Hand to Mouth in a Neandertal: Right-Handedness in Regourdou 1 Virginie Volpato1, Roberto Macchiarelli2,3, Debbie Guatelli-Steinberg4, Ivana Fiore5, Luca Bondioli5, David W. Frayer5,6* 1 Department of Paleoanthropology and Messel Research, Senckenberg Research Institute Frankfurt, Frankfurt am Main, Germany, 2 De´partement de Pre´histoire, UMR 7194, MNHN, Paris, France, 3 De´partement Ge´osciences, Universite´ de Poitiers, Poitiers, France, 4 Department of Anthropology, The Ohio State University, Columbus, Ohio, United States of America, 5 Soprintendenza al Museo Nazionale Preistorico Etnografico ‘‘L. Pigorini’’, Rome, Italy, 6 Department of Anthropology, University of Kansas, Lawrence, Kansas, United States of America Abstract We describe and analyze a Neandertal postcranial skeleton and dentition, which together show unambiguous signs of right- handedness. Asymmetries between the left and right upper arm in Regourdou 1 were identified nearly 20 years ago, then confirmed by more detailed analyses of the inner bone structure for the clavicle, humerus, radius and ulna. The total pattern of all bones in the shoulder and arm reveals that Regourdou 1 was a right-hander. Confirmatory evidence comes from the mandibular incisors, which display a distinct pattern of right oblique scratches, typical of right-handed manipulations performed at the front of the mouth. Regourdou’s right handedness is consistent with the strong pattern of manual lateralization in Neandertals and further confirms a modern pattern of left brain dominance, presumably signally linguistic competence. These observations along with cultural, genetic and morphological evidence indicate language competence in Neandertals and their European precursors. Citation: Volpato V, Macchiarelli R, Guatelli-Steinberg D, Fiore I, Bondioli L, et al. (2012) Hand to Mouth in a Neandertal: Right-Handedness in Regourdou 1. PLoS ONE 7(8): e43949. doi:10.1371/journal.pone.0043949 Editor: Carles Lalueza-Fox, Institut de Biologia Evolutiva – Universitat Pompeu Fabra, Spain Received June 7, 2012; Accepted July 27, 2012; Published August 22, 2012 Copyright: 2012 Volpato et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. Funding: The TNT European project, French CNRS, Re´gion Poitou-Charentes, Nespos Society, and Leakey Foundation provided financial support. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript. Competing Interests: The authors have declared that no competing interests exist. * E-mail: frayer@ku.edu Introduction Various studies have identified striations on the labial (lip) face of Neandertal anterior teeth, beginning with Henri-Martin’s initial observations on the upper incisors from La Quina 5 [1]. These scratches are commonly found in Neandertals from Europe [2–6] and in their likely ancestors from Sima de los Huesos at Atapuerca [7–9] and Mauer [10]. Except for two left-handed individuals, Krapina [KDP] 4 and Hortus 8, dated ,130,000 yrs and ,35,000 yrs, respectively, all specimens show a preponderance of right-handed striations [9]. This yields a ratio of 27 right- handed: 2 left-handed (93%: 7%), approximating the high frequencies of right-handedness found in all modern populations world-wide [9,11–13]. Experimental evidence suggests these scratches were produced inadvertently as items were clenched and processed by stone tools between the canines and incisors [7,14]. Despite earlier contentions that manipulative scratches do not appear in modern populations [15], some recent human hunter-gatherer populations show similar striations [16,17]. However, compared to Neandertals and their European prede- cessors, striations are more rare in later populations. In all studies where scratches have been quantified, the marks occur only on anterior teeth, which are isolated or in jaws not directly associated with other skeletal elements. Here, we describe for the first time, to our knowledge, a Neandertal dentition, preserving multiple teeth with distinct right-handed scratches, associated with a non- pathological upper limb skeleton exhibiting unambiguous right- handedness. Regourdou 1 was discovered in 1957 in a collapsed rock shelter near Lascaux in the Dordogne of southwestern France. In the deposits were remains of a partial skeleton (Figure 1) along with fragmentary pedal remains of a second individual [18,19]. Associated with La Quina type Mousterian [20] and considered to be from OIS 4, it is one of the oldest Neandertal skeletons from Western Europe [21,22]. The skeleton (Regourdou 1) was likely part of a burial [18,21], but the bones were disordered and incomplete, due to taphonomic conditions and early excavation procedures. Regourdou 1 consists of mandible and primarily the parts of the upper torso with scattered vertebra, the upper 1/2 of the sacrum, elements from the right and left ilia, and some bones of the lower limb, notably for the right side [23,24]. Based on mandibular tooth wear and the closed medial clavicle epiphyses Regourdou 1 was probably between 23–30 years old at death, using modern epiphyseal closure data [25]. Sex is generally considered ‘indeterminate’ based on the small size of the long bones [19,24], but the anatomical proportions of the body and alae in the superior sacrum [26,27] suggest that it was a male. Despite the small size of the posterior teeth [20], Regourdou 19s canine breadth is large and at 10.0 mm it is greater than all European Wu¨ rm Neandertal females (mean = 8.2 mm, range: 7.5–9.7 mm, n = 10) and at the top of the range for Neandertal males (mean = 9.5 mm, range: 8.8–10.1 mm, n = 12). Like PLOS ONE | www.plosone.org 1 August 2012 | Volume 7 | Issue 8 | e43949