Williams et al. 2017.1 3 Dental Anthropology 2017 │ Volume 30 │ Issue 01 Expression of Nonmetric Dental Traits in Western European Neanderthals Frank L’Engle Williams 1,* , Rebecca L. George 2 , Marie-Antoinette de Lumley 3,4 , Gaël Becam 3,* 1 Department of Anthropology, Georgia State University, Atlanta, GA 30303 2 Department of Anthropology, University of Nevada, Reno, NV 89557 3 UMR 7194 CNRS, HNHP, MNHN/UPVD/CERP de Tautavel, France Neanderthals are known for their taurodont molar roots, shovel-shaped incisors as well as the expres- sion of dental morphological features that are rela- tively uncommon in modern humans (Patte, 1959; Zubov, 1992; Bailey, 2000, 2002, 2004, 2006; Hublin et al., 2012). However, differences may also exist between Neanderthal sites, irrespective of their relationship to modern humans. Small family groups and a relatively limited degree of genetic variation may have characterized Neanderthals (Lalueza-Fox et al., 2011; Kelso and Prüfer, 2014). Using observations of dental morphology from a relatively large sample of Neanderthals (n = 12) from Western Europe, we examine whether ecoge- ography or chronology (or neither) more adequate- ly accounts for the variation observed. Six individ- uals from a single site (Hortus) allow for a compar- ison of within and between fossil assemblages to investigate the expression of nonmetric dental traits. We also note which features are commonly expressed or are unusual in this sample of Nean- derthals. NEANDERTHALS OF WESTERN EUROPE Several regions of Western Europe appear to have been core areas of Neanderthal habitation, even when colder conditions prevailed (Hublin and Roebroeks, 2009). Neanderthal tools, faunal re- mains, and osteological remnants in these regions are situated primarily between Marine Isotope Stages (MIS) 5 to 3 (Soressi et al., 2007), and are evidenced as far north as Belgium to the terminus of MIS 3 (Würm II) at about 36 kya as demonstrat- ed by the remains from Spy cave (Semal et al., 2005, 2009; Toussaint et al., 2011; Pirson et al., 2014). The earliest Neanderthal discovered in 1829- 1830, Engis 2, is probably also from MIS 3 based on associated Mousterian artifacts and fauna (Twiesselmann, 1971), although it could be young- er, between 26,830 ± 430 and 30,460 ± 210 years BP (Toussaint et al., 2011). Scladina 1-4A, also located along the Meuse River Basin of Belgium, has been dated to MIS 5 or earlier (Pirson et al., 2014), attest- ABSTRACT Neanderthals of Western Europe lived across distinct ecogeographic zones from Marine Iso- tope Stage 7 to 3. Differences in dental morphology from seven Western European sites are compared in terms of ecogeography and chronology. Neanderthals (n = 12) along a north-south gradient were examined. These included the Meuse River Basin of Belgium (Engis 2 and Scladina 1-4A ), Southwest France (Pech de l'Azé 1 and Roc de Marsal), the Pyrenees (Malarnaud and Montmaurin), and the Mediterranean (Hortus). Montmaurin is the oldest, followed by Scladina 1-4A and Malarnaud, whereas the others are younger. Dental casts were prepared from Neanderthal permanent and deciduous dentition. These were de- scribed and scored, according to the ASUDAS. Comparisons of dental traits with respect to ecogeo- graphic regions and chronological categories were constructed. Unusual dental features observed include the anterior fovea, entoconulid, metaconulid, and Cara- belli’s cusp. Dental traits that distinguish ecogeographic regions are the expression of the M1 hypocone and metaconule, whereas the hypoconulid and Carabelli’s cusp separate chronological categories. Dif- ferences are present for the entoconulid and metaconulid in both comparisons. Neither chronology nor ecogeography fully explains the results. Similarities in dental traits exist between Roc de Marsal, Pech de l’Azé 1 and Engis 2, and secondarily within the Hortus assemblage. *Correspondence to: Dr. Frank L’Engle Williams, Department of Anthropol- ogy, Georgia State University e-mail: frankwilliams@gsu.edu Dr. Gaël Becam, UMR 7194 CNRS, HNHP, MNHN/ UPVD/CERP de Tautavel, France gael.becam@cerptautavel.com Keywords: Engis 2, Hortus, Malarnaud, Montmaurin, Pech de l’Azé 1, Roc de Marsal , Scladina 1-4A 4 Dental Anthropology 2017 │ Volume 30 │ Issue 01 ing to the probable recolonization of the Meuse River Ba- sin of Belgium after the climate deterioration of MIS 4 (Hublin and Roebroeks, 2009; Toussaint et al., 2011; Di Modica et al., 2016). Scladina 1-4A is probably between 80 -87 kya based on chronostratigraphic analyses using the Greenland Record as context, although it could be as old as 127 kya (Pirson et al., 2014; Toussaint et al., 2014). Additional Neanderthal sites in Western Europe in- clude those from the Perigord such as Roc de Marsal, which has been dated to 60-70 kya and is close to the ter- minus of MIS 4 (Guérin et al., 2012), and Pech de l’Azé 1 and has been dated to 51-41 kya (Soressi et al., 2007). Fur- ther south along the slopes of the northern Pyrenees in the Ariégeoises region is the site of Malarnaud, which is suggested to be from the Riss-Würm interval (MIS 5). Al- so from the northern Pyrenees region is Montmaurin cave which has been dated to MIS 7 (Crégut-Bonnoure et al., 2010). The Mediterranean region may have been at least par- tially isolated from the Pyrenees and other areas by glaci- er formations in the Massif Central and in particular, the Languedoc Mountains. Between the Mediterranean coast and the piedmont region south of this glaciation is the cave of Hortus yielding between 20 and 36 Neanderthal children and adults, and dated to MIS 3 (Würm II) (Lumley, 1973, 1976). Six of the Hortus remains are exam- ined here, including Hortus II, Hortus IV, Hortus V, Hor- tus VI, Hortus VIII, and Hortus XI. Dental traits in these disparate ecogeographic zones may vary as a function of distance. Alternatively, indi- viduals may differ with respect to time period, whereby older sites (Montmaurin, Scladina 1-4A and Malarnaud) may be distinct from the younger sites of Engis 2, Hortus, Pech de l’Azé 1 and Roc de Marsal. The explanatory pow- er of these models is explored here with respect to the dental morphology of Western European Neanderthals. MATERIALS AND METHODS A total of 12 Neanderthals from seven sites are sam- pled in this study. Two individuals are represented solely by deciduous molars, two preserve both deciduous and permanent teeth and seven include only the permanent molars. Table 1 lists the teeth examined by site and indi- vidual. A macroscopic investigation of the original fossil den- tition was conducted by the first author (FLW) at the Musée de l'Homme, the Centre Européen de Recherches Préhistoriques de Tautavel, the Musée national de Pré- histoire, Les Eyzies-de-Tayac, the Université de Liège, and the Scladina Cave Archaeological Centre. The fossil teeth and Scladina resin casts were molded using polyvi- nylsiloxane (Coltène-Whaledent). Dental casts were creat- ed from the dental molds using centrifuged epoxy resin and hardener (Buehler) at the Bioarchaeology Lab of Georgia State University, and were examined by one ob- server (RLG) using the Arizona State University Dental Anthropology System (ASUDAS; Turner et al., 1991) and a nonmetric data scoring sheet developed by R.L. George. The dental scoring system of the ASUDAS is a replicable method in which the teeth are scored at different intervals to account for the expression of a feature. Deciduous ante- rior teeth were scored according to the Hanihara (1961) standards for deciduous dentition. The macroscopic ob- servations of the original Neanderthal fossil dentition were included where necessary. However, only traits visi- ble on the dental casts were scored and recorded using Fossil Teeth examined Ecogeography Chronology Engis 2 di1, dc, dm1, dm2, dm1, dm2, M1 Meuse River Basin (Belgium) Younger Hortus II M1 Mediterranean (France) Younger Hortus IV M1, M2 Mediterranean (France) Younger Hortus V M1, M2, M3 Mediterranean (France) Younger Hortus VI M3 Mediterranean (France) Younger Hortus VIII M1 Mediterranean (France) Younger Hortus XI M3 Mediterranean (France) Younger Malarnaud M1 Pyrenees (France) Older Montmaurin M1, M2, M3 Pyrenees (France) Older Pech de l'Azé dm1, dm2, dm1, dm2 Southwest France Younger Roc de Marsal dm1, dm2, dm1, dm2 Southwest France Younger Scladina 4A dm1, dm2, M2, M1, M2 Meuse River Basin (Belgium) Older Table 1. Neanderthal dentition used in morphology descriptions. 5 Dental Anthropology 2017 │ Volume 30 │ Issue 01 the ASUDAS. RESULTS Engis 2 Engis 2 preserves a relatively complete immature calvari- um, isolated maxilla and several teeth, and has been aged to between four and five years (Tillier, 1983; Minugh- Purvis, 1988; Toussaint et al., 2011; Williams, 2013; Wil- liams and Cofran, 2016). The dental remains of Engis 2 ex- amined here include the left maxillary deciduous incisors (di1 and di2), which are shoveled (Hanihara grade 3), and a right maxillary deciduous canine characterized by a weak- ly developed tuberculum dentale (Hanihara grade 5). The right maxillary deciduous molars (dm1 and dm2) are de- scribed as well as the right mandibular deciduous first mo- lar (dm1) and right permanent first molar (M1). The right maxillary deciduous first molar (dm1) exhib- its small wear facets on the cusp tips, although much of the original morphology is preserved. There are a total of five cusps and the metacone can be described as only weakly developed (ASUDAS grade 3). Similarly, the hypocone is also small (ASUDAS grade 3) on dm1. The right maxillary deciduous molar (dm2) has a lack of substantial attrition. On dm2 there are four cusps, and the metacone and hypocone are visibly larger than on dm1 (ASUDAS grade 4). A small, but noticeable Carabelli’s trait can be observed on the second deciduous molar (dm2). However, it is not an independent cusp and its distal bor- der does not contact the lingual groove between the proto- cone and the hypocone (ASUDAS grade 5). The right mandibular deciduous second molar (dm2) exhibits a well-developed anterior fovea which is bordered by a pronounced mesial margin (ASUDAS grade 4). The deciduous second molar (dm2) has five cusps inclusive of a prominent hypoconulid (ASUDAS grade 5). Furthermore, a rather large metaconulid can also be observed within the lingual groove between the metaconid and entoconid (ASUDAS grade 4) on dm2. Two large roots are present. The lower right first permanent molar (M1) was proba- bly unerupted or in the process of eruption given the ab- sence of substantial root development (1-2 mm in length) and the virtual lack of attrition with the possible exception of a tiny wear facet on the metaconid. The right permanent first molar (M1) of Engis 2 has six cusps with a weakly de- veloped hypoconulid (ASUDAS grade 1). Lingual to the hypoconulid, a small entoconulid, or sextum tuberculum, is represented as a second lingual groove (ASUDAS grade 1). Furthermore, a small, but noticeable metaconulid exists within the lingual groove between the metaconid and the entoconid (ASUDAS grade 2). The groove pattern on the permanent first molar (M1) can be described as conforming to the “Y” pattern. A weakly developed protostylid can be observed as a small pit on the buccal surface of the proto- conid (ASUDAS grade 1). Hortus II Hortus II, shown in Figure 1, is represented by right and left mandibular permanent first molars and has been aged to 7-8 years (Lumley, 1973). Hortus II is the accompanying lower jaw of the maxilla identified as Hortus III (Lumley, 1973; Minugh-Purvis, 1988). There is only minimal to mod- erate wear on the permanent first molars (M1). The left first molar (M1) of Hortus II (1262) preserves an anterior fovea (ASUDAS grade 3) (Lumley, 1973). The protoconid is large, pointed and separated from the hypoconid by a large buc- cal groove. A large, but short distobuccal groove separates the hypoconid and hypoconulid (Lumley 1973). Similarly, the lingual cusps are separated by a large lingual groove. The groove pattern of the first molar (M1) can be character- ized as exhibiting a “Y” form. The medial ridge of the met- aconid is deflected distally, although it does not approach the entoconid in size (ASUDAS grade 2). A small, well- developed metaconulid is evident (ASUDAS grade 2) (see Fig. 1). Hortus IV Hortus IV consists of a mandibular corpus and the inferior ascending ramus of a young adult aged 18-25 years (Lumley, 1973). Hortus IV preserves the left permanent first and second molars (M1 and M2) and the right perma- nent second incisor (I2), canine (C) and first and second molars (M1 and M2) in situ. The corpus is badly damaged, but the teeth are well preserved and only minimally worn. Heavier wear facets exist along the buccal occlusal surface Figure 1. On the right mandibular permanent first molar (M1) of Hortus II, a number of dental features can be ob- served, such as a metaconulid (a) and an anterior fovea (b). 6 Dental Anthropology 2017 │ Volume 30 │ Issue 01 of both molar rows in contrast to the lingual surfaces (Lumley, 1973). An anterior fovea exists on all the molars (Fig. 2; Lumley, 1973). On the right permanent first molar, the anterior fovea is represented as a large and elon- gated groove distal to a pronounced mesial ridge (ASUDAS grade 4) (see Fig. 2). The same could be said for the left permanent second molar (M2). How- ever, the right permanent second molar (M2) exhibits a slightly less elongated anterior fovea than its anti- mere (ASUDAS grade 3). The groove pattern of the right permanent second molar (M2) evidences an “X” classification. On the second molars (M2), a total of six cusps can be observed and a mid-trigonid crest connects the distal borders of the protoconid and metaconid (ASUDAS grade 1). The buccal surface of the protoconid of the right second molar (M2) exhibits a small protostylid (ASUDAS grade 1). A prominent hypoconulid can be found on the molars (ASUDAS grade 4), although on the right second molar (M2) it is smaller (ASUDAS grade 3). An entoconulid can be observed lingual to the hypoconulid on the second molars (M2); on the right side, the two cuspules are similar in size (ASUDAS grade 3) (see Fig. 2). On the left second molar (M2), the entoconulid is somewhat smaller than the hypoconulid (ASUDAS grade 1). A small metaconulid is evident within the lingual groove between the metaconid and the entoconid on the first and second molars (M1 and M2) (ASUDAS grade 2). Hortus V There are several isolated teeth associated with Hor- tus V, and the age has been estimated to be 18-25 years (Lumley, 1973). The left (730) and right (988) permanent first molars (M1) exhibit moderate wear, heavier on the buccal than on the lingual cusps (Lumley, 1973) and dentine exposures can be ob- served. On the right (988), the metaconid is rather large and pyramidal. The hypoconulid is positioned buccally and distally. Five cusps are visible on the left (730) first molar (M1) and a small protostylid is evi- dent within the buccal groove separating the proto- conid and the hypoconid (ASUDAS grade 1). A well- developed hypoconulid is visible on the first molars (M1), although it is larger on the right (ASUDAS Figure 2. The left (a) and right (b) mandibular permanent first and second molars (M1 and M2) of Hortus IV exhibit well-demarcated anterior foveae. Variation in the development of the entoconulid (Cusp 6) is evident such that it is comparatively smaller vis-à-vis the hypoconulid on the left (c) compared to the relative size of the entoconulid on the right (d) permanent second molars (M2). 7 Dental Anthropology 2017 │ Volume 30 │ Issue 01 grade 3) than on the left (ASUDAS grade 2). The left (693) second molar (M2) exhibits less attrition than the first molars (M1). Taurodont roots are present and the mesial ones are joined together as are the distal ones (Lumley, 1973). A small meta- conulid is nestled within the lingual groove between the metaconid and the entoconid (ASUDAS grade 2), which is unusual on a permanent second molar (Turner et al., 1991). The right (796) and left (695) third molars (M3) present a well preserved and complex morphology. A prominent anterior fovea can be observed on the right (796) third molar (M3), and it is positioned ra- ther mesially (ASUDAS grade 4). A slightly smaller anterior fovea is evident on the left (695) third molar (M3) (ASUDAS grade 3). The right (796) third molar (M3) preserves a groove pattern whereby a “+” clas- sification is evident. There are six cusps on both third molars (M3) and a mid-trigonid crest can be observed (ASUDAS grade 1) on the right (796) third molar (M3). A hypoconulid exists on both third mo- lars (M3), and on the left (695) it is bifurcated and much larger (ASUDAS grade 5) than on the right (796) (ASUDAS grade 3). Both third molars exhibit an entoconulid, albeit the expression varies, such that the left (695) expression (ASUDAS grade 2) is more pronounced than the right (796) (ASUDAS grade 1). Both third molars (M3) also exhibit a pro- nounced metaconulid (ASUDAS grade 2) which is rare for this dental element (Turner et al., 1991). Hortus VI Hortus VI is represented by a left mandibular third molar (M3) from a young adult estimated to be 22-30 years (Lumley, 1973). Much of the original morphol- ogy is preserved (Lumley, 1973). An anterior fovea is present and well-defined (ASUDAS grade 3) (Lumley, 1973). A complex morphology characteriz- es the occlusal surface, and there are a total of six cusps, the first four of which are in close proximity to one another corresponding to the “+” classifica- tion. A weak protostylid is represented as a small fovea within the buccal groove between the proto- conid and hypoconid (ASUDAS grade 1), although it is closer to the superior buccal surface of the hy- poconid. A moderately sized hypoconulid is evident (ASUDAS grade 3) as is an entoconulid, represented as a small fovea on the distal edge of the occlusal surface (ASUDAS grade 3). The hypoconulid and entoconulid are similarly-sized. Hortus VIII Hortus VIII is a well-preserved right maxillary per- manent first molar (M1) which has been aged to 26- 34 years (Lumley, 1973). The tooth is somewhat tri- angular. A prominent metacone exists (ASUDAS grade 5) that rivals the size of the strongly- pronounced hypocone (ASUDAS grade 5), and a large distal groove separates the two. Hortus XI The right maxillary permanent third molar (M3) of Hortus XI has been aged to 45-50 years (Lumley, 1973). The tooth is rectangular in shape. Striations on the occlusal, buccal, and mesial surfaces may possibly derive from paramasticatory behavior (Lumley, 1973). Much of the surface morphology has been destroyed and dentine exposures are evi- dent on the mesial cusps. The metacone is rather small in comparison (ASUDAS grade 3). A weakly developed parastyle is evidenced as a fovea lodged within the buccal groove between the paracone and metacone (ASUDAS grade 1). Malarnaud Malarnaud preserves a right permanent first molar (M1) in situ and crypts for all other permanent teeth are visible, with the exception of the left third molar crown (M3); the right third molar (M3) crown can be partially observed deeply embedded within its crypt, and thus far from eruption. Malarnaud has been aged to around 14 years (Petite-Marie et al., 1971), but may be younger given the minimal wear on the right first permanent molar (M1). A well- developed anterior fovea is present (ASUDAS grade 3). A mid-trigonid crest is evident between the pro- toconid and the metaconid (ASUDAS grade 1). The cusp number is five. A deflecting wrinkle is present (ASUDAS grade 1). A buccal groove between the protoconid and metaconid can be observed (ASUDAS grade 4), although the feature does not approach the development of a protostylid with a free apex. Montmaurin This young adult is represented by a nearly com- plete mandibular corpus and has right and left per- manent molars rows (M1-M3) in pristine condition, in situ, with minimal wear. The wear facets are most pronounced on the left first molar (M1) buccal cusps, followed by those on the right antimere. The anteri- or dentition and premolars are represented by crypts. A clearly demarcated anterior fovea is evi- dent on all three molars, although on the first and second molars (M1 and M2) it is larger (ASUDAS grade 2), in comparison to the third molar (M3) (ASUDAS grade 1) (Fig. 3). The anterior fovea is accentuated further by a strongly developed mid- trigonid crest (ASUDAS grade 1) on the right and left first and second molars (M1 and M2), and on the left third molar (M3). The morphology of all the mo- 8 Dental Anthropology 2017 │ Volume 30 │ Issue 01 lars is complex, particularly the third molars (M3), which can be described as crenulated. On the right first molar (M1), the groove pattern corresponds to the “Y” configuration. On the left second molar (M2), a “+” classification is evident, whereas on the right antimere, only the protoconid and entoconid are in contact indicative of the “X” groove pattern. There are five cusps on the first and second molars (M2 and M2) and the left third molar (M3), and four cusps on the right third molar (M3). A pit can be observed on the buccal surface of the protoconid of the right first molar (M1) (ASUDAS grade 1), and is expressed as a more pronounced groove that curves distally on the buccal surface of the protoconid of the right second molar (M2) (ASUDAS grade 2), though, neither can be described as a true protostylid. The hypoconulid can be characterized as large (ASUDAS grade 5), at least on the left first molar (M1), and, though it is pre- sent on the left second and third molars (M2 and M3), it is much smaller (ASUDAS grade 1). The molars on the right each exhibit a smaller hypoconulid com- pared to their counterparts on the left. Pech de l’Azé I The remains of Pech de l’Azé I include a nearly com- plete cranium and mandible of a young child that has been aged to between 2.5 and 3 years (Heim, 1976; Minugh-Purvis, 1988; Williams, 2013; Williams and Cofran, 2016). All of the deciduous teeth are fully erupted, although, the maxillary lateral incisors are missing postmortem. Dental attrition is minimal such that the deciduous molars preserve much of the orig- inal occlusal morphology. Both of the maxillary deciduous first molars (dm1) exhibit four cusps (ASUDAS grade 4). The right and left deciduous first molars (dm1) exhibit a relatively small metacone, although, a free apex is present (ASUDAS grade 3). The hypocone is weakly developed and is represented as a small ridge on the distobuccal aspect of the deciduous first molars (dm1) on both sides (ASUDAS grade 1). On the deciduous second molars (dm2), five cusps are present. In comparison to the deciduous first molars (dm1), the metacone is quite large on the second molars (dm2) (ASUDAS grade 4). The hy- pocone is much larger on the deciduous second mo- lars (dm2) compared to those expressed on the first molars (dm1). However, the left deciduous second molar (dm2) expresses a large hypocone (ASUDAS grade 4), while on the right antimere it is slightly smaller (ASUDAS grade 3). A prominent Carabelli’s trait is present on the deciduous second molars (dm2) as an independent cusp with a free apex (ASUDAS grade 7) (Fig. 4). A weak metaconule is present with- in the distal groove separating the hypocone and metacone on both deciduous second molars (dm2), although it is somewhat larger on the left (ASUDAS grade 2) than on the right (ASUDAS grade 1). The mandible of Pech de l’Azé 1 is remarkably complete, although only on the right side are the teeth well preserved. The deciduous teeth are fully erupted with minimal attrition. The permanent first molar crown (M1) can be observed deeply in its crypt and the interior occlusal surface of the crown is com- plex. On the right deciduous first molar (dm1) of Pech de l’Azé 1, an elongated and pronounced anterior fovea is evident, bordered by a prominent mesial ridge (ASUDAS grade 4). Four cusps can be ob- served on the right deciduous first molar (dm1). The right deciduous second molar (dm2) exhibits a protostylid, although it is only weakly developed, consisting primarily of a raised fovea that is low and flat with a small indentation that is deep, nestled Figure 3. A well-developed, oblong anterior fovea can be observed on all left permanent molars (M1, M2 and M3, from left to right) of Montmaurin, albeit its expression differs. Note the absence of the entoconulid and metaconulid (Cusps 6 and 7, respectively). 9 Dental Anthropology 2017 │ Volume 30 │ Issue 01 within the buccal groove separating the protoconid and hypoconid (ASUDAS grade 1). There are a total of five cusps evident on the right deciduous second molar (dm2), including a pronounced hypoconulid (ASUDAS grade 5). In addition, there is a large met- aconulid with a free apex situated within the lingual groove that separates the metaconid and entoconid (ASUDAS grade 4) on the right second deciduous molar (dm2). Roc de Marsal The relatively complete skeleton of the Roc de Mar- sal child, has been aged to approximately three years (Heim, 1976; Minugh-Purvis, 1988; Williams, 2013; Williams and Cofran, 2016). The deciduous dentition is fully erupted, and only minimal attrition charac- terizes the deciduous molars. The permanent first molar crowns (M1 and M1) can be seen fully devel- oped and embedded within the crypts. On the right and left maxillary deciduous first molars (dm1), wear facets are visible on the hy- pocone. There are four cusps on both the right and left deciduous first molars (dm1). A large distal groove separates the metacone, which is rather small, but is represented as an independent cuspule (ASUDAS grade 2) (see Fig. 4) from the hypocone, which is poorly-developed (ASUDAS grade 1) on both deciduous first molar (dm1) antimeres. On the hypocone of the deciduous second molar (dm2) a large wear facet is evident, stronger on the left than the right, and a deep cleft separates the hy- pocone and the trigone. There are five cusps evident on this dental element. These include a metacone, which is pronounced in size on the right (ASUDAS grade 5), but much smaller on the left antimere (ASUDAS grade 3). The hypocone on the second deciduous molar (dm2) is larger on the left (ASUDAS grade 4) than on the right (ASUDAS grade 3). A weak metaconule exists on both decidu- ous second molars (dm2), although, like the hy- pocone, it is larger on the left (ASUDAS grade 2) than on the right (ASUDAS grade 1). On the decidu- ous second molar (dm2), a well-developed and inde- pendent Carabelli’s cusp is evident (ASUDAS grade 7) on both sides (see Fig. 4). Small wear facets exist on the buccal cusps of the mandibular deciduous first molars (dm1), although they are deeper on the right than on the left. On the deciduous first molar (dm1) an elongated and deep anterior fovea is present, bordered by a well- developed mesial ridge (ASUDAS grade 4) on right and left sides. On the right, the cusp number is four, whereas the left deciduous first molar (dm1) pre- sents five cusps. Similar to the mandibular deciduous first molar (dm1), small wear facets can be observed on the buc- cal cusps of the deciduous second molars (dm2), al- though they are more extensive on the right than the left. The metaconid central ridge is deflected in a distal direction, although, it does not reach the ento- conid (ASUDAS grade 2) on the left second decidu- ous molar (dm2). On the right, the deflecting wrinkle is more weakly developed (ASUDAS grade 1). On the right deciduous second molar (dm2), a proto- stylid can be observed as a full cusp, represented on the buccal surface of the protoconid as a buccal groove (ASUDAS grade 6). A hypoconulid exists on right and left deciduous second molars (dm2). How- ever, it is medium in size on the right (ASUDAS grade 3) and small on the left (ASUDAS grade 2). On the left deciduous second molar (dm2), a small meta- conulid can be observed (ASUDAS grade 2). Scladina 1-4A Scladina 1-4A comprises right and left demi- Figure 4. The Roc de Marsal right maxillary decidu- ous first and second molars (dm1 and dm2) exhibit a number of traits, including a small hypocone (a) on the deciduous first molar (dm1), and a pronounced Carabelli’s cusp (b) on the deciduous second molar (dm2). 10 Dental Anthropology 2017 │ Volume 30 │ Issue 01 mandibles, a right maxillary fragment as well as 11 teeth from a single older child, aged to 8 to 11 years (Smith et al., 2007, 2014; Toussaint et al. 2011, 2014; Williams, 2013; Toussaint, 2014; Williams and Cofran, 2016). The deciduous teeth examined here include the right maxillary molars (dm1 and dm2). Three permanent molars are also represented. These include the right maxillary second molar (M2) and the right mandibular first and second molars (M1 and M2). Scores for these teeth using the ASUDAS are reported by Toussaint (2014); the two sets of scores present more similarities than differences. The right maxillary deciduous first molar (dm1) of Scladina 4A-7 is substantially worn and exhibits a small metacone (ASUDAS grade 3) (Toussaint, 2014). The cusp number of the deciduous first molar (dm1) is three. The right maxillary deciduous second molar (dm2) of Scladina 4A-5 is moderately worn (Toussaint, 2014). Dentine exposures exist on the protocone and hypocone surfaces. The metacone is larger on the deciduous second molar (dm2) (ASUDAS grade 4) than on dm1. The hypocone can be described as rather large (ASUDAS grade 4) (Toussaint, 2014). The right maxillary permanent second molar (M2) of Scladina 4A-3 is unerupted and the tooth crown is rectangular in shape (Toussaint, 2014). The paracone is large, and both the metacone and hypocone are present, although they differ in grade. Similar to the deciduous second molar (dm2), the permanent sec- ond molar (M2) has a well-developed metacone (ASUDAS grade 4), although the hypocone can be characterized as relatively smaller in size (ASUDAS grade 3). Between the hypocone and metacone of the permanent second molar (M2), a metaconule can be observed, although it is small (ASUDAS grade 3). The protocone features a small “Y” indentation on the lingual surface indicative of a rudimentary Cara- belli’s trait (ASUDAS grade 3). The mandibular permanent first and second mo- lars (M1 and M2) of Scladina 4A-1 are associated with the right demi-mandible. There are wear facets pre- sent on the permanent first molar (M1); wear is greatest on the protoconid, followed by the hypoco- nid. On the permanent first molar (M1), a fovea ex- ists within the buccal groove separating the proto- conid and the hypoconid. However, it cannot be de- scribed as a true protostylid (ASUDAS grade 1). A hypoconulid can be observed, and is rather large (ASUDAS grade 4) (Toussaint, 2014). The permanent second molar (M2) is not fully erupted, and exhibits a complex occlusal morpholo- gy (Toussaint, 2014). A tall mesial ridge borders an elongated and deep anterior fovea on the permanent second molar (M2) (ASUDAS grade 4). The groove pattern indicative of the “+” configuration can be discerned. There are six cusps on the second molar (M2), including a small hypoconulid (ASUDAS grade 2). The six cusps also include a tuberculum sex- tum, although it is substantially smaller than the hy- poconulid (ASUDAS grade 1). ECOGEOGRPAHY There are a number of differences in dental mor- phology across geographic regions (Table 2). For example, in the Mediterranean (Hortus), the meta- cone and hypocone are larger on maxillary decidu- ous and permanent first molars (dm1 and M1) than in the other regions. On the mandibular teeth, indi- viduals from Hortus differ from Montmaurin of the Pyrenees in the expression of an anterior fovea, pro- tostylid, entoconulid, and metaconulid on the per- manent third molar (M3). The presence of a metacon- ule on maxillary deciduous and permanent first mo- lars (dm1 and M1) is unique to the Meuse River Basin of Belgium, as is cusp number on the mandibular deciduous and permanent first molars (dm1 and M1). The Pyrenees and Southwest France do not exhibit any dental traits that are lacking representation in the other ecogeographic regions (Table 2). Dental traits that are expressed in all of the eco- geographic regions include the anterior fovea on the deciduous and permanent first molars (dm1 and M1), which is found on a total of 16 teeth (Table 2). Addi- tionally, a cusp number of 5+ and the presence of a hypoconulid on the deciduous and permanent sec- ond molars (dm2 and M2) are expressed in all ecoge- ographic regions (Table 2). Dental traits that appear in the Mediterranean, Pyrenees, and the Meuse River Basin of Belgium, but not in Southwest France include the development of the protostylid and hypoconulid on the deciduous and permanent first molars (dm1 and M1), and a pro- nounced anterior fovea on the deciduous and per- manent second molars (dm2 and M2) (Table 2). The Pyrenees and Mediterranean sites share 11 traits in common, versus five traits shared between the Med- iterranean and Southwest France. The Pyrenees and Southwest France share four traits in common (Table 2). The dental traits that distinguish ecogeographic regions include the expression of the M1 hypocone and metaconule on the maxillary molars, and the entoconulid and metaconulid on mandibular molars. The expression of the metacone, anterior fovea, and mandibular molar cusp numbers are the most simi- lar across ecogeographic regions. Chronology All of the dental features present in the older period 11 Dental Anthropology 2017 │ Volume 30 │ Issue 01 Tooth type Trait Trait presence Mediterran- ean Meuse River Basin Pyrenees Southwest France dm1 & M1 (n=7) Cusp number Score 5+ present (n=1) Metacone Score 4-5 present (n=1) Hypocone Score 4-5 present (n=1) dm2 & M2 (n=7) Cusp number Score 4+ present (n=1) present (n=4) Metacone Score 4-5 present (n=3) present (n=3) Hypocone Score 2-5 present (n=3) present (n=4) Carabelli Score 5-7 present (n=1) present (n=4) Cusp 5 Score 1-5 present (n=1) present (n=4) dm1 & M1 (n=16) Anterior fovea Score 2-4 present (n=2) present (n=1) present (n=2) present (n=3) Cusp number Score 6+ present (n=1) Mid-trigonid crest Score 1 present (n=1) present (n=3) Protostylid Score 1-7 present (n=1) present (n=2) present (n=1) Cusp 5 Score 3-5 present (n=3) present (n=1) present (n=3) Cusp 6 Score 1-5 present (n=1) Cusp 7 Score 1-4 present (n=3) present (n=1) dm2 & M2 (n=11) Anterior fovea Score 2-4 present (n=2) present (n=2) present (n=2) Cusp number Score 5+ present (n=2) present (n=2) present (n=1) present (n=1) Mid-trigonid crest Score 1 present (n=2) present (n=2) Protostylid Score 1-7 present (n=1) present (n=1) present (n=2) Cusp 5 Score 1-5 present (n=2) present (n=2) present (n=1) present (n=3) Cusp 6 Score 1-5 present (n=2) present (n=1) Cusp 7 Score 1-4 present (n=4) present (n=1) present (n=2) M3 (n=5) Anterior fovea Score 2-4 present (n=3) Cusp number Score 5+ present (n=3) present (n=1) Mid-trigonid crest Score 1 present (n=1) present (n=1) Protostylid Score 1-7 present (n=1) Cusp 5 Score 1-5 present (n=3) present (n=1) Cusp 6 Score 1-5 present (n=3) Cusp 7 Score 1-4 present (n=2) Table 2. Dental trait comparison by ecogeographical region. 12 Dental Anthropology 2017 │ Volume 30 │ Issue 01 are also found in the younger one, with the excep- tion of the Hortus assemblage which lacks Carabel- li’s trait where it could be observed (Hortus VIII) (Table 3), corroborating de Lumley (1973). Dental features that are present in late MIS 4 and MIS 3, but absent in Scladina 1-4A, include the presence of a metaconule and a pronounced expression of the hy- pocone and metacone. On the maxillary deciduous and permanent second molars (dm2 and M2), Scladi- na 1-4A lacks a cusp number of 4+ and a well- developed Carabelli’s feature, as noted by Toussaint (2014). The older time period also lacks more than five cusps on mandibular deciduous and permanent first molars (dm1 and M1), and on the mandibular deciduous and permanent second molars (dm2 and M2), a metaconulid is not observed. On the third mo- lar (M3), Montmaurin of the older period differs from the younger Hortus remains, by lacking a pro- nounced anterior fovea, and any expression whatso- ever of a protostylid, entoconulid, and metaconulid (Table 3). On the maxillary molars, no obvious difference is found between older and younger periods, with the possible exception of Carabelli’s trait (Table 3). The dental features which may differentiate these broad chronological categories of older and younger include the expression of the hypoconulid, the ento- conulid, and metaconulid (Table 3). The dental traits that are the most similar across chronological peri- ods include the expression of the anterior fovea and Tooth type Trait Trait presence Older Younger dm1 & M1 (n=7) Cusp number Score 5+ present (n=1) Metacone Score 4-5 present (n=1) Hypocone Score 4-5 present (n=1) dm2 & M2 (n=7) Cusp number Score 4+ present (n=5) Metacone Score 4-5 present (n=2) present (n=4) Hypocone Score 2-5 present (n=2) present (n=5) Carabelli Score 5-7 present (n=5) Cusp 5 Score 1-5 present (n=1) present (n=4) dm1 & M1 (n=16) Anterior fovea Score 2-4 present (n=2) present (n=6) Cusp number Score 6+ present (n=1) Mid-trigonid crest Score 1 present (n=3) present (n=1) Protostylid Score 1-7 present (n=2) present (n=2) Cusp 5 Score 3-5 present (n=4) present (n=3) Cusp 6 Score 1-5 present (n=1) Cusp 7 Score 1-4 present (n=4) dm2 & M2 (n=11) Anterior fovea Score 2-4 present (n=3) present (n=3) Cusp number Score 5+ present (n=2) present (n=4) Mid-trigonid crest Score 1 present (n=2) present (n=2) Protostylid Score 1-7 present (n=1) present (n=3) Cusp 5 Score 1-5 present (n=2) present (n=6) Cusp 6 Score 1-5 present (n=1) present (n=2) Cusp 7 Score 1-4 present (n=7) M3 (n=5) Anterior fovea Score 2-4 present (n=3) Cusp number Score 5+ present (n=1) present (n=3) Mid-trigonid crest Score 1 present (n=1) present (n=1) Protostylid Score 1-7 present (n=1) Cusp 5 Score 1-5 present (n=1) present (n=3) Cusp 6 Score 1-5 present (n=3) Cusp 7 Score 1-4 present (n=2) Table 3. Dental trait comparison by site chronology. 13 Dental Anthropology 2017 │ Volume 30 │ Issue 01 protostylid on the mandibular molars, and cusp number and hypocone size on the maxillary first molars (Table 3). DISCUSSION Anterior fovea The anterior fovea appears with near ubiquity in the Neanderthals (Bailey, 2006), and other Pleistocene humans (Hrdlička, 1924; Turner et al., 1991). Howev- er, variation is also observed. For example, the ante- rior fovea is not as deep or distinct in Hortus II as is observed on Montmaurin and Malarnaud. Addition- ally, the anterior fovea of the first and second molars (M1 and M2) of Montmaurin is smaller than that ob- served on the first molar (M1) of Malarnaud. It is prominent on the deciduous first molars (dm1) of Pech de l’Azé 1 and Roc de Marsal. Bailey (2006) notes an inter-correlation between the presence of an anterior fovea and the mid-trigonid crest. This rela- tionship was noted in some individuals, such as Hortus IV, Hortus V, and Montmaurin, but not oth- ers (Table 2). Entoconulid and metaconulid The entoconulid and metaconulid, also known as Cusp 6 and Cusp 7, respectively, are routinely ab- sent in the Pyrenees sample, whereas these accesso- ry cusps are present in the Meuse River Basin of Bel- gium category (Table 2). Although the entoconulid is absent in the individuals from Southwest France, the metaconulid is present. Both of these cusps are found in Hortus for the second molars (M2); on the first molars (M1), only the metaconulid can be ob- served. Bailey (2006) observes rather low frequencies of Cusp 6 and Cusp 7 (20-40%), with the exception of the entoconulid in the second and third molars (M2 and M3), which approaches 50%. In this study, the metaconulid occurs more often than does the ento- conulid. Hypocone The hypocone of the maxillary first molars (dm1 and M1) differentiates the ecogeographic groupings (Table 3). This contrast is based primarily on the re- duced expression of this feature in Southwest France (Pech de l’Azé 1 and Roc de Marsal) and the Meuse River Basin of Belgium (Engis 2) contrasting to the enlarged (M1) hypocone in Hortus VIII. Southwest France and the Meuse River Basin of Belgium are aligned in other features that suggest more similari- ty than expected from such distant locations. Bailey (2006) notes that the hypocone is usually present and enlarged in Neanderthals. This is certainly true of Hortus VIII who expresses a large hypocone (Lumley, 1973), but not the deciduous molars of this study. Carabelli’s cusp Carabelli’s trait is noted often in this study, particu- larly on the deciduous second molars where its strongest expression is observed. Bailey (2006) finds that it also occurrs relatively consistently in Nean- derthals in both first (68%) and second molars (50%), corroborating the observations noted for Southwest France and the Meuse River Basin of Belgium, but not in Hortus (Table 2). Additional considerations Zubov (1992) reports a prevalence of the mid- trigonid crest (epicristid) on Neanderthal mandibu- lar molars. Only in the Pyrenees is there a consistent presence of this dental trait across the molar row; although, it is also noted in the Meuse River Basin of Belgium (see Table 2). However, it appears to be En- gis 2, presumably from the younger MIS 3, rather than Scladina 1-4A (MIS 5), which exhibits a mid- trigonid crest. Bailey (2006) reports the expression of at least five cusps on the molars is nearly ubiquitous in Ne- anderthals, an observation corroborated in this study (Table 2). The rarity of the expression of a de- flecting wrinkle in Neanderthals (Bailey, 2006) is supported here as it is present only in Hortus II, Ma- larnaud, and Roc de Marsal. In addition, the “Y” groove pattern for M1 is relatively widespread in Neanderthals as reported in Bailey (2006), and is found in Engis 2, Hortus II, Scladina 1-4A, and Montmaurin. Bailey (2006) suggests the “X” groove pattern for M2 is relatively common, which in this study is expressed in Hortus IV and Montmaurin. It is evident that the Meuse River Basin of Bel- gium is not distinct (Table 2), at least in the younger time period, as it shares several traits in common with Neanderthals from Southwest France. The two sites closest in proximity (Mediterranean and Pyre- nees) are not the most similar, suggesting ecogeo- graphic distance fails to account for all the differ- ences among regions. The fact that all of the traits examined are represented in the younger category (Table 3) suggests that chronology does explain some of the variation, at least for mandibular traits, although a greater number of individuals are includ- ed in the younger category. For the maxillary mo- lars, only Carabelli’s trait differentiates the older/ younger categories (Table 3). CONCLUSIONS The expression of the anterior fovea is common in Neanderthals (Lumley, 1973; Bailey, 2000, 2006). It is often found on mandibular deciduous and perma- nent molar rows, and its presence on M1 increases 14 Dental Anthropology 2017 │ Volume 30 │ Issue 01 the chance that it will be present on both M2 and M3 (Fig. 3). The anterior fovea is expressed on decidu- ous mandibular molars, such as in Pech de l'Azé 1, Roc de Marsal, and Engis 2, as is a prominent Cara- belli’s cusp. Other features such as the expression of the entoconulid and metaconulid are found on the permanent molars in the Hortus assemblage and in Engis 2, but not in the Pyrenees. Maxillary molar hypocone size and the presence of a metaconule dif- fer between Mediterranean Hortus VIII on the one hand, and Southwest France and the Meuse River Basin of Belgium on the other. Montmaurin and Ma- larnaud from the Pyrenees region resemble one an- other in the expression of traits on mandibular M1 and are dissimilar to Scladina 1-4A. Likewise, Hor- tus II and Hortus IV resemble one another, although Hortus II and Hortus V are younger (Phase V) than Hortus IV (Phase IV) (Lumley, 1973). However, they are not identical, and Hortus IV and Hortus V also share traits to the exclusion of Hortus II, suggesting ecogeography alone does not account for the varia- tion in the expression of dental features. Chronology may account for the absence in MIS 7 to 5 of 6+ cusps in mandibular molars, the expression of a well -developed entoconulid and metaconulid, and a pro- nounced Carabelli’s feature, all of which are ob- served in late MIS 4/MIS 3. ACKNOWLEDGMENTS It is with much gratitude that the following curators are thanked for their permission to examine the Ne- anderthal fossils featured here: Aurélie Fort and Lili- ana Huet, Musée de l'Homme; Marie-Antoinette de Lumley, Gaël Becam, and Tony Chevalier, Centre Européen de Recherches Préhistoriques de Tautavel; Jean-Jacques Cleyet-Merle, Musée National de Pré- histoire Les Eyzies-de-Tayac; Valentin Fischer, Uni- versité de Liège, and Dominique Bonjean and Michel Toussaint, Scladina Cave Archaeological Centre. The image utilized in Figure 4 featuring Roc de Marsal belongs to the collections of the Musée National de Préhistoire, les Eyzies, Dordogne, France, and was used with the permission of Jean-Jacques Cleyet- Merle. 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