehringsdorf4
Krapina MdD
Malarnaud
Krapina MdE
Spec
75
77
79
80
81
82
84
85
86
87
IndAge
7·0
7·2
7·8
8·0
8·8
10·4
12·2
12·4
13·2
13·5
FDI
uldm1
uldm2
urM1
ulM1
uldc
uldc
ulM1
llM1
urdm1
lrdm1
lldm2
lldm1
lrdm2
uldm2
uldm1
urdm2
lrI1
llI1
ulI1
urdc
uldm1
urdm2
ulM1
urM1
uldm2
urdm1
lldm2
lrI1
lrI2
lrdc
urdi2
uldm2
urdm2
uldm1
llM1
ulM1
urM1
llM1
lrI1
urI2
urI1
uldi2
llI1
ulI1
ulM1
urI2
ulM2
llC
llI2
urI1
uldm2
lrdm2
lrM2
lldm2
llI1
ulC
lrM1
lrC
lrP1
llI1
llI2
lrI2
lrI1
llP2
llM1
llI2
llP1
llM2
llC
lrM1
llM1
llM2
FunctAge
TthHt
AttScor
AttClass
%Dent
5·30
4·20
0·10
0·10
5·00
5·20
0·30
0·40
5·50
5·40
4·50
5·40
4·50
4·40
5·50
4·40
0·50
0·50
"0·10
5·20
5·80
4·70
0·60
0·60
4·70
5·80
5·00
1·60
0·80
6·20
7·00
5·50
5·50
6·60
1·50
1·40
1·40
1·70
2·60
0·80
2·00
8·00
2·60
2·00
4·00
2·40
"2·20
0·60
3·40
3·60
8·10
8·20
"1·40
8·20
4·20
"1·00
4·10
2·40
2·20
6·00
5·20
5·20
6·00
1·80
6·10
5·40
2·40
0·60
2·60
6·90
7·20
1·70
5·80
6·15
8·10
7·80
6·30
7·30
7·65
6·30
5·35
4·60
6·20
5·35
5·65
5·90
5·40
5·50
10·60
10·65
12·65
6·70
4·50
4·65
6·75
6·35
4·00
4·90
7·25
10·05
11·75
5·70
5·40
5·75
5·95
5·75
6·45
7·35
7·15
5
4
1
0
4
4
2
2
5
4
4
4
4
3
5
4
0
0
0
4
6
6
1
2
5
7
4
3
1
6
6
4
4
5
2
2
1
2
2
1
3
5
3
2
2
0
0
0
4
0
5
5
1
5
4
0
3
0
0
4
4
4
4
2
3
4
1
1
4
4
3
1
DentAdv
DentSlght
EnamTrace
None
DentSlght
DentSlght
EnamMid
EnamMid
DentAdv
DentSlght
DentSlght
DentSlght
DentSlght
EnamAdv
DentAdv
DentSlght
None
None
None
DentSlght
DentStrng
DentStrng
EnamTrace
EnamMid
DentAdv
EnamRing
DentSlght
EnamAdv
EnamTrace
DentStrng
DentStrng
DentSlght
DentSlght
DentAdv
EnamMid
EnamMid
EnamTrace
EnamMid
EnamMid
EnamTrace
EnamAdv
DentAdv
EnamAdv
EnamMid
EnamMid
None
None
None
DentSlght
None
DentAdv
DentAdv
EnamTrace
DentAdv
DentSlght
None
EnamAdv
None
None
DentSlght
DentSlght
DentSlght
DentSlght
EnamMid
EnamAdv
DentSlght
EnamTrace
EnamTrace
DentSlght
DentSlght
EnamAdv
EnamTrace
8·43
1·84
-
9·10
10·85
11·60
5·05
9·10
12·35
-
11·35
-
13·25
11·00
12·70
6·50
6·25
7·60
-
10·15
14·50
8·10
-
6·15
6·10
9·75
4·90
6·40
11·15
6·15
6·95
7·55
-
5·80
2·67
-
13·75
1·88
1·93
1·56
2·47
0·55
4·19
1·59
-
3·12
19·87
13·50
-
9·04
38·26
-
2·04
-
17·89
26·12
1·00
-
7·34
-
19·13
-
5·48
-
2·97
8·65
-
10·96
6·85
-
0·25
11·19
-
2·06
0·55
0·52
-
700 M. Skinner
Appendix continued
Group
MP
MP
MP
SpecName
Puymoyen 1
Puymoyen 2
La Quina 9
Spec
89
90
92
IndAge
16·8
18·6
18·6
FDI
FunctAge
TthHt
AttScor
AttClass
llP1
lrI1
llC
llI1
llP2
llI2
ulM3
llP1
llM1
ulM1
ulM2
llP2
llM2
lrM2
lrM3
lrM1
lrP1
lrC
llM2
llM1
llM3
llP2
llP1
3·50
7·30
3·70
7·30
2·90
6·50
"2·20
6·80
10·50
10·40
4·20
6·20
5·00
6·80
"0·40
12·30
8·60
8·80
6·80
12·30
"0·40
8·00
8·60
4·85
9·35
12·50
9·05
6·30
9·90
6·35
4·95
3
5
3
5
2
5
0
2
4
4
2
1
2
2
1
4
2
4
3
5
3
4
4
EnamAdv
DentAdv
EnamAdv
DentAdv
EnamMid
DentAdv
None
EnamMid
DentSlght
DentSlght
EnamMid
EnamTrace
EnamMid
EnamMid
EnamTrace
DentSlght
EnamMid
DentSlght
EnamAdv
DentAdv
EnamAdv
DentSlght
DentSlght
-
6·30
7·50
6·25
6·40
7·00
6·80
6·55
5·10
12·70
6·85
6·00
6·70
6·75
5·00
%Dent
-
7·27
1·50
9·79
-
7·18
-
1·26
0·62
-
1·90
-
1·35
-
11·02
-
2·88
3·93
*EUP: Early Upper Paleolithic; LUP: Late Upper Paleolithic; MP: Middle Paleolithic; ul: upper left; ur: upper right; ll: lower left; lr: lower
right; i: primary incisor; c: primary canine; m: primary molar; I: permanent incisor; C: permanent canine; M: permanent molar; 1–8: tooth
number.