35 Dental Anthropology 2021 │ Volume 34│ Issue 01 The Meuse River basin of central Belgium extends along a semi-continuous karstic uplift featuring numerous cliff walls, rock formations and at least 3,000 caverns. More than 250 of these caves pre- serve the remains of prehistoric humans. Although these caves have been known for centuries, formal exploration of the sites commenced in the winter of 1829-1830 and has continued to the present (Polet, 2011). Close to 200 of these funerary sites have been radiocarbon dated to the Late Neolithic peri- od (Toussaint et al., 2001). Many of these are collec- tive burials and contain five to 15 individuals (Polet, 2011), however, some are larger, such as the caves of Bois Madame and Sclaigneaux (Dumbruch, 2003; De Paep & Polet, 2007). Only eight percent of these funerary sites contain be- tween 55 and 60 individuals (Polet, 2011). Hundreds of skeletal fragments and dental ele- ments have been investigated from Hastière Caverne M (Hastière M), Hastière Trou Garçon C (Trou Garçon), Sclaigneaux, Bois Madame and Maurenne Caverne de la Cave (Maurenne) (Figure 1), and adults of both sexes and children are repre- sented, suggesting familial, kin or descent groups used the caves for burial. These five cave deposits are all radiocarbon dated to the Late Neolithic (Table 1). However, Maurenne is associated with three dates from the terminus of the Late Neolithic (4,160 ± 45; 3,950 ± 70; 3,830 ± 90 years BP), and one date, 4,635 ± 45 years BP, from the Middle Neolith- Permanent Molar Trait Expression in the Late Neolithic Cave Burials of the Meuse Basin, Belgium Frank L’Engle Williams 1* and Rebecca L. George 2 1 Georgia State University 2 University of Nevada, Reno ABSTRACT At least 250 cave burials along the Meuse river basin of Belgium yield prehistoric remains, and most date from the Late Neolithic period. Several of these collective burials have been radiocarbon dated, including the early/late Neolithic deposits of Hastière Caverne M and Hastière Trou Garçon C and the final/late Neolithic caves of Sclaigneaux and Bois Madame. An additional cave burial, Mau- renne Caverne de la Cave, has been radiocarbon dated to the Middle Neolithic and final/late Neolithic periods, circa 4,635 to 3,830 years BP, which encompasses the entire range of dates for the other collec- tive burials. Individuals (n = 127) are represented by fragmentary gnathic remains with in situ dental elements. Although the remains have been studied in detail, researchers have yet to compare dental morphology across cave sites. Arizona State University Dental Anthropology System (ASUDAS) scores of permanent molar morphology are employed to examine whether differences within and between the cave burials exist, and whether chronology and geography can account for the variation in traits. Af- firming our expectations, the final/late Neolithic cave of Sclaigneaux, the most geographically distant cave burial, and secondarily Hastière Caverne M, possibly the earliest site, emerge as the most distinc- tive. The final/late Neolithic sites of Sclaigneaux and Bois Madame exhibit the greatest variability of trait expression. This research contributes to the understanding of the relatedness of early farming com- munities, and these findings bear on the mobility and continuity of human groups in the Meuse basin of Belgium during the terminus of the Neolithic before the onset of the Bronze Age in northern Europe. *Correspondence to: Frank L’Engle Williams Department of Anthropology Georgia State University P.O. Box 3998 Atlanta, GA, 30302-3998 United States frankwilliams@gsu.edu Keywords: Hastière Caverne M, Hastière Trou Garçon C, Sclaigneaux, Bois Madame, Maurenne Caverne de la Cave mailto:frankwilliams@gsu.edu 36 Dental Anthropology 2021 │ Volume 34│ Issue 01 Figure 1. Map of Belgium showing the location of five Late Neolithic collective burials along the Meuse River system. Table 1. Radiocarbon dates associated with five Neolithic collective burials of Belgium, arranged by site and by dis- tance from Hastière rockshelter; dating was conducted using Accelerated Mass Spectrometry (AMS) at Oxford University, UK (OxA) and the University of Groningen (GrA), and conventional methods at the University of Louvain, Belgium (Lv). a early/late Neolithic; b Middle Neolithic; c final/late Neolithic Collective burial Sample number Dates in years BP Reference Hastière M AMS OxA-6558 4,345 ± 60a Bronk-Ramsey et al. (2002) Trou Garçon AMS OxA-6853 4,220 ± 45a Bronk-Ramsey et al. (2002) Maurenne AMS OxA-9025 4,635 ± 45b Bronk-Ramsey et al. (2002) Maurenne AMS OxA-9026 4,160 ± 45c Bronk-Ramsey et al. (2002) Maurenne Lv-1483 3,950 ± 70c Toussaint (2007) Maurenne Lv-1482 3,830 ± 90c Toussaint (2007) Bois Madame AMS OxA 10831 4,075 ± 38c Dumbruch (2003) Bois Madame AMS OxA 10830 3,910 ± 40c Dumbruch (2003) Sclaigneaux GrA-32975 4,155 ± 35c De Paepe & Polet (2007) 37 Dental Anthropology 2021 │ Volume 34│ Issue 01 ic, implying its use for more than 800 years (Vanderveken, 1997; Bronk-Ramsey et al., 2002; Toussaint, 2007). The Maurenne burial is adjacent to Hastière rockshelter formation (see Figure 1). Two other collective burials at this site include Hastière M and Trou Garçon. Hastière M is one of the oldest Late Neolithic cave sites and dates to 4,345 ± 60 years BP, followed by Trou Garçon, which has yielded a date of 4,220 ± 45 years BP (Bronk- Ramsey et al., 2002; Toussaint, 2007). These two can be described as early/late Neolithic. Two large, well-studied final/late Neolithic cave burials are Sclaigneaux and Bois Madame. Sclaigneaux is associated with a single radiocarbon date of 4,155 ± 35 years BP (De Paepe, 2007; De Paepe & Polet 2007). At Bois Madame in the Burnot Valley, two dates have been obtained. Both of these derive from the boundary of the fourth millennium prior to the Bronze Age, 4,075 ± 38 years BP and 3,910 ± 40 years BP, suggesting the collective burial of Bois Madame may have been utilized for more than 150 years (Bronk-Ramsey et al., 2002; Dum- bruch, 2003, 2007). Funerary context Given the scarcity of habitation sites, these prehis- toric peoples are primarily known from their re- mains in funerary caves and rockshelters. A range of burial practices has been inferred, including cre- mation, burial, a simple deposition of individuals on cave floors and cu-marks with flint implements. Comingled remains comprise a majority of the fu- nerary deposits (Toussaint et al., 2001; Toussaint, 2007; Polet, 2011). At some caves, such as Bois Madame, the bones are found in a haphazard or- der as if the individuals were left unburied and later disturbed by human or non-anthropogenic agents (Dumbruch, 2003). The mixture of individu- als within these collective burials could have arisen from bioturbation. However, the deliberate move- ment, occasional regrouping and comingling of bodies is more likely to be the result of burial rites, reburial and/or adding additional individuals (Toussaint et al., 2001; Toussaint, 2007). Comparing individuals across cave burials Although several well-preserved Late Neolithic crania are present, most individuals are represent- ed by fragmentary gnathic remains with associated molars in situ, permitting an investigation of varia- tion within and between sites in nonmetric dental trait expression using the Arizona State University Dental Anthropology System (ASUDAS) (Turner et al., 1991; Scott and Irish, 2017). Prior studies of the inhabitants of these Late Neolithic caves have found a lack of differentiation in diet (Garcia Mar- tín, 1999; Semal et al., 1999), internment behavior (Vanderveken, 1997; Toussaint et al., 2001, 2003) and stature was estimated to be largely unimodal (Orban et al., 2000). However, chronological dis- tinctions are apparent from radiocarbon dating. On the basis of chronology, we expect the early/late Neolithic sites to be more similar to each other in dental morphological expression than to the final/ late cave burials, and vice versa. The three final/ late Neolithic dates from Maurenne suggest this collective burial is more likely to resemble later sites than earlier ones. It is also possible that differences in dental mor- phology will be patterned with respect to geogra- phy. Based on distance, individuals from Hastière rockshelter (Hastière M and Trou Garçon) and Maurenne should be more similar to one another, and secondarily to Bois Madame, whereas Sclaigneaux should be the most distinctive (see Figure 1). Materials and Methods A total of 127 individuals from the five caves were examined (Vanderveken, 1997; Toussaint et al., Table 2. Neolithic samples by cave, element, and number of individuals. Neolithic cave site Maxillae Mandibles Total Hastière M 10 10 20 Trou Garçon 6 1 7 Maurenne 9 21 30 Bois Madame 13 15 28 Sclaigneaux 12 30 42 Total 50 77 127 38 Dental Anthropology 2021 │ Volume 34│ Issue 01 2001; Dumbruch, 2003; De Paep, 2007; Toussaint, 2007; Williams and Polet, 2017; Table 2). Gnathic fragments were chosen on the basis of complete- ness and only relatively unworn crowns were ex- amined. No isolated teeth were included to avoid errors in attribution. Given the lack of anterior teeth preserved in situ, and the inconsistent preser- vation of premolars, only molars were observed. Preference was given to young adults and subadults with relatively unworn cusps of perma- nent molars to increase the likelihood of accurate scoring, and included Smith (1984) wear stages 1 to 4. Individuals who exhibited substantial attrition, exceeding stage 4 (Smith, 1984), were excluded from the analysis (Turner et al., 1991; Scott and Irish, 2017). Dental cast preparation Dental casts were created from dental impressions of the original Neolithic material housed at the Royal Belgian Institute of Natural Sciences in Brus- sels. To create the dental molds, the dentition was cleaned and a thin layer of dental molding materi- al, polyvinylsiloxane (President Jet Plus Regular Body, Coltène-Whaledent) was applied to the oc- clusal surface of the molars and allowed to air dry. Dental casts were created at Georgia State Univer- sity by pouring centrifuged epoxy resin and hard- ener (Buehler) onto the dental impressions, which were placed into putty crucibles—stabilized with hardener (Buehler)—to catch the excess mixture. The casts dried for 24 hours before extraction. Analysis Dental morphology has been shown to be highly heritable (Turner et al., 1991; Scott and Turner, 1997; Irish, 2006; Hanihara, 2008; Scott and Irish, 2017; Scott et al., 2018). Dental casts, supplemented with photographic images, were scored by a single observer (RLG) to avoid issues of interobserver error (Turner and Scott, 1997; Hardin and Legge, 2013). Previously conducted intraobserver error analyses on 34 dental morphological traits found trait agreement at levels of 0.621 or above (McHugh, 2012). Since single-sided gnathic frag- ments were available for the great majority of the individuals, dental antimeres could not be exam- ined to identify the maximum expression of any trait. It is possible that some of the maxillary and mandibular fragments belonged to the same indi- viduals. However, given the preservation of the remains, pairing these elements was not possible. If some elements are indeed associated, then the total sample size of 127 would be smaller. Since 77 man- dibular and 50 maxillary fragments are included, a potential minimum number of individuals (MNI) is 77 (Table 2). These associations are likely irrele- vant in the current study as maxillary and mandib- ular molar traits are discussed separately Another potential problem from the lack of matching elements might have arisen from inad- vertently scoring antimeres from the same individ- ual. However, this is unlikely for several reasons. First, only in situ molars rather than isolated ele- ments were scored. Second, the range of dental attrition and dental ages suggests each gnathic fragment can be considered unique. Therefore, each fragment was treated as an individual (Hardin and Legge, 2013) as shown in Table 2, and the dental morphological traits were discussed in- dependently. Score frequencies for each trait with respect to each cave site were calculated. Statistical analyses were not attempted due to the small and idiosyncratic sample sizes. Results All scores ascribed to individuals are presented in the context of the ASUDAS. Maxillary molars Metacone For M1, individuals from Hastière M and Trou Gar- çon often exhibit a metacone with a score of 4 (see Table 3). Fewer individuals have a larger metacone with a score of 5. In contrast, individuals from Maurenne and Bois Madame frequently present a metacone with a score of 5 and have a lower fre- quency of individuals with a score of 4. Sclaigneaux shows an equal prevalence of individ- uals with metacone scores of 4 and 5. For the second molar (M2), the dominant pattern across sites is a score of 3 or 4, though there is some variation in expression (Table 3). For in- stance, Hastière M and Trou Garçon are nearly di- vided equally between these two scores, whereas the final/late Neolithic burials present a greater tendency for a metacone with a score of 4. Individ- uals from Bois Madame show a greater range of expression in their metacone scores as they range from 2 to 5. Although the sample size for M3 is limited, the individual from Hastière M has a large metacone with a score of 5, whereas individuals from both Maurenne and Bois Madame exhibit a smaller cusp and have scores of 3. The other two sites are inter- mediate and have scores of 4 for the M3 metacone (Table 3). 39 Dental Anthropology 2021 │ Volume 34│ Issue 01 Table 3. Frequencies of maxillary traits. Frequency of score Site n Trait & Tooth 0 1 2 3 4 5 6 7 Hastière M 8 Metacone (M1) 0.875 0.125 Trou Garçon 5 0.800 0.200 Sclaigneaux 10 0.500 0.500 Maurenne 8 0.250 0.750 Bois Madame 11 0.273 0.727 Hastière M 5 Metacone (M2) 0.600 0.400 Trou Garçon 4 0.500 0.500 Sclaigneaux 6 0.167 0.833 Maurenne 3 0.333 0.667 Bois Madame 8 0.125 0.125 0.375 0.375 Hastière M 1 Metacone (M3) 1.000 Trou Garçon 2 0.500 0.500 Sclaigneaux 3 0.333 0.667 Maurenne 1 1.000 Bois Madame 2 1.000 Hastière M 7 Hypocone (M1) 0.429 0.571 Trou Garçon 4 0.500 0.500 Sclaigneaux 10 0.400 0.600 Maurenne 8 0.125 0.875 Bois Madame 11 0.273 0.364 0.364 Hastière M 6 Hypocone (M2) 0.167 0.667 0.167 Trou Garçon 2 1.000 Sclaigneaux 4 0.500 0.500 Maurenne 3 0.333 0.667 Bois Madame 8 0.125 0.750 0.125 Hastière M 1 Hypocone (M3) 1.000 Trou Garçon 1 1.000 Sclaigneaux 3 0.333 0.333 0.333 Maurenne 4 1.000 Bois Madame 2 0.500 0.500 Hastière M 5 Metaconule (M1) 0.800 0.200 Trou Garçon 3 0.667 0.333 Sclaigneaux 8 1.000 Maurenne 6 0.833 0.167 Bois Madame 9 0.667 0.333 Hastière M 8 Metaconule (M2) 0.500 0.250 0.125 0.125 Trou Garçon 4 0.500 0.250 0.250 Sclaigneaux 4 1.000 Maurenne 2 0.500 0.500 Bois Madame 7 0.714 0.286 Hastière M 1 Metaconule (M3) 1.000 Trou Garçon 2 0.500 0.500 Sclaigneaux 2 1.000 Maurenne 1 1.000 Bois Madame 2 0.500 0.500 Hastière M 3 Carabelli's Trait (M1) 0.333 0.667 Trou Garçon 2 0.500 0.500 Sclaigneaux 3 0.333 0.333 0.333 Maurenne 2 1.000 Bois Madame 5 0.200 0.200 0.200 0.200 0.200 Hastière M 3 Carabelli's Trait (M2) 1.000 Trou Garçon 1 1.000 Sclaigneaux 2 0.500 0.500 Maurenne 3 1.000 Bois Madame 2 0.500 0.500 Hastière M 4 Parastyle (M1) 1.000 Trou Garçon 3 1.000 Sclaigneaux 3 0.667 0.333 Maurenne 2 1.000 Bois Madame 8 1.000 Hastière M 5 Parastyle (M2) 1.000 Trou Garçon 3 0.667 0.333 Sclaigneaux 4 0.750 0.250 Bois Madame 7 1.000 40 Dental Anthropology 2021 │ Volume 34│ Issue 01 Hypocone The M1 hypocone is primarily scored as a 4 or 5 nearly evenly across three of the sites (see Table 3). Most individuals from Maurenne, though, are scored as 5 and more than a quarter of the M1 sam- ples from Bois Madame (27.3%) exhibit a smaller hypocone and are characterized by scores of 3. For M2, the hypocone tends to be expressed most strongly at Maurenne and Sclaigneaux as most individuals at these sites have scores of 4. For Bois Madame and Hastière M, a smaller hypocone with a score of 3 is the most frequent expression, with considerable variation (see able 3). The M3 hypocone is variably expressed at Sclaigneaux and Bois Madame. In comparison, the M3 hypocone is most frequently larger at Trou Gar- çon and Maurenne with scores of 4 (see Table 3). Metaconule (Cusp 5) The metaconule is absent at Sclaigneaux across the molars (see Table 3). This is not the case at the oth- er sites with the exception of M3 in which individu- als from Hastière M and Maurenne also lack Cusp 5. For M1, three individuals from Bois Madame present small metaconules with a score of 2. The early/late Neolithic cave burials of Hastière M and Trou Garçon both exhibit substantial variation in the expression of the metaconule across the molars (see Table 3). Variation at the early/late Neolithic sites is particularly marked for the M2 at Hastière M where the expression of Cusp 5 ranges from ab- sent in half of the individuals to moderately ex- pressed with scores of 1-3 in the other half. Trou Garçon is mostly associated with scores of 1 and 2. Maurenne and Bois Madame are similar in their low to absent expression of the metaconule on M2 and M3 (Table 3). In contrast, a prominent meta- conule is expressed on the M3 of Hastière M 29, presenting a score of 5 (Figure 2). Carabelli’s trait Carabelli’s trait is relatively well represented across these Neolithic sites on M1 and M2 but is absent entirely on M3 (see Table 3). However, there is considerable variation within and between buri- als (Figures 3 and 4). For M1, Bois Madame exhibits the strongest expression of this trait, with one indi- vidual having a prominent Carabelli’s cusp with a score of 7. Bois Madame present the greatest de- gree of variation, with expressions ranging from 2- 5. One individual from Sclaigneaux has a large Carabelli’s trait with a score of 5 and another is even larger with a score of 6 (Figure 4). In compari- son, this trait on M1 is expressed as a 1 or absent altogether at the early/late Neolithic sites of Hastière M and Trou Garçon (see Table 3). For M2, Hastière M and Trou Garçon C lack ex- pressions of Carabelli’s trait while the final/late sites of Sclaigneaux and Bois Madame show sub- stantial variation ranging from scores 1-3 (see Ta- ble 3). Maurenne resembles the Hastière M and Trou Garçon, in lacking evidence of a Carabelli’s trait on M2 (see Table 3). Parastyle As at other locations worldwide (Scott et al. 2018), the expression of a parastyle is rarely observed in these Neolithic collective burials and is completely absent on M3 (see Table 3). However, a large M1 parastyle is scored as a 3 on Sclaigneaux 119. A smaller M2 parastyle is scored as a 2 on Sclaigneaux 99. In addition, a limited expression of a parastyle is noted for one M2 from Trou Garçon (I.G. 3873) characterized as a buccal pit (score of 1). Mandibular molars Anterior fovea The anterior fovea on M1 is most frequently ex- pressed as a score of 1 across the cave burials when it is present (Table 4; Figure 5). There is one indi- vidual, Maurenne 92, who presents a larger anteri- or fovea with a score of 3. Groove pattern The groove pattern for M1 is primarily the Y pat- tern, with the exception of one individual from Hastière M and another from Sclaigneaux that ex- hibit an X pattern. The near ubiquity of the Y pat- tern, particularly at the final/late Neolithic cave burials, is further evidenced by the relatively large number of individuals with this configuration. This includes all of the Maurenne (n = 12) and Bois Madame (n = 8) assemblages, and nine out of 10 individuals from Sclaigneaux (see Table 4). The groove patterns for M2 and M3 are more variable (see Table 4). For M2, the groove pattern for the early/late Neolithic cave burial of Trou Garçon presents as an X. Individuals from Hastière M most often exhibit the plus groove pattern with some expression of the Y pattern. At Maurenne, all three groove pattern variants are evident (see Table 4). For M3, Hastière M 10 exhibits an X groove pat- tern, as do most individuals from Maurenne. The final/late Neolithic sites exhibit more varia- bility in groove patterning for both M2 and M3 than is observed for these teeth in the earlier cave buri- als. All three configurations are visible at Sclaigneaux, although the Y pattern is the least 41 Dental Anthropology 2021 │ Volume 34│ Issue 01 Figure 2. Hastière Trou Garçon C 20Z, a right M1 shows (a) a large metaconule or Cusp 5 (ASUDAS score = 2) and (b) a pit form of Carabelli’s trait (ASUDAS score = 1); scale bar = 1 cm. Figure 3. Bois Madame, BM Mx 11, a right maxil- lary fragment, demonstrates a large Carabelli’s trait (ASUDAS score = 7), identified by a white arrow on M1; scale bar = 1 cm. Figure 4. Sclaigneaux 119, a left M1, exhibits (a) a pro- nounced Carabelli’s cusp (ASUDAS score = 6), and (b) a large metacone (ASUDAS score = 4); scale bar = 1 cm. Figure 5. Bois Madame BM Md 32, a left M1, shows (a) an anterior fovea (ASUDAS score = 1) and (b) a protostylid (ASUDAS score = 1), both of which are commonly found across cave sites; scale bar = 1 cm. 42 Dental Anthropology 2021 │ Volume 34│ Issue 01 Table 4. Frequencies of mandibular traits. Frequency of score Site n Trait & tooth 0 1 2 3 4 5 6 X Y + Hastière M 2 Anterior Fovea (M1) 1.000 Sclaigneaux 4 0.750 0.250 Maurenne 8 0.625 0.250 0.125 Bois Madame 4 0.500 0.500 Hastière M 1 Groove Pattern (M1) 1.000 Sclaigneaux 10 0.100 0.900 Maurenne 12 1.000 Bois Madame 8 1.000 Hastière M 4 Groove Pattern (M2) 0.250 0.750 Trou Garçon 1 1.000 Sclaigneaux 15 0.400 0.067 0.533 Maurenne 7 0.143 0.143 0.714 Bois Madame 8 0.375 0.250 0.375 Hastière M 1 Groove Pattern (M3) 1.000 Sclaigneaux 5 0.200 0.600 0.200 Maurenne 3 0.667 0.333 Bois Madame 2 0.500 0.500 Hastière M 4 Cusp Number (M1) 0.750 0.250 Sclaigneaux 11 0.091 0.727 0.182 Maurenne 12 0.333 0.583 0.083 Bois Madame 9 0.778 0.222 Hastière M 5 Cusp Number (M2) 0.600 0.400 Trou Garçon 1 1.000 Sclaigneaux 12 0.750 0.250 Maurenne 8 1.000 Bois Madame 7 0.714 0.286 Hastière M 1 Cusp Number (M3) 1.000 Sclaigneaux 7 0.429 0.429 0.143 Maurenne 3 1.000 Bois Madame 2 0.500 0.500 Hastière M 5 Mid-Trigonid Crest (M2) 1.000 Trou Garçon 1 1.000 Sclaigneaux 12 0.917 0.083 Maurenne 7 0.857 0.143 Bois Madame 5 1.000 Hastière M 1 Mid-Trigonid Crest (M3) 1.000 Sclaigneaux 9 0.889 0.111 Maurenne 3 1.000 Bois Madame 1 1.000 Hastière M 5 Protostylid (M1) 0.400 0.600 Trou Garçon 1 1.000 Sclaigneaux 10 1.000 Maurenne 13 0.538 0.462 Bois Madame 7 0.143 0.857 Hastière M 4 Protostylid (M2) 0.500 0.500 Trou Garçon 1 1.000 Sclaigneaux 9 0.222 0.778 Maurenne 8 0.250 0.750 Bois Madame 5 0.400 0.600 Hastière M 1 Protostylid (M3) 1.000 Sclaigneaux 7 0.429 0.571 Maurenne 2 0.500 0.500 Bois Madame 1 1.000 Hastière M 2 Hypoconulid (M1) 0.500 0.400 Sclaigneaux 11 0.091 0.091 0.182 0.091 0.091 0.455 Maurenne 13 0.308 0.308 0.154 0.231 Bois Madame 9 0.333 0.222 0.444 Hastière M 5 Hypoconulid (M2) 0.600 0.200 0.200 Trou Garçon 1 1.000 Sclaigneaux 13 0.769 0.154 0.077 Maurenne 8 1.000 Bois Madame 7 0.714 0.143 0.143 Hastière M 1 Hypoconulid (M3) 1.000 Sclaigneaux 7 0.429 0.286 0.143 0.143 Maurenne 3 1.000 Bois Madame 2 0.500 0.500 Hastière M 4 Entoconulid (M1) 0.750 0.250 Sclaigneaux 11 0.818 0.182 Maurenne 12 0.917 0.083 Bois Madame 9 0.778 0.111 0.111 Hastière M 1 Entoconulid (M3) 1.000 Sclaigneaux 7 0.857 0.143 Maurenne 3 1.000 Bois Madame 2 1.000 43 Dental Anthropology 2021 │ Volume 34│ Issue 01 prevalent on M2 and the most frequent expression on M3 (see Table 4). All three groove patterns are present at Bois Madame for M2 as they are at Sclaigneaux and Maurenne. However, only at Sclaigneaux are the three groove patterns present on M3. Cusp number Only five or six cusps are observed on M1 at the early/late Neolithic cave burial of Hastière M and the final/late site of Bois Madame, whereas Sclaigneaux and Maurenne both present 4-6 cusps. However, the predominant number is five cusps across the cave burials (see Table 4). This pattern differs for M2 in which four cusps is the most frequently observed. For the individuals from Maurenne (n = 8) and the individual from Trou Garçon, this is the only pattern observed for M2. In comparison, there are some M2 from Hastière M, Sclaigneaux and Bois Madame that present five cusps (see Table 4). For M3, there are primarily four cusps, with the exception of Maurenne and Sclaigneaux in which the expression of four and five cusps are equally represented (Table 4). Furthermore, at Sclaigneaux, more variation is observed for M3 cusp number which includes the expression of four, five and six cusps. Mid-trigonid crest The mid-trigonid crest is eliminated for M1 since no presence was recorded across sites for this mo- lar. The mid-trigonid crest is also largely absent on M2 and M3 at these Neolithic cave burials. One ex- ception is at Sclaigneaux where it is present, alt- hough rarely, on both M2 and M3. The only other site where a mid-trigonid crest is observable is at Maurenne and only the M2 of Maurenne 18 (see Table 4). Protostylid A buccal pit (score of 1) is common at these Neo- lithic cave deposits and across the mandibular mo- lar row (Figure 5). At Sclaigneaux, the buccal pit is found on all individuals examined (n = 10). Simi- larly, a buccal pit is more often present than absent on M1 at Hastière M and Bois Madame. In contrast, at Maurenne a buccal pit on M1 is more often ab- sent than present; this feature is also absent in the single individual from Trou Garçon (Table 4). On M2, a buccal pit is visible at all sites and is more often expressed than not, particularly at Sclaigneaux and Maurenne (see Table 4). One ex- ception is Trou Garçon 3, where a protostylid is scored as a 3. Any variation of the protostylid is less frequent- ly exhibited on M3 than on the other molars. At Sclaigneaux, it is expressed as a buccal pit across the molar row. A much stronger expression of a protostylid is evidenced on one individual, Mau- renne 15, where it is scored as a 6. Hypoconulid (Cusp 5) For M1, Hastière M exhibits a moderate to large hypoconulid, expressed at scores of 3 and 4. Sclaigneaux presents the greatest degree of varia- tion in the expression of the hypoconulid, ranging across the full spectrum of scores from 0-5, alt- hough the majority of individuals from this site are skewed towards the higher end of the scoring spec- trum. This variation is similar at Bois Madame and Maurenne where the scores range from 0-5. How- ever, most individuals from Bois Madame exhibit a larger hypoconulid with a correspondingly higher score and nearly a third of the individuals from Maurenne lack a cusp 5 entirely (see Table 4). For M2, the hypoconulid is more often absent than present across cave burials, and at Maurenne and Trou Garçon it is absent altogether. When it is expressed, the final/late Neolithic caves of Sclaigneaux and Bois Madame both show greater variation and the presence of a larger cusp 5. For example, when the hypoconulid is expressed at Hastière M, it ranges in score from 1-3. At the fi- nal/late Neolithic sites of Sclaigneaux and Bois Madame, a larger hypoconulid is evident, reflected in one individual from each site scoring a 4 and 5, respectively (see Table 4). Like M2, the variation in M3 is more variable than observed in M1, especially for the final/late Neolithic cave burials. The hypoconulid is com- pletely absent in the one individual from Hastière M and the three individuals from Maurenne. In contrast, at the final/late Neolithic cave of Sclaigneaux, the greatest extent of variation is ob- served, with scores ranging from a low of 1 to a high of 5. Bois Madame has similar variability of expression of the hypoconulid, with scores extend- ing from 0-4. Entoconulid (Cusp 6) An entoconulid is expressed on M1 across the cave burials but at low frequencies. However, its ex- pression varies. The most common expression of the entoconulid, or cusp 6, is a score of 2, as ob- served at Hastière M, Maurenne and Bois Mad- 44 Dental Anthropology 2021 │ Volume 34│ Issue 01 ame. Sclaigneaux presents an entoconulid with a score of 1. At the other extreme is Bois Madame in which a larger entoconulid is scored as a 3. Thus, the final/late Neolithic caves of Sclaigneaux and Bois Madame are distinct in the expression of cusp 6 as compared to the other sites. The entoconulid on M2 was eliminated from the results because it is not observed across the sites. For M3, the entoconulid is entirely absent with the exception of Sclaigneaux. This final/late Neolithic cave burial presents one individual (Sclaigneaux 19) out of seven with an entoconulid on M3 that is scored as a 2. Metaconulid (Cusp 7) Frequencies for the metaconulid (Cusp 7) are ex- cluded since only a single tooth fully expressed this trait in the available Neolithic sample, the left M1 of Boise Madame BM Md 13 (Figure 6). Discussion Based on an earlier study of deciduous molar mor- phology (Williams et al., 2018), it was predicted that the early/late Neolithic cave burial of Hastière M would be distinctive and should differ from the final/late sites of Sclaigneaux and Bois Madame. Although this prediction was confirmed for some traits, the deciduous molar morphology of Hastière M is more distinctive compared to the permanent molars. The observation that deciduous molars are better at identifying relatedness (Paul and Stojan- owski, 2017) may also apply to these Neolithic cave burials. It was also anticipated that Sclaigneaux— situated about 35 km from the Hastière rockshel- ter—would be distinct if differences in morpholo- gy can be explained by geographic distance (Figure 1). Sclaigneaux does differ from the other cave bur- ials in some respects, for example, showing the Y groove pattern for M2. However, like the other fi- nal/late Neolithic site of Bois Madame, Sclaigneaux is quite variable in the expression of traits. These findings suggest variability is more pronounced in the final/late than the early/late Neolithic. The final/late Neolithic sites exhibit greater variation in the expression of traits, partic- ularly the hypoconulid, protostylid, parastyle and Carabelli’s trait across the molar row. However, the sample sizes are also substantially larger at the final/late Neolithic sites. This is particularly true of Sclaigneaux. It is unknown the extent to which the uneven sample sizes influenced the results. It was expected that the two early/late Neolithic cave burials of Hastière M and Trou Garçon should resemble one another as they are similar chrono- logically and geographically. Yet there is no con- vincing evidence that they are similar. In fact, it appears that Trou Garçon resembles the final/late Neolithic sites of Bois Madame and secondarily Maurenne more than these individuals resemble Hastière M. Trou Garçon has a greater number of whole crania available but is represented by a smaller number of individuals compared to the other sites (Table 2). The limited sample size for Trou Garçon precludes definitive statements on its relationship to the other cave burials. However, Trou Garçon individuals are at times extreme in the expression of traits which separates this site from the others, such as a large protostylid on M2 in Trou Garçon 3. Meanwhile, Hastière M is an outlier in other ways, such as the pronounced met- aconule on M3 in Hastière M 29. The prediction that Maurenne would resemble the final/late Neolithic sites of Sclaigneaux and Bois Madame more than Hastière M was largely confirmed by the results. For this reason, it is more likely that the individuals buried at Maurenne are primarily associated with the three final/late Neo- lithic radiocarbon dates. The single Middle Neo- lithic date obtained from Maurenne may be an ex- ception. Supporting this assertion is the observa- tion of similarities between Maurenne and Bois Madame. Three of the dates for the former and the two dates for the latter overlap one another and the two burial chambers are about 10 km from one another suggesting, perhaps, closer contact existed between these two groups than between the earlier Figure 6. Left mandibular fragment of BM Md 13, presents the only fully expressed metaconulid (Cusp 7) observed (ASUDAS score = 2), demarcat- ed by the white arrow on M1; scale bar = 1 cm. 45 Dental Anthropology 2021 │ Volume 34│ Issue 01 and the more geographically distant individuals living close to Sclaigneaux cave. There are also similarities between the caves, such as the large prevalence of a protostylid and Carabelli’s trait, and the near absence of a meta- conulid. There are most frequently five cusps on M1 but often four on M2 and M3. The lack of dis- crete differences in these Belgian Neolithic caves is supported by archaeological evidence that sug- gests common lifeways, an undifferentiated econo- my and phenotypic homogeneity. Carbon and ni- trogen isotopes imply similarities in diet across the Late Neolithic period in which terrestrial resources were relied upon more than aquatic ones (Semal et al., 1999). The dental microwear of several Late Neolithic caves suggests similarities in diet which comprised a large amount of vegetable fiber (Garcia-Martín, 2000), but fish may have also been consumed (Toussaint et al., 2001). Stature regres- sion formulae from available Neolithic long bones and the first metatarsal indicate that most of the individuals were of short stature. It is also possible that the majority of the long bones come from a single sex (female) as the sample lacks a bimodal distribution of values typical of recent Belgians of both sexes (Orban et al., 2000). Comparison with other prehistoric burials A number of studies have been conducted using dental morphology as a proxy for affinity at Neo- lithic and other prehistoric sites. Studies of kinship within and across burials and cemeteries rely on phenotypic similarity as a proxy for genetic rela- tionships and rare traits are often utilized to identi- fy familial relations (Bentley, 1991; Howell and Kintigh, 1996; Alt et al., 1997; Jacobi, 1997; Cor- ruccini and Shimada, 2002; Stojanowski & Schillaci, 2006; Pilloud, 2009; Lukacs & Pal, 2013). Familial, and possibly sibling relations among a triple burial at Dolní Věstonice from the Upper Paleolithic of the Czech Republic were evidenced by a sharing of groove pattern, number of cusps, accessory cusps and the presence of an entoconulid and parastyle for at least two of the three individual for each trait (Alt et al., 1997). The Neolithic cave burials of Bel- gium probably do not represent individuals from the same family as noted at Dolní Věstonice. In fact, it appears that there is a greater degree of var- iation within the Belgian Meuse Neolithic burials than between them. Dental traits of early Neolithic Mediterranean sites The dental morphology of several burial sites in the Mediterranean region have been explored. For example, at early Neolithic Çatalhöyük in Turkey, the protostylid, Carabelli’s cusp, groove pattern, the hypoconulid, entoconulid, hypocone and de- flecting wrinkle are significantly different from expected (Pilloud, 2009; Pilloud and Larsen, 2011). Iberian and Italian Neolithic burials differ in Cara- belli’s trait and the protostylid among other dental traits (López-Onaindia & Subirà, 2017). The proto- stylid on M2 and M3, the hypoconulid of M1 and M2, and the entoconulid on M2 and to a lesser ex- tent, groove pattern and cusp number on M2, are suggested to be the most informative in separating Iberian from Italian Neolithic burials (López- Onaindia et al., 2018). The Neolithic cave burials of Belgium exhibit substantial variation in all of these traits, particularly the size of the hypocone and the expression of Carabelli’s trait, and remarkable uni- formity in the presence of a protostylid. Dental morphology of Late Neolithic cave burials of Eurasia Numerous Late Neolithic collective burials exist across Eurasia, such as the Late Neolithic- Chalcolithic collective tombs of Catalonia in which natural crevices and recesses include adults of both sexes and all ages with few grave goods (López- Onaindia et al., 2018). However, the dental mor- phology of only a few Late Neolithic sites have been studied in detail. An important exception concerns those surrounding Lake Baikal, Siberia where an increasingly greater percentage of Cara- belli’s trait occurs during the Neolithic period (Waters-Rist et al., 2016). Compared to the Late Neolithic collective burials of Belgium, a lower ex- pression of this trait is observed and only at Bois Madame and Sclaigneaux is a large Carabelli’s cusp evident (Table 3). Hastiére M and Bois Mad- ame have higher frequencies of a Y groove pattern on M2 (0.250) compared to those observed in Late Neolithic Siberians (0.140) (Waters-Rist et al., 2016), although Sclaigneaux has a much lower value of 0.067 (see Table 4). For cusp number of M2, 71.4% of the Siberian Late Neolithic peoples of Lake Baikal exhibit 5+ cusps whereas the Late Neolithic burials from Belgium can be characterized as ex- pressing fewer cusps on the second mandibular molar. In fact, mostly only four cusps are observed on M2. However, Hastiére M, and to a lesser extent, Bois Madame and Sclaigneaux, show some expres- sion of five cusps on M2, ranging from 0.400 to 0.250 (see Table 4). Expression of a protostylid on M1 is present in half of Late Neolithic peoples of Lake Baikal, Siberia (Waters-Rist et al., 2016), whereas for this temporal period in the Meuse Riv- 46 Dental Anthropology 2021 │ Volume 34│ Issue 01 er basin of Belgium is it present more often than it is absent, and at Sclaigneaux it is observed in 100% of individuals (n = 10) (Table 4). More than a quar- ter of individuals (27%) of the Late Neolithic of Siberia exhibit an entoconulid (Cusp 6) on M1 (Waters-Rist et al., 2016). Comparable frequencies for the collective burials of Late Neolithic Belgium for this trait exist at Hastiére M and to a lesser ex- tent, Bois Madame (Table 4). Unlike their counter- parts to the east who exhibit a low occurrence of a metaconulid (Cusp 7) on M1 at 6.5% (Waters-Rist et al., 2016), at the Late Neolithic caves of Belgium, it is nearly absent with the exception of BM Md 13 from Bois Madame (Figure 6). Conclusions The five well-studied collective burials examined are somewhat discrete in terms of chronology based on radiocarbon dates. Although only limited samples are available for each cave burial, it ap- pears that our predictions were confirmed. Hastière M is only partly distinct from the other cave deposits in the expression of traits, corrobo- rating an analysis of deciduous molar morphology from the Late Neolithic caves of the Belgian Meuse basin (Williams et al., 2018). The final/late collec- tive burials of Sclaigneaux and Bois Madame ex- hibit a greater range of expression of the hypocon- ulid, entoconulid, protostylid, Carabelli’s cusp, metacone and metaconulid. Although differences between the final/late Neolithic cave burials of Sclaigneaux and Bois Madame and the others from Hastière rockshelter are evidenced by dental mor- phology, these sites likely represent ephemeral communities that experienced only limited conti- nuity over time and were perhaps bounded as a function of distance, and to a lesser degree, by chronology. Alternatively, this lack of partitioning of discrete dental traits per burial location may signal that internment was not strictly kin-based as is observed at Neolithic Çatalhöyük (Pilloud & Larsen, 2011), though larger sample sizes to con- duct statistical analyses would be necessary for an investigation into potential kin relations based on dental morphology. In this study, the very low fre- quencies of a metaconulid and the mid-trigonal crest characterize the burials. Furthermore, the ex- pression of Carabelli’s cusp on M1 and M2 joins Sclaigneaux and Bois Madame and secondarily Hastiére M and Trou Garçon. The greater degree of variation observed for the final/late Neolithic cave burials of Sclaigneaux and Bois Madame may have been the result of a slow but steady influx of peo- ples, perhaps along waterways, from other loca- tions as a prelude to the population restructuring that occurred concomitantly with the onset of the Bronze Age. This seems to be the case at other loca- tions in Eurasia (Subirà et al., 2014; Waters-Rist et al., 2016; López-Onaindia et al., 2018). Acknowledgments Permission to examine these Neolithic remains in Belgium was kindly provided by Patrick Semal, Chief of the Scientific Heritage Service, Royal Bel- gian Institute of Natural Sciences. Many thanks to William Anderson and Laura Aday for assisting in the creation of the epoxy resin dental casts. At the Royal Belgian Institute of Natural Sciences, we thank Caroline Polet for generously assisting with the Neolithic collections, and Laurence Cammaert who skillfully created the topographical map of Belgium featured in Figure 1, which we use with permission. We thank Marin Pilloud and the anon- ymous reviewers for valuable comments that sig- nificantly improved the manuscript. Support for this research was provided by Fulbright-Belgium and the Commission for Educational Exchange between the USA, Belgium, and Luxembourg. REFERENCES Alt, K. W., Pichler, S., Vach, W., Klima, B., Vlček, E. & Sedlmeier, J. (1997). Twenty-five thousand- year-old triple burial from Dolní Věstonice: an Ice-Age family? American Journal of Physical Anthropology, 102, 123-131. Bailey, S. E. (2008). Inter- and intra-specific varia- tion in Pan tooth crown morphology: implica- tions for Neandertal taxonomy. In J. D. Irish & G. C. Nelson (Eds.), Technique and application in dental anthropology (pp. 293-316). 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