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Patterns of size variation and correlation in the dentition of the red colobus monkey (Colobus badius).

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 54:139- 146 (1981)
Patterns of Size Variation and Correlation in the Dentition
of the Red Colobus Monkey (Colobus badius)
LARRY R. COCHARD
Department of Anthropology, University of Wisconsin,
Madison, Wisconsin 53 706
KEY WORDS
Growth fields, Tooth-crown size, Colobus
badius
ABSTRACT
There have been numerous studies on variability and correlation
in dental crown size, but the significance of the resulting patterns remains unclear.
Regions of low variation and high correlation have been hypothesized to represent
the poles of Butler’s morphological fields, to be related to absolute tooth size, or to
be related to morphological complexity of the teeth and functional efficiency.
Variation and correlation of tooth lengths and breadths were investigated in 138
red colobus monkeys to further assess the relations among size associations, crown
morphology, and absolute tooth size. In the maxilla and mandible, the postcanine
teeth are the most highly correlated and least variable, followed by the incisors,
then the canines. There are also lower correlations between premolars and molars
than within either group. While there appears to be a relation between degree of
morphological differentiation and levels of correlation and variation, there are no
notable differences in the correlation of opponents along the dental arcade, which is
the most important functional consideration. This suggests that different levels of
correlation and variation within upper or lower teeth are “artifacts” of tooth
dimensions that contribute to different geometric designs in different tooth groups
as the germs develop. This morphological effect is coupled with the influence of
integration fields, indicated by higher variability and lower correlations of the
third molar, the largest or most molarized tooth. It is concluded that there are wide
functional tolerances in occlusion with respect to the gross dimensions of dental
crowns and their interrelationships.
Teeth are one of the most important components of an animal’s adaptation to its environment. Because of this and because of their
abundance in the fossil record, their high genetic component of variation, and their organization as serially homologous organs, variation
and covariation in tooth size have received
much attention. Such studies are important for
diagnosing sexual differences or species differences in the fossil record (Gingerich, 1974;
Gingerich and Schoeninger, 1979) and for making inferences about developmental processes
(Van Valen, 1962). Patterns of variability and
correlation have also been described in terms of
dental growth fields (Dahlberg, 1945; Henderson, 1975;Suarez and Bernor, 1972;Van Valen,
1962), primarily to provide a framework for
understanding evolutionary changes in dentitions. Butler (1939) first applied the concept of
0002-948318115401-0139$01.70
0 1981 ALAN R. LISS, INC.
a developmental field to teeth to explain the
detailed similarities of adjacent teeth, differences in form in the incisor, canine, and postcanine regions of the dentition, and evolutionary changes in gradients of size and form in
the three different regions.
While much attention has been devoted to
patterns of variation and correlation in tooth
size, their developmental and/or functional
significance is not fully understood. The traditional view is that regions of low variation are
the poles, or areas of strongest effect, of Butler’s
morphological fields (Dahlberg, 1945; Schuman and Brace, 1954). Others consider the
highest correlations between adjacent teeth to
represent the polar areas of morphological
fields (Suarez and Bernor, 1972; Henderson,
Received November 1, 1977; accepted July 9.1980.
140
L.R. COCHARD
1975). Olson and Miller (1958) and Kurten
(1953)consider integration (correlation)within
a dentition to be largely a function of relative
tooth size, which is related to Butler’sconcept of
functional fields. Gingerich and Schoeninger
(1979) recently advanced the hypothesis that
patterns of variation in size are related to
occlusal complexity of the teeth. Among a large
series of primate species, they found the lowest
coefficients of variation located in the first or
second molar. They concluded that because of
the more complex morphology of the postcanine
teeth, they must be less variable to occlude with
the same functional efficiency as canines or
incisors.
In the present study, patterns of variation
and correlation in tooth-crown size were investigated in a n arboreal Old World monkey, CoZobus badius, to further our understanding of
factors t h a t influence tooth-crown size. By
comparing these patterns to the fields responsible for each morphological class, i.e., incisors,
canines, premolars, and molars (Dahlberg,
1945), and to Butler’s morphological fields, the
hypothesis was tested that the fields influencing tooth-crown size are closely related to the
fields determining tooth morphology. The red
colobus monkey is suitable for such a study
because of an enlarged third molar, which will
aid in determining the effects of absolute size.
Hominoids, with their reduced third molars,
have usually been the object of study for
crown-size fields.
In this paper, “growth field” is used only in a
descriptive sense, since t h e specific developmental mechanisms for the patterns are unknown.
MATERIALS AND METHODS
The sample ofCoZobus badius consisted of 85
female and 53 male a d u l t monkeys from
Liberia. They were killed for food by natives
occupying the right bank of the Cess River between the villages of Potogle in the north and
Kpeaple in the south. The teeth were measured
by Leutenegger (1971, 1976) with a sliding
caliper, calibrated to 0.1 mm and with arms
ground to fine points. An average error of
0 . 2 m m was obtained on repeated measurements. The data consist of mesiodistal and
buccolingual diameters of all teeth on the right
side of the maxillary and mandibular dentitions. Only right teeth were measured because
there are no significant differences between the
size of right and left teeth (Ashton and Zuckerman, 1950). If a right tooth was missing, its
antimere was substituted. Mesiodistal diame-
ter (length) was defined as the maximum distance between the mesial and distal contact
points, parallel to the tooth row. Buccolingual
diameter (breadth) was measured as the
maximum distance between the buccal and lingual surfaces, perpendicular to the mesiodistal
diameter. Both dimensions of the canines and
breadth of the incisors were taken at the base of
the crowns.
Matrices of correlation coefficients ( r )for all
possible pairs of teeth were computed for
lengths and breadths. The correlation matrices
for each variable were compiled separately for
males and females and for maxillary and mandibular teeth. For each matrix, the hypothesis
tested was that correlations for all pairs of adjacent teeth are samples of a common parameter
(Snedecor and Cochran, 1967:186). Fisher’s z
scores were used for tests of significance in
comparing individual correlation coefficients.
The patterns of correlation and the distribution
of the coefficients of variation were compared
with each other, with the posterior shift in
molarization due to a n enlarged third molar,
and with the division of the dental arcade into
regions corresponding to the four morphological classes of teeth. The results were compared
with other studies of variation and/or correlation of tooth size in mammals. All data were
processed on a Univac 1110 computer using
programs from the Statjob Series of the madison Academic Computing Center, University of
Wisconsin. Since some teeth were missing or
damaged, a bivariate subsample method was
employed for treating missing data; each correlation coefficient was computed for data pairs
in which values for both variables in the pair
are present. Thus, individual sample sizes for
the correlations range from 76 to 85 for females
and from 40 to 51for males. Wherever male and
female data have been combined, a weighted
average for the means of each sex was computed.
RESULTS
Correlation matrices for the buccolingual
breadths of maxillary and mandibular teeth
are given in Table 1and for mesiodistal lengths
in Table 2. Values for males are found above the
diagonals and for females below. Correlations
greater than r = 0.27 (n = 51) and r = 0.21
(n = 85) are significantly different from zero
(P < .05) for males and females, respectively.
Although the correlations tend to be higher for
females, of 112 male-female comparisons for
individual correlation coefficients, eight are
significantly different a t the 5% level, which
141
SIZE ASSOCIATIONS IN THE DENTITION OF COLOBUS BADIUS
T A B M 1 . Correlation matrices for the buccolingual widths of the maxillary and
mandibular dentitions of Colobus badiuq intradentition correlations for males
( n = 46-51) are above the diagonals, for females f n = 78-85], below
Maxilla
1'
11
I'
C'
p:'
p4
M'
M'
M"
.73*
.50
.48
.55
.50
52
.50
I'
C'
Pt
P
MI
M'
MJ
.49
.45
.52
.36
.32
.49
.34
58
.66
.71
.44
.67
.61
.57
.71
.37
.60
.63
.59
.73
.86
.40
.60
.69
56
.75
.83*
.90
.54
.50
.49
.46
.45
.48
.45
.45
.46
.38
.39
.I4
.49
.49
.46
.65
.68
.80
.67
.64*
.84
P4
M,
MI
M3
.46
.29
.20
.33
52
56
.29
.29
.36
.52
.89
.59
.39
.37
.40
.49
.79
.85
Mandible
1,
.74
11
12
C,
P,
P4
MI
MI
M3
12
.79
53
.27
.43
.48
.44
.44
.59
.31
.44
.46
.45
.51
C,
P:,
.30
.18*
.26
.47
.39
.40
.40
.35
.28
.35
.57
.32
.27
.24
*Significantly different from the opposite sex a t P
=
.29
.35
.40
55
57
.58
52
.80
,653
.78
.05.
TABLE 2. Correlation matrices for the mesiodistal lengths of the maxillary and
mnndibular dentitions of Colobus badius; intradentition correlations for males
f n = 41-51) are aboue the diagnoals, for females f n = 78-85), below
I'
I'
12
C1
PJ
P
M'
M'
MJ
12
50
Maxilla
C'
.48
.32
.65
.52
.38
.47
.55
.45
.25
.55
.38
.42
.30
.27
.20
11
1,
C,
,563
23
.24
p:'
P4
M'
M'
M I
.25
.43
.46
.43
.28*
.68*
.26
.29
26
.48
.47
.26
.41
.35
.57
.60
.66
.18
.30
.25
.35
24
.53
.I2
.20*
.61*
.84*
.43
.44
.43
.68*
50
.45
.42
.48
.49
.46
.73
.43
.51
p:,
P4
Mi
M:!
M3
.31
.31
.35
.29
.42
.26
.26
.38
.22
.08*
-.08*
.53
.48
.32
.I3
.30
.72
.33
.17
.17
.05
.44
.24
.37
Mandible
1,
2,
p,
P4
M,
M,
M3
.69
.01
.I4
.24
.45
.32
.14
.12
.27
.30
.28
26
.15
.37
.22
.31
.25
.32
'Significantly different from the opposite sex at P
.53*
.28*
.40
.31
=
-05
.28
.20
.20
.30
.66
.34
.37
142
L.R. COCHARD
0.9
0.8
0.8
0.7
0.7
0.6
0.6
0.5
0.5
0.4
0.4
0.3
0.2
L'
h
0.6
0.5
0.4
0.3
0.2
I
I'
'
l2
' ' ' ' ' '
C
P3
P4
M'
M2
0 .1
M3
Fig. 1. Correlations (rj for the mesiodistal length (L) and
buccolingual breadth (B) of adjacent maxillary teeth.
Fig. 2. Correlations (r)for the mesiodistal length (L) and
buccolingual breadth (B) of adjacent mandibular teeth.
approximates the expected sampling error. In
each table, the highest correlations are found
between adjacent teeth and generally decrease
inversely with the number of intervening
teeth. But in Table 2, a n irregularity is evident
in the matrix for the lengths of male mandibular teeth. The correlations between the third
mandibular premolar (PJ and all teeth distal to
its hover around zero, including a value of
r = 0.08 for P, and P,. This is probably an artifact of the rotation of the sectorial PB,since its
mesiodistal length was measured and not its
maximum length. The spurious correlations of
P:, were not included in any calculations nor
considered in discussion of the results.
Figures 1and 2 plot correlations for adjacent
teeth and indicate two general patterns for the
distribution of the values. Though the correlations are generally high between adjacent
teeth, the hypothesis that they are identical is
rejected at the 5% level for both variables in
both jaws and both sexes. The correlations are
not equally distributed along the tooth row, but
generally increase both mesially and distally
from the canine region. This is also evident in
comparing the lowest correlation, involving the
canine and either the second incisor or the third
premolar, with the highest value among the
molars. The difference is significant (P < .05)
in seven of eight cases. Superimposed upon this
trichotomy of the anterior, canine, and postcanine teeth, there appears to be a relation
between the patterns of correlations and each
morphological class, i.e., there is a further distinction between the premolars and molars.
Figures 1 and 2 indicate that in six of seven
plots (the spurious correlation of male P:,length
disregarded), the peaks fall within the morphological classes while the lows occur between
them. The correlations of P3-P4 and the molars
Ml-M2 are both significantly higher (P < ,051
than for P 4 M 1 in two cases: female maxillary
and female mandibular lengths.
Similar to the general patterns for correlations between adjacent teeth, the coefficients
of variation for length and breadth (Fig. 3 and
4) distinguish the canine region from the incisors and postcanine teeth. The coefficients
gradually increase from the first or second
molar to the central incisor, but in each case the
gradation is interrupted by elevated values for
the canine and/or first premolar. The patterns
of variation and correlation in this sample are
not only concordant in their distributions, but
also in magnitude. Those teeth that are most
highly integrated in size are least variable in
size; thus the sequence of increasing variability
is postcanine teeth, incisors, and canine region.
Correlations between the size of occluding
teeth were also calculated (Table 3). They are
similar in magnitude to the correlations between adjacent teeth but reveal no major differences between the three regions of the dentition. While the values for the length of the
143
SIZE ASSOCIATIONS IN THE DENTITION OF COLOBUS BADZUS
i
4
.\‘
___ d
-0
.-- -.
-c,
c. v.
c. v.
3
i
1
I’
I*
C
P3
P4
M’
M2
M3
Fig. 3. Coefficientsofvariation (C.V.) for the mesicdistal
length (L) and buccolingual breadth (B) of maxillary teeth.
Fig. 4. Coefficients of variation (C.V.) for the mesicdistal
length (L)and buccolingual breadth (B)ofmandibularteeth.
upper and lower canines in both sexes are low,
the coefficients for the canine breadths are
comparable to those of the other teeth. For
lengths, the values in the center of the molar
region are slightly higher than for incisors, but
the reverse is true for breadths. One important
point is the difference in values between the
second and third molars. For breadths, the values are a little higher for the third molars, but
in both males and females, the correlations between lengths are significantly lower (P < ,051
for the third molars.
The distribution of the correlation coefficients and coefficients of variation relate well
to the regions of morphological differentiation
but not to the absolute size of the postcanine
teeth. The third molar is larger than the second, with 3% more surface area in the upper
and 28% more in the lower tooth (sexes combined). But in general, the size of the third
molars are more variable, more weakly intercorrelated than both the first upper and lower
molars (M,-M1) and the second pair (M2-MS),
and more weakly correlated with M2 than M2
is with M1.
pared schematically in Figure 5 with Butler’s
(1939) fields of incisiform, caniniform, and
molariform differentiation and with the fields
responsible for each morphological class (Dahlberg, 1945). The darkest areas are the poles of
the fields, indicated here as teeth with the lowest amount of size variation or the highest
intercorrelation in size of adjacent and occluding teeth. The third molar is the polar tooth of
Butler’s postcanine field, since the concept of
molarization includes absolute size in addition
to the number of cusps on a tooth. The fields for
each morphological class have no poles because
they are qualitatively different.
While the incisor, canine, and postcanine regions appear independent with respect to the
patterns, the poles within the postcanine teeth
are not concordant, suggesting that fields influencing variation and correlation are somewhat independent of the relative size, or degree
of molarization, of t h e poscanine teeth.
Gingerich and Schoeninger (1979) also concluded that patterns of variation do not represent the stable poles of Butler’s fields and found
no relation between size variation and time of
eruption. They proposed their occlusal complexity-functional efficiency theory, that teeth
with more complex morphology must be less
variable than more simple teeth to occlude a t
the same functional level. While it seems likely
that there is some relation among occlusal
complexity, size restraints, and functional efficiency, covariation in the size of occludingteeth
DISCUSSION AND CONCLUSIONS
The continuous distribution of correlations
and coefficients of variation along the dental
arcade suggests some type of common influence
regulating the size of developing tooth crowns.
To help determine the significance of these patterns, the field effects of the results are com-
144
L.R. COCHARD
would seem to be a more important limiting
factor in occlusion than absolute variation. A
greater amount of variability in any region of
the dentition could be tolerated a s long as small
or large teeth occlude with teeth that deviate
from the mean in the same direction.
While the incisor, canine, and postcanine
teeth differ in levels of variation in both jaws of
the red colobus, there are no outstanding differences in the covariation of occluding teeth
along the dental arcade. The correlations between opposing molars are generally no higher
than for opposing incisors, even though the latter are more variable in size and have a more
simple edge to edge occlusion. Size correlations
between the lengths of upper and lower canines
are low, but this may be a sampling artifact. In
other mammals, including t h e orangutan,
brown bear, red fox, and bobcat, the corre-
TABLE 3. Correlations between homologous
dimensions of occluding teeth
Length
11-1,
1'-12
C'-C,
PI-P,,
P-P,
MI-M,
M'-M2
M'-M,,
Breadth
males
females
males
females
.55
.23*
.23
.45
.50
.67
.63
.37
.64
.57*
.36
.52
.46
.67
.70
.50
.59
.63
.59
.44
.49
.54
.51
.67
.61
.67
.49
*Significantly different from the opposite sex at P
l n c i s vizat i o n ,
c a n i i ization,
and nolariza t ion:
Morp rological c l a s s :
=
.40
.50
.52
.59
.64
.05
I1
...
j -1
12
lations for the length of opposing canines are
among the highest in the dentitions (Kurten,
1953). The same is true for the canines of the
gorilla, even though they are also among the
most variable teeth in length and breadth
within either sex (Suarez and Bernor, 1972).
The occlusal complexity-functional efficiency
theory also implies t h a t t h e same size
restraints that influence the first and second
molars should influence the third molar in the
red colobus, because it is the largest tooth; but
it is more variable, and the correlations between opponents or with M2 are generally
lower for the third molar than for the other
molars.
At least two factors may account for the metric patterns in this study: (1)size relations are
"artifacts" of the action of morphological fields,
and (2)a field for integration of the teeth is
acting independently of all other fields. Although the metric patterns do not correlate
well with the absolute size of the postcanine
teeth, they do correlate well with the degree of
morphological differentiation among the four
classes of teeth. This is particularly evident in
the patterns of correlations. The hypothesis
t h a t tooth germs a r e controlled by developmental regimes that influence both morphology of the crowns and size relations with
other teeth may be described in relation to
growth and development as follows: Whether or
not a tooth germ differentiates into an incisor,
canine, premolar, or molar depends on the
amount of growth of the germ in three dimensions. In this sense, the concept of form involves
a n element of size in each dimension. The
C
P3
1-
P4
M1
I.'..:'.:;.;:*.;:
M2
M3
::I
Co r r e l a t i o n (adj.) :
Correlation (opp.):
Size variation:
Fig. 5. Crown-size fields in Colobus badius compared with the fields for the morphological classes (Dahlberg,'45)and for
incisivization, caninization, and molarization (Butler, '39).Darkened areas represent the poles of the fields. Size correlations
of adjacent (adj.)and occluding (opp.)teeth are shown separately.
SIZE ASSOCIATIONS IN THE DENTITION OF COLOBUS EADIUS
growth of a tooth germ in breadth, for example,
will be coordinated with growth in other dimensions to produce a specific geometric shape.
It would seem likely, then, t h a t the final
breadths of a molar series would share an element in common that is not present in the
breadths of teeth genetically programmed to
participate in a different geometric design. The
common element may be manifest in size correlations, canalization (and the resultant variability), or any process that is responsible for
the final size of the teeth. Of course, this morphological effect is operating concurrently with
a number of other factors that may influence
size relations, such as proximity of tooth germs,
integration fields, or factors that affect the size
of an entire dentition.
This morphological hypothesis is supported
by factor analyses of tooth-crown size. In a n
analysis of tooth-crown size in eight primate
species, the projecting canines of the nonhuman species were isolated in the terminal
factor solutions,while the incisiform cenines of
the human samples were grouped with the incisors (Henderson, 1975). Premolars and molars show some independence in factor analyses
of crown size in human populations (Lombardi,
1975; Suarez and Williams, 19731,even though
both are grinding teeth. Even the shape of tooth
crowns is not independent of morphology. In
human populations, correlations between the
shape (length/breadth) of adjacent dental
crowns are 1% higher within tooth groups
than between (Garn et al., 1967).
The higher variability and lower correlations
of M3 may be due to its position at the periphery
of the tooth row, and hence, its lower integration, even though it is a larger tooth. Data from
Swindler (1976) indicate that many macaque
and baboon species have enlarged third molars
that are more variable in size than the second
molars. In addition to fields responsible for
tooth form, there may be a field for integration
operating independently and wholly within the
constraints for minimal functional efficiency.
Integration fields may also be responsible for
the higher coefficients of variation in the canines. A small canine morphological field
wedged between two much larger fields may
result in some developmental instability. But
as mentioned before, the magnitude of variation along the tooth row is not critical. Developmental influences or disturbances that vary
by region in the dentition can be tolerated since
“. . . most of the factors influencing dentition
size operate equally on both the maxillary and
mandibular dentitions” (Lavelle, 1976:122).
145
Even though distinct patterns of correlation
and variation emerge, the relationship between functional efficiency, size associations,
and the complex interplay of the teeth in mastication remains enigmatic. Explaining patterns
of tooth size as artifacts of morphological fields
or integration fields implies that they have little functional significance or that there is a
wide functional tolerance i n relation to the
gross dimensions of dental crowns. Olson and
Miller (1958:199) applied a n elaborate correlation technique to 83 measurements of the upper
and lower molars and P4 of the South American
owl monkey, Aotus triuirgatus,and found that
integration among the teeth is “. . . low when
compared to integration found in such functional systems as the limbs or axial structures
of various vertebrates.” Even integration between the parts of upper and lower teeth that
meet in occlusion is low in the owl monkey. In
the present study, none of the correlations,
either between adjacent or occluding teeth, are
particularly high, indicating that little of their
variance is shared. Much more work on mechanisms controlling tooth development and
occlusal relations is necessary.
ACKNOWLEDGMENTS
I would like to thank Dr. Walter Leutenegger
for permission to use the data and for commenting on the manuscript, Mr. James Kelly for
assistance in preparing the data, and Ms Susan
Larson and Dr. James Cheverude for their
helpful suggestions. I would also like to thank
Dr. Richard Osborne, Dr. Kenneth Bennett,
and a n anonymous reviewer for their comments on an earlier draft of the paper.
LITERATURE CITED
Ashton, EH, and Zuckerman, S (1950)The influence of geographic isolation on the skull of the green monkey (Cercopithecus aethiops sabueusi: I. A comparison between the
teeth of the St. Kitts and the African green monkey. Proc.
Roy. SOC.,137B:212238.
Butler, PM 11939)Studies of the mammalian dentition: Differentiation of the post-canine dentition. Proc. Zool. SOC.,
London, 109E: 1-36.
Dahlberg, AA (1945) The changing dentition of man. J. Am.
Dent. Assoc., 32:67&690.
Garn, SM, Lewis, AB, and Kerewsky, RS (1967) Shape similarities throughout the dentition. J. Dent. Res.,461481.
Gingerich, PD (1974) Size variability of the teeth in living
mammals and the diagnosis of closely related sympatric
fossil species. J. Paleontol., 48:89&903.
Gingerich, PD, and Schoeninger, MJ (1979)Patterns of tooth
size variability in the dentition of primates. Am. J. Phys.
Anthropol., 51r457-466.
Henderson, AM (1975) Dental Field Theory: An Application
to Primate Dental Evolution. Ph.D. dissertation, Univ. of
Colorado.
146
L.R. COCHARD
Kurt&, B (1953)On the variation and population dynamics
of fossil and recent mammal populations. Acta 2001.Fennica, 76:l-122.
Lavelle, CLB (1976)Odontometric study of African monkey
teeth. Acta anat., 96:115-127.
Leutenegger, W (1971) Metric variability of the postcanine
dentition in colobus monkeys. Am. J. Phys. Anthropol.,
35: 91- 100.
Leutenegger, W (1976) Metric variability in the anterior
dentition of African colobines. Am. J. Phys. Anthropol.,
4545-52.
Lombardi, AV (1975) A factor analysis of morphogenetic
fields in the human dentition. Am. J. Phys. Anthropol.,
42~99-104.
Olson, EC, and Miller, RL (1958)Morphological Integration.
Chicago: University of Chicago Press.
Schuman, EL, and Brace, CL (1954)Metric and morphologic
variations in the dentition of the Liberian chimpanzee;
comparison with anthropoid a nd human dentitions.
Human Biol., 26:239-268.
Snedecor, GW, and Cochran, WG (1967)Statistical Methods.
Ames, Iowa: Iowa State University Press.
Suarez, BK, and Bernor, R (1972)Growth fields in the dentition of the Gorilla. Folia primat., 18:35&367.
Suarez, BK, and Williams, BJ (1973) Dental growth fields
and premolar morphology. J. Dent. Res., 52:632.
Swindler, DR (1976) Dentition of Living Primates. New
York: Academic Press.
Van Valen, L (1962) Growth fields in the dentition ofPeromyscus. Evolution, 16272-277.
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correlation, badius, dentition, colobus, patterns, monkey, variation, red, size
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