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Brief communication High-resolution assessment of the dental developmental pattern and characterization of tooth tissue proportions in the late Upper Paleolithic child from La Madeleine France.

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AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 138:493–498 (2009)
Brief Communication: High-Resolution Assessment
of the Dental Developmental Pattern and
Characterization of Tooth Tissue Proportions
in the Late Upper Paleolithic Child From
La Madeleine, France
Priscilla Bayle,1,2* José Braga,1 Arnaud Mazurier,3 and Roberto Macchiarelli2,4
1
Centre d’Anthropobiologie et Imagerie Anatomique, FRE 2960 CNRS, Université Paul Sabatier, Toulouse, France
Département de Préhistoire, UMR 5198-USM 204, MNHN Paris, France
3
Etudes Recherches Matériaux, Poitiers, France
4
Département Géosciences, Université de Poitiers, Poitiers, France
2
KEY WORDS
dental mineralization; tissue proportions; lower deciduous dentition; late Upper
Paleolithic; microtomography
ABSTRACT
Affinities and differences in dental
maturational patterns between modern humans and
Neanderthals remain a matter of discussion. In particular,
deciduous teeth are rare for Late Pleistocene humans, and
few entire sequences have been detailed for their developmental status. Here, we report the results from the 3D virtual reconstruction and structural analysis of the deciduous lower dentition (nine teeth in situ) of the child from La
Madeleine (LM4), France, the first Upper Paleolithic specimen detailed so far by means of high-resolution microtomography (lCT). With respect to the modern dental developmental standards, age at death of this individual is now
more likely estimated within the interval 3–4 years. LM4
lacks the slight discrepancy between a proportionally
advanced stage of mineralization of the deciduous first
molar and a relatively delayed maturational level of the
incisors, which is found in Neanderthals (Bayle et al.:
J Hum Evol 56 [2009] 66–75). By using a Bayesian
approach, we calculated the probability that its maturational
sequence is found within the extant human variation as
represented by a tomographic (CT) reference sample of 45
children scored according to the same protocol (Liversidge
and Molleson: Am J Phys Anthropol 123 [2004] 172–180).
Results show that the specific sequence of this Magdalenian individual is found three times in the comparative
sample included in this study. LM4 absolute tooth size and
relative dental tissue proportions are close to the modern
human figures (characterized by proportionally reduced
dentine volumes) and lie systematically below the values
shown by the Neanderthal child from Roc de Marsal,
France (OIS 5a). Am J Phys Anthropol 138:493–498,
2009. V2009 Wiley-Liss, Inc.
In his extensive radiographic study of dental developmental patterns in Upper Pleistocene and Holocene
populations, Tompkins (1996a) recorded a high level of
similarity in relative maturation of the permanent teeth
between Neanderthals (OIS 5e-3) and Upper Paleolithic
humans (34-11 kya BP) from Europe and Middle East.
The author (Tompkins, 1996a,b) also noted that only
some minor differences characterize both fossil groups
with respect to the extant human condition. Notably,
Neanderthals and anatomically modern fossil humans
share a relative delay in mineralization of the incisors
and the third premolar associated with a general
advancement of the molars, particularly of the M3.
Recent high-resolution analysis of the immature specimen from Roc de Marsal, France (OIS 5a; Bayle et al.,
2009) shows that a relative developmental delay of the
incisors and a relative advancement of the first molars
also characterize the deciduous lower Neanderthal
dentition.
Deciduous teeth are rare for late Pleistocene humans
(Hillson and Trinkaus, 2002: 356) and few entire sequences of the primary dentition have been qualitatively
and quantitatively assessed so far for their developmental
status (Legoux, 1966; Tompkins, 1991; Henry-Gambier,
2001; Hillson, 2002; Hillson and Santos Coelho, 2002).
By means of high-resolution microtomography (lCT),
here, we detail the maturational condition of the mandibular deciduous dentition of the late Upper Paleolithic
child from La Madeleine, France, and calculate the
Bayesian probabilities that its dental mineralization
sequence is found within extant human variation represented by the comparative deciduous sample included in
this study. Based on recent evidence about the influence
of variations in tooth tissue proportions on dental developmental patterning (Smith et al., 2007a; Bayle et al.,
C 2009
V
WILEY-LISS, INC.
C
Grant sponsor: EU FP6 Marie Curie Actions (EVAN); Grant number: MRTN-CT-2005-019564; Grant sponsor: GDR (CNRS Projects);
Grant number: 2152; Grant sponsor: TNT.
*Correspondence to: Priscilla Bayle, Centre d’Anthropobiologie et
Imagerie Anatomique, Université Paul Sabatier, 37 allées Jules
Guesde, F-31000 Toulouse, France. E-mail: bayle@cict.fr
Received 3 September 2008; accepted 21 November 2008
DOI 10.1002/ajpa.21000
Published online 23 January 2009 in Wiley InterScience
(www.interscience.wiley.com).
494
P. BAYLE ET AL.
2009; Olejniczak et al., 2008), we also quantified and
compared enamel, dentine, and pulp proportions of each
tooth.
MATERIALS AND METHODS
The skeleton of the La Madeleine child (LM4) was discovered in 1926 in Tursac, Dordogne, in a final Magdalenian ochred burial rich of goods and personal ornaments
(Capitan and Peyrony, 1928; VanHaeren and d’Errico,
2001). A recent radiocarbon analysis (14C AMS) of the
burial provided an age of 10,190 6 100 years BP
(Gambier et al., 2000). Skeletal and dental age at death
estimates suggest the interval of 2–4 years for this late
Upper Paleolithic child (Heim, 1991; Gambier et al.,
2000).
Although the LM4 mixed lower dentition also consists
of eight isolated permanent elements, its restored mandible currently bears in situ only nine deciduous teeth,
the right canine being the only missing tooth (Heim,
1991).
In 2007, the LM4 mandible has been detailed by
means of high-resolution lCT at the Centre de Microtomographie of the University of Poitiers (equipment
X8050-16 Viscom AG; camera 1004 3 1004). Scans were
made according to the following parameters: 95 kV, 395
lA current, 32 integrations/projection, and 1,800 projections (each 0.28). The final volume was reconstructed in
984 3 984 format with an isotropic voxel size of 50.9
lm3 using the software DigiCT v.1.15 (DIGISENS).
A semiautomatic segmentation with manual corrections was carried out by means of AMIRA v.4.1.2 (Mercury Computer Systems) and Artecore v.1.0 (NESPOS
Society). Threshold values between segmented components have been found according to the methodology
developed by Spoor et al. (1993). For individual measurements, crowns were digitally isolated from roots following Olejniczak et al. (2008), and surface rendering was
performed using triangulation and constrained smoothing from the volumetric data (marching cube algorithm;
Lorensen and Cline, 1987).
Assessment of the deciduous developmental pattern,
which is based on the 3D independent evaluation of each
virtually extracted element, follows a scoring system
adapted from Liversidge and Molleson (2004; for details,
see Bayle et al., 2009). The sequence of this Magdalenian
individual (first scored by P.B. and A.M. and thus independently validated by J.B. and R.M.) has been compared to those assessed (by P.B. and one independent
observer) following the same scoring method on a computed tomographic (CT) reference sample of 45 living
individuals (29 males and 16 females), mainly of West
European origins, aged 2–5 years and distributed as follows: 2–3 years 5 14 individuals; 3–4 5 19; 4–5 5 12
(record from Braga and Treil, 2007; Bayle et al., 2009;
and original unpublished data).
The statistical analysis of the dental mineralization
sequence was realized by using the Bayesian approach
developed by Braga and Heuzé (2007), already adopted
to characterize the developmental status of the Roc de
Marsal Neanderthal dentition (Bayle et al., 2009).
Accordingly, the LM4 sequence can be represented by a
rearrangement of its five elements (di1-dm2) into a finite
number of combinations, each being composed of two
subsets: the first comprises between 1 and 4 dental elements sampled from the entire set, whereas the second
comprises the remaining elements. In this case, 30 comAmerican Journal of Physical Anthropology
binations are possible. The probability of observing each
of the 30 combinations in the reference sample is calculated by using Bayes’s rule of conditional probability
(Vieland, 1998; Aitken and Taroni, 2004), with teeth
being considered as statistically dependent units and
prior probabilities uniform.
The following 12 linear, surface, and volumetric variables describing tooth tissue proportions were digitally
measured: Vt is the total tooth volume (mm3); Ve, the volume of the enamel cap (mm3); Vd, the total volume of the
dentine (mm3); Vp, the total volume of the pulp (mm3);
Vcdp, the volume of the coronal dentine (including the
coronal aspect of the pulp chamber) (mm3); Vcd, the volume of the coronal dentine (excluding the coronal aspect
of the pulp chamber) (mm3); Vcp, the volume of the coronal pulp (mm3); Vc, the total volume of the crown (mm3);
Vcdp/Vc, the percent of coronal volume that is dentine
and pulp; Sedj, the surface area of the enamel-dentine
junction (mm2); AET, the average enamel thickness
(mm); and RET is the scale-free relative enamel thickness (for methodological details, see Kono, 2004; Macchiarelli et al., 2006; Olejniczak et al., 2008).
Intra- and interobserver tests for accuracy of the
measures were run by two observers (P.B and A.M.). As
a whole, recorded differences are less than 5%, which is
compatible with previous results on similar sets of 2–3D
virtually assessed variables (e.g., Kono, 2004; Suwa and
Kono, 2005; Olejniczak and Grine, 2006; Bayle et al.,
2009).
As the LM4s lower posterior dentition (as well as the
only preserved left canine) is virtually free from wear,
for the molars, the average values have been calculated
for all the variables listed earlier. Conversely, because of
some occlusal dentine patches appearing on the incisors,
the highest value within each tooth pair has been
retained in the quantitative analysis.
To evaluate the influence of variation in tooth tissue
proportions on maturational patterns of the primary
dentition, LM4 has been compared with the Neanderthal
child of Roc de Marsal (left and right average estimates;
Bayle, 2008; Bayle et al., 2009) and to the average values from two modern individuals (MH) of European origin aged 2–4 years, whose unworn lower dentition has
been detailed by means of the same analytical procedures (Bayle, 2008).
RESULTS
The virtually reconstructed deciduous lower dentition
of the La Madeleine 4 (LM4) late Upper Paleolithic child
is shown in Figure 1. Together with the preservation
quality of the dental elements, it also reveals the interventions of restoration on the mandibular body, which
has been reinforced anteriorly by means of a transversally placed metal bar (Heim, 1991).
As recognized by Skinner (1996), the lower left canine
presents a form of localized hypoplasia. Our virtual measurement of this spot, which is set on the midlabial aspect of the crown (see Fig. 1), ranges between 1.47 and
1.52 mm2.
The individual assessment of the developmental stage
of the nine deciduous teeth is shown in Table 1. Three
incisors and the dm1s present a beginning of root resorption (stage r), the left di2 still displaying full apical closure (h2). Compared with the canine root (g), root length
of both dm2s is complete, while apices are not closed
(h1).
DENTAL DEVELOPMENT IN THE LA MADELEINE CHILD
TABLE 1. Individual assessment of the developmental stage of
the nine deciduous teeth of the LM4 child mandible
Left arcade
Tooth
di1
di2
dc
dm1
dm2
Developmental
stagea
Root resorption
initiated (r)
Apical dentine
edge is sharp
and apex is only
just visible (h2)
Root length is
almost complete,
but apical edges
are slightly
converging (g)
Root resorption
initiated (r)
Root length
complete, with
apical walls
converging, but
apex is still open
(h1)
Right arcade
Developmental
stagea
Root resorption
initiated (r)
Root resorption
initiated (r)
Tooth
di1
di2
–
Root resorption
initiated (r)
Root length
complete, with
apical walls
converging, but
apex is still open
(h1)
dm1
dm2
a
Adapted from Liversidge and Molleson (2004); see Bayle et al.
(2009).
With respect to the modern standards of age-related
dental development published by Moorrees et al. (1963),
Ubelaker (1989), and Liversidge and Molleson (2004), as
well as to the reference sample of immature dentitions
available in our files (see earlier), the most likely age at
death interval estimated for this child is between 3 and
4 years.
Within the modern comparative sample of 45 cases
considered in this study, the maturational sequence displayed by LM4 (r-r-g-r-h1) is found three times, in two
male and one female individuals, all aged 3.5 years. In
the reference series, this pattern is the fourth more common among the 25 sequences represented.
For the Bayesian analysis of the LM4 sequence (see
Fig. 2), all theoretically possible probabilities associated
to the formed combinations (n 5 30) have been calculated, the results being distributed as follows: 30% of
probabilities are superior to 0.75 and 70% comprised
between 0.25 and 0.75. Although the value of 0.75 does
not represent an absolute cutoff in a continuous probability distribution ranging from 0 to 1, in such kind of
analyses, probabilities greater than this formal threshold
indicate very likely events, whereas lower values are
more likely associated to random events (Braga and
Heuzé, 2007).
Comparative dental tissue proportions in LM4, Roc de
Marsal (RdM; Bayle, 2008; Bayle et al., 2009), and modern humans (MH; Bayle, 2008) are shown in Table 2.
Although in both fossil specimens the canines and the
molars are virtually free from occlusal dental wear, on
the Magdalenian child, and to a minor extent on Roc de
Marsal, the central incisors show from minimal to extensive dentine patches (stage from 3 to 4 of the scoring diagram elaborated by Smith, 1984), and some minimal
dentine patches also emerge on the lateral incisors
(stage 2/3; Smith, 1984).
For the total tooth volume (Vt), all values displayed by
LM4 are close to the modern figures and lie below the
Neanderthal condition as represented by Roc de Marsal,
which systematically shows larger teeth, especially the
495
incisors. Even in terms of tooth position within the dental arcade, a similar pattern is shown also by the surface
area of the enamel-dentine junction (Sedj). However,
these differences are not homogeneously distributed
among enamel, dentine, and pulp in terms of relative
volumes. In facts, although the Neanderthal deciduous
enamel is absolutely (AET) and relatively (RET) thinner
than Magdalenian and modern enamel, the absolute volumes (Ve) are similar. Conversely, dentine and pulp volumes are significantly larger in Roc de Marsal (for estimates on permanent Neanderthal molars, see Macchiarelli et al., 2008; Olejniczak et al., 2008). This clearly
results from the percent of coronal volume that is dentine and pulp (Vcdp/Vc), the values ranging from 56 to
65% in the Moderns (LM4 and MH) and from 67 to 76%
in the Neanderthal child.
DISCUSSION AND CONCLUSIONS
According to Tompkins (1996a), although dental developmental patterns have continued to slightly evolve
through the Late Pleistocene to the present, both Neanderthals and fossil modern humans are hardly distinguishable from post-Pleistocene populations because of
significant variation in mineralization. Nonetheless, the
high-resolution analysis of the mixed dentition of the
Neanderthal child from Roc de Marsal (Bayle et al.,
2009) showed that neither its deciduous nor its permanent mandibular sequences were precisely found within
a modern reference sample made of 32 and 343 living
children, respectively. In both cases, the most influential
factor affecting the probabilistic analysis is represented
by a slight discrepancy in Roc de Marsal between the
stage of mineralization of the first molar, which is proportionally advanced, and the maturational level
reached by its incisors, which are proportionally delayed.
Although such discrepancies between the Neanderthal
child and a comparative sample of limited size and narrow geographic origin should not be interpreted as necessarily reflecting deep biological differences, it is noteworthy that the deciduous mineralization sequence displayed by the late Upper Paleolithic immature from La
Madeleine (LM4) is found three times in a similar sample of 45 extant individuals, with a rank and a probability distribution suggesting that this pattern is relatively
common, at least in Holocene Western European populations. Also, on radiographic ground, the probabilistic
analysis of the deciduous mineralization sequence displayed by the penecontemporary immature specimen
GE2 from Grimaldi, Italy (Henry-Gambier, 2001) fits
that of LM4 and, again, differs from the Neanderthal
condition. The same is true also for the deciduous mineralization sequence displayed by the Middle Paleolithic
anatomically modern child Qafzeh 13, Israël (Tillier,
1999), which fits the extant human standards.
Even if the canine of the Magdalenian specimen is
affected by localized enamel hypoplasia and some occlusal wear is found on the incisors of both fossil children,
present comparative estimates of relative tooth size,
inner morphology, and volumetric dental tissue proportions point to a closer structural resemblance between
LM4 and the modern human condition, which differs
from the Neanderthal one (Mazurier and Macchiarelli,
2005; Macchiarelli et al., 2006, 2007, 2008; Smith et al.,
2007a; Bayle, 2008; Bayle et al., 2009; Olejniczak et al.,
2008). Accordingly, we suggest that some maturational
characteristics of the deciduous and permanent NeanderAmerican Journal of Physical Anthropology
Fig. 1. Microtomographic-based 3D virtual reconstruction (volume rendered in
transparency) of the LM4 child mandible (in
lateral oblique view) from the late Upper
Paleolithic site of La Madeleine, France,
bearing 10 deciduous teeth. Traces of restoration of the specimen are evident. Scale bar
is 1 cm.
Fig. 2. The maturational sequence
of the LM4 deciduous lower dentition (five elements shown in the
top row). Developmental stages
(r-r-g-r-h1) adapted from Liversidge
and Molleson (2004; details in
Bayle et al., 2009). Distinctly for
any of the 30 calculated combinations (i.e., 100% of theoretically
possible), each cell shows the probability (p) that, given the maturational combination (or unit) below
the line, the specific LM4 developmental subset shown above the line
is found within a comparative reference sample of 45 modern humans
(see Materials and Methods). The
LM4 sequence is found three times
within the comparative sample.
497
DENTAL DEVELOPMENT IN THE LA MADELEINE CHILD
TABLE 2. Linear, surface, volumetric estimates and dental tissue proportions of the La Madeleine (LM4) lower deciduous teeth
compared with those of the Neanderthal child of Roc de Marsal (Bayle, 2008; Bayle et al., 2009) and to the average values from
two modern individuals (MH) showing a similar dental developmental stage (Bayle, 2008)
di1
LM4
MH
LM4/MH
RdM
LM4/RdM
di2
LM4
MH
LM4/MH
RdM
LM4/RdM
dc
LM4
MH
LM4/MH
RdM
LM4/RdM
dm1
LM4
MH
LM4/MH
RdM
LM4/RdM
dm2
LM4
MH
LM4/MH
RdM
LM4/RdM
Vt
(mm3)
Ve
(mm3)
Vd
(mm3)
Vp
(mm3)
Vcdp
(mm3)
Vcd
(mm3)
Vcp
(mm3)
Vc
(mm3)
Vcdp/
Vc (%)
Sedj
(mm2)
AET
(mm)
RET
71.20a
62.43
1.14
125.35a
0.57
10.16a
10.37
0.98
12.72a
0.80
55.03
45.50
1.21
100.63
0.55
6.01
6.56
0.92
12.00
0.50
18.94
16.76
1.13
40.53
0.47
17.50
14.72
1.19
37.42
0.47
1.44
2.03
0.71
3.11
0.46
29.10a
27.13
1.07
53.25a
0.55
65a
62
1.05
76a
0.86
31.66
34.07
0.93
60.48
0.52
0.32a
0.30
1.05
0.21a
1.53
12.04a
11.90
1.01
6.12a
1.97
100.75a
95.84
1.05
159.01a
0.63
16.96a
15.87
1.07
18.49a
0.92
75.67
68.98
1.10
123.04
0.62
8.12
10.99
0.74
17.48
0.46
25.86
20.54
1.26
44.47
0.58
24.12
18.26
1.32
40.55
0.59
1.74
2.28
0.76
3.92
0.44
42.82a
36.41
1.18
62.96a
0.68
60a
56
1.07
71a
0.86
47.11
44.87
1.05
67.61
0.70
0.36a
0.35
1.02
0.27a
1.32
12.17a
12.92
0.94
7.72a
1.58
210.01
183.25
1.15
281.37
0.75
32.87
32.38
1.02
33.16
0.99
150.80
120.23
1.25
187.56
0.80
26.34
30.64
0.86
60.65
0.43
59.76
52.87
1.13
79.57
0.75
56.16
47.54
1.18
70.35
0.80
3.60
5.33
0.67
9.22
0.39
92.63
85.25
1.09
112.73
0.82
65
62
1.04
71
0.91
70.76
71.70
0.99
96.05
0.74
0.46
0.45
1.03
0.35
1.35
11.88
12.03
0.99
8.03
1.48
285.34
284.39
1.00
405.38
0.70
56.82
52.97
1.07
55.11
1.03
196.14
183.47
1.07
281.80
0.70
32.38
47.95
0.68
68.47
0.47
101.26
91.01
1.11
142.22
0.71
89.99
78.78
1.14
122.18
0.74
11.27
12.23
0.92
20.05
0.56
158.08
143.98
1.10
197.33
0.80
64
63
1.01
72
0.89
99.78
103.67
0.96
135.26
0.74
0.57
0.51
1.11
0.41
1.40
12.22
11.36
1.08
7.81
1.57
517.80
446.61
1.16
563.52
0.92
129.94
101.54
1.28
102.14
1.27
327.75
251.62
1.30
359.12
0.91
60.11
93.45
0.64
102.26
0.59
178.02
164.03
1.09
204.67
0.87
163.96
150.59
1.09
177.46
0.92
14.06
13.44
1.05
27.22
0.52
307.96
265.57
1.16
306.81
1.00
58
62
0.94
67
0.87
152.85
150.17
1.02
191.07
0.80
0.85
0.68
1.26
0.53
1.59
15.11
12.35
1.22
9.07
1.67
Vt, total tooth volume (mm3); Ve, volume of the enamel cap (mm3); Vd, total volume of the dentine (mm3); Vp, total volume of the
pulp (mm3); Vcdp, volume of the coronal dentine (including the coronal aspect of the pulp chamber) (mm3); Vcd, volume of the coronal dentine (excluding the coronal aspect of the pulp chamber) (mm3); Vcp, volume of the coronal pulp (mm3); Vc, total volume of
the crown (mm3); Vcdp/Vc, percent of coronal volume that is dentine and pulp; Sedj, surface area of the enamel-dentine junction
(mm2); AET, average enamel thickness (mm); RET, scale-free relative enamel thickness.
a
Estimates affected by occlusal wear.
thal dentition (Legoux, 1965, 1966; Wolpoff, 1979; Tillier,
1996; Tompkins, 1996a; Granat and Heim, 2003; Smith
et al., 2007a; Bayle et al., 2009; but see Guatelli-Steinberg et al., 2005 for maturational similarities to some
modern human groups) may result from the influence on
dental developmental patterning of absolute and relative
differences with respect to the modern human condition
in tissue proportions between front and cheek teeth.
The high-resolution microtomographic investigation
and 3D characterization of immature human fossil specimens are bringing new critical evidence to the understanding of changes in timing and patterning of dental
mineralization and eruption occurred through the Pleistocene (Macchiarelli et al., 2006, 2007; Smith et al.,
2007a,b, in press; Bayle et al., 2009). Nonetheless, to
assess the relationships between dental developmental
patterns and tooth size endostructural variation in anatomically modern fossil humans and to precise the behavioral implications of changes in dental maturational patterns through the Late Pleistocene (Tompkins, 1996a),
the subtle quantitative assessment of Upper Paleolithic
dental sequences deserves greater attention in the future.
ACKNOWLEDGMENTS
The French Musée National de Préhistoire, Les
Eyzies-de-Tayac, kindly granted access to the original
fossil record (courtesy of J.-J. Cleyet-Merle). The UMR
CNRS 5199 PACEA-LAPP (M. Bessou, B. Maureille, and
P. Murail) provided comparative materials for high-resolution analyses. We acknowledge the ESRF beamline
ID17, Grenoble (A. Bravin and C. Nemoz), the Centre de
Microtomographie at the Univ. of Poitiers (P. Sardini),
and the NESPOS Society (www.nespos.org) for technical
collaboration. Thanks to the CHU Pellegrin, Bordeaux
(V. Dousset, C. Douws, and C. Thibaut), and to the Hospital Necker, Paris, for access to their CT datasets (data
anonymized and numbered exclusively storing the information of age and gender). Thanks to M. Coquerelle for
collaboration on the CT record and to L. Bondioli for discussion. The present version was greatly improved by
comments from the editor and two anonymous
reviewers. The original lCT record of the La Madeleine
mandible is available at the NESPOS website https://
www.nespos.org/display/openspace/Home.
LITERATURE CITED
Aitken C, Taroni F. 2004. Statistics and the evaluation of evidence for forensic scientists, 2nd ed. Chichester: Wiley.
Bayle P. 2008. Analyses quantitatives par imagerie à haute résolution des séquences de maturation dentaire et des proportions des tissus des dents déciduales chez les Néanderthaliens
American Journal of Physical Anthropology
498
P. BAYLE ET AL.
et les Hommes modernes. Ph.D. dissertation, University Toulouse III – Paul Sabatier.
Bayle P, Braga J, Mazurier A, Macchiarelli R. 2009. Dental developmental pattern of the Neanderthal child from Roc de
Marsal: a high-resolution 3D analysis. J Hum Evol 56:66–75.
Braga J, Heuzé Y. 2007. Quantifying variation in human dental
developmental sequences. An Evo-Devo perspective. In: Bailey
SE, Hublin J-J, editors. Dental perspectives on human evolution: state of the art research in dental anthropology. Berlin:
Springer. p 247–261.
Braga J, Treil J. 2007. Estimation of pediatric skeletal age using
geometric morphometrics and three-dimensional cranial size
changes. Int J Legal Med 121:439–443.
Capitan L, Peyrony D. 1928. La Madeleine: son gisement, son
industrie, ses œuvres d’art. Paris: Librairie Emile Nourry.
Gambier D, Valladas H, Tisnerat-Laborde N, Arnold M, Besson
F. 2000. Datation des vestiges humains présumés du Paléolithique supérieur par la méthode du carbone 14 en spectrométrie de masse par accélérateur. Paléo 12:201–212.
Granat J, Heim J-L. 2003. Nouvelle méthode d’estimation de
l’âge dentaire des Néandertaliens. L’Anthropologie 107:171–
202.
Guatelli-Steinberg D, Reid DJ, Bishop TA, Larsen CS. 2005. Anterior tooth growth periods in Neandertals were comparable
to those of modern humans. Proc Natl Acad Sci USA
102:14197–14202.
Heim J-L. 1991. L’enfant magdalénien de La Madeleine.
L’Anthropologie 95:611–638.
Henry-Gambier D. 2001. La sépulture des enfants de Grimaldi
(Baoussé Roussé, Italie). Anthropologie et palethnologie funéraire des populations de la fin du Paléolithique supérieur.
Paris: CTHS, Réunion des Musées Nationaux.
Hillson SW. 2002. The dental age-at-death. In: Zilhao J, Trinkaus E, editors. Portrait of the artist as a child—the Gravettian human skeleton from the Abrigo do Lagar Velho and its
archaeological context. Lisbon: Instituto Portugues de Arqueologia. p 242–245.
Hillson SW, Santos Coelho JM. 2002. The dental remains. In:
Zilhao J, Trinkaus E, editors. Portrait of the artist as a
child—the Gravettian human skeleton from the Abrigo do
Lagar Velho and its archaeological context. Lisbon: Instituto
Portugues de Arqueologia. p 342–355.
Hillson SW, Trinkaus E. 2002. Comparative dental crown metrics. In: Zilhao J, Trinkaus E, editors. Portrait of the artist as
a child—the Gravettian human skeleton from the Abrigo do
Lagar Velho and its archaeological context. Lisbon: Instituto
Portugues de Arqueologia. p 356–364.
Kono R. 2004. Molar enamel thickness and distribution patterns
in extant great apes and humans: new insights based on a 3dimensional whole crown perspective. Anthropol Sci 112:121–
146.
Legoux P. 1965. Détermination de l’âge dentaire de l’enfant
néandertalien du Roc-Marsal. Rev Fr Odonto-Stomat 10:1–24.
Legoux P. 1966. Détermination de l’âge dentaire de fossiles de
la lignée humaine. Paris: Maloine S.A.
Liversidge HM, Molleson T. 2004. Variation in crown and root
formation and eruption of human deciduous teeth. Am J Phys
Anthropol 123:172–180.
Lorensen WE, Cline HE. 1987. Marching cubes: a high-resolution 3D surface construction algorithm. Comput Graph (ACM)
21:163–169.
Macchiarelli R, Bondioli L, Debénath A, Mazurier A, Tournepiche J-F, Birch W, Dean C. 2006. How Neanderthal molar
teeth grew. Nature 444:748–751.
Macchiarelli R, Bondioli L, Mazurier A. 2008. Virtual dentitions:
touching the hidden evidence. In: Irish JD, Nelson GC, editors. Technique and application in dental anthropology. Cambridge: Cambridge University Press. p 426–448.
Macchiarelli R, Mazurier A, Volpato V. 2007. L’apport des nouvelles technologies à l’étude des Néandertaliens. In: Vander-
American Journal of Physical Anthropology
meersch B, Maureille B, editors. Les Néandertaliens. Biologie
et cultures. Paris: Editions du CTHS. p 169–179.
Mazurier A, Macchiarelli R. 2005. Anterior deciduous dentition
in Neanderthals: topographic variation in enamel thickness
and inner structural morphology. Bull Mém Soc Anthropol
Paris ns 17:16.
Moorrees CFA, Fanning EA, Hunt EE. 1963. Formation and
resorption of three deciduous teeth in children. Am J Phys
Anthropol 19:99–108.
Olejniczak AJ, Grine FE. 2006. Assessment of the accuracy
of dental enamel thickness measurements using microfocal
X-ray computed tomography. Anat Rec A 288:263–275.
Olejniczak AJ, Smith TM, Feeney RNM, Macchiarelli R, Mazurier A, Bondioli L, Rosas A, Fortea J, de la Rasilla M, GarciaTabernero A, Radovcic J, Skinner MM, Toussaint M, Hublin
J-J. 2008. Dental tissue proportions and enamel thickness in
Neandertal and modern human molars. J Hum Evol 55:12–
23.
Skinner M. 1996. Developmental stress in immature hominines
from Late Pleistocene Eurasia: evidence from enamel hypoplasia. J Archaeol Sci 23:833–852.
Smith BH. 1984. Patterns of molar wear in hunter-gatherers
and agriculturalists. Am J Phys Anthropol 63:39–56.
Smith TM, Reid DJ, Olejniczak AJ, Bailey S, Glantz M, Viola
TB, Hublin J-J. Dental development and age at death of a
Middle Paleolithic juvenile hominin from Obi-Rakhmat
Grotto, Uzbekistan. In: Condemi S, Schrenk F, Weniger G,
editors. Neanderthals, their ancestors and contemporaries.
Dordrecht: Springer (in press).
Smith TM, Tafforeau P, Reid DJ, Grün R, Eggins S, Boutakiout
M, Hublin J-J. 2007b. Earliest evidence of modern human life
history in North African early Homo sapiens. Proc Natl Acad
Sci USA 104:6128–6133.
Smith TM, Toussaint M, Reid DJ, Olejniczak AJ, Hublin J-J.
2007a. Rapid dental development in a Middle Paleolithic Belgian Neanderthal. Proc Natl Acad Sci USA 104:20220–20225.
Spoor CF, Zonneveld FW, Macho GA. 1993. Linear measurements of cortical bone and dental enamel by computed tomography: applications and problems. Am J Phys Anthropol
91:469–484.
Suwa G, Kono RT. 2005. A micro-CT based study of linear
enamel thickness in the mesial cusp section of human molars:
reevaluation of methodology and assessment of within-tooth,
serial, and individual variation. Anthropol Sci 113:273–289.
Tillier A-M. 1996. The Pech de l’Azé and Roc de Marsal children
(Middle Paleolithic, France): skeletal evidence for variation in
Neanderthal ontogeny. Hum Evol 11:113–119.
Tillier A-M. 1999. Les enfants moustériens de Qafzeh. Interprétation phylogénétique et paléoauxologique. Paris: CNRS
Editions.
Tompkins RL. 1991. Relative dental development in Upper
Pleistocene fossil hominids and recent humans. Ph.D. dissertation, University of New Mexico.
Tompkins RL. 1996a. Relative dental development of Upper
Pleistocene Hominids compared to human population variation. Am J Phys Anthropol 99:103–118.
Tompkins RL. 1996b. Human population variability in relative
dental development. Am J Phys Anthropol 99:79–102.
Ubelaker DH. 1989. Human skeletal remains: excavation, analysis, and interpretation, 2nd ed. Washington, DC: Taraxacum
Press.
VanHaeren M, d’Errico F. 2001. La parure de l’enfant de La
Madeleine (fouilles Peyrony). Un nouveau regard sur
l’enfance au Paléolithique supérieur. Paléo 13:201–237.
Vieland V. 1998. Bayesian linkage analysis, or: how I learned to
stop worrying and love the posterior probability of linkage.
Am J Hum Genet 63:947–954.
Wolpoff MH. 1979. The Krapina dental remains. Am J Phys
Anthropol 50:67–114.
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