18      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

Deciduous Molar Morphology from the Neolithic Caves of the 
Meuse River Basin, Belgium 
 

Frank L’Engle Williams
1*

,
 
Rebecca L. George

2
, and Caroline Polet

3
 

1 
Department of Anthropology, Georgia State University, Atlanta, GA 30303 USA 

2 
Department of Anthropology, University of Nevada, Reno, NV 89557 USA 

3 
Directorate Earth & History of Life, Royal Belgian Institute of Natural Sciences,  

1000 Brussels, Belgium 

There are nearly 200 karstic caves of the Meuse 
River Basin of Belgium that preserve collective bur-
ials dating to the late Neolithic (Semal et al., 1999; 
Toussaint et al., 2001; Toussaint, 2007; Polet, 2011). 
Since habitation sites are rare, these funerary caves 
and rockshelters provide the principal source of 
information about these prehistoric farmers of the 
late Neolithic and the transition to the Bronze Age 
(Toussaint, 2007; Polet, 2011). The mortuary prac-
tices of Neolithic peoples from this region vary 
considerably. Some tombs contain a single burial, 
whereas others include two or more individuals 
(Toussaint et al., 2001), although the great majority 
are collective burials (Polet, 2011). The bones of 
multiple individuals are comingled in some caves. 
Burials are rarely found in full articulation, except 
in cases where single individuals are interred 
(Toussaint, 2007). At some caves, there is regroup-
ing of bones into elements, such as circles of crania 
and bundles of long bones (Toussaint, 2007). Some 
individuals are completely macerated as evidenced 
by flint tool use (Polet, 2011), whereas others are 
cremated remains (Toussaint, 2007). At Bois Mad-
ame, a site in the Burnot Valley (Figure 1), it is un-
clear whether individuals were buried as the bones 
are found in an unordered manner (Dumbruch, 
2003). 

There may be several explanations to account 

for commingling within the collective burials, in-
cluding the actions of burrowing and scavenging 
animals, geological or hydrological effects, recent 
human activity from grave robbers and cave ex-
plorers, or the intentional manipulation of the re-
mains by those who deposited the deceased. Inten-
tional manipulation may have several motivations, 
including burial rites, secondary reburial and cre-
ating space for additional bodies (Dumbruch, 2003; 
Toussaint, 2007).  

About 40% of nearly 600 individuals excavated 
from 34 sites along the Meuse river system are 
subadults (Toussaint et al., 2001; Toussaint, 2007). 
At Bois Madame  nearly a third of the individuals 
(33%) are identified as children (Dumbruch, 2003, 
2007). More recent excavations with improved 
techniques are able to capture additional subadult 
remains, raising the proportion of children to 50% 
(Toussaint, 2007).  

Four cave sites contain numerous subadult re-
mains, including Hastière Caverne M, Sclaigneaux, 
Bois Madame and Maurenne Caverne de la Cave, 

ABSTRACT  The karstic caves of the Meuse River Basin in Belgium preserve nearly 200 collective burials 
dating to the late Neolithic period. Among these, the cave burials of Hastière Caverne M, Sclaigneaux, 
Bois Madame and Maurenne Caverne de la Cave are represented by numerous individuals and radio-
carbon dated to circa 4,635 to 3,830 years B.P. Dental casts from mandibular and maxillary deciduous 
molars are scored using multiple methods to provide a regional overview of the prevalence and expres-
sion of deciduous molar crown traits, and to compare frequencies between cave burial sites with a focus 
on temporal differentiation. Carabelli’s trait varies from a small pit to a full cusp, the largest of which 
are found at Hastière Caverne M. The hypoconulid ranges from moderately large to very large. A meta-
conulid is absent or small. Although the results are contingent on idiosyncratic preservation, differences 
in the frequencies of expression of Carabelli’s trait, a pronounced hypoconulid, and the presence of a 
metaconule and protostylid separate the earlier cave burial at Hastière Caverne M from the final/late 
Neolithic sites of Sclaigneaux and Bois Madame.   

*Correspondence to:   
Frank L’Engle Williams, PhD 
Department of Anthropology 
Georgia State University 
Atlanta, GA 30303 USA 
frankwilliams@gsu.edu 

Keywords: Hastière Caverne M; Sclaigneaux; Bois Madame; Maurenne Caverne de la Cave  



19      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

radiocarbon dated to circa 4,635 to 3,830 years B.P. Two of 
the four burials are from Hastière rockshelter and include 
Hastière Caverne M and Maurenne Caverne de la Cave 
(see Figure 1). Hastière Caverne M has been radiocarbon 
dated to 4,345 ± 60 (AMS OxA-6558; Bronk-Ramsey et al., 
2002) and can be considered early/late Neolithic.    

Both Sclaigneaux and Bois Madame can be considered 
final/late Neolithic. The collective burial at Sclaigneaux 
cave is dated to 4,155 ± 35 (Paepe, 2007). Bois Madame is 
one of the latest sites in the sample, and the two dates ob-
tained, 4,075 ± 38 (AMS OxA 10831) and 3,910 ± 40 (AMS 
OxA 10830), suggest the cave was in use during a relative-
ly narrow time frame (Dumbruch, 2003, 2007; Toussaint, 
2007).  

Maurenne Caverne de la Cave from the Hastière rock-
shelter is associated with four dates spanning ~800 years, 
including 4,635 ± 45 (AMS OxA-9025), which is consid-
ered middle Neolithic (Toussaint, 2007) 4,160 ± 45 (AMS 
OxA-9026), 3,950 ± 70 (Lv-1483) and 3,830 ± 90 (Lv-1482), 
all of which are final/late Neolithic  (Bronk-Ramsey et al., 
2002; Toussaint, 2007). Since Maurenne Caverne de la 
Cave includes such a wide range of radiocarbon dates, it 
cannot necessarily be considered “early” or “final” late 
Neolithic. 

Given the disparate funerary contexts that exist at Ne-
olithic Belgian sites, the number of deciduous molars, and 
the high heritability of dental morphology (Turner et al., 
1991; Scott and Irish, 1997, 2017; Irish, 2006; Pilloud and 
Larsen, 2011; Paul and Stojanowski, 2015, 2017; Pilloud et 
al., 2016; Scott et al., 2018), it is possible that differences in 
dental remains from each cave burial will parallel the 
chronological distinctions between sites (Bronk-Ramsey et 
al., 2002; Paepe, 2007; Toussaint, 2007). Previous studies 
of the osteological remains have concentrated on dental 
microwear, isotopic signatures, stature estimation, patho-
logical conditions, and funerary ritual (Semal et al., 1999; 
García-Martin, 2000; Orban et al., 2000; Toussaint et al., 
2001; Toussaint, 2007; Polet, 2011). The expression of non-
metric deciduous dental traits from these cave burials has 
not been previously explored. The aim of the study is to 
provide a regional review of the incidence and expression 
of crown traits on the maxillary and mandibular decidu-
ous molars, and to examine the frequencies of traits across 
cave burials with respect to temporal variation.   
     On the basis of chronology, we expect the early/late 
Neolithic site of Hastière Caverne M to be distinct from 
the final/late Neolithic sites of Sclaigneaux and Bois Mad-
ame. We anticipate Maurenne Caverne de la Cave to pre-
sent the most variation in dental morphology given its 
relatively broad time frame. Ecogeography may also ex-
plain the results; the two cave burials from Hastière rock-
shelter (Hastière Carverne M and Maurenne Caverne de 
la Cave) may resemble each other and differ from 
Sclaigneaux and Bois Madame (Figure 1).  
 
 

 
 
 
Materials  
Deciduous molars were examined from Hastière Caverne 
M, Sclaigneaux, Bois Madame and Maurenne Caverne de 
la Cave for a total of 27 individuals (Table 1). These 
gnathic remains are isolated such that each fragmentary 
molar and adjoining alveolus can be considered the sole 
remains from a given individual. Although there are hun-
dreds of isolated teeth, only in situ deciduous molars were 
considered. The stages (1-8) created by Smith (1983, 1984) 
were used to characterize the wear on the deciduous mo-
lars (Tables 2-5).  
 
 

 

Figure 1. Map of Belgium shows the location of 
Hastière rockshelter (Hastière Caverne M and Mau-
renne Caverne de la Cave), Bois Madame and 
Sclaigneaux. Sclaigneaux is approximately 35 km 
northeast whereas Bois Madame is circa 15 km north of 
Hastière rockshelter (drawing: ADIA ©). 

Cave burial Maxillae Mandibles Individuals 

Hastière 
Caverne M 

3 2 5 

Sclaigneaux 8 1 9 

Bois Madame 2 4 6 

Maurenne 
Caverne  
de la Cave 

5 2 7 

Total 18 9 27 

Table 1. Neolithic samples  



20      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

Identifier Preservation Dental wear (Smith, 1984) 

Hastière 38 Nearly complete maxilla extending from the left unerupted 
crown of M1 to the empty M1 crypt on the right 

dm1 = stage 4; dm2 = stage 3 

Hastière 381 Right maxillary alveolar fragment with dm1 and dm2 dm1 = stage 4; dm2 = stage 3 

Hastière 39 Small maxillary alveolar fragment with dm1 and dm2 dm1 = stage 3; dm2 = stage 2 

Hastière 18 Mandibular corpus extending from the left ramus base to the 
right dm1; only the left dm1 and dm2 are preserved 

dm1 and dm2 = stage 2 

Hastière 19 Partial mandibular corpus extending from the left dm2 to the 
right damaged dm1 

dm2 = stage 3 

Table 2. Preservation and wear at Hastière Caverne M, an early/late Neolithic cave burial 

Identifier Preservation Dental wear (Smith, 1984) 

Sclaigneaux 115 Relatively complete lower maxilla and dental arcade holding dm1 and dm2 on 
both sides 

dm1 = stage 5; dm2 = stage 4 

Sclaigneaux 116 Relatively complete lower maxilla with dm1 and dm2 preserved on both sides dm1 and dm2 = stage 3 

Sclaigneaux 117 Left maxillary fragment with dm1 and dm2 dm1 and dm2 = stage 3 

Sclaigneaux 118 Left maxillary fragment, preserving dm2 and M1 dm2 = stage 5 

Sclaigneaux 119 Left maxillary fragment and palate, preserving dm1, dm2 and M1 dm1 = stage 6; dm2 = stage 4 

Sclaigneaux 122 Left maxillary fragment with the deciduous molars preserved; dm1 is slightly 
chipped 

dm1 = stage 7; dm2 = stage 4 
  

Sclaigneaux 124 Small right maxillary fragment holding dm1, dm2 and M1 dm1 = stage 5; dm2 = stage 3 

Sclaigneaux 125 Well-preserved right maxilla, with dm1 and dm2 dm1 = stage 5; dm2 = stage 4 

Sclaigneaux 82 Small left corpus fragment holding dm1 and dm2 dm1 = stage 5; dm2 = stage 4 

Table 3. Preservation and wear at Sclaigneaux, a final/late Neolithic cave burial 

Identifier Preservation Dental wear (Smith, 1984) 

BM mx 26 Left fragmentary maxilla, preserving dm1 and dm2 dm1 and dm2 = stage 4 

BM mx 27 Left partial maxilla including most of the palate and alveolus, along with dm1, dm2 
and M1 

dm1 = stage 5; dm2 = stage 4 

BM Md 27 Right mandibular corpus fragment with a partial ramus including dm1, dm2 and M1 dm1 = stage 5; dm2 = stage 4 

BM Md 28 Right mandibular fragment with a complete ramus, and dm1 and dm2 dm1 = stage 5; dm2 = stage 4 

BM Md 32 Left corpus and ascending ramus holding dm2 and M1 dm2 = stage 4 

BM Md 37 Right corpus fragment with dm1 and dm2 dm1 = stage 4; dm2 = stage 3 

Table 4. Preservation and wear at Bois Madame, a final/late Neolithic cave burial  

Table 5. Preservation and wear at Maurenne Caverne de la Cave, a cave burial with one middle and three final/late Neolithic dates 

Identifier Preservation Dental wear (Smith, 1984) 

Maurenne 22 Right maxillary fragment holding dm1, dm2 and the unerupted crown of M1 dm1 = stage 3; dm2 = stage 2 

Maurenne 23 Right maxillary fragment from the mesial margin of the M1 crypt to di1 dm2 = stage 7; dm2 = stage 4 

Maurenne 24 Right maxillary fragment extending from the crypt of the M1 crown to di1 
with only dm1 (obscured by matrix) and dm2 

dm2 = stage 2 

Maurenne 25 
  

Maxillary alveolus extending from a fully formed M2 crown to the canine 
crypt, with dm1, dm2 and M1 

dm1 = stage 6; dm2 = stage 4 

Maurenne 26 
  

Maxillary alveolus, from the unerupted crown of M1 to di1, preserving dm1 
and dm2 

dm1 = stage 4; dm2 = stage 2 

Maurenne 82 Nearly complete mandible holding dm1 and dm2 on both sides dm1 = stage 4; dm2 = stage 3 
  

Maurenne 85 Corpus fragment extending from the base of the left ascending ramus to right 
dm1 crypt holding the left dm1, dm2 and M1 

dm1 and dm2 = stage 4 
  



21      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

Methods 
Dental impressions were created by the first author 
using a thin layer of polyvinylsiloxane (President Jet 
Plus Regular Body, Coltène-Whaledent) applied to the 
occlusal surface of in situ molars curated at the Royal 
Belgian Institute of Natural Sciences. Dental casts were 
created at Georgia State University using centrifuged 
epoxy resin and hardener (Beuhler), which was poured 
on the dental molds nestled within putty crucibles af-
fixed beforehand with hardener (Beuhler). The casts 
dried for 24 hours before extraction. 
     Dental casts were scored by the second author using 
Hanihara (1961) and supplemented with scores for the 
hypoconulid (Cusp 5) on dm2 and the metaconule 
(Cusp 5) on dm2 from the ASUDAS (Turner et al., 1991; 
Scott and Irish, 2017), following Paul and Stojanowski 
(2015). The maxillary deciduous molars were scored 
for crown pattern of dm1 and dm2, Carabelli’s trait on 
dm2, and the presence of a metaconule (Cusp 5) on 
dm2. For the mandibular deciduous molars, only dm2 

is considered following Hanihara (1961) and the traits 
scored included the protostylid, hypoconulid (Cusp 5), 
metaconulid (Cusp 7), the central ridge of the metaco-
nid (CRM), and the distal trigonid crest (DTC). To vali-
date the scores, photographs of the original material 
were consulted.  
 
Results 
Maxillary Traits 
Crown pattern for dm1 is noted in two individuals and 
varies considerably (Table 6). Specifically, at 

Sclaigneaux, one individual exhibits a large protocone 
and paracone, and is classified as a score of 2. In con-
trast, an individual from Maurenne Caverne de la 
Cave presents all four cusps but the hypocone and 
metacone are relatively modest in size corresponding 
to a score of 4- (Hanihara, 1961). The crown pattern for 
dm2 is uniformly classified as a score of 4 (Hanihara, 
1961). Carabelli’s trait varies from a small pit in some 
individuals from Sclaigneaux to a large independent 
cusp on the dm2 of Hastière Caverne M 39 (Figure 2). It 
is present at all sites except Bois Madame. A metacon-

 
Neolithic time period Site 

ID 
No. 

Crown 
Pattern 

dm1 

 Crown 
Pattern 

      dm2 

Carabelli's 
trait 
dm2 

Metaconule 
(Cusp 5) dm2 

early/late Hastière 38  4 5 0 

Hastière 381  4   

Hastière 39  4 7 1 

final/late Sclaigneaux 124 2 4 1 0 

Sclaigneaux 115  4 1  

Sclaigneaux 122  4   

Sclaigneaux 118  4   

Sclaigneaux 117  4 6 0 

Sclaigneaux 116  4 3 0 

Sclaigneaux 119  4   

Sclaigneaux 125  4 1 0 

Bois Madame 26   4     

Bois Madame 27   4     

middle and final/late Maurenne 24  4 2 0 

Maurenne 25  4   

Maurenne 23  4   

Maurenne 22 4- 4 5 0 

Maurenne 26  4 4  

Table 6. Maxillary deciduous molar traits 

Figure 2. The only metaconule observed is a small 
cuspule on the right dm2 of Hastèire Caverne M 39 
(a); this individual also exhibits a prominent Cara-
belli’s cusp (b). Scale bar = 1 cm. 



22      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

ule is expressed only on Hastière Caverne M 39 
(Figure 2) and is absent at Sclaigneaux and Mau-
renne Caverne de la Cave. In the Bois Madame sam-
ple, it could not be observed (see Table 6).  
 
Mandibular Traits 
A protostylid is present only on the dm2 of a single 
individual from Hastière Caverne M (Figure 3). The 
metaconulid expression varies from a larger feature 
in Sclaigneaux (Hanihara score 3) and Maurenne 
Caverne de la Cave (Hanihara score 2) to its absence 
or low expression at Hastière Caverne M and Bois 
Madame (Table 7). A central ridge of the metaconid 
(CRM), or Cusp 7, is noted at all cave burials except 
Bois Madame, and a distal trigonid crest (DTC) is 
absent at Hastière Caverne M and Maurenne 
Caverne de la Cave, but present at Bois Madame 
(Figure 4). Where it could be examined, the hy-
poconulid (Cusp 5) was scored as either prominent 
(ASUDAS grade 5) such as at Hastière Caverne M 
and Sclaigneaux, or large (ASUDAS grade 4) as at 
Maurenne Caverne de la Cave, or both as is the case 
at Bois Madame (Figure 5).  

 

Figure 3. The expression of a protostylid (white 
arrow) is visible on the left dm2 of Hastière 
Caverne M 19. Scale bar = 1 cm. 

 
Neolithic 

time period 

 
Site 

ID 
No. 

Protostylid 
dm2 

Metaconulid 
(Cusp 7) 

dm2 

CRM 
dm2 

DTC 
dm2 

Hypoconulid 
(Cusp 5) 

dm2 

early/late Hastière 19 1 1  0  

Hastière 18  0 2 0 5+ 

final/late Sclaigneaux 82  3 2  5 

Bois Madame 37  0   5 

Bois Madame 32  1    

Bois Madame 28     4 

Bois Madame 27    1  

middle and 
final/late 

Maurenne 82   2 2 0 4 

Maurenne 85   0       

Table 7. Mandibular deciduous molar traits 

Figure 4. A small meta-
conulid (a) and a large 
hypoconulid (b) can be 
observed on the right 
dm2 of Maurenne 82. 
Scale bar = 1 cm. 

Figure 5. The largest and 
most distinctive hy-
poconulid (cusp 5) is 
visible on the right dm2 
of Hastière Caverne M 
18, identified by a white 
arrow. Scale bar = 1 cm. 



23      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

Frequencies of traits per Neolithic cave site 
For dm2, the crown pattern of Hastière Caverne M did 
not differ from the other collective burials. However, 
Hastière Caverne M was partially distinct from the oth-
er sites for Carabelli’s trait, which was strongly ex-
pressed where it was demonstrably visible (Table 8). 
Furthermore, there is more variation in the expression 
of a metaconule on dm2 at Hastière Caverne M, and 
this is the only site known to express a protostylid 
(Figure 3). Hastière Caverne M also differs in the ex-
pression of the metaconulid (Figure 4) when compared 
to the ones from the final/late collective burial of 
Sclaigneaux, and to a lesser extent Maurenne Caverne 
de la Cave (see Table 8). The final/late Neolithic cave 
burial of Bois Madame differs from Hastière Caverne 
M in exhibiting a distal trigonid crest (DTC), albeit of a 
low expression. Bois Madame is also distinct from the 

early/late Neolithic site of Hastière Caverne M for ex-
hibiting greater variation in the expression of the hy-
poconulid (Cusp 5) on dm2 (see Table 8).  
 
Discussion 
The morphology of the deciduous teeth has been exam-
ined in studies of modern humans (Hanihara, 1961; 
Edgar and Lease, 2007; Pilloud and Larsen, 2011), Pleis-
tocene Homo (Smith and Tillier, 1989; Bailey and Hub-
lin, 2006; Martinón-Torres et al., 2012; Hershkovitz et 
al., 2016; Zubova et al., 2016) and the African apes 
(Hardin and Legge, 2013). Because primary crown for-
mation time is shorter, environmental pressures are 
reduced, resulting in a tendency of the deciduous den-
tition to preserve the ancestral condition more often 
than permanent successors (Paul and Stojanowski, 
2017; Scott et al., 2018). Indeed, deciduous teeth have 

   Grade 

Site N Trait 0 1 2 3 4 4- 5 6 7 

Sclaigneauxb 1 Crown Pat-
tern (dm1) 

  1.00       

Maurennec 1      1.00    

Hastièrea 3 

Crown Pat-
tern (dm2) 

    1.00     

Sclaigneauxb 8     1.00     

Bois Madameb 2     1.00     

Maurennec 5     1.00     

Hastièrea 2 
Carabelli's 

(dm2) 

      0.50  0.50 

Sclaigneauxb 5  0.60  0.20    0.20  

Maurennec 3   0.33  0.33  0.33   

Hastièrea 2 
Metaconule 

Cusp 5 (dm2) 

0.50 0.50        

Sclaigneauxb 4 1.00         

Maurennec 2 1.00         

Hastièrea 1 
Protostylid 

(dm2) 
 1.00     

   

Hastièrea 2 

Metaconulid 
Cusp 7 (dm2) 

0.50 0.50        
Sclaigneauxb 1    1.00      

Bois Madameb 2 0.50 0.50        
Maurennec 2 0.50  0.50       

Hastièrea 1 

CRM (dm2) 

   1.00       
Sclaigneauxb 1    1.00       
Maurennec 1   1.00       

Hastièrea 2 

DTC (dm2) 

1.00         
Bois Madameb 1   1.00               

Maurennec 1 1.00         

Hastièrea 1 

Hypoconulid 
Cusp 5 (dm2) 

      1.00   
Sclaigneauxb 1       1.00   

Bois Madameb 2     0.50  0.50   
Maurennec 1     1.00     

Table 8. Pooled frequencies 

a early/late Neolithic; b final/late Neolithic; c middle and final/late Neolithic  



24      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

been shown to exhibit greater efficacy in indicating 
relatedness than the permanent dentition (Kitagawa 
et al., 1995; Paul and Stojanowski, 2017), and are bet-
ter at distinguishing groups than metric traits 
(Sciulli, 1977). Although deciduous and permanent 
teeth may not differ in proxies of environmental sta-
bility, such as fluctuating asymmetry (Guatelli-
Steinberg et al., 2006), the primary dentition has few-
er cases of agenesis or supernumerary teeth com-
pared to adult successors (Scott et al., 2018).  

The entire primary molar row (including the de-
ciduous and permanent teeth) may reflect an under-
lying unified mechanism of expression. At the same 
time, the deciduous dentition appears to be gov-
erned by partially distinct genetic and developmen-
tal processes than the permanent teeth, such that the 
presence of a protostylid on dm2 (see Figure 4) does 
not necessarily imply that this trait will appear on 
any of the permanent molars (Scott et al., 2018). Fur-
thermore, Carabelli’s trait tends to be more preva-
lent and more strongly expressed in dm2 compared 
to M1 (Kaul and Prakash, 1981; Bermúdez de Castro, 
1989; Edgar and Lease, 2007; Scott et al., 2018). To 
the degree to which Carabelli’s trait and the proto-
stylid are informative about biological relationships, 
it would suggest that Hastière Caverne M does in-
deed differ from the other cave burials. In a study of 
early Neolithic Çatalhöyük, Pilloud (2009) found 
that for the deciduous dentition, Carabelli’s trait of 
dm2 and the presence of a protostylid on dm2 signifi-
cantly separated groups, and this appears to be true 
among the late Neolithic cave burials from Belgium. 
     Prehistoric deciduous teeth have been scarcely 
examined given a historical preference for the per-
manent dentition (Scott et al., 2018). An informative 
study by Scuilli (1977) described the deciduous den-
tal morphology of prehistoric Amerindian hunter/
gatherer/fishers and early Mississippian cultivators 
of the Ohio Valley. The crown form on dm1 in the 
prehistoric Amerindian remains is most frequently 
four cusps (paracone, protocone, metacone and hy-
pocone) like at Maurenne Caverne de la Cave (see 
Table 6). The crown form on dm1 of an individual 
from Sclaigneaux including only the two mesial 
cusps was rarely found in 58 individuals from 12 
sites (Sciulli, 1997). The dm2 presents four cusps in 
both prehistoric Amerindian and Neolithic Belgian 
cave sites (Sciulli, 1977; see Table 8). However, Cara-
belli’s trait is extremely rare among prehistoric Am-
erindians, whereas it is present and expressed 
strongly in three of the four Neolithic cave sites from 
Belgium, only being absent from Bois Madame (see 
Table 6). In comparison to Sciulli (1977) a metacon-
ulid (Cusp 7) on dm2 was found at lower frequencies 

compared to the results from this study, although 
the constraints of the small sample sizes must be 
taken into consideration (see Table 8). Each of the 
Neolithic cave burials exhibits a hypoconulid (Cusp 
5) which compares to 97% of the prehistoric Amerin-
dians who exhibit five or more cusps on dm2 
(Sciulli, 1977).  
 
Conclusions 
The deciduous molars from the Neolithic caves of 
Belgium present considerable variation in the ex-
pression of traits. Crown pattern varies where it can 
be observed. Carabelli’s trait is found at Hastière 
Caverne M, as well as the final/late Neolithic cave 
site of Sclaigneaux and at Maurenne Caverne de la 
Cave, although its expression varies. The individuals 
preserving dm2 generally exhibit a large or very 
large hypoconulid (Cusp 5). 

Given the wide range of radiometric dates from 
Maurenne Caverne de la Cave, it was expected to 
exhibit the greatest variability. Like Sclaigneaux, 
Maurenne Caverne de la Cave does present substan-
tial variation in the expression of Carabelli’s cusp 
and in the metaconulid (Cusp 7) compared to 
Hastière Caverne M. The resemblance of the two 
collective burials from Hastière rockshelter (Hastière 
Caverne M and Maurenne Caverne de la Cave) is 
not particularly strong although the number of indi-
viduals involved is severely constrained. 

We expected to observe differences between the 
early/late collective burial of Hastière Caverne M 
and the final/late Neolithic sites of Sclaigneaux and 
Bois Madame. Hastière Caverne M does exhibit the 
most pronounced expression of Carabelli’s cusp on 
dm2, and this trait is found nearly universally in the 
sample (see Figure 2). Hastière Caverne M also has 
the largest hypoconulid (see Figure 5), and this cave 
burial is the only assemblage to express a protostylid 
on dm2 (see Figure 3) and a metaconule on dm2 (see 
Figure 2). Since no other site presents these distinc-
tions, it appears that the deciduous dental morphol-
ogy of the early/late Neolithic cave assemblage of 
Hastière Caverne M does indeed differ from the fi-
nal/late Neolithic collective burials of Sclaigneaux 
and Bois Madame.  
     Population movement or displacement and/or 
secular changes may explain some of the differences 
in the frequencies of traits if the cave burials repre-
sent a single group of closely related peoples. Alter-
natively, these populations may have had only a 
limited amount of regional gene flow during the late 
Neolithic period. Previous studies indicate that de-
ciduous dental morphology approximates, to a 
greater extent than the secondary dentition, the ge-



25      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

netic relationships among individuals (Kitagawa et 
al., 1995; Paul and Stojanowski, 2017). To the degree 
to which this is also true of these Neolithic cave buri-
als, it can be assumed the people represented at 
Hastière Caverne M were relatively isolated several 
centuries prior to a partial restructuring of the re-
gional population associated with the Bronze Age.  
 
Acknowledgments 
Permission to examine the Neolithic remains from 
Belgium and to conduct this study of the deciduous 
dentition was kindly provided by Patrick Semal, 
Chief of the Scientific Heritage Service, Royal Bel-
gian Institute of Natural Sciences. We are indebted 
to Laurence Cammaert from the ADIA (Association 
pour la Diffusion de l'Information Archéologique) 
who created Figure 1 which we utilize with permis-
sion. We are grateful to two anonymous reviewers 
who significantly improved the manuscript, and to 
attendees of the 2017 annual meeting of WeBig 
(Western Bioarchaeology Group) for their helpful 
insights. Funding for this research was awarded to 
FLW by Fulbright-Belgium and the Commission for 
Educational Exchange between the USA, Belgium, 
and Luxembourg.  
 
REFERENCES 
Bailey S.E., Hublin J.-J. (2006). Dental remains from 

the Grotte du Renne at Arcy-sur-Cure 
(Yonne). Journal of Human Evolution, 50, 485–
508. 

Bermúdez de Castro J.M. (1989). The Carabelli trait 
in human prehistoric populations of the Ca-
nary Islands. Human Biology, 61, 117–131. 

Bronk-Ramsey C., Higham T.F.G., Owen D.C., Pike 
W.G., Hedges R.E.M. (2002). Radiocarbon 
dates from the Oxford AMS system: datelist 
31. Archaeometry, 44(3) Supplement 1:1–149. 

Dumbruch I. (2003). Edute du site de l’abri-sous-
roche du “Bois-Madame”, Néolithique, à 
Arbre, dans la vallée du Burnot (Province de 
Namur). Etude anthropologique et ar-
chéologique, Volume I et II. MA thesis, Uni-
versité Libre de Bruxelles. 

Dumbruch I. (2007). Le Site de l'Abri-sous-Roche du 
"Bois-Madame" à Arbre (Province de Na-
mur, Belgique). Archæologia Mosellana, 7, 609
–612. 

Edgar H.J.H., Lease L.R. (2007). Correlations be-
tween deciduous and permanent tooth mor-
phology in a European American sample. 
American Journal of Physical Anthropology, 
133, 726–734. 

García-Martin C. (2000). Reconstitution du régime 
alimentaire par l’étude des micro-traces 
d’usure dentaire. MA thesis, Université Li-
bre de Bruxelles. 

Guatelli-Steinberg D., Sciulli P.W., Edgar H.J.H. 
(2006). Dental fluctuating asymmetry in the 
Gullah: tests of hypotheses regarding devel-
opmental stability in deciduous vs. perma-
nent and male vs. female teeth. American 
Journal of Physical Anthropology, 129, 427–434. 

Hanihara K. (1961). Criteria for classification of 
crown characters of the human deciduous 
dentition. Journal of the Anthropological Society 
of Nipon, 69, 27–45. 

Hardin A.M., Legge S.S. (2013). Geographic variation 
in nonmetric dental traits of the deciduous 
molars of Pan and Gorilla. International Jour-
nal of Primatology, 34, 1000–1019. 

Hershkovitz I., Smith P., Sarig R., Quam R., Rodri-
guez L., Garcia R., Arsuaga J.L., Barkai R., 
Gopher A. (2016). Middle Pleistocene dental 
remains from Qesem Cave (Israel). American 
Journal of Physical Anthropology, 144, 575–592. 

Irish J.D. (2006). Who were the ancient Egyptians? 
Dental affinities among Neolithic through 
postdynastic peoples. American Journal of 
Physical Anthropology, 22, 529–543. 

Kitagawa Y., Manabe Y., Oyamada J., Rokutanda A. 
(1995). Deciduous dental morphology of the 
prehistoric Jomon people of Japan – compar-
ison of nonmetric characters. American Jour-
nal of Physical Anthropology, 97, 101–111.  

Martinón-Torres M., de Castro J.M.B., Gomez-Robles 
A., Prado-Simon L., Arsuaga J.L. (2012). 
Morphological description and comparison 
of the dental remains from Atapuerca-Sima 
de los Huesos site (Spain). Journal of Human 
Evolution, 62, 7–58.Bottom of Form 

Orban R., Polet C., Semal P., Leguebe A. (2000). La 
stature des Néolithiques mosans. Bulletin de 
l'Institut Royal des Sciences Naturelles de Bel-
gique, Série Sciences de la Terre, 70, 207–222. 

Paepe M. de (2007). Studie van de laat-neolithische 
menselijke resten uit een collectief graf te 
Sclaigneaux (provincie Namen, B.). MA the-
sis, Universiteit Gent. 

Paul K.S., Stojanowski C.W. (2015). Performance 
analysis of deciduous morphology for de-
tecting biological siblings. American Journal of 
Physical Anthropology, 164, 97–116.  

Paul K.S., Stojanowski C.W. (2017). Comparative 
performance of deciduous and permanent 
dental morphology in detecting biological 
relatives. American Journal of Physical Anthro-
pology, 157, 615–629.  

Pilloud M.A. (2009). Community structure at Neo-
lithic Çatalhöyük: biological distance analy-
sis of household, neighborhood, and settle-
ment. Ph.D. dissertation. The Ohio State 
University. 

Pilloud M.A., Larsen C.S. (2011). "Official" and 



26      

 

Dental Anthropology  2018 │ Volume 31 │ Issue 02 

"practical" kin: inferring social and commu-
nity structure from dental phenotype at Ne-
olithic Catalhoyuk, Turkey. American Journal 
of Physical Anthropology, 145, 519–530. 

Pilloud M.A., Edgar H.J.H., George R., Scott G.R. 
(2016). Dental morphology in biodistance 
analysis. In M. Pilloud & J. Hefner (Eds.), 
Biological distance analysis: forensic and bioar-
chaeological perspectives (pp. 109–133). San 
Diego: Elsevier. 

Polet C. (2011). Les squelettes néolithiques décou-
verts dans les grottes du basin mosan. In N. 
Cauwe, A. Hauzeur, I. Jadin, C. Polet & B. 
Vanmontfort (Eds.), 5200-2000 av. J.-C. prem-
iers agriculteurs en Belgique (pp. 85–94). Édi-
tions du Cedarc.  

Sciulli P.W. (1977). A descriptive and comparative 
study of the deciduous dentition of prehis-
toric Ohio Valley Amerindians. American 
Journal of Physical Anthropology, 47, 71–80. 

Scott G.R., Turner C.G. (1997). The anthropology of 
modern human teeth. Cambridge: Cambridge 
University Press. 

Scott G.R., Irish J.D. (2017). Human tooth crown and 
root morphology. Cambridge: Cambridge Uni-
versity Press. 

Scott G.R., Turner C.G., Townsend G.C., Martinón-
Torres M. (2018). The anthropology of modern 
human teeth, 2nd Edition. Cambridge: Cam-
bridge University Press. 

Semal P., García Martín C., Polet C., Richards M.P. 
(1999). Considération sur l'alimentation des 
Néolithiques du Bassin mosan: usures den-
taires et analyses isotopiques du collagène 
osseux. Notae Praehistoricae, 19, 127–135. 

Smith B.H. (1983). Dental attrition in hunter-
gatherers and agriculturalists. Ph.D. disser-
tation, University of Michigan. 

Smith B.H. (1984). Patterns of molar wear in hunter-
gatherers and agriculturalists. American Jour-
nal of Physical Anthropology, 63, 39–56. 

Smith P., Tillier A.-M. (1989). Additional infant re-
mains from the Mousterian Strata, Kebara 
Cave (Israel). In O. Bar-Yosef & B. Vander-
meersch (Eds.), Investigation in South Levan-
tine prehistory (pp. 323–335). Oxford, Eng-
land: British Archaeological Reports Interna-
tional Series 497. 

Toussaint M. (2007). Les sépultures Néolithiques du 
basin mosan Wallon et leurs relations avec 
les bassins de la Seine et du Rhin. Archaeolo-
gia Mosellana, 7, 507–549. 

Toussaint M., Orban R., Polet C., Semal P., 
Bocherens H., Masy P., García Martín C. 
(2001). Apports récents sur l’anthropologie 
des Mésolithiques et des Néolithiques mo-
sans. Anthropologica et Préhistorica, 112, 91–

105. 
Turner C.G., II, Nichol C., Scott G.R. (1991). Scoring 

procedures for key morphological traits of 
the permanent dentition: The Arizona State 
University Dental Anthropology System. In 
M. A. Kelley & C. S. Larsen (Eds.), Advances 
in dental anthropology (pp. 13–31). New York: 
Wiley-Liss. 

Vanderveken S. (1997). Etude anthropologique des 
sépultures néolithiques de Maurenne et 
Hastière (province de Namur). MA thesis, 
Université Libre de Bruxelles. 

Zubova A.V., Stepanov A.D., Kuzmin Y.V. (2016). 
Comparative analysis of a Stone Age human 
tooth fragment from Khaiyrgas Cave on the 
Middle Lena (Yakutia, Russian Federation). 
Anthropological Science, 124, 135–143.