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Dental reduction in post-pleistocene Nubia.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 70349-363 (1986)
Dental Reduction in Post-Pleistocene Nubia
JAMES M. CALCAGNO
Department of Sociology and Anthropology, Loyola University of Chicago,
Chicago, Illinois 60626
KEY WORDS
Agriculturalist
Tooth Size, Mesolithic, Agriculturalist, Intensive
ABSTRACT
Tooth size changes among Nubian archaeological populations
dating from the Mesolithic through the Christian era, a period of approximately 12,000years, are analyzed. Standard length and breadth dimensions
of all permanent teeth from several cultural horizons are combined to form
three large samples: Mesolithic, 10000-7000 B.C.; Agriculturalist, 3300-1100
B.C. (A-group,C-group, Pharaonic); and Intensive Agriculturalist, A.D. 0-1400
(Meroitic, X-Group, Christian). Such information not only fills a void in the
knowledge of Nubian skeletal biology, but also provides a much needed African
reference point for the comparison of tooth size data among anatomically
modern Homo sapiens from various regions of the world.
Changes in mean tooth size and associated t-tests reveal strong and significant reduction in dental size between the Mesolithic and Agriculturalist samples, followed by a continued although diminished trend of reduction for only
the molar teeth between the two Agriculturalist groups. These patterns are
best observed by examining tooth breadths, which are considered as the most
reliable indicator of tooth sue. Previously published odontometrics of the
Nubian Mesolithic are briefly compared to the findings of this diachronic
analysis of Nubian dental change.
Despite the abundance of archaeological
and skeletal analyses conducted on the various cultural phases of post-Pleistocene Nubia, basic odontometric data spanning the
entire sequence have been conspicuously absent. This is not unusual, since a lack of tooth
size data among recent populations has been
a deficiency in physical anthropology as a
whole and not just in the study of Nubia.
Nearly two decades ago, C. Loring Brace
brought the undesirably small body of available dental data to the attention of anthropologists, with the hope of stimulating others
to conduct work on the “systematic appraisal
of the teeth of modern Homo sapiens” (Brace,
1967:815).This plea, in tandem with a similar appeal four years later for further analysis of an area “being pursued by altogether
too few scholars” (Brace and Mahler, 1971:
202), apparently produced the intended results, for the number of relevant studies in
the short time since Brace’s comments
greatly outnumber the previous works. However, the continent of M i c a still remains
0 1986 ALAN R. LISS, INC.
virtually unexplored in this regard. By providing tooth size data, which until now were
seriously lacking in the rapidly expanding
body of Nubian research, this paper is intended to contribute information to help alleviate various shortcomings in the study of
Nubia, while simultaneously supplying an
African reference point of dental evolution
among populations of anatomically modern
Homo sapiens. Possible mechanisms accounting for observed changes in tooth sue are not
discussed at this time, to better focus upon
the presentation of these dental data.
THE POPULATIONS
Geographically, Nubia is generally considered as the land between the first and fourth
cataracts of the Nile River (Carlson, 1974;
Carlson and Van Gerven, 1979),although the
southern limit has shifted over time to as far
as the confluence of the White and Blue Niles
Received M a y 13,1985; revision accepted January 30,1986.
350
J.M. CALCAGNO
at the onset of the Christian era (Adams,
1977). However, the first cataract (near Aswan) has always sharply marked the northern border, while the eastern and western
boundaries have been restricted to the mile
or two of arable and habitable land on each
side of the Nile.
The ongoing systematic analysis of Nubian
archaeological and skeletal material began
in response to the enlargement of the original Aswan Dam in 1908 and the subsequent
flooding of the Nile River (Adams, 1977). Today, as a consequence of three major archaeological campaigns (1907-1911, 1929-1934,
1959-19691, as well as numerous expeditions
in between, Nubia is often considered as one
of the best known archaeological areas in the
world (Adams, 1966, 1967, 1977; Carlson,
1974; Carlson and Van Gerven, 1979).
All skeletal material (except for four individuals from a Mesolithic site in southern
Egypt) analyzed in this study was excavated
from the relatively small region in northern
Sudan that is still considered as part of Lower
Nubia. Measurements for the Mesolithic
sample were taken at the University of Colorado (Boulder)and Southern Methodist University (Dallas), with data from postMesolithic phases being gathered at the
Universitetets Anthropologiske Laboratorium in Copenhagen, Denmark. Each of these
collections was recovered during the early-tomid 1960’s as part of the High Dam Campaign (see Greene and Armelagos, 1972;
Nielsen, 1970; Wendorf, 1968).
Three samples are compared in this diachronic examination of tooth size in Nubia:
an early Mesolithic group (10000-7000 B.C.),
an “Agriculturalist” population (3300-1100
B.C.), and a more “Intensive Agriculturalist” phase (A.D. 0-1400). The Agriculturalist
period displays greater sedentism and increased reliance upon domesticated resources relative to its Nubian predecessors,
with subsistence being characterized by
floodplain farming and animal husbandry
(Adams, 1967; Butzer, 1971; Nielsen, 1970),
fishing (Adams and Nordstrom, 1963), and
hunting and gathering (Butzer, 1971). However, Butzer (1971)notes that the presence of
numerous Mesolithic subsistence traits indicates a general unwillingness by the A-group
people to assimilate a totally Neolithic economy into their lifestyle. Included within the
Agriculturalist phase is the A-group, Cgroup, and Pharaonic horizons (refer to Adams, 1977; Carlson and Van Gerven, 1979;
and Calcagno, 1984, for lengthy discussions
of the continuity vs. migrationism controversy regarding Nubian culture history). Although some prefer that the Pharaonic
horizon be excluded from the former grouping owing to the possibility of this group
being represented solely by Egyptian colonists (Adams, 1977; Nielsen, 1970),the small
sample size from this horizon only slightly
and insignificantly alters the dental means
when included among the Agriculturalists
(Calcagno, 1984). Thus, to satisfy both sides
of this issue, Pharaonic measurements are
included in this study, with occasional reference also made to results obtained when only
the A and C horizons make up this phase.
The Agriculturalist phase may be viewed
as transitional between the Mesolithic and
the “Intensive Agriculturalists,” which unlike previous periods of Nubian prehistory,
represent a total reliance upon agriculturalism. Due to their close archaeological, skeletal, and economic affinities, the Meroitic, Xgroup, and Christian periods may be safely
combined to form this final sample. In addition, with regard to the dentition, few comparisons of tooth size or variation show
significant change beween these three horizons (Calcagno, 1986), and the proper merging of such samples permits a more lucid and
understandable analysis of closely related
populations employing a similar subsistence
pattern.
THE MEASUREMENTS
Maximum crown length (mesiodistal),and
the maximum crown breadth (buccolingual,
labiolingual) measured perpendicular to the
length, were taken following the procedure
recommended by Frayer (1978) for all permanent teeth whose size was relatively unaffected by attrition. Maximum dimensions
were preferred because they tend to be more
easily and consistently measured, thereby
helping to reduce the amount of intraobserver error, which was under 1.3% in this
study. When tooth dimensions from both the
right and left sides of the jaw were obtained,
the two figures were averaged to provide a
single measurement for a given tooth per
individual.
Although both mesiodistal and buccolingual dimensions are reported in this study,
the latter appears to be a better measure of
the underlying genetic component of tooth
size. When maximum dimensions are recorded, occlusal and interproximal wear can
DENTAL REDUCTION IN POST-PLEISTOCENENUBIA
significantly diminish mesiodistal lengths
(Biggerstaff, 1979; Frayer, 1978; Johanson,
1974; and others), while at the same time not
affecting the buccolingual breadths, which
are measured below the original occlusal surface Wan Reenen, 1982).The amount of attrition within a given specimen varies with
each tooth class, depending upon such factors
as size, morphology, and location of the tooth
in the jaws (Van Reenen, 1982). Incisors are
especially vulnerable to the decreasing reliability of mesiodistal metrics, because their
maximum length is measured essentially at
the unworn occlusal surface, which when
erupted is sharp but then tends to be rapidly
worn down. (Nubian dental remains characteristically display considerable amounts of
wear, and although unworn or slightly worn
teeth were the focus of analysis, less codidence is held in the incisor lengths than in
any other dimension.) Van Reenen (1982)has
also shown that males sometimes exhibit
greater reductions in mesiodistal metrics owing to attrition than do females, which would
bias an analysis of patterns of tooth size
change between the sexes within a group.
And finally, in addition to the effects of attrition, higher heritabilities (Alvesalo and Tigerstadt, 1974) and correlations between
relatives (Sofaer, 1973) have been documented for buccolingual dimensions relative
to mesiodistal diameters. Thus, for several
reasons buccolingual dimensions appear to
be less influenced by environmental factors,
and therefore should be weighed more heavily when analyzing tooth size in human
populations.
Sexes are analyzed as separate samples to
determine if males and females experienced
similar patterns of dental change. Also, not
delimiting samples by sex may result in an
inaccurate diachronic depiction of tooth size
trends if a disproportionate number of one
sex is found in one sample relative to others
(e.g., what might appear to be dental reduction over time could be the result of relatively fewer males, with larger teeth, in later
samples). The sex of each specimen was independently determined by the author on the
basis of cranial morphology, with the decision then being compared to the judgement
of previously published reports by Greene
and Armelagos (1972), Wendorf (1968), and
Nielsen (1970). Rarely was there disagreement with the assessment of these investigators, who often originally had more
skeletal parts (including pelves) associated
351
with skulls from which to base their sex assignment. If uncertainty still existed after
comparing decisions, the specimen was excluded from the analysis. Dental size was
never used to sex an individual.
RESULTS
Descriptive statistics reveal the closeness
in the dental dimensions of the Agriculturalists and Intensive Agriculturalists, relative to the Mesolithic (Table 1). Percent
differences in mean tooth size between chronologically adjacent groups clearly display
that the bulk of dental reduction that occurred in Nubia took place between the Mesolithic and Agriculturalist period. This fact
is readily observable in Figure 1, which
graphically illustrates differences in breadth
measurements between these phases. The
average amount of dental reduction seen in
Agriculturalist males relative to Mesolithic
males is 6.1%, as opposed to only 0.5% between the Agriculturalist and Intensive Agriculturalist males. Females closely approximate these statistics, showing a 6.4% reduction in mean tooth size between the first two
phases, followed by a 0.7% drop in the final
phase. Another interesting pattern found in
this table concerns the variables displaying
minor increases in dental size in the Intensive Agriculturalist population, most of
which are anterior tooth measurements.
Most striking is the comparison of male maxillary dimensions, where each of the six anterior tooth dimensions show slight increases
in size, while the reverse is true for the posterior set. An almost identical pattern is observed in the female mandibular variables.
Thus, as a cautionary note to the general
conclusion that teeth of the Agriculturalist and Intensive Agriculturalist samples
exhibit little change in size, it should be
added that the posterior teeth (premolars and
molars) are showing relatively greater decreases than the anterior teeth (incisors and
canines). This pattern will be examined later
in greater detail.
Significance tests on the dental means
reinforce that which can be intuitively derived from the descriptive statistics (Table 1).
When comparing the Mesolithic with the Agriculturalists, significant decreases in tooth
size dominate all possible comparisons, irrespective of sex. For males, 30 of the 32 variables significantly reduce over this time
frame, females show significant reductions
in 26 categories, and all variables except
Males only
I1 LT
Meso
Ag
In Ag
I1 BR
Meso
Ag
In Ag
I2 LT
Meso
Ag
In Ag
I2 BR
Meso
Ag
In Ag
C1 LT
Meso
Ag
In Ag
C1 BR
Meso
Ag
In Ag
P3 LT
Meso
Ag
In Ag
P3 BR
Meso
Ag
In Ag
P4 LT
Meso
Ag
In Ag
5.7-7.1
5.4-6.7
5.5-6.6
7.5-9.3
6.6-8.9
6.5-8.5
6.9-8.2
6.2-8.2
6.2-8.0
0.39
0.30
0.26
0.53
0.31
0.34
0.26
0.34
0.39
0.41
0.50
0.35
0.44
0.53
0.47
0.40
0.45
0.44
0.47
0.56
0.50
0.38
0.41
0.51
6.43
5.99*
5.96
6.31
6.04*
6.09
6.86
6.33*
6.31
7.51
6.87*
6.94
8.40
7.65*
7.58
7.50
7.13*
7.02
8.96
8.13*
8.11
7.37
7.25
7.09*
6.7-8.1
6.6-8.2
5.9-8.9
8.4-9.7
6.8-9.8
7.1-9.0
6.8-8.3
6.2-8.7
6.3-7.8
6.4-7.5
5.7-7.1
5.3-7.1
5.4-7.2
5.5-6.7
5.3-6.6
5.2-6.4
4.9-6.2
4.9-5.9
Range
0.35
0.35
0.26
S
Mandibular
5.81
5.44*
5.44
Xmm)
-
(N)
1.6
2.3
9.3
0.2
4.9
1.5
8.9
0.9
+
8.5
1.0
7.7
0.3
4.3
+0.8
6.8
0.5
6.4
0.0
(%)
Diff
6.98
6.80
6.75
10.09
9.53*
9.49
7.63
7.17*
7.09
8.74
8.39*
8.43
8.18
7.68*
7.83
7.00
6.48*
6.57
7.47
6.85*
7.10
7.60
7.15*
7.18
9.33
8.90
8.91
Wmm)
-
0.28
0.40
0.35
0.45
0.51
0.54
0.30
0.43
0.40
0.52
0.48
0.65
0.39
0.44
0.37
0.34
0.35
0.50
0.51
0.51
0.79
0.27
0.27
0.42
0.78
0.71
0.62
S
6.5-7.3
5.8-7.9
5.8-7.7
9.4-10.8
8.1-10.5
8.2-10.6
7.2-8.1
6.1-8.3
6.0-7.8
7.8-9.6
7.4-9.4
7.0-9.6
7.7-8.9
6.7-8.8
7.1-8.8
6.6-7.7
5.9-7.0
5.2-7.6
6.4-8.4
5.8-7.8
5.7-9.5
7.1-8.0
6.6-7.8
6.7-8.2
8.2-10.6
7.8-10.1
7.9-10.4
Range
Maxillary
(N)
TABLE 1. Dental descriptive statistics and percent differences between successive phases of the Mesolithic (Meso), Agriculturist (Ad, and Intensive
Agriculturalist (In Ag) groups
2.6
0.7
5.6
0.4
6.0
1.1
4.0
+0.5
6.1
+2.0
7.4
+ 1.4
8.3
+3.6
5.9
+0.4
4.6
+0.1
(a)
Diff
Females only
I t LT
Meso
Ag
In Ag
I1 BR
Meso
Ag
In Ag
I2 LT
Meso
Ag
In Ag
I2 BR
Meso
Ag
In Ag
P4 BR
Meso
Ag
In Ag
M1 LT
Meso
Ag
In Ag
M1 BR
Meso
Ag
In Ag
M2 LT
Meso
Ag
In Ag
M2 BR
Meso
Ag
In Ag
M3 LT
Meso
Ag
In Ag
M3 BR
Meso
Ag
In Ag
0.44
0.40
0.40
0.51
0.37
0.29
5.95
5.73
5.79
6.62
5.99*
6.01
0.58
0.62
0.67
11.17
10.33*
10.15
0.39
0.33
0.31
0.79
0.85
0.86
11.18
10.60*
10.37
6.31
5.64*
5.66
0.65
0.60
0.58
11.43
10.64*
10.44*
0.34
0.35
0.25
9.4-12.8
8.6-12.6
7.3-11.9
0.58
0.68
0.68
11.39
10.93*
10.76
5.48
5.13*
5.21
10.3-12.4
9.3-11.9
9.0-11.5
0.43
0.46
0.41
11.75
10.89*
10.68*
5.7-7.7
4.9-6.8
5.4-6.5
5.3-7.0
4.8-6.4
5.0-6.7
5.7-7.1
5.0-6.3
4.9-6.1
4.8-6.1
4.4-5.8
4.7-5.6
10.0-12.3
8.8-11.7
8.0-11.4
10.3-12.8
9.1-12.4
9.2-12.4
10.8-12.6
9.6-12.4
9.5-11.7
11.1-12.6
9.8-12.7
9.9-12.1
0.49
0.60
0.48
11.85
11.41*
11.26
8.5-10.0
7.4-9.6
6.7-9.5
0.46
0.48
0.57
9.16
8.57*
8.40
(16)
(40)
(30)
(15)
(36)
(24)
(14)
(28)
(22)
(12)
(25)
(16)
9.5
+0.3
3.7
+ 1.0
10.6
+0.4
6.4
+1.6
7.5
1.7
5.2
2.2
6.9
1.9
4.0
1.6
7.3
1.9
3.7
1.3
6.4
2.0
6.83
6.31*
6.25
7.15
6.82
6.70
7.44
6.89*
7.19
9.21
8.64*
8.46
12.01
11.21*
11.05
9.16
8.72*
8.71
12.75
11.80*
11.62
10.59
9.79*
9.73
12.60
11.78*
11.61
11.08
10.49*
10.42
10.32
9.63*
9.60
0.58
0.43
0.35
0.59
0.52
0.51
0.58
0.34
0.35
0.38
0.46
0.47
0.67
0.72
0.99
0.81
0.73
0.85
0.54
0.66
0.70
0.54
0.65
0.57
0.41
0.49
0.56
0.67
0.53
0.39
0.44
0.63
0.56
5.6-7.9
5.4-7.1
5.3-6.7
6.4-8.4
5.7-8.1
5.7-7.4
6.2-8.4
6.3-7.8
6.5-7.7
8.5-9.7
7.7-9.6
7.4-9.1
10.6-12.8
9.6-12.6
8.0-13.0
7.5-10.7
5.8-10.4
6.8-10.9
11.9-13.6
10.3-13.1
10.1-13.4
9.6-11.5
8.5-12.2
7.9-11.2
11.6-13.1
10.4-12.6
10.1-12.6
9.9-12.4
9.3-11.9
9.3-11.3
9.5-11.2
8.2-11.0
8.3-10.8
(continued)
7.6
1.0
4.6
1.8
7.4
+4.4
6.2
2.1
6.7
1.4
4.8
0.1
7.5
1.5
7.6
0.6
6.5
1.4
5.3
0.7
6.7
0.3
C1 LT
Meso
Ag
I n Ag
C1 BR
Meso
Ag
In Ag
P3 LT
Meso
Ag
In Ag
P3 BR
Meso
Ag
I n Ag
P4 LT
Meso
Ag
I n Ag
P4 BR
Meso
Ag
In Ag
M1 LT
Meso
Ag
In Ag
M1 BR
Meso
Ag
In Ag
M2 LT
Meso
Ag
In Ag
6.8
+LO
7.6
0.1
6.1-7.9
5.9-7.9
6.3-7.8
9.2-10.4
7.9-10.5
7.9-10.2
0.46
0.48
0.40
7.37
6.87*
6.94
9.91
9.16*
9.15
6.1-7.9
5.8-7.9
5.9-7.7
7.8-9.7
6.3-9.6
6.6-8.5
6.2-7.8
6.1-8.0
6.2-7.5
0.51
0.49
0.42
0.56
0.61
0.44
0.43
0.41
0.34
0.58
0.60
0.47
7.05
6.84
6.85
8.58
7.77*
7.68
7.02
6.88
6.80
8.08*
11.11
10.81
10.75
4.2
0.5
7.6
3.5
8.9-11.0
8.1-11.0
7.7-11.2
0.62
0.67
0.69
10.34
9.55*
9.22
(19)
(100)
(48)
9.3-11.9
8.9-12.3
8.8-12.0
0.66
0.68
0.64
5.3
1.9
11.2-13.0
10.0-12.8
9.9-12.4
10.83
10.38*
10.33
7.0
2.6
0.60
0.55
0.55
(16)
(73)
(31)
10.7-12.2
9.3-11.9
9.0-11.9
0.48
0.54
0.58
11.26
10.47*
10.20*
12.10
11.46*
11.24*
4.7
1.7
9.8-11.5
9.3-11.6
8.6-11.3
0.51
0.54
0.60
10.75
10.24*
10.07
(17)
(75)
(32)
9.9-12.2
9.4-12.1
9.5-12.2
0.69
0.63
0.59
7.97
2.7
0.6
7.8
0.1
9.2-10.9
8.0-10.5
8.1-10.4
0.59
0.59
0.54
10.02
9.24*
9.23
7.8-9.7
6.6-9.4
7.1-9.1
8.83
3.1
+ 1.5
6.0-7.5
5.7-7.3
5.9-7.3
0.42
0.37
0.34
6.75
6.54
6.64
0.43
0.59
0.52
7.5
+ 1.3
7.1-9.4
6.5-8.9
6.7-8.7
0.57
0.57
0.44
8.50
7.86*
7.96
7.4-8.8
5.8-8.6
6.4-8.0
4.9
1.2
(%)
Diff
0.36
0.57
0.35
(N)
7.97
7.25*
7.14
Range
7.1-9.0
6.4-8.5
6.7-8.1
S
0.52
0.47
0.38
2.0
1.2
(%I
Maxillary
7.79
7.41*
7.32
(20)
i77j
(43)
(N)
X(mm)
6.5-7.7
5.7-7.2
5.9-7.2
Range
Dig
0.36
0.41
0.28
S
Mandibular
7.00
6.48*
6.62
X(mm)
TABLE 1. Dental descriptive statistics and percent differences between successive phases ofthe Mesolithic (Meso), Agriculturist (A&, and Intensive
Agriculturalist (In A d groups (continued)
8.3
0.2
+LO
8.6
0.0
8.5
+2.3
9.3
0.7
3.6
0.6
9.1
0.4
5.7-7.1
5.0-6.7
4.9-6.6
5.3-7.2
4.8-6.7
5.0-6.7
5.7-7.7
4.9-7.1
5.3-7.1
6.5-8.3
5.7-8.7
5.9-7.8
7.4-9.3
5.8-8.9
6.4-8.5
6.1-8.2
5.8-8.2
5.9-8.0
7.8-9.7
6.3-9.8
6.6-9.0
0.39
0.36
0.32
0.51
0.38
0.40
0.42
0.39
0.38
0.46
0.49
0.36
0.46
0.58
0.47
0.51
0.49
0.44
0.55
0.61
0.52
3.9
5.9
+0.8
4.8-6.4
4.4-6.2
4.7-5.9
9.1
2.7
9.1-11.9
7.9-11.6
8.0-11.3
0.75
0.74
0.66
10.82
9.84*
9.57*
0.37
0.38
0.28
5.0
1.7
8.6-11.5
7.7-12.4
8.1-12.0
0.78
0.98
0.87
8.2
1.1
10.52
9.99*
9.82
9.5-12.2
8.8-11.7
8.8-11.4
0.71
0.59
0.51
10.97
10.07*
9.96
Males and females combined
I1 LT
Meso
5.62
5.29*
Ag
5.33
In Ag
I1 BR
6.36
Meso
5.83*
Ag
5.82
In Ag
I2 LT
6.12
Meso
5.88*
Ag
5.94
In Ag
I2 BR
6.74
Meso
6.16*
Ag
6.16
In Ag
C1 LT
7.28
Meso
6.66*
Ag
6.81
In Ag
C1 BR
8.20
Meso
7.44*
Ag
7.39
In Ag
P3 LT
7.25
Meso
6.99*
Ag
In Ag
6.95
P3 BR
Meso
8.76
7.96*
Ag
In Ag
7.93
M2 BR
Meso
Ag
In Ag
M3 LT
Meso
Ag
In Ag
M3 BR
Meso
Ag
In Ag
10.00
9.35%
9.34
7.49
7.01*
7.03
8.62
8.09*
8.24
7.99
7.53*
7.63
6.91
6.39*
6.42
7.32
6.83*
6.91
7.52
7.01*
7.19
9.26
8.74
8.72
11.47
10.82*
10.61
8.96
8.44*
8.24
11.97
11.36*
10.97*
6.4-8.4
5.7-8.1
5.7-9.5
0.56
0.51
0.70
0.44
0.58
0.56
0.41
0.48
0.41
0.55
0.59
0.62
0.49
0.47
0.45
(continued)
6.5
0.1
9.2-10.8
7.9-10.5
7.9-10.6
(30)
(115)
(87)
6.4
+0.3
6.1
+ 1.9
5.8
+ 1.3
7.5
+0.5
6.4
+ 1.2
6.8
+2.6
5.6
0.2
5.7
1.9
5.8
2.4
5.1
3.4
6.1-8.1
5.9-8.3
6.0-7.8
7.1-9.6
6.5-9.4
6.7-9.6
7.1-9.0
6.4-8.8
6.7-8.8
5.6-7.9
5.4-7.1
5.2-7.6
6.2-8.4
6.3-7.8
6.5-8.2
0.45
. ~.
0.33
0.38
0.48
0.40
0.46
8.2-10.6
7.7-10.1
7.4-10.4
10.3-12.7
8.6-12.9
8.3-12.2
0.77
0.79
0.81
0.59
0.58
0.59
7.7-9.8
6.7-10.6
5.6-10.0
10.6-13.1
9.8-12.7
9.8-12.2
0.59
0.66
0.77
0.82
0.66
0.56
7.8-10.0
6.6-9.6
6.7-9.5
0.54
0.59
0.57
0.71
0.68
0.58
0.51
0.54
0.53
0.68
0.73
0.69
0.71
0.66
0.59
0.84
0.96
0.90
0.69
0.72
0.72
8.98
8.33*
8.21
11.44
11.12*
11.07
11.48
10.69*
10.49*
11.09
10.66*
10.57
11.17
10.37*
10.23
10.82
10.32*
10.11
10.99
10.10*
9.88*
9.1-12.3
7.9-11.7
8.0-11.4
8.6-12.8
7.7-12.6
7.3-12.0
9.5-12.4
8.8-11.9
8.8-11.5
9.3-12.8
8.9-12.4
8.8-12.4
10.7-12.6
9.3-12.4
9.0-11.9
9.9-12.6
9.4-12.7
9.5-12.2
6.2-8.1
6.1-8.2
5.9-8.9
0.44
0.45
0.47
7.18
7.07
6.97
*Significant difference (at a = ,051 between successive phases.
P4 LT
Meso
Ag
In Ag
P4 BR
Meso
Ag
In Ag
M1 LT
Meso
Ag
In Ag
M1 BR
Meso
Ag
In Ag
M2 LT
Meso
Ag
In Ag
M2 BR
Meso
Ag
In Ag
M3 LT
Meso
Ag
In Ag
M3 BR
Meso
Ag
In Ag
(N)
(37)
(192)
(102)
(37)
(194)
(102)
(32)
(196)
(105)
(35)
(204)
(109)
(29)
(153)
(80)
(31)
(156)
(85)
(36)
(155)
(95)
(37)
(163)
(98)
8.1
2.2
4.6
2.0
7.2
1.4
3.9
0.8
6.9
1.9
2.8
0.4
7.2
1.4
1.5
1.4
(%)
Range
X(mm)
s
Diff
-
Mandibular
0.76
0.78
0.93
0.71
0.71
0.84
9.06
8.59*
8.48
11.74
11.02*
10.83
0.75
0.69
0.71
0.58
0.67
0.68
10.47
9.67*
9.49
12.37
11.58*
11.30*
0.56
0.54
0.58
0.61
0.55
0.52
0.53
0.64
0.58
0.37
0.41
0.35
S
12.36
11.61*
11.44*
10.91
10.36*
10.26
10.18
9.44*
9.44
6.87
6.68
6.70
Wmm)
-
10.3-12.8
8.6-12.9
8.0-13.0
7.5-10.7
5.8-10.6
5.6-10.9
10.6-13.6
9.8-13.1
9.8-13.4
8.9-11.5
8.1-12.2
7.7-11.2
11.2-13.1
10.0-12.8
9.9-12.6
9.8-12.4
9.3-11.9
8.6-11.3
9.2-11.2
8.0-11.0
8.1-10.8
6.0-7.5
5.7-7.9
5.8-7.7
Range
Maxillary
(36)
(169)
(104)
(36)
(169)
(103)
(37)
(177)
(114)
(36)
(184)
(110)
(29)
(147)
(100)
(32)
(151)
(104)
(32)
(142)
(91)
(31)
(146)
(89)
(N)
TABLE 1. Dental descriptive statistics and percent differences between successive phases of the Mesolithic (Meso), Agriculturist (Ad, and Intensive
Agriculturalist (In Ag) groups (continued)
6.1
1.7
5.2
1.3
6.4
2.4
7.6
1.9
6.1
1.5
5.0
1.0
7.3
0.0
2.8
+0.3
(%)
Diff
DENTAL REDUCTION IN POST-PLEISTOCENE NUBIA
mandibular P4 length significantly decrease
in the combined sample (Table 2).
As previously discussed, breadth dimensions appear to be better indicators of genetic
changes in tooth size and should be viewed
with added scrutiny. In males, each of the 16
mandibular and maxillary breadths decrease
at the .001 level of significance, except for
the maxillary canine (which nonetheless still
significantly decreases, but at the .05 level).
The same pattern holds true for females,
where 15 of the 16 breadths decrease at the
.001 level of significance, with the maxillary
third molar reducing at the .01 level. When
the samples are judged irrespective of sex,
all breadth dimensions significantly reduce
at the .001 level between the Mesolithic and
Agriculturalist phases. Length dimensions do
not reduce with nearly the consistency and
magnitude as the breadths, and some even
display insignificant increases. This is probably a reflection of environmental factors
such as dental wear that can greatly influence and bias tooth length. Regardless, there
can be no doubt that tooth size in the Agriculturalist phase significantly and markedly
decreased from the Mesolithic in Nubia.
Although tooth size is very similar among
the Agriculturalists and Intensive Agriculturalists, t-tests on the dental means reveal
that of the significant reduction that has occurred, all of it involves the posterior teeth.
No incisor or canine dimensions, among
males or females, exhibit a significant decrease in size. In fact, one anterior tooth dimension actually increases significantly, and
the majority of the remaining variables insignificantly increase. In marked contrast,
significant decreases occur in 3 of the 20 posterior dimensions among the males (15%),4
posterior variables among females t20%),and
5 posterior teeth in the combined sample
(25%).Almost all of the remaining variables
display insignificant decreases, and no posterior dimensions significantly increase in
size. Thus, whereas the anterior teeth remain relatively constant in size from the Agriculturalist to Intensive Agriculturalist
periods, the posterior teeth exhibit a continued, though slight, trend for reduction.
This lessened trend of posterior tooth reduction may be better understood by examining only breadth dimensions, which
indicate that a more specific pattern of molar
reduction is evident in the comparison of the
Agriculturalist phases. Of the six molar
357
breadths, two (mandibular first and second)
significantly reduce in males, while four
(mandibular first and third, maxillary first
and second)significantly decrease in both the
females and combined sample. At the same
time, none of the four premolars significantly
decrease in males, females, or combined
sexes. Hence, rather than simply noting a
general pattern of posterior tooth reduction,
a more specific trend for molar reduction is
manifested when analyzing dental breadths.
When Pharaonic measurements are excluded from the Agriculturalist sample (as in
studies by Carlson, 1974, 1976; Carison and
Van Gerven, 1977), the above patterns are
accentuated, because mean tooth size is
slightly larger and more distant from the
Intensive Agriculturalist sample but still
very much smaller than the Mesolithic teeth
(see Table 2). Once again, most teeth significantly reduce from the Mesolithic to the Agriculturalist period (28 of 32 variables for
males, 26 of 32 for females, and 31 of 32 for
the combined sample). All 16 breadth dimensions decrease at the .001 level of significance for males and the total sample, and
only the maxillary third molar of females,
which is significant at the .01 level, fails to
do the same. In comparing the two Agriculturalist stages, no anterior tooth dimensions
show significant decreases, as opposed to 7 of
20 (35%)posterior dimensions for males, 6 of
20 (30%)for females, and 10 of 20 (50%)for
the combined sample. Five of the six molar
breadths decrease (all except the maxillary
third molar) among the males, while four
molar breadths (mandibular first and third,
maxillary first and second) from both the female and combined samples reduce in size.
Again then, a continued decline in molar size
only is evidenced from the Agriculturalists
to the Intensive Agriculturalists.
Posterior tooth areas-individual and
summed
Estimates of the occlusal area of the posterior teeth of both jaws display a great amount
of reduction through time, especially from
the Mesolithic to Agriculturalist phase, as
would be anticipated from the analysis of
individual length and breadth dimensions.
For the maxilla, the second molar displays
the greatest amount of reduction in area
throughout the entire sequence (Table 3).
Maxillary R42 decreases by 16.9% from the
Mesolithic to the Intensive Agriculturalists
./
E
E
-
C
I1
1
a
I1
ME50
c1
1
+
P3
AG
0
P4
M1
INT AG
MZ
0
12
ME50
c1
+
P3
AG
0
P4
INT AG
M1
M2
Female M a n d i b u l a r B r e a d t h s
0
12
Male Mandibular B r e a d t h s
3
M3
l
E
E
rn
8
c
I
v
l3
c1
MESO
P3
+
AG
0
P4
M1
INT A G
M2
Fern a I e M a x i I Ia ry B r e a d t h s
n
12
M3
d
0
ME50
+
AG
0
INT A G
3
b
I1
Male Maxillary B r e a d t h s
DENTAL REDUCTION IN POST-PLEISTOCENE NUBIA
TABLE 2. Summary of Student's t-test (twetailed)
results on dental means of Nubian Mesolithic (Meso),
Agriculturalist (Ad, and Intensive Agriculturalist
phases (Int Ag) (number and percentage of 32 variables
that significantly (0.05)reduce between each group)
Males
With Pharaonic tradition
30 (94)
Meso/Ag
3 (9)
AgAnt Ag'
MesoDnt Ag
28 (88)
Without Pharaonic tradition
MesoiAg
28 (88)
7 (22)
AgAnt Ag'
MesoDnt Ag
27 (84)
Percentage of:
Females
Combined
26 (81)
4 (13)'
27 (84)
31 (97)
5 (16)
31 (97)
26 (81)
6 (19)'
27 (84)
31 (97)
10 (31)
31 (97)
'All reductions involve posterior teeth only.
'One significant increase in this category is excluded.
TABLE 3. Maxillary dental areas (in mm') and
percentage differences between successive groups (in
parentheses) for the Mesolithic (Meso), Agriculturalist
( A d . and Intensive Agriculturalist (In A d Dhases
Meso
Males
P3
P4
M1
M2
M3
Summed
Females
P3
P4
M1
M2
M3
Summed
Combined
P3
P4
M1
M2
M3
Summed
(%)
Ag
(%)
In Ag
77.75
72.21
140.42
135.29
110.36
536.03
(11.8)
(9.01
(11.6)
(14.4)
(11.3)
(11.9)
68.57
65.73
124.15
115.81
97.94
472.20
(1.3)
(1.1)
(2.4)
(2.5)
(1.3)
(1.9)
67.69
65.01
121.04
112.92
96.67
463.43
72.85
67.68
130.27
123.84
103.03
497.67
(13.2)
(10.5)
(9.4)
(12.4)
(11.2)
(11.2)
63.27
60.59
118.02
108.49
91.49
444.86
(+0.7)
(i1.3)
(3.7)
(6.8)
(4.2)
(3.2)
63.71
61.37
113.69
101.12
87.67
427.56
75.21
70.02
135.35
129.73
106.69
517.00
(12.5)
(9.7)
(10.7)
(13.6)
(11.2)
(11.6)
65.85
63.23
120.92
112.13
94.79
456.92
(+0.2)
(+0.4)
(2.7)
(4.3)
(2.7)
(2.3)
65.95
63.46
117.64
107.35
92.26
446.66
for males, 19.2% for females, and 17.9% for
the combined sexes. Although the premolars
reduce to a similar degree from the Mesolithic to Agriculturalist period, little change
in premolar area occurs thereafter. The third
and fourth premolars of the female and combined samples actually exhibit slight increases in occlusal area, unlike any other
area dimensions, while the males show mod-
359
est decreases. At the same time, molar area
decreased by an average of 2.1% in males,
4.9% in females, and 3.2% when the sexes
were combined. In the mandible, it was the
third molar that reduced the most from beginning to end, decreasing by 16.4%in males,
17.9% in females, and 16.7% in both sexes
(Table 4). The mandibular premolars displayed a little more reduction than in the
maxilla, but were still reducing at a slower
pace than the molars. Once again then, it
appears that all teeth generally reduced from
the Mesolithic to the Agriculturalist phase,
whereas only the molars continued that trend
into the Intensive Agriculturalist period.
Wolpoff (1971) has proposed that the selective intensity governing posterior tooth area
exceeds that which acts upon either the
length or breadth dimensions of a tooth, and
consequently the summed areas of the posterior teeth are the best metric measure of
dental function. Hence, the summed posterior area may be considered as a single functional unit. Because of this possibility that
the evolutionary forces may be acting on the
total occlusal area of the posterior dentition
as a single genetic and functional entity,
summed posterior areas are examined here.
As might be expected, trends in summed
posterior area largely conform to those already examined. The Mesolithic sample possesses by far the largest summed areas, as
evidenced by the fact that the females of this
group have larger tooth sums than the males
of the other two groups. Male mandibular
areas reduce 11.51% from the Mesolithic to
Agriculturalist phase, and maxillary areas
display a similar decrease of 11.91%. The
amount of female reduction in posterior tooth
size is slightly smaller, with nearly identical
figures of 11.26%and 11.21%in the mandible
and maxilla, respectively. Combining both
sexes into a single sample also shows a reduction of greater than 10%in both the lower
(10.69%)and upper (11.62%)jaws. Relative to
this comparison, the degree of tooth area reduction from the Agriculturalist to Intensive
Agriculturalist phase drops markedly, to an
average level of under 3%for both the mandible and maxilla in the combined sample.
-
DISCUSSION
This study substantiates previous reports
(Brace and Mahler, 1971;Greene et al., 1967)
Fig. 1. Tooth breadth (in mm) profiles comparing the Mesolithic. Agriculturalist, and Intensive Agriculturalist phases for (a) male mandibular, 01)male maxillary, (c) female mandibular,
(d) female maxillary dimensions.
360
J.M. CALCAGNO
TABLE 4. Mandibular dental areas (in mm') and percentage differences between successive groups (in
parentheses) for the Mesolithic (Meso), Agriculturalist (Ad, and Intensive Agriculturalist (In Ag) phases.
Meso
Males
P3
P4
M1
M2
M3
Summed
Females
P3
P4
M1
M2
M3
Summed
Combined
P3
P4
M1
M2
M3
Summed
(%)
(%I
In Ag
(%I
67.64
67.63
140.39
131.12
125.04
531.82
(14.3)
(8.3)
(11.5)
(11.1)
(12.1)
(11.5)
57.95
62.02
124.20
116.51
109.95
470.63
(1.4)
(3.6)
(3.0)
(3.4)
(4.3)
(3.3)
57.15
59.80
120.49
112.56
105.20
455.20
60.35
61.82
125.32
119.06
113.94
480.49
(11.6)
(9.8)
(9.0)
(12.2)
(13.4)
(11.3)
53.36
55.77
113.99
104.54
98.71
426.37
(1.3)
(2.8)
(3.3)
(1.3)
(4.5)
(2.8)
52.68
54.22
110.25
103.15
94.28
414.58
63.74
64.56
131.77
124.12
119.18
503.37
(12.5)
(8.6)
(9.4)
(10.9)
(12.1)
(10.7)
55.75
59.00
119.38
110.64
104.80
449.57
(1.0)
(2.8)
(2.4)
(2.1)
(4.6)
(2.7)
55.20
57.36
116.58
108.27
99.96
437.37
of large dental dimensions of the Nubian Mesolithic, and documents a substantial amount
of dental reduction thereafter. The teeth of
the Nubian Mesolithic are considerably
larger than those of the European Upper Paleolithic (see Frayer, 1978), despite the
greater age of the latter sample. However,
tooth size in the Intensive Agriculturalist
phase is comparable t o that of certain modern Chinese and Southeast Asian populations that are considered as having some of
the smallest dentitions in the world (Brace
and Hinton, 1981; Brace et al., 1984). Thus,
regardless of the relatively massive teeth
characteristic of Mesolithic Nubia, there is
no reason to suspect that modern day descendants in that region still possess large teeth
relative to other contemporary groups.
Some discrepancy exists between the Mesolithic dimensions of this study and those
reported nearly two decades ago by Greene
et al. (1967) for remains recovered at Wadi
Halfa. Two factors may account for this fact.
First, and more obvious, sample sizes greatly
vary, and the intention of Greene et al. (1967)
was most likely to provide a descriptive analysis of site 6B36 rather than a complete representation of Mesolithic tooth size in Nubia.
The possibility of sampling error may be reflected by the fact that the mandibular
summed posterior area is much larger than
the maxillary total published in that article,
which is the opposite of what generally occurs. Thus, because sample size in the previous study ranges from five to ten for the
individual posterior dimensions,plus the fact
that all specimens came from a single site,
the present analysis should be a more reliable portrayal of tooth size in the Nubian Mesolithic. Greene et al. (1967) reported the
Nubian sample as having a larger dentition
than that found in the Skhfil Neanderthals,
but the metrics obtained from this study are
considerably smaller, and caution should be
used when comparing Mesolithic Nubia to
Neanderthal samples. Between the two Nubian studies, mandibular and maxillary
summed posterior areas drop from 549.0 to
506.1 mm2 and 534.2 to 516.9 mm2, respectively. At 535.4 mm2 for the mandible and
562.1 mm2 for the maxilla (Brace and Mahler, 19711, Neanderthal posterior summed
areas are considerably larger than those of
the Nubian Mesolithic. Moreover, if compared to the Krapina Neanderthals as reported by Wolpoff (19791, which display a
mandibular posterior summed area of 577.7
mm2 and a maxillary figure of 615.3 mm2,
the teeth of the Nubian Mesolithic fail to
even come close in size.
A second factor possibly accounting for the
discrepancy between the figures of Greene et
al. (1967) and the present analysis may relate to the fact that length dimensions are
36 1
DENTAL REDUCTION IN POST-PLEISTOCENE NUBIA
incorporated into the posterior summed
areas, which may be the major source of interobserver error. A comparison of length and
breadth averages between these two studies
reveals a much greater similarity in breadth
dimensions (Table 5). Because the sample
sizes are so different, this may again be
merely an artifact of sampling error. However, it may not be totally coincidental that
breadths can be more consistently measured
and display an average difference of only
1.7%between studies, as opposed to 4.4% for
lengths. Thus, if two researchers possess
slightly different standards regarding the degree of wear needed to prohibit the measurement of a tooth, the primary consequences
are varying lengths and even greater dissimilarity in posterior summed areas, since the
bias that is due to technique is multiplied
into each of the five posterior teeth.
In an attempt t o alleviate this problem,
only breadths may be compared between populations. When this is done, a much closer
correspondence between the dimensions of
Greene et al. (1967) and this study is found.
For example, the summed posterior breadth
for the male maxilla is 57.5 mm in the former
study and 57.8 mm in the present analysis,
and the combination of all breadths produces
an identical figure of 81.1 mm for both studies. Similar results for posterior summed
breadths are obtained for the female maxilla
(55.0 mm, vs. 55.5 mm in this study) and
mandible (50.7 mm, vs. 50.5 mm in this
study). Only the male mandibular figures
vary markedly, with the posterior breadths
of Greene et al. (1967) adding to 55.5 mm as
compared to 52.6 mm in the present analysis.
Thus, although the increased similarity of
tooth size estimates may be coincidental since
there is disparity in the number of specimens
forming the samples of these two studies,
once again breadth dimensions seem increasingly reliable as measures of tooth size for
both intraobserver and interobserver comparisons.
SUMMARY AND CONCLUSIONS
The bulk of post-Pleistocene dental reduction in Nubia occurred from approximately
12,000 years ago to the second millenia B.C.
During this time, virtually every tooth displayed a marked decrease in length, breadth,
and occlusal area, regardless of whether the
sexes were being analyzed as separate or
combined samples. Dental reduction is perhaps best exemplified by viewing changes in
tooth breadth, which was deemed as the most
reliable indicator of tooth size among the various dimensions. These displayed decreases
in size at the .001 level of significance in 15
of 16 variables for both sexes (with each remaining variable also reducing, but at a
lower level of significance).
TABLE 5. Comparative Nubian Mesolithic tooth means (in mm) ofpresent study and Greene
et al. (1967)
LT
I1 Present
Greene
Diff
I2 Present
Greene
Diff
C 1 Present
Greene
Diff
P3 Present
Greene
Diff
P4 Present
Greene
Diff
M1 Present
Greene
Diff
M2 Present
Greene
Diff
M3 Present
Greene
Diff
Mandible
(N)
BR
5.6
(20)
5.7
(4)
1.8%
6.1 (29)
6.2
(3)
1.6%
7.3
(28)
7.4
1.4%
7.3
7.6
3.9%
7.2
7.7
6.5%
11.4
12.1
5.8%
11.1
11.8
5.9%
10.8
11.8
8.5%
(4)
(30)
(5)
(37)
(5)
(31)
(9)
(35)
(8)
(37)
(6)
6.4
6.3
1.6%
6.7
6.6
1.5%
8.2
8.1
1.2%
8.8
8.9
1.1%
9.0
9.2
2.2%
11.5
11.5
0.0%
11.2
11.5
2.6%
11.0
11.5
4.5%
Maxilla
(N)
LT
(N)
BR
(N)
(25)
(4)
9.3
9.8
5.1%
7.3
7.6
3.9%
8.0
7.6
5.0%
7.5
7.6
1.3%
6.9
7.4
6.8%
10.9
11.3
3.5%
10 5
11.1
5.4%
9.1
9.4
3.3%
(26)
(5)
7.5
7.3
2.7%
6.9
6.8
1.4%
8.6
8.5
1.2%
10.0
9.7
3.0%
10.2
10.0
2.0%
12.4
(28)
(5)
(31)
(3)
(32)
(4)
(31)
(5)
(36)
(5)
(29)
(91
(32)
(8)
(37)
(6)
(25)
(5)
(281
(4)
(28)
(9)
(31)
(8)
(32)
(8)
(36)
(9)
(36)
(10)
12.3
(30)
(5)
(31)
(4)
(30)
(9)
(32)
(8)
(29)
(8)
0.8%
12 4
(37)
12.3
(9)
0.8%
11.7 (36)
11.8 (10)
0.8%
362
J.M. CALCAGNO
A much different pattern of dental change
is revealed from the Agriculturalist to Intensive Agriculturalist phase. Whereas tooth
lengths and breadths dropped by more than
6%between the Mesolithic and Agriculturalist phase, the magnitude of tooth reduction
roughly amounts to only one-tenth of that
figure between the two Agriculturalist
phases, and the rate of change is considerably slowed as well (Calcagno, 1984). However, these statistics conceal the fact that an
interesting shift occurred in the latter comparison, which involves more than just a levelling off in the size of all teeth. The posterior
teeth, and particularly the molars, exhibit a
continuation (though of reduced intensity) of
the trend of dental reduction documented
since the Mesolithic, whereas the anterior
teeth show insignificant changes in size,
bringing an end to at least a 10,000-year
course of declining anterior tooth size. In fact,
every significant reduction in tooth size from
the Agriculturalist to Intensive Agriculturalist phase involves a posterior tooth dimension. In summary, post-Pleistocene changes
in the Nubian dentition are characterized by
the marked reduction of all teeth from the
Mesolithic to the Agriculturalist phase,
which is followed by a levelling off of anterior
tooth size together with a lower but continued rate of molar reduction for posterior teeth
between the two Agriculturalist phases.
Through this study, it was also revealed
that although tooth size in Mesolithic Nubia
is certainly massive by modern standards
and is also larger than that of the European
Upper Paleolithic, it is perhaps misleading
to consider it as comparable to some Neanderthals. A more accurate analogy is with
recent aboriginal populations of Australia,
whose dentitions are among the largest
known for recent human groups (Brace and
Ryan, 1980;Brace et al., 1984).Following the
Mesolithic, tooth size in Nubia greatly reduces to a state that, by the time of the Intensive Agriculturalists, could be considered as
average relative to modern populations.
ACKNOWLEDGMENTS
The assistance of Dr. David W. Frayer,
throughout all phases of this research, is
deeply appreciated. Funding for museum
fieldwork was provided by an NSF Doctoral
Dissertation Improvement Grant (BNS8114920) and a Marshall Award from the
American Scandinavian Foundation.
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