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Krapina and Other Neanderthal Clavicles: a Peculiar Morphology? Krapina and Other Neanderthal Clavicles : A Peculiar Morphology? Jean-Luc Voisin To cite this version: Jean-Luc Voisin. Krapina and Other Neanderthal Clavicles : A Peculiar Morphology?. Periodicum Biologorum, 2006, 108 (3), pp.331-339. halshs-00352689 HAL Id: halshs-00352689 https://halshs.archives-ouvertes.fr/halshs-00352689 Submitted on 13 Jan 2009 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. PERIODICUM BIOLOGORUM UDC 57:61 VOL. 108, No 3, 331–339, 2006 CODEN PDBIAD ISSN 0031-5362 Original scientific paper Krapina and Other Neanderthal Clavicles: A Peculiar Morphology? Abstract JEAN-LUC VOISIN The clavicle is the less studied element of the shoulder girdle, even if it is USM 103 a very important bone for human evolution because it permits all move- Institut de Paléontologie Humaine ments outside the parasagittal plan. In this work, clavicle curvatures are 1 rue René Panhard 75013 Paris studied by projecting them on a cranial and a dorsal plan, which are perpen- E-mail: [email protected] dicular. In cranial view, there is no difference within the genus Homo, and [email protected] Neanderthal clavicles are not more S-shaped than modern human ones. On the contrary, the dorsal view allows to distinguish two human groups. The first includes Homo habilis, Homo ergaster and Neanderthal. Their clavi- Key words: Clavicle, Shoulder, cles are characterized by two curvatures, an inferior one at the acromial end Evolution, Homo habilis, Homo ergaster, and a superior one at the sternal end. The second group includes only mod- Homo antecessor, Neanderthal ern human, whose clavicles are characterized by the presence of the inferior curvature only. The shape of the clavicle in dorsal view is associated to the position of the scapula in regard to the thorax. Two curvatures are associated to a high scapula, and, on the contrary, a unique curvature is associated to a scapula in a low position in regard to the thorax. Moreover, the two curva- tures of the modern human clavicle in dorsal view constitute an apomor- phic character in regard to the other human species. INTRODUCTION ven if the clavicle is less studied than the other two bones of the Eshoulder complex, it is not out of interest because it allows all movements of the upper limbs outside the parasagittal plane. Most works on this bone are strictly anthropological and give a good over- view of its morphological variations (1–15). On the contrary, just a very few studies compare Primate clavicles from the point of view of com- parative anatomy or functional morphology (16–26). This latter re- mark explains in part why fossil clavicles are so neglected and limited to descriptions although some authors have tried to go further (25, 27–30). In this work we describe clavicle morphology within the genus Homo, from Homo habilis to modern human, and try to interpret differ- ences and similarities in adaptive and evolutionary ways. MATERIALS AND METHODS Materials Our samples include the following fossils: Homo habilis, Homo Received April 27, 2006. ergaster, Homo antecessor, Neanderthal and Upper Palaeolithic remains (Table 1) and modern human (MH) or Homo sapiens sapiens from sev- J.-L. Voisin Krapina and Other Neanderthal Clavicles: A Peculiar Morphology? TABLE 1 Fossil clavicles studied; * Original remains; R: right and L: left; Upper Palaeolithic humans include Abri Pataud, Omo I KSH and all Taforalt remains. Middle Palaeolithic Upper Palaeolithic humans (UP) Neanderthal (Nd) Régourdou (R) Abri Pataud (R)* Omo I KSH (L) Qafzeh 9 (L) Régourdou (L) Taforalt (Taf) Kebara 2 (R) Taf V-6* (R) Taf XIIIa* (L) Kebara 2 (L) Taf XXIII* (L) Taf VIII-3bis* (L) La Ferrassie I (R)* Taf XIX-3a* (L) Taf XXVa* (R) La Ferrassie I (L)* Neanderthal (R) Taf V-24* (L) Taf IX-39* (L) Krapina 153 (L)* Taf XI-AR* (R) Taf XIIIb* (R) Krapina 142 (R)* TafXVa*(L) Taf XVIIa* (R) Krapina 143 (R)* Taf XVc* (L) Taf XVI-15* (R) Krapina 154 (L)* Taf XVII-26* (R) Taf XVI* (R) Krapina 145 (R)* Taf XVIII-6* (R) Taf XIV* (L) Krapina 145 (R)* Taf 24-5* (R) Taf XIX-3* (R) Krapina 155 (L)* Taf XXVc* (L) Krapina 149 (R)* Krapina 144 (R)* Lower Palaeolithic Krapina 156 (L)* Homo antecessor Homo ergaster La Chapelle-aux-Saints (L)* Gran Dolina ATD6-50 KNM-WT 15000 Homo habilis OH 48 eral parts of the world (33 clavicles). This material be- longs to the Laboratoire d’Anthropologie Biologique du Musée de l’Homme, Paris (France), the Institut de Paléon- tologie Humaine, Paris (France) and the Croatian Natu- ral History Museum, Zagreb (Croatia). Most fossil clavicles used in this study are well pre- served even if some are eroded at their extremities, and only La Chapelle-aux-Saints, Krapina 149 right, 145 right, 144 right, 155 left, 156 left and Qafzeh 9 left are not complete (31–33). Methods Due to its complexity, the morphology of the clavicle will be approached in regard to its curvatures. When pro- jected on two perpendicular plane, one cranial and one Figure 1. Masurements of clavicle curvatures (34). Example on a right dorsal, the clavicle morphology can be decomposed in el- clavicle of Pan troglodytes. ementary curvatures, as shown on Figure 1. The middle arc of curvature is estimated according to Olivier’s method (34) as the proportion between the length Dorsal plane: of the chord and the height of the curvature (Figure. 1): • The acromial curvature (inferior one): e’ / h’.100 Cranial plane: • The sternal curvature (superior one): f ’ / g’.100 • The acromial curvature (external one):e/h.100 The total length of the clavicle is measured with a cal- • The sternal curvature (internal one):f/g.100 liper square as the greatest length of the bone. 332 Period biol, Vol 108, No 3, 2006. Krapina and Other Neanderthal Clavicles: A Peculiar Morphology? J.-L. Voisin The measurements and distribution of the variables TABLE 2 have been computed with ViStat 6.4® (35). Graphics Values of the external and internal curvatures in Upper showing the range of variation of each variable are repre- Palaeolithic remains, Neanderthal, Homo antecessor, Homo ergaster and Homo habilis. #: The incompleteness of the bone sented by the mean and +/– two times standard deviation. make impossible to know if a curvature exist or not. RESULTS Individual Internal External curvature curvature Curvatures in cranial view Upper Abri Pataud 9.7 12.1 Palaeolithic In superior view, all fossil clavicles studied here are Omo I KSH (L) 13.1 14.56 distributed within the range of variation of modern hu- Qafzeh 9 (L) 10.7 / man ones (Figure 2, Table 2). This result means that Taf V-6 (R) 19.1 16.3 Neanderthal clavicles, in superior view are less S-shaped Taf XXIII (L) 10.9 9.9 than classically described (27, 29, 31, 36, 37) and show no Taf XIX-3a (L) 14.0 10.6 differences with modern human ones in cranial view Taf V-24 (L) 16.3 13.2 (Table 3). This result is confirmed by other recent works Taf XI-AR (R) 13.4 11.2 (24–26). Taf XVa (L) 18.0 13.8 Some of the older remains display nevertheless some Taf XVc (L) 18.9 16.5 features of their own, even if their range of variation stays Taf XVII-26 (R) 19.3 12.7 within that of modern human clavicles. In KNNM-WT Taf XVIII-6 (R) 16.3 15.0 15000 (left and right) and OH 48 clavicles the internal Taf XIIIa (L) 11.9 11.4 curvature is more pronounced than the external one (Ta- Taf VIII-3bis (L) 16.5 13.2 ble 2). In the case of OH48, this morphology may be due Taf XXVa (R) 17.1 10.5 to its state of conservation, because, as Napier (38) show- Taf IX-39 (L) 17 10.5 ed it, this clavicle lack a great part of is acromial extremity Taf XIIIb (R) 16.5 12.8 and thus its external curvature is underestimated. On the Taf XVIIa (R) 14.3 14.4 contrary, the KNM-WT 15000 clavicles is nearly com- plete (39). Taf XVI-15 (R) 13.1 8.2 Taf XVI (R) 20.8 16.8 Curvatures in dorsal view Taf XIV (L) 16.2 12.2 Taf 24-5 (R) 14.7 11.6 In dorsal view, modern human clavicles can be classi- fied into three morphological groups, or types (Figure 3 Taf XIX-3 (R) 20.5 18.2 Taf XXVc (L) 20.2 14.0 Neanderthal Régourdou (R) 10.0 14.5 Régourdou (L) 11.9 13.2 Kebara (L) 11.1 16.5 Kebara (R) 11.3 9.5 La Ferrassie 1 (R) 13.2 14.2 La Ferrassie 1 (L) 12.3 17.7 Neanderthal (R) 13.2 16.7 Krapina 153 (L) 10.8 10.3 Krapina 142 (R) 17.4 16.7 Krapina 143 (R) 13.9 12.1 Krapina 154 (L). 14.1 11.4 Krapina 149 (R) 20.0 / Krapina 145 (R) / 11.3 Krapina 144 (R) 12.0 / Krapina 155 (L) 18.1 / Krapina 156 (L) / 14.7 La Chapelle-aux-Saints (L) / 12.2 Mean 13.5 13.6 Standard deviation 3.0 2.6 Homo KNM-ER15 000 (R) 13.1 14.6 ergaster KNM-ER15 000 (L) 15.3 14.7 Figure 2.
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