. .. -introducción,

. .. Métodos, 2.6-Precisión estadística (accuracy) y márgenes de error, vol.257

. .. -resultados,

. .. -discusión, 4.4-Modularidad intra-vertebral. Hipótesis dos-módulos

. .. -conclusiones,

, Selección evolutiva e integración morfológica en humanos modernos 239

. Este, Apéndice: Artículos, p.334, 2019.

R. R. Ackermann, Morphological integration and the interpretation of fossil hominin diversity, Evolutionary Biology, vol.36, pp.149-156, 2009.

R. R. Ackermann and J. M. Cheverud, Phenotypic covariance structure in tamarins (genus Saguinus): a comparison of variation patterns using matrix correlation and common principal component analysis, American Journal of Physical Anthropology, vol.111, issue.4, pp.489-501, 2000.

D. C. Adams, Evaluating modularity in morphometric data: challenges with the RV coefficient and a new test measure, Methods in Ecology and Evolution, vol.7, pp.565-572, 2016.

D. C. Adams, F. J. Rohlf, and D. E. Slice, Geometric morphometrics: ten years of progress following the 'revolution', Italian Journal of Zoology, vol.71, issue.1, pp.5-16, 2004.

D. C. Adams and M. L. Collyer, On the comparison of the strength of morphological integration across morphometric datasets, Evolution, vol.70, pp.2623-2631, 2016.

D. C. Adams, M. L. Collyer, A. Kaliontzopoulou, and E. Sherratt, Geomorph: Software for geometric morphometric analyses, 2017.

L. M. Adams and W. J. Moore, Biomechanical appraisal of some skeletal features associated with head balance and posture in the Hominoidea, Acta anatomica, vol.92, issue.4, pp.580-584, 1975.

L. Aiello and C. Dean, An introduction to human evolutionary anatomy, 1990.

D. M. Alba, Fossil apes from the vallès-penedès basin, Evolutionary Anthropology: Issues, News, and Reviews, vol.21, issue.6, pp.254-269, 2012.

G. H. Albrecht, Multivariate analysis and the study of form, with special reference to canonical variate analysis, American Zoologist, vol.20, issue.4, pp.679-693, 1980.

Z. Alemseged, F. Spoor, W. H. Kimbel, R. Bobe, D. Geraads et al., A juvenile early hominin skeleton from Dikika, Ethiopia, Nature, vol.443, issue.7109, pp.296-301, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00838289

F. Ankel, Vertebral morphology of fossil and extant primates, The functional and evolutionary biology of primates, vol.295, pp.223-240, 2017.

F. Ankel-simons, Primate anatomy: an introduction, 2010.

J. D. Archibald, Extinction and radiation: how the fall of dinosaurs led to the rise of mammals, 2011.

B. Arensburg, The vertebral column, thoracic cage and hyoid bone, Le squelette moustérien de Kébara, pp.113-147, 1991.

C. Éditions-du,

M. Arlegi, A. Gómez-olivencia, L. Albessard, I. Martínez, A. Balzeau et al., The role of allometry and posture in the evolution of the hominin subaxial cervical spine, Journal of Human Evolution, vol.104, pp.80-99, 2017.
URL : https://hal.archives-ouvertes.fr/hal-02106776

M. Arlegi, A. Gómez-robles, and A. Gómez-olivencia, Morphological integration in the gorilla, chimpanzee, and human neck, American Journal of Physical Anthropology, vol.166, issue.2, pp.408-416, 2018.

P. Arnold, F. Forterre, J. Lang, and M. S. Fischer, Morphological disparity, conservatism, and integration in the canine lower cervical spine: insights into mammalian neck function and regionalization, Mammalian Biology-Zeitschrift für, 2016.

. Säugetierkunde, , vol.81, pp.153-162

P. Arnold, B. Esteve-altava, and M. S. Fischer, Musculoskeletal networks reveal topological disparity in mammalian neck evolution, BMC Evolutionary Biology, vol.17, p.251, 2017.

J. L. Arsuaga and I. Martínez, The chosen species: the long march of human evolution, 2005.

J. L. Arsuaga, I. Martínez, L. J. Arnold, A. Aranburu, A. Gracia-téllez et al., Neandertal roots: Cranial and chronological evidence from Sima de los Huesos, Science, vol.344, issue.6190, pp.1358-1363, 2014.

J. L. Arsuaga, J. M. Carretero, C. Lorenzo, A. Gómez-olivencia, A. Pablos et al., Postcranial morphology of the middle Pleistocene humans from Sima de los Huesos, vol.112, pp.11524-11529, 2015.

R. J. Asher, K. H. Lin, N. Kardjilov, and L. Hautier, Variability and constraint in the mammalian vertebral column, Journal of Evolutionary Biology, vol.24, pp.1080-1090, 2011.

E. H. Ashton and S. Zuckerman, , 1956.

, Proceedings of the Zoological Society of London, vol.126, pp.315-326

W. R. Atchley and D. Anderson, Ratios and the statistical analysis of biological data, Systematic Zoology, vol.27, issue.1, pp.71-78, 1978.

K. M. Bagnall, P. F. Harris, and P. R. Jones, A radiographic study of the human fetal spine. 2. The sequence of development of ossification centres in the vertebral column, Journal of Anatomy, vol.124, p.791, 1977.

J. B. Bailey, Neural spine elongation in dinosaurs: sailbacks or buffalo-backs?, Journal of Paleontology, vol.71, issue.6, pp.1124-1146, 1997.

C. Barrey, J. Jund, O. Noseda, and P. Roussouly, Sagittal balance of the pelvis spine complex and lumbar degenerative diseases. A comparative study about 85 cases, European Spine Journal, vol.16, pp.1459-1467, 2007.

M. Bastir, A. Rosas, and H. D. Sheets, The morphological integration of the hominoid skull: a partial least squares and PC analysis with implications for European Middle Pleistocene mandibular variation, Modern Morphometrics in Physical Anthropology, pp.265-284, 2005.

M. Bastir, D. G. Martínez, W. Recheis, A. Barash, M. Coquerelle et al., Differential growth and development of the upper and lower human thorax, PloS one, vol.8, issue.9, p.75128, 2013.

W. Bateson, Materials for the study of variation treated with especial regard to discontinuity in the origin of species, 1894.

E. Been, The anatomy of the lumbar spine of Homo neanderthalensis and its phylogenetic and functional implications. Doctoral dissertation (Unpublished), 2005.

E. Been, S. Peleg, A. Marom, and A. Barash, Morphology and function of the lumbar spine of the Kebara 2 Neandertal, American Journal of Physical Anthropology, vol.142, issue.4, pp.549-557, 2010.

E. Been, A. Gómez-olivencia, and P. A. Kramer, Lumbar lordosis of extinct hominins, American Journal of Physical Anthropology, vol.147, issue.1, pp.64-77, 2012.

E. Been and L. Kalichman, Lumbar lordosis, The Spine Journal, vol.14, issue.1, pp.87-97, 2014.

E. Been, A. Gómez-olivencia, and P. A. Kramer, Brief Communication: Lumbar, 2014.

E. Been, S. Shefi, M. Soudack, L. R. Zilka, A. Barash et al., Cervical lordosis and the orientation of the foramen magnum, implications to human evolution, American Journal of Physical Anthropology, vol.153, p.75, 2014.

E. Been, S. Shefi, L. R. Zilka, and M. Soudack, Foramen magnum orientation and its association with cervical lordosis: a model for reconstructing cervical curvature in archeological and extinct hominin specimens, Advances in Anthropology, vol.4, issue.3, p.133, 2014.

E. Been, A. Gómez-olivencia, S. Shefi, M. Soudack, M. Bastir et al., Evolution of Spinopelvic Alignment in Hominins, The Anatomical Record, vol.300, pp.900-911, 2017.

E. Been and J. Bailey, The Association Between Spinal Posture and Spinal Biomechanics in Modern Humans: Implications for Extinct Hominins, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.283-299, 2019.

Y. Benjamini and Y. Hochberg, Controlling the false discovery rate: a practical and powerful approach to multiple testing, Journal of the Royal Statistical Society. Series B, Statistical methodology, vol.57, issue.1, pp.289-300, 1995.

C. V. Bennett and A. Goswami, Does developmental strategy drive limb integration in marsupials and monotremes?, Mammalian Biology, vol.76, issue.1, pp.79-83, 2011.

R. S. Benton, Morphological evidence for adaptations within the epaxial region of the primates, The baboon in medical research, vol.2, pp.201-216, 1967.

R. L. Berg, The ecological significance of correlation pleiades, Evolution, vol.14, issue.2, pp.171-180, 1960.

L. R. Berger, D. J. De-ruiter, S. E. Churchill, P. Schmid, K. J. Carlson et al., Australopithecus sediba: a new species of Homo-like australopith from South Africa, Science, vol.328, issue.5975, pp.195-204, 2010.

L. R. Berger, J. Hawks, D. J. De-ruiter, S. E. Churchill, P. Schmid et al.,

M. M. Skinner, Homo naledi, a new species of the genus Homo from the Dinaledi Chamber, South Africa. Elife, vol.4, p.9560, 2015.

J. Biegert, The evaluation of characteristics of the skull, hands and feet for primate taxonomy, Classification and human evolution, pp.116-145, 1963.

N. T. Boaz, R. L. Ciochon, Q. Xu, and J. Liu, Mapping and taphonomic analysi of the Homo erectus loci at Locality 1 Zhoukoudian, China. Journal of Human Evolution, vol.46, issue.5, pp.519-549, 2004.

L. Bolk, Over de ligging en verschuiving van het Foramen magnum bij de Primaten. Verslag vd Gew. Verg. der Wis-en Natuurkundige afdeeling vd Koninklijke Academie van Wetenschappen, p.26, 1909.

A. Bonmatí, A. Gómez-olivencia, J. L. Arsuaga, J. M. Carretero, A. Gracia et al., Middle Pleistocene lower back and pelvis from an aged human individual from the Sima de los Huesos site, vol.107, pp.18386-18391, 2010.

F. L. Bookstein, P. D. Gingerich, and A. G. Kluge, Hierarchical linear modeling of the tempo and mode of evolution, Paleobiology, vol.4, issue.2, pp.120-134, 1978.

F. L. Bookstein, Foundations of morphometrics, Annual Review of Ecology and Systematics, vol.13, issue.1, pp.451-470, 1982.

F. L. Bookstein, A statistical method for biological shape comparisons, Journal of Theoretical Biology, vol.107, issue.3, pp.475-520, 1984.

F. L. Bookstein, Size and shape": a comment on semantics, Systematic zoology, vol.38, pp.173-180, 1989.

F. L. Bookstein, Thin-plate splines and the atlas problem for biomedical images, International Conference on Information Processing in Medical Imaging, pp.326-342, 1991.

F. L. Bookstein, L. F. Marcus, M. Corti, A. Loy, and G. J. Naylor, Combining the tools of geometric morphometrics, Advances in morphometrics, pp.131-151, 1996.

F. L. Bookstein, Integration, disintegration, and self-similarity: characterizing the scales of shape variation in landmark data, Evolutionary Biology, vol.42, pp.395-426, 2015.

F. L. Bookstein, , 2017.

F. L. Bookstein, P. D. Sampson, A. P. Streissguth, and H. M. Barr, Measuring "dose" and "response" with multivariate data using partial least squares techniques, Communications in Statistics-Theory and Methods, vol.19, issue.3, pp.765-804, 1990.

B. M. Boszczyk, A. A. Boszcyk, and R. Putz, Comparative and functional anatomy of the mammalian lumbar spine, The Anatomical Record, vol.264, pp.157-168, 2001.

C. Boulay, C. Tardieu, J. Hecquet, C. Benaim, B. Mouilleseaux et al., Sagittal alignment of spine and pelvis regulated by pelvic incidence: standard values and prediction of lordosis, European Spine Journal, vol.15, pp.415-422, 2006.

M. Boule, L'homme fossile de la Chapelle aux Saints, Annales de Paléontologie, vol.6, pp.1-70
URL : https://hal.archives-ouvertes.fr/halshs-01488527

D. M. Bramble and D. E. Lieberman, Endurance running and the evolution of Homo, Nature, vol.432, issue.7015, p.345, 2004.

G. Bräuer, Osteometrie, Anthropologie. Hand-buch der vergleichenden Biologie des Menschen, pp.160-232, 1988.

P. Broca, Sur la direction du trou occipital, Description du niveau occipital et du goniomètre occipital. Bulletins et Mémoires de la Société d'Anthropologie de Paris, vol.7, pp.649-668, 1872.

R. Broom and J. T. Robinson, A new type of fossil man, Nature, vol.164, issue.4164, p.322, 1949.

L. T. Brown, The mechanics of the lumbosacral and sacro-iliac joints, JBJS, vol.19, issue.3, p.775, 1937.

F. Brown, J. Harris, R. Leakey, and A. Walker, Early Homo erectus skeleton from west lake Turkana, Nature, issue.6031, pp.788-792, 1985.

P. Brown, T. Sutikna, M. J. Morwood, R. P. Soejono, E. W. Saptomo et al., A new small-bodied hominin from the Late Pleistocene of Flores, Indonesia, Nature, issue.7012, pp.1055-1061, 2004.

E. Bruner and G. Manzi, Allometric analysis of the skull in Pan and Gorilla by geometric morphometrics, Riv. Antropol, vol.79, pp.45-52, 2001.

E. A. Buchholtz, Vertebral and rib anatomy in Caperea marginata: implications for evolutionary patterning of the mammalian vertebral column, Marine Mammal Referencias, p.300, 2011.

, Science, vol.27, pp.382-397

E. A. Buchholtz, Flexibility and constraint: patterning the axial skeleton in mammals. From clone to bone: the synergy of morphological and molecular tools in palaeobiology, pp.230-253, 2012.

E. A. Buchholtz, Crossing the frontier: a hypothesis for the origins of meristic constraint in mammalian axial patterning, Zoology, vol.117, pp.64-69, 2014.

E. A. Buchholtz and C. C. Stepien, Anatomical transformation in mammals: developmental origin of aberrant cervical anatomy in tree sloths, Evolution & Development, vol.11, pp.69-79, 2009.

E. A. Buchholtz, H. G. Bailin, S. A. Laves, J. T. Yang, M. Y. Chan et al., Fixed cervical count and the origin of the mammalian diaphragm, Evolution & Development, vol.14, pp.399-411, 2012.

E. A. Buchholtz, K. L. Wayrynen, and I. W. Lin, Breaking constraint: axial patterning in Trichechus (Mammalia: Sirenia), Evolution & Development, vol.16, pp.382-393, 2014.

R. Bürger, Constraints for the evolution of functionally coupled characters: a nonlinear analysis of a phenotypic model, Evolution, vol.40, issue.1, pp.182-193, 1986.

A. C. Burke, C. E. Nelson, B. A. Morgan, and C. Tabin, Hox genes and the evolution of vertebrate axial morphology, Development, vol.121, pp.333-346, 1995.

W. P. Cane, The ontogeny of postcranial integration in the common tern, Sterna hirundo, Evolution, vol.47, pp.1138-1151, 1993.

E. Carbonell, J. B. De-castro, J. L. Arsuaga, J. C. Diez, A. Rosas et al., Lower Pleistocene hominids and artifacts from Atapuerca-TD6 (Spain), pp.826-830, 1995.

A. Cardini, Integration and modularity in Procrustes shape data: is there a risk of spurious results?, Evolutionary Biology, vol.46, issue.1, pp.90-105, 2019.

J. M. Carretero, C. Lorenzo, and J. L. Arsuaga, Axial and appendicular skeleton of Homo antecessor, Journal of Human Evolution, vol.37, issue.3-4, pp.459-499, 1999.

D. R. Carrier, The evolution of locomotor stamina in tetrapods: circumventing a mechanical constraint, Paleobiology, vol.13, issue.3, pp.326-341, 1987.

A. J. Cave, The morphology of the mammalian cervical pleurapophysis, 1975.

J. Y. Chen, The sudden appearance of diverse animal body plans during the Cambrian explosion, International Journal of Developmental Biology, vol.53, issue.5-6, pp.733-751, 2009.

J. Y. Chen, J. Dzik, G. D. Edgecombe, L. Ramsköld, and G. Q. Zhou, A possible Early Cambrian chordate, Nature, vol.377, issue.6551, pp.720-722, 1995.

J. M. Cheverud, Phenotypic, genetic, and environmental morphological integration in the cranium, Evolution, vol.36, issue.3, pp.499-516, 1982.

J. Cheverud, Quantitative genetics and developmental constraints on evolution by selection, Journal of Theoretical Biology, vol.110, pp.155-171, 1984.

J. M. Cheverud, A comparison of genetic and phenotypic correlations, Evolution, vol.42, issue.5, pp.958-968, 1988.

J. M. Cheverud, Morphological integration in the saddle-back tamarin (Saguinus fuscicollis) cranium, The American Naturalist, vol.145, issue.1, pp.63-89, 1995.

J. M. Cheverud, Developmental integration and the evolution of pleiotropy, 1996.

, American Zoologist, vol.36, pp.44-50

J. M. Cheverud, J. J. Rutledge, and W. R. Atchley, Quantitative genetics of development: genetic correlations among age-specific trait values and the evolution of ontogeny, Evolution, vol.37, pp.895-905, 1983.

J. M. Cheverud, G. P. Wagner, and M. M. Dow, Methods for the comparative analysis of variation patterns, Systematic Zoology, vol.38, pp.201-213, 1989.

L. M. Chevin, R. Lande, and G. M. Mace, Adaptation, plasticity, and extinction in a changing environment: towards a predictive theory, PLoS biology, vol.8, p.1000357, 2010.

H. Choi, E. Keshner, and B. W. Peterson, Comparison of cervical musculoskeletal kinematics in two different postures of primate during voluntary head tracking, KSME international journal, vol.17, issue.8, pp.1140-1147, 2003.

B. Christ, R. Huang, and J. Wilting, The development of the avian vertebral column, Anatomy and Embryology, vol.202, issue.3, pp.179-194, 2000.

A. Cil, M. Yazici, A. Uzumcugil, U. Kandemir, A. Alanay et al., The evolution of sagittal segmental alignment of the spine during childhood Spine, vol.30, pp.93-100, 2005.

R. J. Clarke, The history of research in human evolution in Africa and what lessons have been learned, Human origin sites and the World Heritage Convention in Africa, vol.33, pp.44-67, 2012.

R. J. Clarke, Australopithecus from Sterkfontein Caves, South Africa, The paleobiology of Australopithecus, pp.105-123, 2013.

R. J. Clarke, Excavation, reconstruction and taphonomy of the StW 573, 2019.

, Australopithecus prometheus skeleton from Sterkfontein Caves, South Africa, Journal of Human Evolution, vol.127, p.41

R. J. Clarke and P. V. Tobias, Sterkfontein Member 2 foot bones of the oldest South African hominid, Science, vol.269, issue.5223, pp.521-524, 1995.

R. J. Clarke and K. Kuman, The skull of StW 573, a 3.67 Ma Australopithecus prometheus skeleton from Sterkfontein Caves, South Africa, Journal of Human Evolution, vol.134, p.102634, 2019.

M. N. Coleman, What does geometric mean, mean geometrically? Assessing the utility of geometric mean and other size variables in studies of skull allometry, 2008.

, American Journal of Physical Anthropology, vol.135, issue.4, pp.404-415

D. C. Cook, J. E. Buikstra, C. J. Derousseau, and D. C. Johanson, Vertebral pathology in the Afar australopithecines, American Journal of Physical Anthropology, vol.60, issue.1, pp.83-101, 1983.

A. W. Crompton and F. A. Jenkins, Mammals from reptiles: a review of mammalian origins, Annual Review of Earth and Planetary Sciences, vol.1, pp.131-155, 1973.

J. N. Darroch and J. E. Mosimann, Canonical and principal components of shape, 1985.

, Biometrika, vol.72, pp.241-252

C. Darwin, The Origin of Species; And, the Descent of Man. Modern library, 1859.

G. R. De-beer, The development on the vertebrate skull, 1937.

M. A. De-lumley, L'homme de Tautavel. Un Homo erectus européen évolué. Homo erectus tautavelensis, l'Anthropologie, vol.119, issue.3, pp.303-348, 2015.

M. C. Dean and B. A. Wood, Basicranial anatomy of Plio-Pleistocene hominids from East and South Africa, American Journal of Physical Anthropology, vol.59, issue.2, 1982.

P. H. Dirks, E. M. Roberts, H. Hilbert-wolf, J. D. Kramers, J. Hawks et al., The age of Homo naledi and associated sediments in the Rising Star Cave, 2017.

A. G. Drake and C. P. Klingenberg, The pace of morphological change: historical transformation of skull shape in St Bernard dogs, Proceedings of the Royal Society of London B: Biological Sciences, vol.275, pp.71-76, 2008.

I. L. Dryden and K. V. Mardia, Statistical analysis of shape, 1998.

G. J. Eble, Contrasting evolutionary flexibility in sister groups: disparity and diversity in Mesozoic atelostomate echinoids, Paleobiology, vol.26, issue.1, pp.56-79, 2000.

G. J. Eble, The macroevolution of phenotypic integration. Phenotypic integration: studying the ecology and evolution of complex phenotypes, pp.253-273, 2004.

N. A. Ebraheim, V. Patil, J. Liu, S. P. Haman, and R. A. Yeasting, Morphometric analyses of the cervical superior facets and implications for facet dislocation, International Orthopaedics, vol.32, issue.1, pp.97-101, 2008.

M. D. Ercoli, F. J. Prevosti, and A. Alvarez, Form and function within a phylogenetic framework: locomotory habits of extant predators and some Miocene Sparassodonta (Metatheria), Zoological Journal of the Linnean Society, vol.165, issue.1, pp.224-251, 2012.

R. Ericsson, R. Knight, and Z. Johanson, Evolution and development of the vertebrate neck, Journal of Anatomy, vol.222, issue.1, pp.67-78, 2013.

Y. Escoufier, Le traitement des variables vectorielles, Biometrics, vol.29, issue.4, pp.751-760, 1973.

J. S. Farris, Methods for computing Wagner trees, Systematic Biology, vol.19, issue.1, pp.83-92, 1970.

B. Favier and P. Dolle, Developmental functions of mammalian Hox genes, Molecular Human Reproduction, vol.3, pp.115-131, 1997.

V. Feipel, T. De-mesmaeker, P. Klein, and M. Rooze, Three-dimensional kinematics of the lumbar spine during treadmill walking at different speeds, European Spine Journal, vol.10, pp.16-22, 2001.

J. Felsenstein, Phylogenies from molecular sequences: inference and reliability, 1988.

, Annual Review of Genetics, vol.22, issue.1, pp.521-565

R. Ferring, O. Oms, J. Agustí, F. Berna, M. Nioradze et al., Earliest human occupations at Dmanisi (Georgian Caucasus) dated to 1.85-1.78 Ma, vol.108, pp.10432-10436, 2011.

J. G. Fleagle, C. C. Gilbert, and A. L. Baden, Primate cranial diversity, American Journal of Physical Anthropology, vol.142, issue.4, pp.565-578, 2010.

A. Fleming, M. G. Kishida, C. B. Kimmel, and R. J. Keynes, Building the backbone: the development and evolution of vertebral patterning, Development, vol.142, pp.1733-1744, 2015.

J. W. Fondon and H. R. Garner, Molecular origins of rapid and continuous morphological evolution, Proceedings of the National Academy of Sciences of the United States of America, vol.101, pp.18058-18063, 2004.

J. Fraipont and M. Lohest, La race humaine de Néanderthal ou de Canstadt en Belgique, Archives de Biologie, vol.7, pp.587-757, 1887.

C. Fromental-ramain, X. Warot, N. Messadecq, M. Lemeur, P. Dollé et al., Hoxa-13 and Hoxd-13 play a crucial role in the patterning of the limb autopod, vol.122, pp.2997-3011, 1996.

H. M. Frost, An Introduction to Biomechanics, 1967.

Q. Fu, M. Hajdinjak, O. T. Moldovan, S. Constantin, S. Mallick et al., An early modern human from Romania with a recent Neanderthal ancestor, Nature, vol.524, p.216, 2015.

C. Fuhlrott, Menschliche Ueberreste aus einen Felsengrotte des Dusselthals. Verhandlungen des naturhistorischen Vereines, vol.60, pp.131-153, 1859.

H. F. Gadow, The Evolution of the Vertebral Column. A Contribution to the Study of Vertebrate Phylogeny, 1933.

F. Galis, Why do almost all mammals have seven cervical vertebrae? Developmental constraints, Hox genes, and cancer, Journal of Experimental Zoology, vol.285, pp.19-26, 1999.

F. Galis, On the homology of structures and Hox genes: the vertebral column, Referencias, vol.305, 1999.

, Homology, pp.80-94

F. Galis and J. A. Metz, Evolutionary novelties: the making and breaking of pleiotropic constraints, Integrative and Comparative Biology, vol.47, pp.409-419, 2007.

F. Galis, D. R. Carrier, J. Van-alphen, S. D. Van-der-mije, T. J. Van-dooren et al., Fast running restricts evolutionary change of the vertebral column in mammals, vol.111, pp.11401-11406, 2014.

D. García-martínez, N. Torres-tamayo, I. Torres-sánchez, F. García-río, A. Rosas et al., Ribcage measurements indicate greater lung capacity in Neanderthals and Lower Pleistocene hominins compared to modern humans, Communications biology, vol.1, issue.1, p.117, 2018.

D. García-martínez, S. Nalla, M. T. Ferreira, R. A. Guichón, M. D. Campo et al., Eco-geographic adaptations in the human ribcage throughout a 3D geometric morphometric approach, American Journal of Physical Anthropology, vol.166, issue.2, pp.323-336, 2018.

S. J. Gaunt, Conservation in the Hox code during morphological evolution, International Journal of Developmental Biology, vol.38, pp.549-552, 2002.

D. L. Gebo, Climbing, brachiation, and terrestrial quadrupedalism: historical precursors of hominid bipedalism, American Journal of Physical Anthropology, vol.101, issue.1, pp.55-92, 1996.

H. Gee, Across the bridge: understanding the origin of the vertebrates, 2018.

S. Gerber, On the relationship between the macroevolutionary trajectories of morphological integration and morphological disparity, PLoS One, vol.8, p.63913, 2013.

C. A. Giglio and J. B. Volpon, Development and evaluation of thoracic kyphosis and lumbar lordosis during growth, Journal of Children's Orthopaedics, vol.1, p.187, 2007.

S. F. Gilbert, Opening Darwin's black box: teaching evolution through developmental genetics, Nature Reviews Genetics, vol.4, issue.9, p.735, 2003.

C. Gomez, E. M. Özbudak, J. Wunderlich, D. Baumann, J. Lewis et al., , 2008.

, Control of segment number in vertebrate embryos, Nature, p.335

A. Gómez-olivencia, Estudios paleobiológicos sobre la columna vertebral y la caja torácica de los humanos fósiles del Pleistoceno, con especial referencia a los fósiles de la Sierra de Atapuerca. Doctoral dissertation (unpublished), 2009.

A. Gómez-olivencia, Back to the old man's back: Reassessment of the anatomical determination of the vertebrae of the Neandertal individual of, Annales de Paléontologie, vol.99, issue.1, pp.43-65, 2013.

A. Gómez-olivencia, The presacral spine of the La Ferrassie 1 Neandertal: a revised inventory, vol.25, pp.19-38, 2013.

A. Gómez-olivencia, J. M. Carretero, J. L. Arsuaga, L. Rodríguez-garcía, R. García-gonzález et al., Metric and morphological study of the upper cervical spine from the Sima de los Huesos site, Journal of Human Evolution, vol.53, issue.1, pp.6-25, 2007.

A. Gómez-olivencia, J. L. Arsuaga, J. M. Carretero, A. Gracia, and I. Martínez, , 2011.

, A complete neck from Sima de los Huesos and the evolution of the cervical spine in Neandertal lineage, Proceedings of the First annual meeting, European Society for the study of Human Evolution, p.41

A. Gómez-olivencia, E. Been, J. L. Arsuaga, and J. T. Stock, The Neandertal vertebral column 1: The cervical spine, Journal of Human Evolution, vol.64, issue.6, pp.608-630, 2013.

A. Gómez-olivencia, C. Couture-veschambre, S. Madelaine, and B. Maureille, , 2013.

, The vertebral column of the Regourdou 1 Neandertal, Journal of Human Evolution, vol.64, issue.6, pp.582-607

A. Gómez-olivencia and J. L. Arsuaga, The vertebral column and thorax in the Middle Pleistocene: the case of the Sima de los Huesos, American Journal of Physical Anthropology, vol.156, issue.S60, p.155, 2015.

A. Gómez-olivencia, M. Arlegi, A. Barash, J. T. Stock, and E. Been, The Neandertal vertebral column 2: The lumbar spine, Journal of Human Evolution, vol.106, pp.84-101, 2017.

A. Gómez-olivencia, A. Barash, D. García-martínez, M. Arlegi, P. Kramer et al., , 2018.

A. Gómez-olivencia and E. Been, The Spine of Late Homo, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.185-211, 2019.

A. Gómez-robles and P. D. Polly, Morphological integration in the hominin dentition: evolutionary, developmental, and functional factors, Evolution, vol.66, pp.1024-1043, 2012.

D. Gommery, Les atlas et les axis des hominides du Plio-Pleistocene: morphologie et systematique. Comptes Rendus de l'Académie des, Sciences-Series IIA-Earth and Planetary Science, vol.325, issue.8, pp.639-642, 1997.

D. Gommery, Axe vertébral, Hominoidea fossiles et posture orthograde: préambule à la bipédie, Primatologie, vol.1, pp.135-160, 1998.

R. Gomulkiewicz and D. Houle, Demographic and genetic constraints on evolution, The American Naturalist, vol.174, issue.6, pp.218-229, 2009.

C. Goodall, Procrustes methods in the statistical analysis of shape, Journal of the Royal Statistical Society: Series B (Methodological), vol.53, issue.2, pp.285-321, 1991.

D. Gorjanovi?-kramberger, Der diluviale Mensch von Krapina in Kroatien: ein Beitrag zur Paläoanthropologie, 1906.

A. Goswami, Cranial modularity shifts during mammalian evolution, The American Naturalist, vol.168, pp.270-280, 2006.

A. Goswami and P. D. Polly, The influence of modularity on cranial morphological disparity in Carnivora and Primates (Mammalia), PloS One, vol.5, p.9517, 2010.

A. Goswami, J. B. Smaers, C. Soligo, and P. D. Polly, The macroevolutionary consequences of phenotypic integration: from development to deep time, Philosophical Transactions Royal Society B, vol.369, 2014.

A. Goswami, W. J. Binder, J. Meachen, and F. R. Keefe, The fossil record of phenotypic integration and modularity: A deep-time perspective on developmental and evolutionary dynamics, Proceedings of the National Academy of Sciences of the United States of America, vol.112, issue.16, pp.4891-4896, 2015.

M. W. Grabowski, J. D. Polk, and C. C. Roseman, Divergent patterns of integration and reduced constraint in the human hip and the origins of bipedalism, Evolution: Referencias 308 International Journal of Organic Evolution, vol.65, pp.1336-1356, 2011.

M. Grabowski and C. C. Roseman, Complex and changing patterns of natural selection explain the evolution of the human hip, Journal of Human Evolution, vol.85, pp.94-110, 2015.

M. Grabowski and A. Porto, How many more? Sample size determination in studies of morphological integration and evolvability, Methods in Ecology and Evolution, vol.8, pp.592-603, 2017.

D. E. Granger, R. J. Gibbon, K. Kuman, R. J. Clarke, L. Bruxelles et al., New cosmogenic burial ages for Sterkfontein Member 2 Australopithecus and Member 5 oldowan, Nature, vol.522, p.85, 2015.
URL : https://hal.archives-ouvertes.fr/hal-02017442

H. Gray, Anatomy of the human body, CM Goss. Lea and Febiger, 1973.

H. Gray, Gray's anatomy: with original illustrations by Henry Carter, 2009.

P. Gunz and K. Harvati, The Neanderthal "chignon": variation, integration, and homology, Journal of Human Evolution, vol.52, issue.3, pp.262-274, 2007.

P. Gunz, S. Neubauer, B. Maureille, and J. J. Hublin, Brain development after birth differs between Neanderthals and modern humans, Current Biology, vol.20, issue.21, pp.921-922, 2010.

P. Gunz, S. Neubauer, L. Golovanova, V. Doronichev, B. Maureille et al., A uniquely modern human pattern of endocranial development. Insights from a new cranial reconstruction of the Neandertal newborn from Mezmaiskaya, Journal of Human Evolution, vol.62, issue.2, pp.300-313, 2012.
URL : https://hal.archives-ouvertes.fr/hal-01842827

M. Haeusler, S. A. Martelli, and T. Boeni, Vertebrae numbers of the early hominid lumbar spine, Journal of Human Evolution, vol.43, issue.5, pp.621-643, 2002.

M. Haeusler, R. Schiess, and T. Boeni, New vertebral and rib material point to modern bauplan of the Nariokotome Homo erectus skeleton, Journal of Human Evolution, vol.61, issue.5, pp.575-582, 2011.

Y. Haile-selassie, B. M. Latimer, M. Alene, A. L. Deino, L. Gibert et al.,

C. O. Lovejoy, An early Australopithecus afarensis postcranium from Woranso-Mille, Ethiopia, Proceedings of the National Academy of Sciences of the United States of America, vol.107, pp.12121-12126, 2010.

Y. Haile-selassie, B. Latimer, C. O. Lovejoy, S. M. Melillo, and M. R. Meyer, Conclusion: Implications of KSD-VP-1/1 for early hominin paleobiology and insights into the chimpanzee/human last common ancestor, The Postcranial Anatomy of Australopithecus afarensis, pp.179-187, 2016.

B. Hallgrímsson, K. Willmore, and B. K. Hall, Canalization, developmental stability, and morphological integration in primate limbs, American Journal of Physical Anthropology, vol.119, pp.131-158, 2002.

B. Hallgrímsson, D. E. Lieberman, W. Liu, A. F. Ford-hutchinson, and F. R. Jirik, , 2007.

, Epigenetic interactions and the structure of phenotypic variation in the cranium

, Evolution & Development, vol.9, issue.1, pp.76-91

B. Hallgrímsson, H. Jamniczky, N. M. Young, C. Rolian, T. E. Parsons et al., Deciphering the palimpsest: studying the relationship between morphological integration and phenotypic covariation, Evolutionary Biology, vol.36, issue.4, pp.355-376, 2009.

Ø. Hammer, D. A. Harper, and P. D. Ryan, PAST: paleontological statistics software package for education and data analysis, Palaeontologia Electronica, vol.4, issue.1, p.9, 2001.

P. Hanot, A. Herrel, C. Guintard, and R. Cornette, The impact of artificial selection on morphological integration in the appendicular skeleton of domestic horses, Journal of Anatomy, vol.232, issue.4, pp.657-673, 2018.
URL : https://hal.archives-ouvertes.fr/hal-02105766

T. F. Hansen, Is modularity necessary for evolvability, Remarks on the relationship between pleiotropy and evolvability, vol.69, pp.83-94, 2003.

T. F. Hansen and D. Houle, Measuring and comparing evolvability and constraint in multivariate characters, Journal of Evolutionary Biology, vol.21, pp.1201-1219, 2008.

S. E. Hartman, Evaluation of some alternative procedures used in numerical systematics, Systematic Zoology, vol.37, pp.1-18, 1988.

L. Hautier, V. Weisbecker, M. R. Sánchez-villagra, A. Goswami, and R. J. Asher, , 2010.

, Skeletal development in sloths and the evolution of mammalian vertebral patterning, Proceedings of the National Academy of Sciences of the United States of America, vol.107, pp.18903-18908

J. Hawks, M. Elliott, P. Schmid, S. E. Churchill, D. J. De-ruiter et al.,

E. M. Feuerriegel, New fossil remains of Homo naledi from the Lesedi Chamber, South Africa. ELife, vol.6, p.24232, 2017.

S. Hayama, M. Nakatsukasa, and Y. Kunimatsu, Monkey performance: the development of bipedalism in trained Japanese monkeys, Kaibogaku zasshi. Journal of Anatomy, vol.67, pp.169-185, 1992.

J. J. Head and P. D. Polly, Evolution of the snake body form reveals homoplasy in amniote Hox gene function, Nature, vol.520, pp.86-89, 2015.

J. L. Heim, Les hommes fossiles de La Ferrassie: Tome I. Le gisement. Les squelettes adultes (crâne et squelette du tronc), 1976.

W. G. Hill and R. Thompson, Probabilities of non-positive definite between-group or genetic covariance matrices, Biometrics, vol.34, pp.429-439, 1978.

L. J. Hlusko, Identifying metameric variation in extant hominoid and fossil hominid mandibular molars, American Journal of Physical Anthropology, vol.118, pp.86-97, 2002.

L. J. Hlusko, Integrating the genotype and phenotype in hominid paleontology, Proceedings of the National Academy of Sciences of the United States of America, vol.101, pp.2653-2657, 2004.

T. W. Holliday, Body size, body shape, and the circumscription of the genus Homo, Current Anthropology, vol.53, pp.330-345, 2012.

E. Holt, P. Dirks, C. Placzek, and L. Berger, The stable isotope setting of Australopithecus sediba at Malapa, South Africa, South African Journal of Science, vol.112, issue.7-8, pp.1-9, 2016.

J. J. Hublin, A. Ben-ncer, S. E. Bailey, S. E. Freidline, S. Neubauer et al., New fossils from Jebel Irhoud, Morocco and the pan-African origin of Homo sapiens, Nature, vol.546, issue.7657, p.289, 2017.

R. B. Huey and A. F. Bennett, Phylogenetic studies of coadaptation: preferred temperatures versus optimal performance temperatures of lizards, Evolution, issue.5, pp.1098-1115, 1987.

K. D. Hunt, Positional behavior in the Hominoidea, International Journal of Primatology, vol.12, pp.95-118, 1991.

J. R. Hutchinson, On the inference of function from structure using biomechanical modelling and simulation of extinct organisms, Biology Letters, vol.8, issue.1, pp.115-118, 2011.

H. Ishida, Y. Kunimatsu, T. Takano, Y. Nakano, and M. Nakatsukasa, , 2004.

F. A. Jenkins, The postcranial skeleton of African cynodonts, Bulletin of the Peabody Museum of Natural History, vol.36, pp.1-216, 1971.

D. C. Johanson, M. Taieb, and Y. Coppens, Pliocene hominids from the Hadar Formation, stratigraphic, chronologic, and paleoenvironmental contexts, with notes on hominid morphology and systematics, 1973.

, American Journal of Physical Anthropology, vol.57, issue.4, pp.373-402

D. C. Johanson, F. T. Masao, G. G. Eck, T. D. White, R. C. Walter et al., New partial skeleton of Homo habilis from Olduvai Gorge, Nature, vol.327, issue.6119, pp.205-209, 1987.

K. E. Jones, K. D. Angielczyk, P. D. Polly, J. J. Head, V. Fernandez et al., Fossils reveal the complex evolutionary history of the mammalian regionalized spine, Science, vol.361, pp.1249-1252, 2018.

K. E. Jones, L. Benitez, K. D. Angielczyk, and S. E. Pierce, Adaptation and constraint in the evolution of the mammalian backbone, BMC Evolutionary Biology, vol.18, issue.1, p.172, 2018.

W. L. Jungers, A. B. Falsetti, and C. E. Wall, Shape, relative size, and sizeadjustments in morphometrics, American Journal of Physical Anthropology, Referencias, p.313, 1995.

C. P. Klingenberg, MorphoJ: an integrated software package for geometric morphometrics, Molecular Ecology Resources, vol.11, issue.2, pp.353-357, 2011.

C. P. Klingenberg, Cranial integration and modularity: insights into evolution and development from morphometric data, Hystrix, the Italian Journal of Mammalogy, vol.24, issue.1, pp.43-58, 2013.

C. Klingenberg, Analyzing fluctuating asymmetry with geometric morphometrics: concepts, methods, and applications, Symmetry, vol.7, issue.2, pp.843-934, 2015.

C. P. Klingenberg and S. D. Zaklan, Morphological integration between developmental compartments in the Drosophila wing, Evolution, vol.54, issue.4, pp.1273-1285, 2000.

C. P. Klingenberg, M. Barluenga, and A. Meyer, Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry, Evolution, vol.56, issue.10, pp.1909-1920, 2002.

C. P. Klingenberg and N. A. Gidaszewski, Testing and quantifying phylogenetic signals and homoplasy in morphometric data, Systematic Biology, vol.59, issue.3, pp.245-261, 2010.

C. P. Klingenberg and J. Marugán-lobón, Evolutionary covariation in geometric morphometric data: analyzing integration, modularity, and allometry in a phylogenetic context, Systematic Biology, vol.62, issue.4, pp.591-610, 2013.

R. Krumlauf, Evolution of the vertebrate Hox homeobox genes, Bioessays, vol.14, issue.4, pp.245-252, 1992.

B. Kurtén, On the variation and population dynamics of fossil and recent mammal populations, Acta Zoologica Fennica, vol.76, pp.1-122, 1953.

R. Lande, Quantitative genetic analysis of multivariate evolution, applied to brain: body size allometry, Evolution, vol.33, pp.402-416, 1979.

O. Larouche, M. L. Zelditch, and R. Cloutier, Modularity promotes morphological divergence in ray-finned fishes, Scientific Reports, vol.8, issue.1, p.7278, 2018.

B. Latimer and C. V. Ward, The thoracic and lumbar vertebrae, The Nariokotome Homo erectus Skeleton, pp.266-293, 1993.

L. Clark, W. E. Penman, and G. G. , The projection of the retina in the lateral geniculate body, Proceedings of the Royal Society of London. Series B, vol.114, 1934.

L. S. Leakey, P. V. Tobias, and J. R. Napier, A new species of the genus Homo from Olduvai Gorge, 1964.

R. E. Leakey and A. C. Walker, Further hominids from the Plio-Pleistocene of, 1985.

K. Fora and K. , American Journal of Physical Anthropology, vol.67, issue.2, pp.135-163

C. A. Lemen, The effectiveness of methods of shape analysis, Fieldiana Zoology, vol.13, p.17, 1983.

K. L. Lewton, Evolvability of the primate pelvic girdle, Evolutionary Biology, vol.39, issue.1, pp.126-139, 2012.

D. Lieberman, The Evolution of the Human Head, 2011.

D. E. Lieberman, C. F. Ross, and M. J. Ravosa, The primate cranial base: ontogeny, function, and integration, American Journal of Physical Anthropology, vol.113, pp.117-169, 2000.

C. López-aguirre, S. J. Hand, D. Koyabu, N. T. Son, and L. A. Wilson, Postcranial heterochrony, modularity, integration and disparity in the prenatal ossification in bats (Chiroptera), BMC Evolutionary Biology, vol.19, issue.1, p.75, 2019.

C. O. Lovejoy, The natural history of human gait and posture: Part 2. Hip and thigh. Gait and Posture, vol.21, pp.113-124, 2005.

C. O. Lovejoy, D. C. Johanson, and Y. Coppens, Elements of the axial skeleton recovered from the Hadar Formation: 1974-1977 collections, American Journal of Physical Anthropology, vol.57, issue.4, pp.631-635, 1982.

C. O. Lovejoy and M. A. Mccollum, Spinopelvic pathways to bipedality: why no hominids ever relied on a bent-hip-bent-knee gait, Philosophical Transactions of the Royal Society B: Biological Sciences, vol.365, pp.3289-3299, 2010.

Z. Lu, D. J. Meldrum, Y. Huang, J. He, and E. E. Sarmiento, The Jinniushan hominin pedal skeleton from the late Middle Pleistocene of China, Homo, vol.62, issue.6, pp.389-401, 2011.

G. J. Maat, B. Matricali, and E. L. Van-meerten, Postnatal development and structure of the neurocentral junction: its relevance for spinal surgery, Spine, vol.21, issue.6, pp.661-666, 1996.

W. P. Maddison, , 1991.

E. Manfreda, P. Mitteroecker, F. L. Bookstein, and K. Schaefer, Functional morphology of the first cervical vertebra in humans and nonhuman primates, The Anatomical Record, vol.289, pp.184-194, 2006.

J. H. Mansfield and A. Abzhanov, Hox expression in the American alligator and evolution of archosaurian axial patterning, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.314, pp.629-644, 2010.

J. D. Marcot and D. W. Mcshea, Increasing hierarchical complexity throughout the history of life: phylogenetic tests of trend mechanisms, Paleobiology, vol.33, issue.2, pp.182-200, 2007.

K. V. Mardia, J. T. Kent, and J. M. Bibby, Multivariate Analysis, 1979.

K. V. Mardia, F. L. Bookstein, and I. J. Moreton, Statistical assessment of bilateral symmetry of shapes, Biometrika, pp.285-300, 2000.

G. Marroig and J. M. Cheverud, A comparison of phenotypic variation and covariation pat terns and the role of phylogeny, ecology, and ontogeny during cranial evolution of New World monkeys, Evolution, vol.55, pp.2576-2600, 2001.

G. Marroig and J. M. Cheverud, Cranial evolution in sakis (Pithecia, Platyrrhini) I: interspecific differentiation and allometric patterns, American Journal of Physical Anthropology: The Official Publication of the American Association of Physical Anthropologists, vol.125, pp.266-278, 2004.

G. Marroig, L. T. Shirai, A. Porto, F. B. De-oliveira, and V. De-conto, The evolution of modularity in the mammalian skull II: evolutionary consequences, Evolutionary Biology, vol.36, pp.136-148, 2009.

S. A. Martelli, The Modern and Fossil Hominoid Spinal Ontogeny, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.247-281, 2019.

H. Martin, L'homme fossile de la Quina, Librairie Octave Doin, 1923.

I. Martínez, La Base del Cráneo y el Hueso Temporal en la Evolución de los Homínidos, con Especial Referencia a los Fósiles de Atapuerca (Burgos). Doctoral Referencias 316 dissertation (Unpublished), 1995.

E. P. Martins, Estimation of ancestral states of continuous characters: a computer simulation study, Systematic Biology, vol.48, issue.3, pp.642-650, 1999.

J. G. Maisey, Heads and tails: a chordate phylogeny, Cladistics, vol.2, issue.4, pp.201-256, 1986.

E. H. Margulies, S. L. Kardia, and J. W. Innis, A comparative molecular analysis of developing mouse forelimbs and hindlimbs using serial analysis of gene expression, SAGE) Genome Research, vol.11, issue.10, pp.1686-1698, 2001.

R. C. Mccarthy, Anthropoid cranial base architecture and scaling relationships, Journal of Human Evolution, vol.40, issue.1, pp.41-66, 2001.

M. A. Mccollum, B. A. Rosenman, G. Suwa, R. S. Meindl, and C. O. Lovejoy, , 2010.

, The vertebral formula of the last common ancestor of African apes and humans, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.314, issue.2, pp.123-134

I. Mcdougall, F. H. Brown, P. M. Vasconcelos, B. E. Cohen, D. S. Thiede et al., New single crystal 40Ar/39Ar ages improve time scale for deposition of the Omo Group, Journal of the Geological Society, vol.169, issue.2, pp.213-226, 2012.

G. R. Mcghee, Theoretical morphology: the concept and its applications, 1999.

D. Melo, G. Garcia, A. Hubbe, A. P. Assis, and G. Marroig, EvolQG-An R package for evolutionary quantitative genetics, vol.4, pp.1-25, 1000.

M. Meyer, Functional biology of the Homo erectus axial skeleton from Dmanisi, Georgia Doctoral dissertation (Unpublished), 2005.

M. R. Meyer, The cervical vertebrae of KSD-VP-1/1, 2016.

, The Postcranial Anatomy of Australopithecus afarensis: New Insights from KSD-VP-1, pp.63-111

M. R. Meyer and M. Haeusler, Spinal cord evolution in early Homo, Journal of Human Evolution, vol.88, pp.43-53, 2015.

M. R. Meyer, S. A. Williams, P. Schmid, S. E. Churchill, and L. R. Berger, The cervical spine of Australopithecus sediba, Journal of Human Evolution, vol.104, pp.32-49, 2017.

M. R. Meyer and S. A. Williams, The Spine of Early Pleistocene Homo, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.153-183, 2019.

C. Springer,

R. A. Mittermeier, D. E. Wilson, and . Rylands, Handbook of the Mammals of the World: Primates. Lynx Edicions, 2013.

P. Mitteroecker, P. Gunz, M. Bernhard, K. Schaefer, and F. L. Bookstein, , 2004.

, Comparison of cranial ontogenetic trajectories among great apes and humans, Journal of Human Evolution, vol.46, issue.6, pp.679-698

P. Mitteroecker and F. Bookstein, The conceptual and statistical relationship between modularity and morphological integration, Systematic Biology, vol.56, pp.818-836, 2007.

P. Mitteroecker, E. Manfreda, F. L. Bookstein, and K. Schaefer, Does the morphology of the human atlas and axis reflect bipedality? A multivariate approach to functional morphology, American Journal of Physical Anthropology, vol.132, issue.S44, pp.172-173, 2007.

A. M. Moigne, M. R. Palombo, V. Belda, D. Heriech-briki, S. Kacimi et al., Les faunes de grands mammifères de la Caune de l'Arago (Tautavel), 2006.

. L'anthropologie, , vol.110, pp.788-831

L. R. Monteiro, Multivariate regression models and geometric morphometrics: the search for causal factors in the analysis of shape, Systematic Biology, vol.48, pp.192-199, 1999.

K. L. Moore and T. V. Persaud, Embriologia clínica, 2008.

M. J. Morwood, P. Brown, T. Sutikna, E. W. Saptomo, K. E. Westaway et al., Further evidence for small-bodied hominins from the Late Pleistocene of Flores, Indonesia, Nature, vol.437, issue.7061, pp.1012-1017, 2005.

J. E. Mosimann, Size allometry: size and shape variables with characterizations of the lognormal and generalized gamma distributions, Journal of the American Statistical Association, vol.65, pp.930-945, 1970.

S. Moyà-solà, M. Köhler, D. M. Alba, I. Casanovas-vilar, and J. Galindo, , 2004.

, Pierolapithecus catalaunicus, a new Middle Miocene great ape from Spain, Science, vol.306, issue.5700, pp.1339-1344

J. Müller, T. M. Scheyer, J. J. Head, P. M. Barrett, I. Werneburg et al., Homeotic effects, somitogenesis and the evolution of vertebral numbers in recent and fossil amniotes, vol.107, pp.2118-2123, 2010.

M. Nakatsukasa, Comparative study of Moroto vertebral specimens, Journal of Human Evolution, vol.55, issue.4, pp.581-588, 2008.

M. Nakatsukasa, Miocene ape spinal morphology: the evolution of orthogrady, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.73-96, 2019.

M. Nakatsukasa, C. V. Ward, A. Walker, M. F. Teaford, Y. Kunimatsu et al., Tail loss in Proconsul heseloni, Journal of Human Evolution, vol.46, issue.6, pp.777-784, 2004.

M. Nakatsukasa, Y. Kunimatsu, Y. Nakano, and H. Ishida, Vertebral morphology of Nacholapithecus kerioi based on KNM-BG 35250, Journal of Human Evolution, vol.52, issue.4, pp.347-369, 2007.

M. Nakatsukasa and Y. Kunimatsu, Nacholapithecus and its importance for understanding hominoid evolution, Evolutionary Anthropology: Issues, News, and Reviews: Issues, News, and Reviews, vol.18, issue.3, pp.103-119, 2009.

M. Nakatsukasa, S. Almécija, and D. R. Begun, The hands of Miocene hominoids, The Evolution of the Primate Hand, pp.485-514, 2016.

T. K. Nalley and N. Grider-potter, Functional morphology of the primate head and neck, American Journal of Physical Anthropology, vol.156, pp.531-542, 2015.

T. K. Nalley and N. Grider-potter, Functional analyses of the primate upper cervical vertebral column, Journal of Human Evolution, vol.107, pp.19-35, 2017.

T. K. Nalley and N. Grider-potter, Vertebral Morphology in Relation to Head Posture and Locomotion I: The Cervical Spine, 2019.

, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, p.319

Y. Narita and S. Kuratani, Evolution of the vertebral formulae in mammals: a perspective on developmental constraints, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.304, pp.91-106, 2005.

C. Naruse, S. Shibata, M. Tamura, T. Kawaguchi, K. Abe et al., New insights into the role of Jmjd3 and Utx in axial skeletal formation in mice, The FASEB Journal, issue.6, pp.2252-2266, 2017.

L. Nevell and B. Wood, Cranial base evolution within the hominin clade, Journal of Anatomy, vol.212, issue.4, pp.455-468, 2008.

P. O'higgins, The study of morphological variation in the hominid fossil record: biology, landmarks and geometry, The Journal of Anatomy, vol.197, issue.1, pp.103-120, 2000.

J. C. Ohman, The first rib of hominoids, American Journal of Physical Anthropology, vol.70, issue.2, pp.209-229, 1986.

E. C. Olson and R. L. Miller, Morphological integration. Chicago, 1958.

A. Pablos, I. Martínez, C. Lorenzo, A. Gracia, N. Sala et al., , 2013.

, Human talus bones from the Middle Pleistocene site of Sima de los Huesos, Journal of Human Evolution, vol.65, issue.1, pp.79-92

G. P. Pal, Weight transmission through the sacrum in man, Journal of Anatomy, vol.162, pp.9-17, 1989.

G. P. Pal and R. V. Routal, A study of weight transmission through the cervical and upper thoracic regions of the vertebral column in man, Journal of Anatomy, vol.148, pp.245-261, 1986.

G. P. Pal and R. V. Routal, Transmission of weight through the lower thoracic and lumbar regions of the vertebral column in man, Journal of Anatomy, vol.152, pp.93-105, 1987.

G. P. Pal, R. V. Routal, and S. K. Saggu, The orientation of the articular facets of the zygapophyseal joints at the cervical and upper thoracic region, The Journal of Anatomy, vol.198, issue.4, pp.431-441, 2001.

A. R. Palmer and C. Strobeck, Fluctuating asymmetry: measurement, analysis, patterns.Annual review of, Ecology and Systematics, vol.17, issue.1, pp.391-421, 1986.

J. M. Parés and A. Pérez-gonzález, Paleomagnetic age for hominid fossils at Atapuerca archaeological site, Spain. Science, vol.269, issue.5225, pp.830-832, 1995.

M. Pavlicev, J. M. Cheverud, and G. P. Wagner, Measuring morphological integration using eigenvalue variance, Evolutionary Biology, vol.36, pp.157-170, 2009.

A. Peacock, Observations on the pre-natal development of the intervertebral disc in man, Journal of Anatomy, vol.85, p.260, 1951.

A. Peacock, Observations on the postnatal structure of the intervertebral disc in man, Journal of Anatomy, vol.86, p.162, 1952.

R. Pickering, P. H. Dirks, Z. Jinnah, D. J. De-ruiter, S. E. Churchill et al., Australopithecus sediba at 1.977 Ma and implications for the origins of the genus Homo, Science, vol.333, issue.6048, pp.1421-1423, 2011.

S. E. Pierce, J. A. Clack, and J. R. Hutchinson, Comparative axial morphology in pinnipeds and its correlation with aquatic locomotory behaviour, Journal of Anatomy, vol.219, pp.502-514, 2011.

M. Pigliucci, Phenotypic integration: studying the ecology and evolution of complex phenotypes, Ecology Letters, vol.6, issue.3, pp.265-272, 2003.

D. Pilbeam, The anthropoid postcranial axial skeleton: comments on development, variation, and evolution, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.302, pp.241-267, 2004.

D. R. Pilbeam and D. E. Lieberman, Reconstructing the last common ancestor of chimpanzees and humans, Chimpanzees and Human Evolution, pp.22-141, 2017.

J. Piveteau, La grotte de Regourdou (Dordogne), Annales de Paléontologie, vol.49, pp.163-194, 1963.

J. M. Polanski, Morphological integration of the modern human mandible during ontogeny, International Journal of Evolutionary Biology, 2011.

J. M. Polanski and R. G. Franciscus, Patterns of craniofacial integration in extant Homo, Pan, and Gorilla, American Journal of Physical Anthropology, vol.131, pp.38-49, 2006.

M. L. Ponssa, Cladistic analysis and osteological descriptions of the frog species in the Leptodactylus fuscus species group (Anura, Leptodactylidae), Journal of Zoological Systematics and Evolutionary Research, vol.46, issue.3, pp.249-266, 2008.

A. Porto, F. B. De-oliveira, L. T. Shirai, V. De-conto, and G. Marroig, The evolution of modularity in the mammalian skull I: morphological integration patterns and magnitudes, Evolutionary Biology, vol.36, pp.118-135, 2009.

A. Porto, L. T. Shirai, F. B. De-oliveira, and G. Marroig, Size variation, growth strategies, and the evolution of modularity in the mammalian skull, Evolution, vol.67, pp.3305-3322, 2013.

H. Preuschoft, Mechanisms for the acquisition of habitual bipedality: are there biomechanical reasons for the acquisition of upright bipedal posture?, Journal of Anatomy, vol.204, issue.5, pp.363-384, 2004.

H. Preuschoft and N. Klein, Torsion and bending in the neck and tail of sauropod dinosaurs and the function of cervical ribs: insights from functional morphology and biomechanics, PloS one, vol.8, issue.10, p.78574, 2013.

. R-core-team, R: a language and environment for statistical computing, 2018.

Y. Rak, A. Ginzburg, and E. Geffen, Gorilla-like anatomy on Australopithecus afarensis mandibles suggests Au. afarensis link to robust australopiths, Proceedings of the National Academy of Sciences of the United States of America, vol.104, pp.6568-6572, 2007.

M. Randau and A. Goswami, Unravelling intravertebral integration, modularity and disparity in Felidae (Mammalia), Evolution & Development, vol.19, pp.85-95, 2017.

M. Randau and A. Goswami, Morphological modularity in the vertebral column of Felidae (Mammalia, Carnivora), BMC Evolutionary Biology, vol.17, p.133, 2017.

M. Randau and A. Goswami, Shape Covariation (or the Lack Thereof) Between Vertebrae and Other Skeletal Traits in Felids: The Whole is Not Always Greater than the Sum of Parts, Evolutionary Biology, vol.45, pp.196-210, 2018.

J. P. Raynal, Essai de datation directe, et la préhistoire en Corrèze. Association pour la Recherche Archéologique en Limousin, 1990.

J. D. Reist, An empirical evaluation of several univariate methods that adjust for size variation in morphometric data, Canadian Journal of Zoology, vol.63, pp.1429-1439, 1985.

M. K. Richardson, S. P. Allen, G. M. Wright, A. Raynaud, and J. Hanken, Somite Referencias 322 number and vertebrate evolution, Development, vol.125, issue.2, pp.151-160, 1998.

R. Riedl and D. Auer, Die Ordnung des Lebendigen: Systembedingungen d. Evolution, 1975.

L. Rios and H. F. Cardoso, Age estimation from stages of union of the vertebral epiphyses of the ribs, American Journal of Physical Anthropology, vol.140, issue.2, pp.265-274, 2009.

L. Ríos, A. Rosas, A. Estalrrich, A. García-tabernero, M. Bastir et al., Possible further evidence of low genetic diversity in the El Sidrón, Neandertal group: Congenital clefts of the atlas, vol.10, p.136550, 2015.

L. Ríos, T. L. Kivell, C. Lalueza-fox, A. Estalrrich, A. García-tabernero et al., Skeletal Anomalies in the Neandertal Family of el Sidrón (Spain) support A Role of Inbreeding in Neandertal Extinction, Scientific Reports, vol.9, issue.1, p.1697, 2019.

L. Ríos, C. Palancar, F. Pastor, S. Llidó, J. A. Sanchís-gimeno et al., , 2017.

, Shape change in the atlas with congenital midline non-union of its posterior arch: a morphometric geometric study, The Spine Journal, vol.17, issue.10, pp.1523-1528

J. T. Robinson, Early hominid posture and locomotion, 1972.

F. J. Rohlf and D. Slice, Extensions of the Procrustes method for the optimal superimposition of landmarks, Systematic Biology, vol.39, pp.40-59, 1990.

F. J. Rohlf and L. F. Marcus, A revolution in morphometrics, Trends in Ecology & Evolution, vol.8, issue.4, pp.129-132, 1993.

F. J. Rohlf and M. Corti, Use of two-block partial least-squares to study covariation in shape, Systematic Biology, vol.49, pp.740-753, 2000.

C. Rolian, Integration and evolvability in primate hands and feet, Evolutionary Biology, vol.36, pp.100-117, 2009.

C. Rolian, Genes, development, and evolvability in primate evolution, Evolutionary Anthropology: Issues, News, and Reviews, vol.23, pp.93-104, 2014.

C. Rolian, D. E. Lieberman, J. Hamill, J. W. Scott, and W. Werbel, Walking, running and the evolution of short toes in humans, Journal of Experimental Biology, vol.323, issue.212, pp.713-721, 2009.

C. Rolian and K. E. Willmore, Morphological integration at 50: patterns and processes of integration in biological anthropology, Evolutionary Biology, vol.36, pp.1-4, 2009.

C. Rolian, D. E. Lieberman, and B. Hallgrímsson, The coevolution of human hands and feet, Evolution, vol.64, pp.1558-1568, 2010.

A. S. Romer, The nature and relationships of the Paleozoic microsaurs, American Journal of Science, vol.248, issue.9, pp.628-654, 1950.

A. S. Romer, Vertebrate Paleontology, 1955.

A. Rosas, C. Martínez-maza, M. Bastir, A. García-tabernero, C. Lalueza-fox et al., Paleobiology and comparative morphology of a late Neandertal sample from El Sidrón, vol.103, pp.19266-19271, 2006.

A. Rosas, A. Estalrrich, A. García-tabernero, M. Bastir, S. Garcia-vargas et al., Les Néandertaliens d'El Sidrón (Asturies, Espagne). Actualisation d'un nouvel échantillon, L'Anthropologie, vol.116, issue.1, pp.57-76, 2012.

A. Rosas, L. Ríos, A. Estalrrich, H. Liversidge, A. García-tabernero et al., The growth pattern of Neandertals, El Sidrón (Spain). Science, issue.6357, pp.1282-1287, 2017.

K. R. Rosenberg, L. Zuné, and C. B. Ruff, Body size, body proportions, and encephalization in a Middle Pleistocene archaic human from northern China, Proceedings of the National Academy of Sciences of the United States of America, vol.103, pp.3552-3556, 2006.

C. F. Ross and M. J. Ravosa, Basicranial flexion, relative brain size, and facial kyphosis in nonhuman primates, American Journal of Physical Anthropology, vol.91, issue.3, pp.305-324, 1993.

E. S. Russell, Form and Function: A Contribution to the History of Animal Morphology Murray, 1916.

G. A. Russo, Prezygapophyseal articular facet shape in the catarrhine thoracolumbar vertebral column, American Journal of Physical Anthropology, vol.142, issue.4, pp.600-612, 2010.

G. A. Russo and E. C. Kirk, Foramen magnum position in bipedal mammals, Journal of Human Evolution, vol.65, issue.5, pp.656-670, 2013.

G. A. Russo and L. J. Shapiro, Reevaluation of the lumbosacral region of Oreopithecus bambolii, Journal of Human Evolution, vol.65, issue.3, pp.253-265, 2013.

G. A. Russo and S. A. Williams, Giant pandas (Carnivora: Ailuropoda melanoleuca) and living hominoids converge on lumbar vertebral adaptations to orthograde trunk posture, Journal of Human Evolution, vol.88, pp.160-179, 2015.

G. A. Russo and E. C. Kirk, Another look at the foramen magnum in bipedal mammals, Journal of Human Evolution, vol.105, pp.24-40, 2017.

A. Russo and K. Kirk, The Hominoid Cranial Base in Relation to Posture and Locomotion, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.15-34, 2019.

M. R. Sánchez-villagra, Developmental palaeontology in synapsids: the fossil record of ontogeny in mammals and their closest relatives, Proceedings of the Royal Society B:Biological Sciences, vol.277, pp.1139-1147, 2010.

W. J. Sanders, Comparative morphometric study of the australopithecine vertebral series Stw-H8/H41, Journal of Human Evolution, vol.34, issue.3, pp.249-302, 1998.

N. Satoh, D. Rokhsar, and T. Nishikawa, Chordate evolution and the three-phylum system, Proceedings of the Royal Society B: Biological Sciences, vol.281, 1794.

B. Z. Saylor, M. Alene, A. Deino, L. Gibert, Y. Haile-selassie et al., The geologic context of Korsi Dora and the partial skeleton, 2016.

, The Postcranial Anatomy of Australopithecus afarensis, pp.13-23

L. Scheuer, S. M. Black, and A. Christie, Developmental Juvenil Osteology, 2000.

D. Schluter, Adaptive radiation along genetic lines of least resistance, Evolution, vol.50, pp.1766-1774, 1996.

A. H. Schultz, Growth and development of the chimpanzee, Control Embryology, vol.28, pp.1-63, 1940.

A. H. Schultz, Growth and development of the orangutan, Control Embryology, vol.29, pp.57-110, 1941.

A. H. Schultz, Conditions for balancing the head in primates, American Journal of Physical Anthropology, vol.29, issue.4, pp.483-497, 1942.

A. H. Schultz, Age changes and variability in gibbons. A morphological study on a population sample of a man-like ape, American Journal of Physical Anthropology, vol.2, issue.1, pp.1-129, 1944.

A. H. Schultz, The position of the occipital condyles and of the face relative to the skull base in primates, American Journal of Physical Anthropology, vol.13, issue.1, pp.97-120, 1955.

A. H. Schultz, Postembryonic age changes, Primatologia, vol.1, pp.887-964, 1956.

A. H. Schultz, , 1961.

A. H. Schultz and W. L. Straus, The numbers of vertebrae in primates, Proceedings of the American Philosophical Society, vol.89, pp.601-626, 1945.

A. H. Schultz, The skeleton of the Hylobatidae and other observations on their morphology, Gibbon and Siamang, vol.2, pp.1-54, 1973.

M. S. ?enyürek, Cranial equilibrium index, American Journal of Physical Anthropology, vol.24, issue.1, pp.23-41, 1938.

L. J. Shapiro, Functional morphology of the primate spine with special reference to orthograde posture and bipedal locomotion. Doctoral dissertation, 1991.

L. J. Shapiro, Functional morphology of the vertebral column in primates, 1993.

L. J. Shapiro, Morphological and functional differentiation in the lumbar spine of lorisids and galagids, American Journal of Primatology, vol.69, issue.1, pp.86-102, 2007.

L. J. Shapiro, C. V. Seiffert, L. R. Godfrey, W. L. Jungers, E. L. Simons et al., Morphometric analysis of lumbar vertebrae in extinct Malagasy strepsirrhines, American Journal of Physical Anthropology, vol.128, issue.4, pp.823-839, 2005.

B. T. Shea, Developmental perspective on size change and allometry in evolution, Evolutionary Anthropology: Issues, News, and Reviews, vol.1, pp.125-134, 1992.

R. M. Shearman and A. C. Burke, The lateral somitic frontier in ontogeny and phylogeny, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.312, pp.603-612, 2009.

L. T. Shirai and G. Marroig, Skull modularity in neotropical marsupials and monkeys: size variation and evolutionary constraint and flexibility, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.314, pp.663-683, 2010.

S. Shou, V. Scott, C. Reed, R. Hitzemann, and H. S. Stadler, Transcriptome analysis of the murine forelimb and hindlimb autopod. Developmental dynamics: an official publication of the, vol.234, pp.74-89, 2005.

M. T. Silcox, J. I. Bloch, D. M. Boyer, M. Godinot, T. M. Ryan et al., Semicircular canal system in early primates, Journal of Human Evolution, vol.56, issue.3, pp.315-327, 2009.
URL : https://hal.archives-ouvertes.fr/hal-00549509

N. Singh, K. Harvati, J. J. Hublin, and C. P. Klingenberg, Morphological evolution through integration: a quantitative study of cranial integration in Homo, Pan, Gorilla and Pongo, Journal of Human Evolution, vol.62, issue.1, pp.155-164, 2012.

E. J. Slijper, Comparative biologic-anatomical investigations on the vertebral column and spinal musculature of mammals. Koninklijke Nederlandse Akademie van Wetenschappen, vol.42, pp.1-128, 1946.

V. Slon, F. Mafessoni, B. Vernot, C. De-filippo, S. Grote et al.,

K. Douka, The genome of the offspring of a Neanderthal mother and a Denisovan father, Nature, vol.561, issue.7721, pp.113-116, 2018.

M. D. Sockol, D. A. Raichlen, and H. Pontzer, Chimpanzee locomotor energetics and the origin of human bipedalism, Proceedings of the National Academy of Sciences of the United States of America, vol.104, pp.12265-12269, 2007.

F. Spoor, Basicranial architecture and relative brain size of Sts 5, South African Journal of Science, vol.93, issue.4, pp.182-186, 1997.

F. Spoor, B. Wood, and F. Zonneveld, Implications of early hominid labyrinthine morphology for evolution of human bipedal locomotion, Nature, vol.369, pp.645-648, 1994.

S. J. Steppan, P. C. Phillips, and D. Houle, Comparative quantitative genetics: evolution of the G matrix, Trends in Ecology & Evolution, vol.17, issue.7, pp.320-327, 2002.

J. T. Stern, Climbing to the top: a personal memoir of Australopithecus afarensis, Evolutionary Anthropology: Issues, News, and Reviews, vol.9, issue.3, pp.113-133, 2000.

T. D. Stewart, , 1962.

, Neanderthals from Iraq. Anthropos, vol.57, pp.779-800

D. S. Strait, Integration, phylogeny, and the hominid cranial base, American Journal of Physical Anthropology, vol.114, pp.273-297, 2001.

D. S. Strait and C. F. Ross, Kinematicdata on primate head and neck posture: implications for the evolution of basicranial flexion and an evaluation of registration planes used in paleoanthropology, American Journal of Physical Anthropology, vol.108, pp.205-222, 1999.

I. Susanna, D. M. Alba, S. Almécija, and S. Moya-sola, The vertebral remains of the late Miocene great ape Hispanopithecus laietanus from Can Llobateres 2 (Valles-Penedes Basin, NE Iberian Peninsula), Journal of Human Evolution, vol.73, pp.15-34, 2014.

R. L. Susman, New hominid fossils from the Swartkrans Formation (1979-1986 excavations): postcranial specimens, American Journal of Physical Anthropology, vol.79, issue.4, pp.451-474, 1989.

T. Sutikna, M. W. Tocheri, M. J. Morwood, E. W. Saptomo, R. D. Awe et al.,

M. Storey, Revised stratigraphy and chronology for Homo floresiensis at, 2016.

, Liang Bua in Indonesia, Nature, vol.532, issue.7599, pp.366-369

T. Sutikna, M. W. Tocheri, J. T. Faith, R. D. Awe, H. J. Meijer et al., The spatio-temporal distribution of archaeological and faunal finds at Liang Bua (Flores, Indonesia) in light of the revised chronology for Homo floresiensis, Journal of Human Evolution, vol.124, pp.52-74, 2018.

H. Tatsuta, K. H. Takahashi, and Y. Sakamaki, Geometric morphometrics in entomology: Basics and applications, Entomological science, vol.21, issue.2, pp.164-184, 2018.

C. M. Ten-broek, A. J. Bakker, I. Varela-lasheras, M. Bugiani, S. Van-dongen et al., Evo-devo of the human vertebral column: on homeotic transformations, pathologies and prenatal selection, Evolutionary Biology, vol.39, pp.456-471, 2012.

L. Testut and A. Latarjet, Anatomía Humana. Salvat, 1979.

D. W. Thompson, On growth and form, 1917.

S. K. Thorpe, J. M. Mcclymont, and R. H. Crompton, The arboreal origins of human bipedalism, Antiquity, vol.88, issue.341, pp.906-914, 2014.

N. Torres-tamayo, D. García-martínez, S. Lois-zlolniski, and I. Torres-sánchez, , p.329

, sloths, manatees and homeotic mutations, vol.2, p.11

P. P. Vidal, W. Graf, and A. Berthoz, The orientation of the cervical vertebral column in unrestrained awake animals, Experimental Brain Research, vol.61, issue.3, pp.549-559, 1986.

C. I. Villamil, Locomotion and basicranial anatomy in primates and marsupials, Journal of Human Evolution, vol.111, pp.163-178, 2017.

C. I. Villamil, Phenotypic integration of the cervical vertebrae in the Hominoidea (Primates), Evolution, vol.72, issue.3, pp.490-517, 2018.

B. Villmoare, Morphological integration, evolutionary constraints, and extinction: a computer simulation-based study, Evolutionary Biology, vol.40, pp.76-83, 2013.

B. Villmoare, J. Fish, and W. Jungers, Selection, morphological integration, and strepsirrhine locomotor adaptations, Evolutionary Biology, vol.38, pp.88-99, 2011.

G. P. Wagner, On the eigenvalue distribution of genetic and phenotypic dispersion matrices: evidence for a nonrandom organization of quantitative character variation, Journal of Mathematical Biology, vol.21, pp.77-95, 1984.

G. P. Wagner, The systems approach: An interface between development and population genetic aspects of evolution, Patterns and Processes in the History of Life, pp.149-165, 1986.

G. P. Wagner, The influence of variation and of developmental constraints on the rate of multivariate phenotypic evolution, Journal of Evolutionary Biology, vol.1, issue.1, pp.45-66, 1988.

G. P. Wagner, Adaptation and the modular design of organisms, Advances in Artificial Life, vol.929, pp.317-328, 1995.

G. P. Wagner, Homologues, natural kinds and the evolution of modularity, 1996.

, American Zoologist, vol.36, pp.36-43

G. P. Wagner and L. Altenberg, Perspective: complex adaptations and the evolution of evolvability, Evolution, vol.50, pp.967-976, 1996.

G. P. Wagner and K. Schwenk, Evolutionarily stable configurations: functional integration and the evolution of phenotypic stability, Evolutionary biology, vol.31, pp.155-217, 2000.

A. Walker, M. R. Zimmerman, and R. E. Leakey, , 1982.

, A in Homo erectus, Nature, vol.296, issue.5854, pp.248-250

A. Walker and . Leakey, The nariokotome Homo erectus skeleton, 1993.

B. Springer,

C. V. Ward, Torso morphology and locomotion in Proconsul nyanzae, American Journal of Physical Anthropology, vol.92, issue.3, pp.291-328, 1993.

C. V. Ward, Interpreting the posture and locomotion of Australopithecus afarensis: where do we stand, American Journal of Physical Anthropology, vol.119, issue.S35, pp.185-215, 2002.

C. V. Ward, Postcranial and locomotor adaptations of hominoids, Handbook of Paleoanthropology, pp.1011-1030, 2007.

C. V. Ward, T. K. Nalley, F. Spoor, P. Tafforeau, and Z. Alemseged, Thoracic vertebral count and thoracolumbar transition in Australopithecus afarensis, Proceedings of the National Academy of Sciences of the United States of America, vol.114, pp.6000-6004, 2017.

A. G. Warrener, K. L. Lewton, H. Pontzer, and D. E. Lieberman, A wider pelvis does not increase locomotor cost in humans, with implications for the evolution of childbirth, PloSone, vol.10, issue.3, p.118903, 2015.

D. M. Wellik and M. R. Capecchi, Hox10 and Hox11 genes are required to globally patternthe mammalian skeleton, Science, vol.301, pp.363-367, 2003.

A. A. White and M. M. Panjabi, Clinical biomechanics of the spine, 1990.

T. D. White and P. A. Folkens, The human bone manual, 2005.

T. D. White, G. Woldegabriel, B. Asfaw, S. Ambrose, Y. Beyene et al., Asa Issie, Aramis and the origin of Australopithecus, Nature, vol.440, issue.7086, p.883, 2006.
URL : https://hal.archives-ouvertes.fr/hal-00140503

T. D. White, B. Asfaw, Y. Beyene, Y. Haile-selassie, C. O. Lovejoy et al., Ardipithecus ramidus and the paleobiology of early hominids, vol.326, pp.64-86, 2009.

K. K. Whitcome, L. J. Shapiro, and D. E. Lieberman, Nature, vol.450, issue.7172, p.1075, 2007.

S. A. Williams, Morphological integration and the evolution of knuckle-walking, Journal of Human Evolution, vol.58, pp.432-440, 2010.

S. A. Williams, Placement of the diaphragmatic vertebra in catarrhines: implications for the evolution of dorsostability in hominoids and bipedalism in hominins, American Journal of Physical Anthropology, vol.148, issue.1, pp.111-122, 2012.

S. A. Williams, Variation in anthropoid vertebral formulae: implications for homology and homoplasy in hominoid evolution, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.318, issue.2, pp.134-147, 2012.

S. A. Williams, K. R. Ostrofsky, N. Frater, S. E. Churchill, P. Schmid et al., The vertebral column of Australopithecus sediba, Science, vol.340, issue.6129, 2013.

S. A. Williams and G. A. Russo, Evolution of the hominoid vertebral column: the long and the short of it, Evolutionary Anthropology: Issues, News, and Reviews, vol.24, issue.1, pp.15-32, 2015.

S. A. Williams, E. R. Middleton, C. I. Villamil, and M. R. Shattuck, Vertebral numbers and human evolution, American Journal of Physical Anthropology, vol.159, pp.19-36, 2016.

S. A. Williams, D. García-martínez, M. Bastir, M. R. Meyer, S. Nalla et al.,

L. R. Berger, The vertebrae and ribs of Homo naledi, Journal of Human Evolution, vol.104, pp.136-154, 2017.

S. A. Williams, M. R. Meyer, S. Nalla, D. García-martínez, T. K. Nalley et al., The vertebrae, ribs, and sternum of Australopithecus sediba, pp.156-233, 2018.

S. A. Williams, A. Gómez-olivencia, and D. R. Pilbeam, Numbers of Vertebrae in Hominoid Evolution, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.97-124, 2019.

S. A. Williams and M. R. Meyer, The Spine of Australopithecus, Spinal Evolution, Morphology, Function, and Pathology of the spine in Hominoid Evolution, pp.125-151, 2019.

B. Wood and B. G. Richmond, Human evolution: taxonomy and paleobiology, The Journal of Anatomy, vol.197, pp.19-60, 2000.

B. Wood and M. Collard, The meaning of Homo. Ludus Vitalis, vol.9, pp.63-74, 2001.

R. E. Wood, T. F. Higham, T. De-torres, N. Tisnérat-laborde, H. Valladas et al., A new date for the Neanderthals from El Sidrón Cave, vol.55, pp.148-158, 2013.

N. M. Young and B. Hallgrímsson, Serial homology and the evolution of mammalian limb covariation structure, Evolution, vol.59, pp.2691-2704, 2005.

N. M. Young, G. P. Wagner, and B. Hallgrímsson, Development and the evolvability of human limbs, Proceedings of the National Academy of Sciences of the United States of America, vol.107, pp.3400-3405, 2010.

M. L. Zelditch, Ontogenetic variation in patterns of phenotypic integration in the laboratory rat, Evolution, vol.42, pp.28-41, 1988.

M. L. Zelditch and C. Carmichael, Ontogenetic variation in patterns of developmental and functional integration in skulls of Sigmodon fulviventer, 1989.

, Evolution, vol.43, issue.4, pp.814-824

L. Aiello and C. Dean, An introduction to Human Evolutionary Anatomy, 1990.

G. H. Albrecht, Multivariate analysis and the study of form with special reference to canonical variate analysis, Am. Zool, vol.20, pp.679-693, 1980.

F. Ankel, The Functional and Evolutionary biology of Primates, pp.223-240, 1972.

J. L. Arsuaga, I. Martinez, O. Blackwell, J. L. Arsuaga, J. Carretero et al., The anatomy of the lumbar spine of Homo neanderthalensis and its phylogenetic and functional implications, Spain. Proc. Natl. Acad. Sci. 112, 11524e11529. Bastir, vol.8, 2005.

L. R. Berger, D. J. De-ruiter, S. E. Churchill, P. Schmid, K. J. Carlson et al., Middle Pleistocene lower back and pelvis from an aged human individual from the Sima de los Huesos site, Spain. Proc. Natl. Acad. Sci. 107, 18386e18391. Bookstein, vol.328, pp.131-151, 1990.

M. Boule, . 1911e1913.-l'homme-fossile-de-la-chapelle-aux-saints, . M. Ann-;-d, and D. E. Lieberman, Endurance running and the evolution of Homo, Anthropologie. Handbuch der vergleichenden Biologie des Menschen. Gustav Fischer, vol.432, pp.160-232, 1988.

F. H. Brown, J. Harris, R. Leakey, and A. Walker, Early Homo erectus skeleton from West Lake Turkana, Nature, vol.316, pp.788-792, 1985.

E. Bruner and G. Manzi, Allometric analysis of the skull in Pan and Gorilla by geometric morphometrics, Riv. Antropol, pp.45-52, 2001.

J. M. Carretero, C. Lorenzo, J. L. Arsuaga, M. N. Coleman, D. C. Cook et al., What does geometric mean, mean geometrically? Assessing the utility of geometric mean and other size variables in studies of skull allometry, Am. J. Phys. Anthropol, vol.37, pp.83-102, 1983.

A. G-omez-olivencia, A. Omez-olivencia, A. ;-g-omez-olivencia, J. M. Carretero, J. L. Arsuaga et al., Conclusion: Implications of KSD-VP-1/1 for Early Hominin Paleobiology and Insights into the Chimpanzee/Human Last Common Ancestor, 582e607. Gommery, D., 1997. Les atlas et les axis des Hominid es du Plio-Pl eistoc ene: morphologie et syst ematique. C.R. Acad. Sci, vol.99, pp.179-187, 2002.

Ø. Hammer, D. A. Harper, and P. D. Ryan, PAST: Paleontological statistics software package for education and data analysis, Palaeontol. Electron, vol.4, issue.1, p.9, 2001.

H. Ishida, Y. Kunimatsu, T. Takana, Y. Nakano, and M. Nakatsukasa, Nacholapithecus skeleton from the Middle Miocene of Kenya, J. Hum. Evol, vol.46, pp.69-103, 2004.

D. C. Johanson, M. Taieb, and Y. Coppens, Stratigraphic, chronologic, and paleoenvironmental contexts, with notes on hominid morphology and systematics, Am. J. Phys. Anthropol, vol.57, pp.137-161, 1982.

Y. Kikuchi, Y. Nakano, M. Nakatsukasa, D. Kunimatsu, D. Shimizu et al., Functional morphology and anatomy of cervical vertebrae in Nacholapithecus kerioi, a middle Miocene hominoid from Kenya, J. Hum. Evol, vol.62, pp.677-695, 2012.

Y. Kikuchi, M. Nakatsukasa, Y. Nakano, Y. Kunimatsu, D. Shimizu et al., Morphology of the thoracolumbar spine of the middle Miocene hominoid Nacholapithecus kerioi from northern Kenya, Advances in morphometrics, vol.88, pp.23-49, 1996.

C. P. Klingenberg, MorphoJ: an integrated software package for geometric morphometrics, 353e357. Klingenberg, C.P., 2015. Analyzing Fluctuating Asymmetry with Geometric Morphometrics: Concepts, Methods, and Applications. Symmetry 7, vol.11, pp.843-934, 2011.

C. P. Klingenberg and S. D. Zaklan, Morphological integration between developmental compartments in the Drosophila wing, Evolution, vol.54, pp.1273-1285, 2000.

C. P. Klingenberg, M. Barluenga, and A. Meyer, Shape analysis of symmetric structures: quantifying variation among individuals and asymmetry, Evolution, vol.56, issue.10, 2002.

B. Latimer and C. V. Ward, The Nariokotome Homo erectus skeleton, pp.266-293, 1993.

C. O. Lovejoy, D. C. Johanson, Y. Coppens, E. Manfreda, P. Mitteroecker et al., Elements of the axial skeleton recovered from the Hadar Formation: 1974e1977 collections, Am. J. Phys. Anthropol, vol.57, 1982.

K. V. Mardia, J. T. Kent, and J. M. Bibby, , 1979.

K. V. Mardia, F. L. Bookstein, and I. J. Moreton, Statistical assessment of bilateral symmetry of shapes, Biometrika, vol.285, issue.300, 2000.

I. Martínez, La base del cr aneo y el hueso temporal en la evoluci on de los homínidos, con especial referencia a los f osiles de la, 1995.

M. Mccollum, B. A. Rosenman, G. Suwa, R. S. Meindl, and C. O. Lovejoy, The vertebral formula of the last common ancestor of african apes and humans, J. Exp. Zool. (Mol. Dev. Evol.), vol.314, pp.123-134, 2010.

I. Mcdougall, F. H. Brown, P. M. Vasconcelos, B. E. Cohen, D. S. Thiede et al., New single crystal 40Ar/39Ar ages improve time scale for deposition of the Omo Group, J. Geol. Soc, vol.169, pp.213-226, 2012.

M. Meyer, Functional biology of the Homo erectus axial skeleton from Dmanisi, 2005.

M. R. Meyer, The cervical vertebrae of KSD-VP-1/1, The Postcranial Anatomy of Australopithecus afarensis: New Insights from KSD-VP-1/1, pp.63-111, 2016.

M. R. Meyer and M. Haeusler, Spinal cord evolution in early Homo, J. Hum. Evol, vol.88, pp.43-53, 2015.

R. A. Mittermeier, A. B. Rylands, and . Wilson, 2013. Handbook of the Mammals of the World. 3. Primates. Lynx Edicions

P. Mitteroecker, P. Gunz, M. Bernhard, K. Schaefer, and F. L. Bookstein, Comparison of cranial ontogenetic trajectories among great apes and humans, 2004.

, Comparative study of Moroto vertebral specimens, 679e698. Nakatsukasa, M, vol.46, pp.581-588, 2008.

M. Nakatsukasa, C. V. Ward, A. Walker, M. F. Teaford, Y. Kunimatsu et al., The study of morphological variation in the hominid fossil record: biology, landmarks and geometry, 777e784. Nakatsukasa, vol.46, pp.103-120, 2000.

A. Pablos, I. Martínez, C. Lorenzo, A. Gracia, N. Sala et al., The orientation of the articular facets of the zygapophyseal joints at the cervical and upper thoracic region, J. Hum. Evol, vol.65, pp.431-441, 2001.

A. R. Palmer and C. Strobeck, Fluctuating asymmetry: measurement, analysis, patterns, Annu. Rev. Ecol. Evol. Syst, pp.391-421, 1986.

R. Pickering, P. H. Dirks, Z. Jinnah, D. J. De-ruiter, S. E. Churchill et al., Australopithecus sediba at, 2011.

D. Pilbeam, . 302b, J. Piveteau, L. Xlix, and H. Preuschoft, Mechanisms for the acquisition of habitual bipedality: are there biomechanical reasons for the acquisition of upright bipedal posture?, 363e384. R Development Core Team, 2011. R: A Language and Environment for Statistical Computing. the R Foundation for Statistical Computing, vol.333, 1963.

Y. Rak, A. Ginzburg, and E. Geffen, Gorilla-like anatomy on Australopithecus afarensis mandibles suggests Au. afarensis link to robust australopiths, Proc. Natl. Acad. Sci, 1990.

L. Ríos and F. V. Cardoso, Age estimation from stages of union of the vertebral epiphyses of the ribs, Am. J. Phys. Anthropol, vol.140, pp.265-274, 2009.

F. J. Rohlf and M. Corti, The use of two block partial least-squares to study covariation in shape, Syst. Biol, vol.49, pp.740-753, 2000.

G. A. Russo, L. J. Shapiro, and W. J. Sanders, Comparative morphometric study of the australopithecine vertebral series Stw-H8/H41, J. Hum. Evol, vol.65, pp.249-302, 1998.

A. H. Schultz, . Karger, . Basel, A. H. Schultz, and W. L. Straus, Primatologia. Handbuch der primatenkunde, Proc. Am. Phil. Soc, vol.89, pp.601-626, 1945.

L. J. Shapiro, C. V. Seiffert, L. R. Godfrey, W. L. Jungers, E. L. Simons et al., Kinematic Data on Primate Head and Neck Posture: Implications for the Evolution of Basicranial Flexion and an Evaluation of Registration Planes Used in Paleoanthropology, Am. J. Phys. Anthropol, vol.128, pp.205-222, 1999.

Y. L. Tsai, M. C. Weng, T. W. Chen, Y. L. Hsieh, C. S. Chen et al., Correlation between the ossification of nuchal ligament and clinical cervical disorders, Poirier et Charpy's Trait e d'Anatomie, vol.28, pp.439-476, 1926.

S. A. Williams, Placement of the diaphragmatic vertebra in Catarrhines: Implications for the Evolution of dorsostability in Hominoids and bipedalism in hominins, Am. J. Phys. Anthropol, vol.148, pp.111-122, 2012.

S. A. Williams, Variation in anthropoid vertebral formulae: implications for homology in hominoid evolution, J. Exp. Zool. (Mol. Dev. Evol.), vol.318, pp.134-147, 2012.

S. A. Williams and G. A. Russo, Evolution of the hominoid vertebral column: The long and the short of it, Evol. Anthropol, vol.24, pp.15-32, 2015.

S. A. Williams, K. R. Ostrofsky, N. Frater, S. E. Churchill, P. Schmid et al., The Vertebral Column of Australopithecus sediba, Science, vol.340, p.1232996, 2013.

S. A. Williams, E. R. Middleton, C. I. Villamil, M. R. Shattuck, M. L. Zelditch et al., Geometric Morphometrics for Biologists: A Primer, Am. J. Phys. Anthropol, vol.159, 2004.

D. C. Adams, Evaluating modularity in morphometric data: Challenges with the RV coefficient and a new test measure, Methods in Ecology and Evolution, vol.7, pp.565-572, 2016.

D. C. Adams and E. Ot-arola-castillo, Geomorph: an R package for the collection and analysis of geometric morphometric shape data, Methods in Ecology and Evolution, vol.4, pp.393-399, 2013.

D. C. Adams and M. L. Collyer, On the comparison of the strength of morphological integration across morphometric datasets, Evolution, vol.70, pp.2623-2631, 2016.

D. C. Adams, M. L. Collyer, A. Kaliontzopoulou, and E. Sherratt, Geomorph: Software for geometric morphometric analyses, 2017.

M. Arlegi, A. Olivencia, L. Albessard, I. Martínez, A. Balzeau et al., The role of allometry and posture in the evolution of the hominin subaxial cervical spine, Journal of Human Evolution, vol.104, pp.80-99, 2017.
URL : https://hal.archives-ouvertes.fr/hal-02106776

P. Arnold, F. Forterre, J. Lang, and M. S. Fischer, Morphological disparity, conservatism, and integration in the canine lower cervical spine: Insights into mammalian neck function and regionalization, Mammalian Biology-Zeitschrift F? ur Säugetierkunde, vol.81, pp.153-162, 2016.

. Arlegi and . Al,

C. Barrey, J. Jund, O. Noseda, and P. Roussouly, Sagittal balance of the pelvis-spine complex and lumbar degenerative diseases. A comparative study about 85 cases, European Spine Journal, vol.16, pp.1459-1467, 2007.

M. Bastir, A. Rosas, and H. D. Sheets, The morphological integration of the hominoid skull: A partial least squares and PC analysis with implications for European Middle Pleistocene mandibular variation, Modern morphometrics in physical anthropology, pp.265-284, 2005.

W. Bateson, Materials for the study of variation treated with especial regard to discontinuity in the origin of species, 1894.

E. Been, A. Olivencia, S. Shefi, M. Soudack, M. Bastir et al., Evolution of spinopelvic alignment in hominins, The Anatomical Record, vol.300, pp.900-911, 2017.

F. L. Bookstein, Integration, disintegration, and self-similarity: Characterizing the scales of shape variation in landmark data, Evolutionary Biology, vol.42, pp.395-426, 2015.

B. M. Boszczyk, A. A. Boszcyk, and R. Putz, Comparative and functional anatomy of the mammalian lumbar spine, The Anatomical Record, vol.264, pp.157-168, 2001.

C. Boulay, C. Tardieu, J. Hecquet, C. Benaim, B. Mouilleseaux et al., Sagittal alignment of spine and pelvis regulated by pelvic incidence: standard values and prediction of lordosis, European Spine Journal, vol.15, pp.415-422, 2006.

D. M. Bramble and D. E. Lieberman, Endurance running and the evolution of Homo, Nature, vol.432, p.345, 2004.

J. M. Cheverud, Developmental integration and the evolution of pleiotropy, American Zoologist, vol.36, pp.44-50, 1996.

A. G. Drake and C. P. Klingenberg, The pace of morphological change: Historical transformation of skull shape in St Bernard dogs, Proceedings of the Royal Society of London B: Biological Sciences, vol.275, pp.71-76, 2008.

Y. Escoufier, Le traitement des variables vectorielles, Biometrics, vol.29, pp.751-760, 1973.

V. Feipel, T. De-mesmaeker, P. Klein, and M. Rooze, Threedimensional kinematics of the lumbar spine during treadmill walking at different speeds, European Spine Journal, vol.10, pp.16-22, 2001.

H. F. Gadow, The evolution of the vertebral column. A contribution to the study of vertebrate phylogeny, 1933.

F. Galis, Why do almost all mammals have seven cervical vertebrae? Developmental constraints, Hox genes, and cancer, Journal of Experimental Zoology, vol.285, pp.19-26, 1999.

F. Galis and J. A. Metz, Evolutionary novelties: The making and breaking of pleiotropic constraints, Integrative and Comparative Biology, vol.47, pp.409-419, 2007.

G. Omez-robles, A. Polly, and P. D. , Morphological integration in the hominin dentition: Evolutionary, developmental, and functional factors, Evolution, vol.66, pp.1024-1043, 2012.

A. Goswami, J. B. Smaers, C. Soligo, and P. D. Polly, The macroevolutionary consequences of phenotypic integration: From development to deep time, Philosophical Transactions Royal Society B, vol.369, 2014.

M. W. Grabowski, J. D. Polk, and C. C. Roseman, Divergent patterns of integration and reduced constraint in the human hip and the origins of bipedalism, Evolution, vol.65, pp.1336-1356, 2011.

L. J. Hlusko, Identifying metameric variation in extant hominoid and fossil hominid mandibular molars, American Journal of Physical Anthropology, vol.118, pp.86-97, 2002.

L. J. Hlusko, Integrating the genotype and phenotype in hominid paleontology, Proceedings of the National Academy of Sciences of the United States of America, vol.101, pp.2653-2657, 2004.

T. W. Holliday, Body size, body shape, and the circumscription of the genus Homo, Current Anthropology, vol.53, pp.330-345, 2012.

K. D. Hunt, Positional behavior in the Hominoidea, International Journal of Primatology, vol.12, pp.95-118, 1991.

T. L. Kivell and D. Schmitt, Independent evolution of knucklewalking in African apes shows that humans did not evolve from a knuckle-walking ancestor, Proceedings of the National Academy of Sciences of the United States of America, vol.106, pp.14241-14246, 2009.

C. P. Klingenberg, Morphometric integration and modularity in configurations of landmarks: Tools for evaluating a priori hypotheses, Evolution & Development, vol.11, pp.405-421, 2009.

B. Kurt-en, On the variation and population dynamics of fossil and recent mammal populations, Acta Zoologica Fennica, vol.76, pp.1-122, 1953.

C. O. Lovejoy and M. A. Mccollum, Spinopelvic pathways to bipedality: Why no hominids ever relied on a bent-hip-bent-knee gait, Philosophical Transactions of the Royal Society B: Biological Sciences, vol.365, pp.3289-3299, 2010.

E. Manfreda, P. Mitteroecker, F. L. Bookstein, and K. Schaefer, Functional morphology of the first cervical vertebra in humans and nonhuman primates, The Anatomical Record, vol.289, pp.184-194, 2006.

G. Marroig, L. T. Shirai, A. Porto, F. B. De-oliveira, and V. De-conto, The evolution of modularity in the mammalian skull II: Evolutionary consequences, Evolutionary Biology, vol.36, pp.136-148, 2009.

I. Martínez, La Base del Cr aneo y el Hueso Temporal en la Evoluci on de los Homínidos, con Especial Referencia a los F osiles de Atapuerca (Burgos) (Unpublished doctoral dissertation), 1995.

M. Meyer, Functional biology of the Homo erectus axial skeleton from Dmanisi, Georgia (Unpublished doctoral dissertation), 2005.

M. R. Meyer, The cervical vertebrae of KSD-VP-1/1, The postcranial anatomy of Australopithecus afarensis: New insights from KSD-VP-1, pp.63-111, 2016.

M. R. Meyer, S. A. Williams, P. Schmid, S. E. Churchill, and L. R. Berger, The cervical spine of Australopithecus sediba, Journal of Human Evolution, vol.104, pp.32-49, 2017.

P. Mitteroecker and F. Bookstein, The conceptual and statistical relationship between modularity and morphological integration, Systematic Biology, vol.56, pp.818-836, 2007.

L. R. Monteiro, Multivariate regression models and geometric morphometrics: The search for causal factors in the analysis of shape, Systematic Biology, vol.48, pp.192-199, 1999.

J. E. Mosimann, Size allometry: Size and shape variables with characterizations of the lognormal and generalized gamma distributions, Journal of the American Statistical Association, vol.65, pp.930-945, 1970.

T. K. Nalley and N. Grider-potter, Functional morphology of the primate head and neck, American Journal of Physical Anthropology, vol.156, pp.531-542, 2015.

T. K. Nalley and N. Grider-potter, Functional analyses of the primate upper cervical vertebral column, Journal of human evolution, vol.107, pp.19-35, 2017.

E. C. Olson and R. L. Miller, Morphological integration. Chicago, 1958.

S. E. Pierce, J. A. Clack, and J. R. Hutchinson, Comparative axial morphology in pinnipeds and its correlation with aquatic locomotory behaviour, Journal of Anatomy, vol.219, pp.502-514, 2011.

D. Pilbeam, The anthropoid postcranial axial skeleton: Comments on development, variation, and evolution, Journal of Experimental Zoology Part B: Molecular and Developmental Evolution, vol.302, pp.241-267, 2004.

. Arlegi and . Al,

J. M. Polanski, Morphological integration of the modern human mandible during ontogeny, International Journal of Evolutionary Biology, 2011.

J. M. Polanski and R. G. Franciscus, Patterns of craniofacial integration in extant Homo, Pan, and Gorilla, American Journal of Physical Anthropology, vol.131, pp.38-49, 2006.

A. Porto, F. B. De-oliveira, L. T. Shirai, V. De-conto, and G. Marroig, The evolution of modularity in the mammalian skull I: Morphological integration patterns and magnitudes, Evolutionary Biology, vol.36, pp.118-135, 2009.

. R-core-team, R: A language and environment for statistical computing, version 3.2. 3. R Foundation for Statistical Computing, 2015.

M. Randau and A. Goswami, Unravelling intravertebral integration, modularity and disparity in Felidae (Mammalia), Evolution & Development, vol.19, pp.85-95, 2017.

M. Randau and A. Goswami, Morphological modularity in the vertebral column of Felidae (Mammalia, Carnivora), BMC Evolutionary Biology, vol.17, p.133, 2017.

F. J. Rohlf and M. Corti, Use of two-block partial least-squares to study covariation in shape, Systematic Biology, vol.49, pp.740-753, 2000.

F. J. Rohlf and D. Slice, Extensions of the Procrustes method for the optimal superimposition of landmarks, Systematic Biology, vol.39, pp.40-59, 1990.

C. Rolian, Integration and evolvability in primate hands and feet, Evolutionary Biology, vol.36, pp.100-117, 2009.

C. Rolian, D. E. Lieberman, and B. Hallgrímsson, The coevolution of human hands and feet, Evolution, vol.64, pp.1558-1568, 2010.

C. Rolian and K. E. Willmore, Morphological integration at 50: Patterns and processes of integration in biological anthropology, Evolutionary Biology, vol.36, pp.1-4, 2009.

G. A. Russo and S. A. Williams, Giant pandas (Carnivora: Ailuropoda melanoleuca) and living hominoids converge on lumbar vertebral adaptations to orthograde trunk posture, Journal of Human Evolution, vol.88, pp.160-179, 2015.

L. Scheuer, S. M. Black, and A. Christie, Developmental juvenil osteology, 2000.

A. H. Schultz and W. L. Straus, The numbers of vertebrae in primates, Proceedings of the American Philosophical Society, vol.89, pp.601-626, 1945.

B. T. Shea, Developmental perspective on size change and allometry in evolution, Evolutionary Anthropology: Issues, News, and Reviews, vol.1, pp.125-134, 1992.

E. J. Slijper, Comparative biologic-anatomical investigations on the vertebral column and spinal musculature of mammals. Koninklijke Nederlandse Akademie van Wetenschappen, vol.42, pp.1-128, 1946.

D. S. Strait and C. F. Ross, Kinematicdata on primate head and neck posture: Implications for the evolution of basicranial flexion and an evaluation of registration planes used in paleoanthropology, American Journal of Physical Anthropology, vol.108, pp.205-222, 1999.

R. H. Tuttle and D. P. Watts, The positional behavior and adaptive complexes of Pan (Gorilla), Primate morphophysiology, locomotor analyses and human bipedalism, pp.261-288, 1985.

I. Varela-lasheras, A. J. Bakker, S. D. Van-der-mije, J. A. Metz, J. Van-alphen et al., Breaking evolutionary and pleiotropic constraints in mammals: On sloths, manatees and homeotic mutations, EvoDevo, vol.2, p.11, 2011.

C. I. Villamil, Phenotypic integration of the cervical vertebrae in the Hominoidea (Primates), Evolution. Accepted Author Manuscript

S. A. Williams, Morphological integration and the evolution of knuckle-walking, Journal of Human Evolution, vol.58, pp.432-440, 2010.

S. A. Williams, E. R. Middleton, C. I. Villamil, and M. R. Shattuck, Vertebral numbers and human evolution, American Journal of Physical Anthropology, vol.159, pp.19-36, 2016.

S. A. Williams, K. R. Ostrofsky, N. Frater, S. E. Churchill, P. Schmid et al., The vertebral column of Australopithecus sediba, Science, vol.340, p.1232996, 2013.

A. A. White, M. Panjabi, . Philadelphia, and . Lippincot, Clinical biomechanics of the spine, 1978.

N. M. Young and B. Hallgrímsson, Serial homology and the evolution of mammalian limb covariation structure, Evolution, vol.59, pp.2691-2704, 2005.

M. L. Zelditch, Ontogenetic variation in patterns of phenotypic integration in the laboratory rat, Evolution, vol.42, pp.28-41, 1988.

R. R. Ackermann, Morphological integration and the interpretation of fossil hominin diversity, Evolutionary Biology, vol.36, pp.149-156, 2009.

D. C. Adams, Evaluating modularity in morphometric data: Challenges with the RV coefficient and a new test measure, Methods in Ecology and Evolution, vol.7, pp.565-572, 2016.

D. C. Adams, M. L. Collyer, A. Kaliontzopoulou, and E. Sherratt, Geomorph: Software for geometric morphometric analyses, 2017.

M. Arlegi, A. Gómez-olivencia, L. Albessard, I. Martínez, A. Balzeau et al., The role of allometry and posture in the evolution of the hominin subaxial cervical spine, Journal of Human Evolution, vol.104, pp.80-99, 2017.
URL : https://hal.archives-ouvertes.fr/hal-02106776

M. Arlegi, A. Gómez-robles, and A. Gómez-olivencia, Morphological integration in the gorilla, chimpanzee, and human neck, American Journal of Physical Anthropology, vol.166, pp.408-416, 2018.

P. Arnold, B. Esteve-altava, and M. S. Fischer, Musculoskeletal networks reveal topological disparity in mammalian neck evolution, BMC Evolutionary Biology, vol.17, p.251, 2017.

P. Arnold, F. Forterre, J. Lang, and M. S. Fischer, Morphological disparity, conservatism, and integration in the canine lower cervical spine: Insights into mammalian neck function and regionalization, Mammalian Biology-Zeitschrift für Säugetierkunde, vol.81, pp.153-162, 2016.

R. J. Asher, K. H. Lin, N. Kardjilov, and L. Hautier, Variability and constraint in the mammalian vertebral column, Journal of Evolutionary Biology, vol.24, pp.1080-1090, 2011.

M. Bastir and A. Rosas, Hierarchical nature of morphological integration and modularity in the human posterior face, American Journal of Physical Anthropology, vol.128, pp.26-34, 2005.

W. Bateson, Materials for the study of variation: treated with especial regard to discontinuity in the origin of species, American Journal of Physical Anthropology, vol.147, pp.64-77, 1894.

Y. Benjamini and Y. Hochberg, Controlling the false discovery rate: A practical and powerful approach to multiple testing, Journal of the Royal Statistical Society. Series B, Statistical Methodology, vol.57, pp.289-300, 1995.

F. L. Bookstein, Size and shape": A comment on semantics, Systematic Zoology, vol.38, pp.173-180, 1989.

F. L. Bookstein, Integration, disintegration, and self-similarity: Characterizing the scales of shape variation in landmark data, Evolutionary Biology, vol.42, pp.395-426, 2015.

G. Bräuer, Osteometrie, Anthropologie. Hand-buch der vergleichenden Biologie des menschen, pp.160-232, 1988.

G. Fischer, :. Stuttgart, G. Buchholtz, and E. A. , Vertebral and rib anatomy in Caperea marginata: Implications for evolutionary patterning of the mammalian vertebral column, Marine Mammal Science, vol.27, pp.382-397, 2011.

E. A. Buchholtz, Flexibility and constraint: patterning the axial skeleton in mammals, From Clone to Bone: The Synergy of Morphological and Molecular Tools in Palaeobiology, pp.230-253, 2012.

E. A. Buchholtz, Crossing the frontier: A hypothesis for the origins of meristic constraint in mammalian axial patterning, Zoology, vol.117, pp.64-69, 2014.

E. A. Buchholtz, H. G. Bailin, S. A. Laves, J. T. Yang, M. Y. Chan et al., Fixed cervical count and the origin of the mammalian diaphragm, Evolution & Development, vol.14, pp.399-411, 2012.

E. A. Buchholtz and C. C. Stepien, Anatomical transformation in mammals: Developmental origin of aberrant cervical anatomy in tree sloths, Evolution & Development, vol.11, pp.69-79, 2009.

E. A. Buchholtz, K. L. Wayrynen, and I. W. Lin, Breaking constraint: Axial patterning in Trichechus (Mammalia: Sirenia). Evolution & Development, vol.16, pp.382-393, 2014.

A. C. Burke, C. E. Nelson, B. A. Morgan, and C. Tabin, Hox genes and the evolution of vertebrate axial morphology, Development, vol.121, pp.333-346, 1995.

W. P. Cane, The ontogeny of postcranial integration in the common tern, Sterna hirundo, Evolution, vol.47, pp.1138-1151, 1993.

J. Cheverud, Quantitative genetics and developmental constraints on evolution by selection, Journal of Theoretical Biology, vol.110, pp.155-171, 1984.

J. M. Cheverud, A comparison of genetic and phenotypic correlations, Evolution, vol.42, pp.958-968, 1988.

J. M. Cheverud, Developmental integration and the evolution of pleiotropy, American Zoologist, vol.36, pp.44-50, 1996.

J. M. Cheverud, J. J. Rutledge, and W. R. Atchley, Quantitative genetics of development: Genetic correlations among age-specific trait values and the evolution of ontogeny, Evolution, vol.37, pp.895-905, 1983.

J. M. Cheverud, G. P. Wagner, and M. M. Dow, Methods for the comparative analysis of variation patterns, Systematic Zoology, vol.38, pp.201-213, 1989.

L. M. Chevin, R. Lande, and G. M. Mace, Adaptation, plasticity, and extinction in a changing environment: Towards a predictive theory, PLoS Biology, vol.8, p.1000357, 2010.

A. Cil, M. Yazici, A. Uzumcugil, U. Kandemir, A. Alanay et al., The evolution of sagittal segmental alignment of the spine during childhood, Spine, vol.30, pp.93-100, 2005.

A. W. Crompton, F. A. Jenkins, and . Jr, Mammals from reptiles: A review of mammalian origins, Annual Review of Earth and Planetary Sciences, vol.1, pp.131-155, 1973.

J. N. Darroch and J. E. Mosimann, Canonical and principal components of shape, Biometrika, vol.72, pp.241-252, 1985.

Y. Escoufier, Le traitement des variables vectorielles, Biometrics, vol.29, pp.751-760, 1973.

B. Favier and P. Dolle, Developmental functions of mammalian Hox genes, Molecular Human Reproduction, vol.3, pp.115-131, 1997.

A. Fleming, M. G. Kishida, C. B. Kimmel, and R. J. Keynes, Building the backbone: The development and evolution of vertebral patterning, Development, vol.142, pp.1733-1744, 2015.

F. Galis, Why do almost all mammals have seven cervical vertebrae? Developmental constraints, Hox genes, and cancer, Journal of Experimental Zoology, vol.285, pp.19-26, 1999.

F. Galis, On the homology of structures and Hox genes: The vertebral column, Homology, vol.222, pp.80-94, 1999.

F. Galis, D. R. Carrier, J. Van-alphen, S. D. Van-der-mije, T. J. Van-dooren et al., Fast running restricts evolutionary change of the vertebral column in mammals, Proceedings of the National Academy of Sciences, vol.111, pp.11401-11406, 2014.

F. Galis and J. A. Metz, Evolutionary novelties: The making and breaking of pleiotropic constraints, Integrative and Comparative Biology, vol.47, pp.409-419, 2007.

S. J. Gaunt, Conservation in the Hox code during morphological evolution, International Journal of Developmental Biology, vol.38, pp.549-552, 1994.

C. A. Giglio and J. B. Volpon, Development and evaluation of thoracic kyphosis and lumbar lordosis during growth, Journal of Children's Orthopaedics, vol.1, pp.187-193, 2007.

A. Gómez-olivencia, M. Arlegi, A. Barash, J. T. Stock, and E. Been, The Neandertal vertebral column 2: The lumbar spine, Journal of Human Evolution, vol.106, pp.84-101, 2017.

A. Gómez-olivencia, J. M. Carretero, J. L. Arsuaga, L. Rodríguez-garcía, R. García-gonzález et al., Metric and morphological study of the upper cervical spine from the Sima de los Huesos site, Journal of Human Evolution, vol.53, pp.6-25, 2007.

A. Gómez-robles and P. D. Polly, Morphological integration in the hominin dentition: Evolutionary, developmental, and functional factors, Evolution: International Journal of Organic Evolution, vol.66, pp.1024-1043, 2012.

A. Goswami, Cranial modularity shifts during mammalian evolution, The American Naturalist, vol.168, pp.270-280, 2006.

A. Goswami, P. D. Polly, A. Goswami, J. B. Smaers, C. Soligo et al., The influence of modularity on cranial morphological disparity in Carnivora and primates (Mammalia), Philosophical Transactions of the Royal Society, B: Biological Sciences, vol.5, 2010.

M. Grabowski and A. Porto, How many more? Sample size determination in studies of morphological integration and evolvability, Methods in Ecology and Evolution, vol.8, pp.592-603, 2017.

M. W. Grabowski, J. D. Polk, and C. C. Roseman, Divergent patterns of integration and reduced constraint in the human hip and the origins of bipedalism, Evolution: International Journal of Organic Evolution, vol.65, pp.1336-1356, 2011.

B. Hallgrímsson, K. Willmore, and B. K. Hall, Canalization, developmental stability, and morphological integration in primate limbs, American Journal of Physical Anthropology, vol.119, pp.131-158, 2002.

T. F. Hansen, Is modularity necessary for evolvability, Remarks on the relationship between pleiotropy and evolvability, vol.69, pp.83-94, 2003.

T. F. Hansen and D. Houle, Measuring and comparing evolvability and constraint in multivariate characters, Journal of Evolutionary Biology, vol.21, pp.1201-1219, 2008.

S. E. Hartman, Evaluation of some alternative procedures used in numerical systematics, Systematic Zoology, vol.37, issue.1, p.18, 1988.

L. Hautier, V. Weisbecker, M. R. Sánchez-villagra, A. Goswami, and R. J. Asher, Skeletal development in sloths and the evolution of mammalian vertebral patterning, Proceedings of the National Academy of Sciences, vol.107, pp.18903-18908, 2010.

S. Hayama, M. Nakatsukasa, and Y. Kunimatsu, Monkey performance: The development of bipedalism in trained Japanese monkeys, Kaibogaku Zasshi. Journal of Anatomy, vol.67, pp.169-185, 1992.

J. J. Head and P. D. Polly, Evolution of the snake body form reveals homoplasy in amniote Hox gene function, Nature, vol.520, pp.86-89, 2015.

W. G. Hill and R. Thompson, Probabilities of non-positive definite between-group or genetic covariance matrices, Biometrics, vol.34, pp.429-439, 1978.

L. J. Hlusko, Identifying metameric variation in extant hominoid and fossil hominid mandibular molars, American Journal of Physical Anthropology, vol.118, pp.86-97, 2002.

F. A. Jenkins, The postcranial skeleton of African cynodonts. Bulletin of the Peabody Museum of Natural History, vol.36, pp.1-216, 1971.

K. E. Jones, K. D. Angielczyk, P. D. Polly, J. J. Head, V. Fernandez et al., Fossils reveal the complex evolutionary history of the mammalian regionalized spine, Science, vol.361, pp.1249-1252, 2018.

W. L. Jungers, A. B. Falsetti, and C. E. Wall, Shape, relative size, and size-adjustments in morphometrics, American Journal of Physical Anthropology, vol.38, pp.137-161, 1995.