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Neanderthal Teeth from Moula-Guercy, Arde`Che, France

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Neanderthal Teeth from Moula-Guercy, Arde`Che, France AMERICAN JOURNAL OF PHYSICAL ANTHROPOLOGY 151:477–491 (2013) Neanderthal Teeth from Moula-Guercy, Arde` che, France Leslea J. Hlusko,1* Joshua P. Carlson,1 Debbie Guatelli-Steinberg,2 Kristin L. Krueger,3 Ben Mersey,1 Peter S. Ungar,4 and Alban Defleur5 1Human Evolution Research Center, University of California Berkeley, 3101 Valley Life Sciences Building, Berkeley, CA 94720 2Department of Anthropology, 4034 Smith Laboratory, The Ohio State University, 174 West 18th Columbus, OH 43210-1106 3Department of Anthropology, Loyola University Chicago, Chicago, IL 60660 4Department of Anthropology, University of Arkansas, Fayetteville, AR 72701 5CNRS UMR 5276, Laboratoire de Geologie de Lyon, Ecole Normale Superieure de Lyon, 46, Allee d’Italie, 69364 Lyon Cedex 07, France KEY WORDS evolution; dental variation; paleontology; Hominidae ABSTRACT Here we describe dental remains from a significant number of linear enamel hypoplasias indi- Neanderthal fossil assemblage from Moula-Guercy, cates that these individuals were stressed during child- France. Our report demonstrates that the Moula-Guercy hood. Molar microwear data suggest that these hominid remains contribute important morphological, Neanderthals did not differ significantly from modern developmental, and behavioral data to understanding humans in terms of the fracture properties of the food Neanderthal evolutionary history. We include gross com- they were consuming. The incisor microwear and macro parative morphological descriptions and enamel surface striations provide evidence that these individuals may microstructure and microwear data. These teeth reveal have been using their anterior teeth as tools, similar to numerous characteristics that are diagnostic of Neander- the practices of several modern human populations such thals and provide no evidence for the presence of any as the Inuit, Ipiutak, and Australian Aboriginals, and other hominid taxa. Enamel growth increment data reminiscent of evidence from other Neanderthals from from the Moula-Guercy specimens yield evidence of a Krapina, Croatia, as well as the 600,000 year old homi- Neanderthal pattern of development, although at the nids from Sima de los Huesos, Spain. Am J Phys lower end of the range of variation. The presence of a Anthropol 151:477–491, 2013. VC 2013 Wiley Periodicals, Inc. Ever since the first extensive study of Neanderthal extended the sequence to a total of 7 m and revealed the skeletal variation (Gorjanovic´-Kramberger, 1906), the first Neanderthal remains (Defleur et al., 1998). Strati- dentition has proven to be highly informative of Nean- graphic analysis led to the division of the sequence into derthal origins, adaptation, and evolution (e.g., Kallay, 20 layers grouped into three climatic phases correspond- 1951; Carbonell, 1965; Dean et al., 1986; Brace et al., ing to marine isotope stages 4–6 (Desclaux and Defleur, 1987; Stringer et al., 1997; Irish, 1998; Bailey, 1997; Defleur et al., 1998; Defleur et al., 2001). The upper 2006aa,b). New discoveries continue to be made in the unit, Phase I, comprises Layers VI–XI. A date of 72 6 12 field, laboratory, and also in museum collections (e.g., ka has been assigned to a volcanic ash (Layer VI) above Walker et al., 2008; Crevecoeur et al., 2010; Smith et al., the hominid level (Sanzelle et al., 2000). Fauna from this 2010; Benazzi et al., 2011; Walker et al., 2011), making Phase includes Mammuthus primigenius and Rangifer Neanderthals one of the most important extinct homi- tardanus. Together with associated micromammals, these nids for studying species paleobiology. Here, we describe taxa indicate a cold climate consistent with MIS 4. The for the first time the Neanderthal fossil dental assem- underlying unit, Phase II, consists of Layers XII–XVI and blage from Moula-Guercy, France. Our analysis includes includes the majority of the archeological material, all of three approaches to understanding these remains: (1) comparative descriptions of the fossil material, (2) enamel surface microstructure, and (3) microwear. Additional Supporting Information may be found in the online ver- Other anatomical components of the skeleton are sion of this article. forthcoming. Grant sponsor: KLK and PU; NSF BCS-0925818 DDIG; DGS; STRATIGRAPHIC, CHRONOLOGICAL, NSF BCS-0607520. ECOLOGICAL, AND ARCHEOLOGICAL *Correspondence to: Leslea J. Hlusko; Human Evolution CONTEXTS Research Center, University of California Berkeley, 3101 Valley Life Sciences Building, Berkeley, CA 94720, USA. Discovered in 1972, Moula-Guercy cave overlooks the E-mail: [email protected] Rhone River 10 km south of the city of Valence in the parish of Soyons, Arde`che, France. Initial work at the Received 1 August 2012; accepted 16 April 2013 site between 1975 and 1982 uncovered 5 m of infill char- acterized by a Mousterian assemblage associated with DOI: 10.1002/ajpa.22291 fauna indicating a cold climate (Defleur et al., 1993aa). Published online 4 June 2013 in Wiley Online Library Subsequent excavations spanning from 1991–1999 (wileyonlinelibrary.com). Ó 2013 WILEY PERIODICALS, INC. 478 L.J. HLUSKO ET AL. the Neanderthal remains, and fauna and charcoal con- waves approximately 100 lm apart (Hillson and Bond, sistent with a more temperate climate. Biostratigraphy of 1997). In the cuspal region of the tooth, striae of Retzius micromammals indicates a date of 100–120 ka for Phase do not emerge onto the enamel surface, such that this II, consistent with MIS 5. The base of the sequence, portion of enamel growth is hidden from a surface view. Phase III, includes Layers XVII–XX. These contain However, the time lateral enamel takes to form repre- micromammal remains indicating a cold, steppe environ- sents approximately 85% or more of the overall enamel ment consistent with MIS 6. formation time of anterior teeth, at least in modern When excavations were halted in 1999, 108 hominid humans (Reid and Dean, 2006). Thus, the number of remains (45 craniodental and 63 postcranial specimens) perikymata on anterior teeth reflects a large percentage had been recovered from Layer XV. The best preserved of the time it took for the crown to form. Perikymata of the elements are two partial crania, a partial femur, counts on Neanderthal teeth tend to fall within the teeth, and many complete hand and foot bones. As range of modern human variation, but are at the low reported previously (Defleur et al., 1993b; Defleur et al., end of that range for particular teeth (the upper incisors 1999), the remains conform to typical Neanderthal mor- and lower canines, Guatelli-Steinberg and Reid, 2008; phology and diagnostic characters include Neanderthal anterior teeth, Ramirez-Rozzi and Bermudez de Castro, apomorphies. These remains also show significant peri- 2004). In addition, in Neanderthals perikymata are more mortem damage consistent with the nutritional exploita- evenly distributed across the enamel surface than they tion of a minimum of six individuals: four juveniles/ are in modern humans (Guatelli-Steinberg et al., 2007; children, two adults (Defleur et al., 1999). Shortly after Guatelli-Steinberg and Reid, 2008). the publication of these remains (Defleur et al., 1999), Linear enamel hypoplasia (LEH) appears as a line, excavations were halted by local officials. As only 30% of groove or furrow in the enamel surface, reflecting peri- the primary layer (Layer XV) had been excavated prior ods of disrupted growth caused by systemic physiological to this, full taphonomic assessment and determination of stress, such as malnutrition and disease (Goodman and the minimum number of individuals is being withheld Rose, 1990; Ten Cate, 1994). Systemic physiological pending resumption and completion of the excavation stress is likely to affect all of the teeth of an individual work. However, we do note that our reassessment does developing at the time of the stress episode (Hillson and not counter the original interpretation of six individuals: Bond, 1997). Growth increments on the surface of the two were 15–16 years of age at death, two were 6–7 root known as periradicular bands also may become years of age at death, and there are two adult specimens accentuated, presumably as a result of growth disrup- representing one large and one small individual (Defleur tion (Smith et al., 2007). et al., 1999). While the enamel microstructure of a tooth is informa- tive of the development and health of an individual, we can add to our reconstruction of an organism’s life his- THE STUDY OF NEANDERTHAL DENTAL tory by investigating the damage to the crown surface REMAINS during life. We have also undertaken an investigation of dental wear—the study of microscopic scratches and pits The gross morphological structure of teeth provides that form on a tooth’s surface as the result of its use. enormous information about an organism’s adaptive re- Such analyses of nonhuman primates and recent human gime and evolutionary history. Given their largely inor- foragers indicate that microwear patterns on incisors ganic content, teeth are often the only remaining body vary with front tooth behaviors and those on molars part we have for many extinct animals. Neanderthals vary with the fracture properties of foods eaten (see are no exception, as numerous studies of their phylogeny Ungar et al., 2007 for review). Dental microwear pat- and paleobiology have relied on tooth crown shape (e.g., terns have also been shown to distinguish fossil homi- Boule, 1921; Bailey, 2006a,b). Trinkaus’ summary of nids, suggesting their value for inferring diet and tooth primitive versus derived
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