The Vindija Neanderthal scapular glenoid fossa: Comparative shape analysis suggests evo-devo changes among Neanderthals Fabio Di Vincenzo a, b , Steven E. Churchill c, d , Giorgio Manzi a, b, * a Dipartimento di Biologia Ambientale, Sapienza Università di Roma, 00185 Roma, Italy b Istituto Italiano di Paleontologia Umana, 00198 Roma, Italy c Department of Evolutionary Anthropology, Duke University, Durham, NC 27708, USA d Institute for Human Evolution, University of the Witwatersrand, Wits 2050, South Africa article info Article history: Received 23 May 2011 Accepted 12 November 2011 Keywords: Human evolution Neanderthal/modern human interactions Geometric morphometrics Shoulder Late Pleistocene Europe abstract Although the shape of the scapular glenoid fossa (SGF) may be influenced by epigenetic and develop- mental factors, there appears to be strong genetic control over its overall form, such that variation within and between hominin taxa in SGF shape may contain information about their evolutionary histories. Here we present the results of a geometric morphometric study of the SGF of the Neanderthal Vi-209 from Vindjia Cave (Croatia), relative to samples of Plio-Pleistocene, later Pleistocene, and recent homi- nins. Variation in overall SGF shape follows a chronological trend from the plesiomorphic condition seen in Australopithecus to modern humans, with pre-modern species of the genus Homo exhibiting inter- mediate morphologies. Change in body size across this temporal series is not linearly directional, which argues against static allometry as an explanation. However, life history and developmental rates change directionally across the series, suggesting an ontogenetic effect on the observed changes in shape (ontogenetic allometry). Within this framework, the morphospace occupied by the Neanderthals exhibits a discontinuous distribution. The Vindija SGF and those of the later Near Eastern Neanderthals (Kebara and Shanidar) approach the modern condition and are somewhat segregated from both northwestern European (Neandertal and La Ferrassie) and early Mediterranean Neanderthals (Krapina and Tabun). Although more than one scenario may account for the pattern seen in the Neanderthals, the data is consistent with palaeogenetic evidence suggesting low levels of gene flow between Neanderthals and modern humans in the Near East after ca. 120e100 ka (thousands of years ago) (with subsequent introgression of modern human alleles into eastern and central Europe). Thus, in keeping with previous analyses that document some modern human features in the Vindija Neanderthals, the Vindija G 3 sample should not be seen as representative of ‘classic’ e that is, unadmixed, pre-contact e Neanderthal morphology. Ó 2011 Elsevier Ltd. All rights reserved. Introduction The morphology of the scapular glenoid fossa (SGF) has long been of interest to scholars of human evolution (Vallois, 1928e46; McCown and Keith, 1939; Stewart, 1962; Oxnard,1967; Endo and Kimura, 1970; Trinkaus, 1983; Churchill and Trinkaus, 1990). Traditional osteometric studies indicate that the SGF in Homo sapiens is more dorso-ventrally expanded and more concave, with higher mean values of the width/height index, compared with that of Homo neanderthalensis (Vallois, 1928e46; Stewart, 1962; Trinkaus, 1983; Churchill and Trinkaus, 1990). The relatively narrow and shallow SGF of the Neanderthals occurs in the context of elongated scapulae (Churchill, 1994; Churchill and Rhodes, 2006) with a tendency to have dorsal sulcate axillary borders (Eickstedt, 1925; Trinkaus, 1977). These differences have been functionally interpreted in light of contrasts between Neanderthals and early modern Europeans in behaviour patterns and their associated loadings during humeral rotation and abduction, having to do with the adoption of innovative throwing-based weapons for hunting at a distance in the Upper Paleolithic (Trinkaus, 1977; Churchill and Trinkaus, 1990; Churchill and Rhodes, 2009; Rhodes and Churchill, 2009). However, the Neanderthal-like narrow SGF occurs at low-to- moderate frequencies even in post-Neolithic modern human pop- ulations, calling into question the inferred relationship between SGF morphology and throwing activities (Trinkaus, 2006; Churchill and * Corresponding author. E-mail address: giorgio.manzi@uniroma1.it (G. Manzi). Contents lists available at SciVerse ScienceDirect Journal of Human Evolution journal homepage: www.elsevier.com/locate/jhevol 0047-2484/$ e see front matter Ó 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.jhevol.2011.11.010 Journal of Human Evolution 62 (2012) 274e285