Postcranial Morphology of the Middle Pleistocene Humans from Sima De Los Huesos, Spain
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POSTCRANIAL MORPHOLOGY OF THE MIDDLE PLEISTOCENE HUMANS FROM SIMA DE LOS HUESOS, SPAIN. Authors: Juan-Luis Arsuagaa,b,1, José-Miguel Carreteroc,a, Carlos Lorenzod,e,a, Asier Gómez-Olivenciaf,g,h,a, Adrián Pablosi,a, Laura Rodríguezc,j, Rebeca García-Gonzálezc, Alejandro Bonmatía,b, Rolf M. Quamk,l,a, Ana Pantoja-Péreza,b, Ignacio Martínezi,a, Arantza Aranburum, Ana Gracia-Téllezn,a, Eva Poza-Reya,b, Nohemi Salaa, Nuria Garcíaa,b, Almudena Alcázar de Velascoa, Gloria Cuenca-Bescóso, José-María Bermúdez de Castroj, Eudald Carbonelld,e,p. Author’s Affiliations: aCentro Mixto UCM-ISCIII de Evolución y Comportamiento Humanos. Avda. Monforte de Lemos, 5. 28029 Madrid, Spain. bDpto. de Paleontología. Fac. Ciencias Geológicas. Universidad Complutense de Madrid, Avda. Complutense s/n, 28040 Madrid, Spain. cLaboratorio de Evolución Humana, Dpto. de Ciencias Históricas y Geografía, Universidad de Burgos, Edificio I+D+i, Plaza Misael Bañuelos s/n, 09001 Burgos, Spain. dÀrea de Prehistòria, Dept. d’Història i Història de l’Art, Univ. Rovira i Virgili, Av. Catalunya, 35, 43002 Tarragona, Spain. eInstitut Català de Paleoecologia Humana i Evolució Social, C/ Marcel·lí Domingo s/n (Edifici W3), Campus Sescelades, 43007 Tarragona, Spain. fDept. Estratigrafía y Paleontología, Facultad de Ciencia y Tecnología, Universidad del País Vasco-Euskal Herriko Unibertsitatea. Apdo. 644, 48080 Bilbao, Spain. gIKERBASQUE. Basque Foundation for Science, 48013 Bilbao, Spain. hUMR 7194, CNRS, Dépt. Préhistoire, Muséum national d'Histoire naturelle. Musée de l’Homme, 17, Place du Trocadéro, 75016 Paris, France. iÁrea de Antropología Física, Depto. de Ciencias de la Vida. Universidad de Alcalá, 28871 Alcalá de Henares, Spain. jCentro Nacional de Investigación sobre la Evolución Humana, Paseo Sierra de Atapuerca, 09002 Burgos, Spain. kDept. of Anthropology, Binghamton University (State University of New York), Binghamton, NY 13902-6000, USA. lDivision of Anthropology, American Museum of Natural History, New York, NY 10024- 5192, USA. mDepartamento Mineralogía y Petrología, Facultad de Ciencia y Tecnología, Universidad del País Vasco-Euskal Herriko Unibertsitatea. Apdo. 644, 48080 Bilbao, Spain. nÁrea de Paleontología. Depto. de Geografía y Geología. Universidad de Alcalá, 28871 Alcalá de Henares, Spain. oPaleontología, Aragosaurus-IUCA and Facultad Ciencias, Universidad de Zaragoza,50009 Zaragoza, Spain. pVisiting Professor, Institute of Vertebrate Paleontology and Paleoanthropology of Beijing, 100044 China. 1Corresponding author: Juan-Luis Arsuaga, Centro Mixto Universidad Complutense de Madrid - Instituto de Salud Carlos III (UCM-ISCIII) de Evolución y Comportamiento Humanos, c/ Monforte de Lemos 5 – Pabellón 14, 28029 Madrid (Spain). Telephone: +34918222834. Email: [email protected] 1-SKELETAL REPRESENTATION Table S1. Complete inventory of SH postcranial remains and MNI represented by each element Minimum Number of Minimum Number of Individuals Elements (MNE) (MNI) AD SubAD Indet Total AD SubAD Indet Total M F ? MF ? Total vertebrae 212 12 Cervical V. 28 28 14 70 5 2 5 12 Thoracic V. 46 49 95 Lumbar V. 19 28 47 Total ribs 58 58 2 118 3 3 1 7 Manubria 4 4 Clavicles 3 2 4 11 20 2 1 2 6 11 Scapulae 6 2 6 13 27 5 2 3 6 16 Humeri 8 2 2 12 24 5 1 1 7 14 Radii 7 3 4 11 25 3 2 2 8 15 Ulnae 6 3 4 12 25 3 3 3 5 14 Carpal bones 131 13 Scaphoid 14 5 19 8 5 13 Lunate 14 7 21 8 4 12 Triquetral 10 6 16 6 4 10 Pisiform 6 3 9 3 3 6 Trapezium 13 3 16 8 2 10 Trapezoid 13 4 17 8 2 10 Capitate 14 5 19 8 3 11 Hamate 10 4 14 7 4 11 Metacarpals 63 10 Mtc I 6 12 18 3 7 10 Mtc II 9 3 12 4 2 6 Mtc III 7 3 10 4 3 7 Mtc IV 3 3 6 2 2 4 Mtc V 11 6 17 6 3 9 Hand phalanges 325 13 Proximal 68 39 19 126 7 6 13 Middle 50 34 11 104 Distal 59 33 3 95 7 5 12 Innominate bones 5 5 4 17 4 31 3 1 1 10 2 17 Sacra 2 3 5 10 2 3 5 10 Coccygeal V. 5 3 8 2 2 4 Femora 6 2 1 23 32 4 1 14 19 Patellae 4 2 5 5 16 2 2 3 3 10 Tibiae 5 2 3 13 23 4 2 7 13 Fibulae 1 1 7 9 18 1 1 3 3 8 Tarsal bones 130 14 Talus 21 3 24 34 4 3 14 Calcaneus 17 9 26 43 2 6 15 Navicular 20 5 25 9 5 14 Cuboid 15 3 18 7 3 10 Medial cuneiform 9 5 14 5 3 8 Inter. cuneiform 13 2 15 8 2 10 Lateral cuneiform 9 3 12 5 1 6 Metatarsals 51 11 Mtt I 10 5 15 6 5 11 Mtt II 7 1 8 4 1 5 Mtt III 6 4 10 5 3 8 Mtt IV 5 4 9 4 3 7 Mtt V 5 4 9 4 3 7 Foot phalanges 230 16 Proximal Ph. I 10 10 20 8 8 16 Distal Ph. I 9 7 16 5 4 9 Proximal II-V 20 35 9 64 Middle II-V 52 16 68 Distal II-V 53 9 62 Total MNE 1523 Final MNI 19 M = male; F = female; AD = adult; SubAD = subadult Table S2. Relative representation and cumulative percentage One Relative Cumulative Anatomical Units (AU) NºAU MNAU Skeleton MNAU % MNAU 1. Neurocrania 17 1 17.0 3.9 3.9 2. Mandibles 20 1 20.0 4.6 8.5 3. Dentition 533 32 16.7 3.9 12.4 4. Cerv. vertebrae 70 7 10.0 2.3 14.7 5. Thor. vertebrae 95 12 7.9 1.8 16.5 6. Lumb. vertebrae 47 5 9.4 2.2 18.7 7. Sacra 10 1 10.0 2.3 21.0 8. Ribs 118 24 4.9 1.1 22.1 9. Claviculae 20 2 10.0 2.3 24.4 10. Scapulae 27 2 13.5 3.1 27.5 11. Prox. humeri 18 2 9.0 2.1 29.6 12. Humeral shafts 37 2 18.5 4.3 33.9 13. Dist. humeri 27 2 13.5 3.1 37.0 14. Prox. ulnae 22 2 11.0 2.6 39.6 15. Ulnar shafts 27 2 13.5 3.1 42.7 16. Dist. ulnae 10 2 5.0 1.2 43.9 17. Prox. radii 24 2 12.0 2.8 46.7 18. Radial shafts 32 2 16.0 3.7 50.4 19. Dist. radii 20 2 10.0 2.3 52.7 20. Carpals 131 16 8.2 1.9 54.6 21. Metacarpals 63 10 6.3 1.4 56.0 22. Hand phalanges 325 28 11.6 2.7 58.7 23. Innominate bones 31 2 15.5 3.6 62.3 24. Prox. femora 35 2 17.5 4.1 66.4 25. Femoral shafts 36 2 18.0 4.2 70.6 26. Dist. femora 22 2 11.0 2.6 73.2 27. Patellae 16 2 8.0 1.9 75.1 28. Prox. tibiae 18 2 9.0 2.1 77.2 29. Tibial shafts 35 2 17.5 4.1 81.3 30. Dist. tibiae 19 2 9.5 2.2 83.5 31. Prox. fibulae 10 2 5.0 1.2 84.7 32. Fibular shafts 20 2 10.0 2.3 87.0 33. Dist. fibulae 19 2 9.5 2.2 89.2 34. Tali 24 2 12.0 2.8 92.0 35. Calcanei 26 2 13.0 3.0 95.0 36. Ant. tarsals 84 10 8.4 1.9 96.9 37. Metatarsals 51 10 5.1 1.2 98.1 38. Foot phalanges 230 28 8.2 1.9 100.0 TOTAL 2369 233 431.2 100 2048.9 MNAU = minimum number of anatomical units = number of bones or bone portions preserved in a sample divided by number of that bone or bone portion in a complete skeleton. Figure S1. Cumulative percentage of the minimum number of anatomical units (MNAU), i.e., number of bones or bone portions preserved in a sample divided by number of that bone or bone portion in a complete skeleton. 1. Neurocrania; 2. Mandibles; 3. Dentition;4. Cerv. vertebrae; 5 Thor. vertebrae; 6. Lumb. vertebrae; 7. Sacra; 8. Ribs; 9. Claviculae; 10. Scapulae; 11. Prox. humeri; 12. Humeral shafts; 13. Dist. humeri; 14. Prox. ulnae; 15. Ulnar shafts; 16. Dist. ulnae; 17. Prox. radii; 18. Radial shafts; 19. Dist. radii; 20. Carpals; 21. Metacarpals; 22. Hand phalanges; 23. Innominate bones; 24. Prox. femora; 25. Femoral shafts; 26. Dist. femora; 27. Patellae; 28. Prox. tibiae; 29. Tibial shafts; 30. Dist. tibiae; 31. Prox. fibulae; 32. Fibular shafts; 33. Dist. fibulae; 34. Tali; 35. Calcanei; 36. Ant. tarsals; 37. Metatarsals; 38. Foot phalanges. 2- GENERAL BODY SIZE AND SHAPE, INTRAPOPULATION VARIATION AND ENCEPHALIZATION 2.1-STATURE Systematic work at the Sima de los Huesos has allowed us to reconstruct 27 complete long bones. Despite the methodological difficulties involved in the estimation of stature in fossil humans (i.e. bone type, body proportions, sex assignment, statistical techniques, etc.) and the lack of consensus on a valid method broadly applicable in all cases, we have used the multiracial and combined-sex formulae proposed by Sjøvold (1) to estimate the mean stature of the SH hominins based on 24 complete long bones (excluding the fibula) (2). Although the differences between the SH and Neandertal samples are not significant, it is likely that the SH hominins were somewhat taller on average than the Neandertals since the SH hominins had, on average, longer limb bones than the Neandertals (2).