Published July 8, 2026
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New associated postcranial remains from the Shungura Formation (Lower Omo Valley, Ethiopia) provide insights on behaviors and ecology of early Homo
Authors/Creators
- 1. Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM), Université de Poitiers and CNRS, 6 rue Michel Brunet, 86000 Poitiers (France) & Department of Anatomy, Midwestern University, Glendale, 19555 N 59th Ave, Glendale, AZ 85308 (United States)
- 2. Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM), Université de Poitiers & CNRS, 6 rue Michel Brunet, 86000 Poitiers (France)
- 3. Division of Integrative Anatomical Sciences and Department of Radiology, Keck School of Medicine, University of Southern California, 1975 Zonal Ave, Los Angeles, CA 90033 (United States) & Department of Geology and Paleontology, Georgian National Museum, 3 Shota Rustaveli Ave, Tbilisi 0105 (Georgia)
- 4. Aix Marseille Université, CNRS, EFS, ADES, 58 Boulevard Charles Livon, 13007 Marseille (France)
- 5. Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM), Université de Poitiers and CNRS, 6 rue Michel Brunet, 86000 Poitiers (France) & Kyoto University, Graduate School of Science, Laboratory of Physical Anthropology, Yoshidahonmachi, Sakyo Ward, Kyoto 606-8501 (Japan) & Histoire Naturelle des Humanités Préhistoriques (HNHP), MNHN/CNRS/UPVD, 17 Place du Trocadéro et du 11 Novembre, 75116 Paris (France)
- 6. Institut PPrime CNRS, École Normale Supérieure de Mécanique et Aérotechnique (ENSMA), Université de Poitiers, 2 Boulevard des Frères Lumière, 86360 Chasseneuil-du-Poitou (France)
- 7. Institute of Human Origin, School of Human Evolution and Social Change, Arizona State University, 900 S Cady Mall, Tempe, AZ 85281 (United States) & Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM), Université de Poitiers and CNRS, 6 rue Michel Brunet, 86000 Poitiers (France) & French Center for Ethiopian Studies, CNRS, Ministry of Europe and Foreign Affairs, Addis Ababa, PO BOX 5554 (Ethiopia)
- 8. French Center for Ethiopian Studies, CNRS, Ministry of Europe and Foreign Affairs, Addis Ababa, PO BOX 5554 (Ethiopia) & Laboratoire Paléontologie Evolution Paléoécosystèmes Paléoprimatologie (PALEVOPRIM), CNRS and Université de Poitiers, 6 rue Michel Brunet, 86000 Poitiers (France)
Description
Blasi-Toccacceli, Alicia, Daver, Guillaume, Brusse, Tiphaine, Jashashvili, Tea, Voisin, Jean-Luc, Pallas, Laurent, Domalain, Mathieu, Surault, Jérôme, Redae, Blade Engda, Boisserie, Jean-Renaud (2026): New associated postcranial remains from the Shungura Formation (Lower Omo Valley, Ethiopia) provide insights on behaviors and ecology of early Homo. Comptes Rendus Palevol 25 (12): 207-247, DOI: 10.5852/cr-palevol2026v25a12, URL: https://doi.org/10.5852/cr-palevol2026v25a12
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- URN
- urn:lsid:zoobank.org:pub:urn:lsid:zoobank.org:pub:6542A9B5-DD38-4729-98CB-81C6A167B2B7
- LSID
- urn:lsid:plazi.org:pub:FFE30B2DFFFF0C198E64C85AFFE5FFEA
References
- ADAMS D. C., COLLYER M. L., KALIONTZOPOULOU A. & BAKEN E. 2022. - Geomorph: Software for geometric morphometric analyses. R package version 4.0.4.
- ALEMSEGED Z. 2003. - An integerated approach to taphonomy and faunal change in the Shungeura Formation (Ethiopia) and its implication for hominid evolution. Journal of Human Evolution 44 (4): 451-478.https://doi.org/10.1016/S0047-2484(03)00012-5
- ALEMSEGED Z., SPOOR F., KIMBEL W. H., BOBE R., GERAADS D., REED D. & WYNN J. G. 2006. - A juvenile early hominin skeleton from Dikika, Ethiopia. Nature 443 (7109): 296-301. https://doi.org/10.1038/nature05047
- ANDRESEN P. R., BOOKSTEIN F. L., COURADSEN K., ERSBOLL B. K., MARSH J. L. & KREIBORG S. 2000. - Surface-bounded growth modeling applied to human mandibles. IEEE Transactions on Medical Imaging 19 (11): 1053-1063. https://doi. org/10.1109/42.896780
- ARIAS-MARTORELL J. 2019. - The morphology and evolutionary history of the glenohumeral joint of hominoids: a review.Ecology and Evolution 9 (1): 703-722.https://doi.org/10.1002/ece3.4392
- ARIAS-MARTORELL J., POTAU J. M., BELLO-HELLEGOUARCH G. & PEREZ-PEREZ A. 2015. - Like father, like son: assessment of the morphological affinities of A.L. 288-1 (A. afarensis), Sts 7 (A. africanus) and Omo 119-73-2718 (Australopithecus sp.) through a three-dimensional shape analysis of the shoulder joint, in BONDIOLI L. (ed.). PLOS ONE 10 (2): e0117408.https://doi. org/10.1371/journal.pone.0117408
- ASFAW B., BEYENE Y., SUWA G., WALTER R. C., WHITE T. D., WOLDEGABRIEL G. & YEMANE T. 1992. - The earliest Acheulean from Konso-Gardula.Nature 360 (6406): 732-735.https:// doi.org/10.1038/360732a0
- ASHTON E. H. & OXNARD C. E. 1964. - Functional adaptation in the primate shoulder girdle. Proceedings of the Zoological Society of London 142 (1): 49-66.https://doi.org/10.1111/j.1469-7998.1964. tb05153.x
- ASHTON E. H., HEALY M. J. R., OXNARD C. E. & SPENCE T. F. 1965. - The combination of locomotor features of the primate shoulder girdle by canonical analysis. Proceedings of the Zoological Society of London 147 (4): 406-429. https://doi. org/10.1111/j.1469-7998.1966.tb02908.x
- BALTER V., BRAGA J., TELOUK P. & THACKERAY J. F. 2012. - Evidence for dietary change but not landscape use in South African early hominins. Nature 489 (7417): 558-560. https:// doi.org/10.1038/nature11349
- BAR-YOSEF O. & GOREN-INBAR N. (eds) 1993. - The lithic assemblages of 'Ubeidiya: a lower palaeolithic site in the Jordan Valley. Institute of Archaeology, Hebrew University of Jerusalem, Qedem, Jerusalem, 280 p.
- BOISSERIE J.-R., GUY F., DELAGNES A., HLUKSO L. J., BIBI F., BEYENE Y. & GUILLEMOT C. 2008. - New palaeoanthropological research in the Plio-Pleistocene Omo Group, Lower Omo Valley, SNNPR (Southern Nations, Nationalities and People Regions), Ethiopia. Comptes Rendus Palevol 7 (7): 429-439. https://doi. org/10.1016/j.crpv.2008.07.010
- BOOKSTEIN F. L. 1991. - Morphometric tools for landmark data: geometry and biology [Orange book].Cambridge New York, Cambridge University Press.
- BRAGA J., THACKERAY J. F. & ZIPFEL B. 2022. - The Kromdraai early hominin-bearing site. A review of recent findings. L'Anthropologie 126 (4): 103054. https://doi.org/10.1016/j. anthro.2022.103054
- BRAMBLE D. M. & LIEBERMAN D. E. 2004. - Endurance running and the evolution of Homo. Nature 432 (7015): 345-352.https:// doi.org/10.1038/nature03052
- CAMPISANO C. J. & FEIBEL C. S. 2008. - Depositional environments and stratigraphic summary of the Pliocene Hadar Formation at Hadar, Afar Depression, Ethiopia, in QUADE J. & WYNN J. G. (eds), The Geology of Early Humans in the Horn of Africa. Geological Society of America, Boulder, Colorado:179-201. https:// doi.org/10.1130/2008.2446(08)
- CARLSON K. J., GREEN D. J., JASHASHVILI T., PICKERING T. R., HEATON J. L., BEAUDET A., STRATFORD D., CROMPTON R., KUMAN K., BRUXELLES L. & CLARKE R. J. 2021. - The pectoral girdle of StW 573 ('Little Foot') and its implications for shoulder evolution in the Hominina. Journal of Human Evolution 158: 102983. https://doi.org/10.1016/j.jhevol.2021.102983
- CARRETERO J. M., ARSUAGA J. L. & LORENZO C. 1997. - Clavicles, scapulae and humeri from the Sima de los Huesos site (Sierra de Atapuerca, Spain). Journal of Human Evolution 33 (2-3): 357- 408. https://doi.org/10.1006/jhev.1997.0128
- CERLING T. E., MANTHI F. K., MBUA E. N., LEAKEY L. N., LEAKEY M. G., LEAKEY R. E., BROWN F. H., GRINE F. E., HART J. A., KALEM P., ROCHE H., UNO K. T. & WOOD B. A. 2013. - Stable isotope-based diet reconstructions ofTurkana Basin hominins. Proceedings of the National Academy of Sciences 110 (26): 10501-10506. https://doi.org/10.1073/pnas.1222568110
- CHURCHILL S. E., HOLLIDAY T. W., CARLSON K. J., JASHASHVILI T., MACIAS M. E., MATHEWS S., SPARLING T. L., SCHMID P., DE RUITER D. J. & BERGER L. R. 2013. - The Upper Limb of Australopithecus sediba. Science 340 (6129).https://doi.org/10.1126/ science.1233477
- CHURCHILL S. E., GREEN D. J., FEUERRIEGEL E. M., MACIAS M. E., MATHEWS S., CARLSON K. J., SCHMID P. & BERGER L. 2018. - The Shoulder, Arm, and Forearm of Australopithecus sediba. PaleoAnthropology: 234-281
- CIGNONI P., CALLIERI M., CORSINI M., DELLEPIANE M., GANOVELLI F. & RANZUGLIA G. 2008. - MeshLab: an Open-Source Mesh Processing Tool, in SCARANO V., DE CHIARA R. & ERRA U. (eds), Eurographics Italian Chapter Conference.The Eurographics Association: 129-136.
- COPPENS Y., CHAVAILLON J. & BEDEN M. 1973. - Resultats de la nouvelle mission de l'Omo (campagne 1972). Decouverte de restes d'Hominides et d'une industrie sur eclats. Comptes Rendus de l'Academie des Sciences, Paris 276: 161-164.
- CORRUCCINI R.S. & CIOCHON R.L. 1976. - Morphometric affinities of the human shoulder. American Journal of Physical Anthropology 45 (1): 19-37. https://doi.org/10.1002/ ajpa.1330450104
- CUNNINGHAM C. B., SCHEUER L. & BLACK S. 2016. - The vertebral colomn, in CUNNINGHAM C., SCHEUER L. & BLACK S. (eds), Developmental Juvenile Osteology. Second Edition. Chap. 7. Academic Press, London: 177-223.
- DAVER G., BERILLON G., JACQUIER C., ARDAGNA Y., YADETA M., MAURIN T., SOURON A., BLONDEL C., COPPENS Y. & BOISSERIE J.-R. 2018. - New hominin postcranial remains from locality OMO 323, Shungura Formation, Lower Omo Valley, southwestern Ethiopia.Journal of Human Evolution 122: 23-32. https://doi.org/10.1016/j.jhevol.2018.03.011
- DESILVA J. M. & GRABOWSKI M. 2020. - Femur, in ZIPFEL B., RICHMOND B. G. & WARD C. V. (eds), Hominin Postcranial Remains from Sterkfontein, South Africa, 1936-1995. Ch. 12. Oxford University Press, New-York: 210-229.
- DI VINCENZO F., CHURCHILL S. E. & MANZI G. 2012. - The Vindija Neanderthal scapular glenoid fossa: comparative shape analysis suggests evo-devo changes among Neanderthals.Journal of Human Evolution 62 (2): 274-285.https://doi.org/10.1016/j. jhevol.2011.11.010
- DIRKS P. H. G. M., KIBII J. M., KUHN B. F., STEININGER C., CHURCHILL S. E., KRAMERS J. D., PICKERING R., FARBER D. L., MERIAUX A.-S., HERRIES A. I. R., KING G. C. P. & BERGER L. R. 2010. - Geological Setting and Age of Australopithecus sediba from Southern Africa. Science 328 (5975): 205-208. https://doi. org/10.1126/science.1184950
- DIRKS P. H., ROBERTS E. M., HILBERT-WOLF H., KRAMERS J. D., HAWKS J., DOSSETO A., DUVAL M., ELLIOTT M., EVANS M., GRUN R., HELLSTROM J., HERRIES A. I., JOANNES-BOYAU R., MAKHUBELA T. V., PLACZEK C. J., ROBBINS J., SPANDLER C., WIERSMA J., WOODHEAD J. & BERGER L. R. 2017. - The age of Homo naledi and associated sediments in the Rising Star Cave, South Africa. eLife 6: e24231. https://doi.org/10.7554/ eLife.24231
- DOMINGUEZ-RODRIGO M., PICKERING T. R., BAQUEDANO E., MABULLA A., MARK D. F., MUSIBA C., BUNN H. T., URIBELARREA D., SMITH V., DIEZ-MARTIN F., PEREZ-GONZALEZ A., SANCHEZ P., SANTONJA M., BARBONI D., GIDNA A., ASHLEY G., YRAVEDRA J., HEATON J. L. & ARRIAZA M. C. 2013. - First partial skeleton of a 1.34-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania, in CURNOE D. (ed.). PLoS ONE 8 (12): e80347. https://doi.org/10.1371/journal. pone.0080347
- DRAPEAU M. S. M. 2008. - Articular morphology of the proximal ulna in extant and fossil hominoids and hominins. Journal of Human Evolution 55 (1): 86-102. https://doi.org/10.1016/j. jhevol.2008.01.005
- DRAPEAU M. S. M., WARD C. V., KIMBEL W. H., JOHANSON D. C. & RAK Y. 2005. - Associated cranial and forelimb remains attributed to Australopithecus afarensis from Hadar, Ethiopia. Journal of Human Evolution 48 (6): 593-642.https://doi.org/10.1016/j. jhevol.2005.02.005
- DWIGHT T. 1905. - The size of the articular surfaces of the long bones as characteristic of sex; an anthropological study. American Journal of Anatomy 4 (1): 19-31. https://doi.org/10.1002/ aja.1000040103
- FEIBEL C. S., BROWN F. H. & MCDOUGALL I. 1989. - Stratigraphic context of fossil hominids from the Omo group deposits: Northern Turkana Basin, Kenya and Ethiopia. American Journal of Physical Anthropology 78 (4): 595-622.https://doi.org/10.1002/ ajpa.1330780412
- FEUERRIEGEL E. M., GREEN D. J., WALKER C. S., SCHMID P., HAWKS J., BERGER L. R. & CHURCHILL S. E. 2017. - The upper limb of Homo naledi. Journal of Human Evolution 104: 155-173. https://doi.org/10.1016/j.jhevol.2016.09.013
- FEUERRIEGEL E. M., VOISIN J.-L., CHURCHILL S. E., HAEUSLER M., MATHEWS S. & SCHMID P. 2019. - Upper limb fossils of Homo naledi from the Lesedi Chamber, rising star system, South Africa. Paleoanthropology 2019: 311-349
- FIFER F. C. 1987. - The adoption of bipedalism by the hominids: a new hypothesis. Human Evolution 2 (2): 135-147. https://doi. org/10.1007/BF02436402
- GAGEY O. 1985. - Etude de l'elevation du membre superieur. Role des ligaments articulaires et des muscles flechisseurs de l'articulation scapulo-humerale. Laboratoire d'Anatomie de la Faculte de Medecine de Paris.
- GEORGIOU L., DUNMORE C. J., BARDO A., BUCK L. T., HUBLIN J.-J., PAHR D. H., STRATFORD D., SYNEK A., KIVELL T. L. & SKINNER M. M. 2020. - Evidence for habitual climbing in a Pleistocene hominin in South Africa. Proceedings of the National Academy of Sciences 117 (15): 8416-8423. https://doi. org/10.1073/pnas.1914481117
- GIBBON R. J., PICKERING T. R., SUTTON M. B., HEATON J. L., KUMAN K., CLARKE R. J., BRAIN C. K. & GRANGER D. E. 2014. - Cosmogenic nuclide burial dating of hominin-bearing Pleistocene cave deposits at Swartkrans, South Africa. Quaternary Geochronology 24: 10-15. https://doi.org/10.1016/j.quageo.2014.07.004
- GOWER J. C. 1975. - Generalized procrustes analysis.Psychometrika 40 (1): 33-51. https://doi.org/10.1007/BF02291478
- GRANGER D. E., GIBBON R. J., KUMAN K., CLARKE R. J., BRUXELLES L. & CAFFEE M. W. 2015. - New cosmogenic burial ages for Sterkfontein Member 2 Australopithecus and Member 5 Oldowan. Nature 522 (7554): 85-88. https://doi.org/10.1038/ nature14268
- GRANGER D. E., STRATFORD D., BRUXELLES L., GIBBON R. J., CLARKE R. J. & KUMAN K. 2022. - Cosmogenic nuclide dating of Australopithecus at Sterkfontein, South Africa. Proceedings of the National Academy of Sciences 119 (27): e2123516119.https:// doi.org/10.1073/pnas.2123516119
- GREEN D. J. 2013. - Ontogeny of the hominoid scapula: the influence of locomotion on morphology: ontogeny of hominoid scapular morphology and locomotion. American Journal of Physical Anthropology 152 (2): 239-260.https://doi.org/10.1002/ ajpa.22353
- GREEN D. J. & ALEMSEGED Z. 2012. - Australopithecus afarensis scapular ontogeny, function, and the role of climbing in Human Evolution. Science 338 (6106): 514-517.https://doi.org/10.1126/ science.1227123
- GREEN D. J., CHIRCHIR H., MBUA E., HARRIS J. W. K., BRAUN D. R., GRIFFIN N. L. & RICHMOND B. G. 2018. - Scapular anatomy of Paranthropus boisei from Ileret, Kenya. Journal of Human Evolution 125: 181-192.https://doi.org/10.1016/j.jhevol.2017.06.013
- HAILE-SELASSIE Y., LATIMER B. M., ALENE M., DEINO A. L., GIBERT L., MELILLO S. M., SAYLOR B. Z., SCOTT G. R. & LOVEJOY C. O. 2010. - An early Australopithecus afarensis postcranium from Woranso-Mille, Ethiopia. Proceedings of the National Academy of Sciences 107 (27): 12121-12126. https://doi.org/10.1073/ pnas.1004527107
- HAMMOND A. S., MAVUSO S. S., BIERNAT M., BRAUN D. R., JINNAH Z., KUO S., MELAKU S., WEMANYA S. N., NDIEMA E. K., PATTERSON D. B., UNO K. T. & PALCU D. V. 2021. - New hominin remains and revised context from the earliest Homo erectus locality in East Turkana, Kenya. Nature Communications 12 (1): 1-12. https://doi.org/10.1038/s41467-021-22208-x
- HARMON E. H. 2013. - Age and Sex Differences in the Locomotor Skeleton of Australopithecus, in REED K. E., FLEAGLE J. G. & LEAKEY R. E. (eds), The Paleobiology of Australopithecus. Springer (Vertebrate Paleobiology and Paleoanthropology; 18), Dordrecht: 263-272. https://doi.org/10.1007/978-94- 007-5919-0_18
- HEATON J. L., PICKERING T. R., CARLSON K. J., CROMPTON R. H., JASHASHVILI T., BEAUDET A., BRUXELLES L., KUMAN K., HEILE A. J., STRATFORD D. & CLARKE R. J. 2019. - The long limb bones of the StW 573Australopithecus Skeleton from Sterkfontein Member 2: Descriptions and Proportions.Journal of Human Evolution 133: 167-197. https://doi.org/10.1016/j. jhevol.2019.05.015
- HEINZELIN J. 1983. - The Omo Group: Archives of the International Omo Research Expedition. Musee royal de l'Afrique centrale. (Annalen; vol. 1), Tervuren, 365 p.
- HRDLICKA A. 1920. - Anthropometry. American Journal of Physical Anthropology 3 (1): 147-173. https://doi.org/10.1002/ ajpa.1330030108
- IBANEZ-GIMENO P., DE ESTEBAN-TRIVIGNO S., JORDANA X., MANYOSA J., MALGOSA A. & GALTES I. 2013. - Functional plasticity of the human humerus: shape, rigidity, and muscular entheses. American Journal of Physical Anthropology 150 (4): 609-617. https://doi.org/10.1002/ajpa.22234
- JOHANSON D. C., MASAO F. T., ECK G. G., WHITE T. D., WALTER R. C., KIMBEL W. H., ASFAW B., MANEGA P., NDESSOKIA P. & SUWA G. 1987. - New partial skeleton of Homo habilis from Olduvai Gorge, Tanzania. Nature 327: 205- 209. https://doi.org/10.1038/327205a0
- JOSS J. & HUSSON F. 2016. - missMDA: A package for handling missing values in multivariate data analysis. Journal of Statistical Software 70 (1): 1-31. https://doi.org/10.18637/jss.v070.i01
- KAGAYA M., OGIHARA N. & NAKATSUKASA M. 2010. - Is the clavicle of apes long? An investigation of clavicular length in relation to body mass and upper thoracic width. International Journal of Primatology 31 (2): 209-217.https://doi.org/10.1007/ s10764-010-9402-x
- KARAKOSTIS F. A., HAEUFLE D., ANASTOPOULOU I., MORAITIS K., HOTZ G., TOURLOUKIS V. & HARVATI K. 2021. - Biomechanics of the human thumb and the evolution of dexterity. Current Biology 31 (6): 1317-1325. https://doi.org/10.1016/j. cub.2020.12.041
- KIMBEL W. H., WALTER R. C., JOHANSON D. C., REED K. E., ARONSON J. L., ASSEFA Z., MAREAN C. W., ECK G. G., BOBE R., HOVERS E., RAK Y., VONDRA C., YEMANE T., YORK D., CHEN Y., EVENSEN N. M. & SMITH P. E. 1996. - Late Pliocene Homo and Oldowan Tools from the Hadar Formation (Kada Hadar Member), Ethiopia. Journal of Human Evolution 31 (6): 549- 561. https://doi.org/10.1006/jhev.1996.0079
- KRAHL V. E. 1947. - The torsion of the humerus: its localization, cause and duration in man. American Journal of Anatomy 80 (3): 275-319. https://doi.org/10.1002/aja.1000800302
- KUMAN K. & CLARKE R. J. 2000. - Stratigraphy, artefact industries and hominid associations for Sterkfontein, Member 5. Journal of Human Evolution 38 (6): 827-847. https://doi.org/10.1006/ jhev.1999.0392
- LAGUE M. R. 2015. - Taxonomic identification of Lower Pleistocene fossil hominins based on distal humeral diaphyseal cross-sectional shape. PeerJ 3: e1084. https://doi.org/10.7717/peerj.1084
- LAGUE M. R. 2019. - Another massive distal humerus of Paranthropus boisei from Koobi Fora, Kenya. American Journal of Physical Anthropology 168: 134-135.
- LAGUE M. R., CHIRCHIR H., GREEN D. J., MBUA E., HARRIS J. W. K., BRAUN D. R., GRIFFIN N. L. & RICHMOND B. G. 2019. - Humeral anatomy of the KNM-ER 47000 upper limb skeleton from Ileret, Kenya: Implications for taxonomic identification. Journal of Human Evolution 126: 24-38.https://doi.org/10.1016/j. jhevol.2018.06.011
- LARSON S. G. 1988. - Subscapularis function in gibbons and chimpanzees: Implications for interpretation of humeral head torsion in hominoids.American Journal of Physical Anthropology 76 (4): 449-462. https://doi.org/10.1002/ajpa.1330760405
- LARSON S. G. 1995. - New characters for the functional interpretation of primate scapulae and proximal humeri.American Journal of Physical Anthropology 98 (1): 13-35. https://doi.org/10.1002/ ajpa.1330980103
- LARSON S. G. 1996. - Estimating humeral torsion on incomplete fossil anthropoid humeri. Journal of Human Evolution 31 (3): 239-257. https://doi.org/10.1006/jhev.1996.0059
- LARSON S. G. 2007. - Evolutionary transformation of the hominin shoulder. Evolutionary Anthropology: Issues, News, and Reviews 16 (5): 172-187. https://doi.org/10.1002/evan.20149
- LARSON S. G. 2009. - Evolution of the Hominin Shoulder: Early Homo, in GRINE F. E., FLEAGLE J. G. & LEAKEY R. E. (eds), The First Humans - Origin and Early Evolution of the Genus Homo. Vertebrate Paleobiology and Paleoanthropology.Springer, Dordrecht: 65-75. https://doi.org/10.1007/978-1-4020-9980-9_7
- LARSON S. G. 2013. - Shoulder Morphology in Early Hominin Evolution, in REED K. E., FLEAGLE J. G. & LEAKEY R. E. (eds), The Paleobiology of Australopithecus. Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands, Dordrecht: 247-261. https://doi.org/10.1007/978-94-007-5919-0_17
- LARSON S. G., JUNGERS W. L., TOCHERI M. W., ORR C. M., MORWOOD M. J., SUTIKNA T., AWE R. D. & DJUBIANTONO T. 2009. - Descriptions of the upper limb skeleton of Homo floresiensis. Journal of Human Evolution 57 (5): 555-570. https:// doi.org/10.1016/j.jhevol.2008.06.007
- LEAKEY L., TOBIAS P. V. & NAPIER J. R. 1964. - A new species of the genus Homo from Olduvai Gorge. Nature 202 (4927): 7-9. https://doi.org/10.1038/202007a0
- LEAKEY M. D. 1971. - Olduvai Gorge: Volume 3, Excavations in Beds I and II, 1960-1963. Cambridge University Press, 378 p.
- LEAKEY R. E. F. & WALKER A. C. 1985. - Further hominids from the Plio-Pleistocene of Koobi Fora, Kenya. American Journal of Physical Anthropology 67 (2): 135-163. https://doi.org/10.1002/ ajpa.1330670209
- LEE E. C. S., ROACH N. T., CLOUTHIER A. L., BICKNELL R. T., BEY M. J., YOUNG N. M. & RAINBOW M. J. 2020. - Three-dimensional scapular morphology is associated with rotator cuff tears and alters the abduction moment arm of the supraspinatus. Clinical Biomechanics 78: 105091. https://doi.org/10.1016/j. clinbiomech.2020.105091
- LEE E. C. S., YOUNG N. M. & RAINBOW M. J. 2023. - A comparative approach for characterizing the relationship among morphology, range-of-motion and locomotor behaviour in the primate shoulder.Proceedings of the Royal Society B: Biological Sciences 290: 20231446. https://doi.org/10.1098/rspb.2023.1446
- LORDKIPANIDZE D., JASHASHVILI T., VEKUA A., DE LEON M. S. P., ZOLLIKOFER C. P. E., RIGHTMIRE G. P., PONTZER H., FERRING R., OMS O., TAPPEN M., BUKHSIANIDZE M., AGUSTI J., KAHLKE R., KILADZE G., MARTINEZ-NAVARRO B., MOUSKHELISHVILI A., NIORADZE M. & ROOK L. 2007. - Postcranial evidence from early Homo from Dmanisi, Georgia. Nature 449 (7160): 305- 310. https://doi.org/10.1038/nature06134
- LOVEJOY C. O., SIMPSON S. W., WHITE T. D., ASFAW B. & SUWA G. 2009. - Careful climbing in the Miocene: the forelimbs of Ardipithecus ramidus and humans are primitive.Science 326 (5949): 70-70e8. https://doi.org/10.1126/science.1175827
- LUDECKE T., KULLMER O., WACKER U., SANDROCK O., FIEBIG J., SCHRENK F. & MULCH A. 2018. - Dietary versatility of Early Pleistocene hominins. Proceedings of the National Academy of Sciences 115 (52): 13330-13335. https://doi.org/10.1073/ pnas.1809439115
- MACIAS M. E. & CHURCHILL S. E. 2015. - Functional morphology of the Neandertal scapular glenoid fossa. The Anatomical Record 298 (1): 168-179. https://doi.org/10.1002/ar.23072
- MACLEAN K. F. E. & DICKERSON C. R. 2020. - Development of a comparative chimpanzee musculoskeletal glenohumeral model: implications for human function. The Journal of Experimental Biology 223 (22): jeb225987.https://doi.org/10.1242/jeb.225987
- MARTIN J. E., TACAIL T., BRAGA J., CERLING T. E. & BALTER V. 2020. - Calcium isotopic ecology of Turkana Basin hominins. Nature Communications 11 (1): 3587. https://doi.org/10.1038/ s41467-020-17427-7
- MARTIN R. 1957. - Lehrbuch der Anthropologie in systematischer Darstellung mit besouderer Berucksichtigung der anthropologischen Methoden. Vol. 1. G. Fischer (Lehrbuch der Anthropologie in systematischer Darstellung), Stuttgart, 518 p.
- MCDOUGALL I. & BROWN F. H. 2006. - Precise 40Ar/ 39Ar geochronology for the upper Koobi Fora Formation, Turkana Basin, northern Kenya. Journal of the Geological Society 163 (1): 205- 220. https://doi.org/10.1144/0016-764904-166
- MCDOUGALL I. & BROWN F. H. 2008. - Geochronology of the pre-KBS tuff sequence, Omo Group, Turkana Basin.Journal of the Geological Society 165 (2): 549-562. https://doi. org/10.1144/0016-76492006-170
- MCDOUGALL I., BROWN F. H., VASCONCELOS P. M., COHEN B. E., THIEDE D. S. & BUCHANAN M. J. 2012. - New single crystal 40Ar/39Ar ages improve time scale for deposition of the Omo Group, Omo -Turkana Basin, East Africa. Journal of the Geological Society 169 (2): 213-226.https://doi.org/10.1144/0016-76492010-188
- MCHENRY H. M. & CORRUCCINI R. S. 1975. - Distal humerus in hominoid evolution.Folia Primatologica; International Journal of Primatology 23 (3): 117-144.
- MCHENRY L. J. 2012. - A revised stratigraphic framework for Olduvai Gorge Bed I based on tuff geochemistry, Five Decades after Zinjanthropus and Homo habilis: landscape Paleoanthropology of Plio-Pleistocene Olduvai Gorge, Tanzania. Journal of Human Evolution 63 (2): 284-299. https://doi.org/10.1016/j. jhevol.2011.04.010
- MELILLO S. M. 2016. - The Shoulder Girdle of KSD-VP-1/1, in HAILE-SELASSIE Y. & SU D. F. (eds), The Postcranial Anatomy of Australopithecus afarensis. Vertebrate Paleobiology and Paleoanthropology. Springer Netherlands, Dordrecht: 113-141. https:// doi.org/10.1007/978-94-017-7429-1_6
- MELILLO S., GUNZ P., COQUEUGNIOT H., RESKE S. & HUBLIN J.-J. 2019. - Structural effects of variation in the human clavicle. American Journal of Physical Anthropology 168 (4): 687-704. https://doi.org/10.1002/ajpa.23787
- MGELADZE A., LORDKIPANIDZE D., MONCEL M.-H., DESPRIEE J., CHAGELISHVILI R., NIORADZE M. & NIORADZE G. 2011. - Hominin occupations at the Dmanisi site, Georgia, Southern Caucasus: raw materials and technical behaviours of Europe's first hominins. Journal of Human Evolution 60 (5): 571-596. https://doi.org/10.1016/j.jhevol.2010.10.008
- MORGAN L. E., RENNE P. R., KIEFFER G., PIPERNO M., GALLOTTI R. & RAYNAL J.-P. 2012. - A chronological framework for a long and persistent archaeological record: Melka Kunture, Ethiopia. Journal of Human Evolution 62 (1): 104-115. https:// doi.org/10.1016/j.jhevol.2011.10.007
- MORREY B. F. & AN K.-N. 1983. - Articular and ligamentous contributions to the stability of the elbow joint. The American Journal of Sports Medicine 11 (5): 315-319. https://doi. org/10.1177/036354658301100506
- MURTAGH F. & LEGENDRE P. 2014. - Ward's hierarchical agglomerative clustering method: clustering criterion and agglomerative algorithm. Journal of Classification 31: 274-295. https://doi. org/10.1007/s00357-014-9161-z
- O'BRIEN K., HEBDON N. & FAITH J. T. 2023. - Paleoecological evidence for environmental specialization in Paranthropusboisei compared to early Homo. Journal of Human Evolution 177: 103325. https://doi.org/10.1016/j.jhevol.2023.103325
- PATTERSON & HOWELLS 1967. - Hominid Humeral Fragment from Early Pleistocene of Northwestern Kenya.
- PATTERSON D. B., BRAUN D. R., ALLEN K., BARR W. A., BEHRENSMEYER A. K., BIERNAT M., LEHMANN S. B., MADDOX T., MANTHI F. K., MERRITT S. R., MORRIS S. E., O'BRIEN K., REEVES J. S., WOOD B. A. & BOBE R. 2019. - Comparative isotopic evidence from East Turkana supports a dietary shift within the genus Homo. Nature Ecology & Evolution 3 (7): 1048-1056. https://doi.org/10.1038/s41559-019-0916-0
- PATTERSON D. B., DU A., FAITH J. T., ROWAN J., UNO K., BEHRENSMEYER A. K., BRAUN D. R. & WOOD B. A. 2022. - Did vegetation change drive the extinction of Paranthropus boisei? Journal of Human Evolution 173: 103154.https://doi.org/10.1016/j. jhevol.2022.103154
- PICKERING T. R., EGELAND C. P., DOMINGUEZ-RODRIGO M., BRAIN C. K. & SCHNELL A. G. 2008. - Testing the "shift in the balance of power" hypothesis at Swartkrans, South Africa: Hominid cave use and subsistence behavior in the Early Pleistocene. Journal of Anthropological Archaeology 27 (1): 30-45. https://doi.org/10.1016/j.jaa.2007.07.002
- PICKERING T. R., HEATON J. L., CLARKE R. J. & STRATFORD D. 2019. - Hominin vertebrae and upper limb bone fossils from Sterkfontein Caves, South Africa (1998-2003 excavations). American Journal of Physical Anthropology 168 (3): 459-480. https://doi.org/10.1002/ajpa.23758
- PLUMMER T. W., OLIVER J. S., FINESTONE E. M., DITCHFIELD P. W., BISHOP L. C., BLUMENTHAL S. A., LEMORINI C., CARICOLA I., BAILEY S. E., HERRIES A. I. R., PARKINSON J. A., WHITFIELD E., HERTEL F., KINYANJUI R. N., VINCENT T. H., LI Y., LOUYS J., FROST S. R., BRAUN D. R., REEVES J. S., EARLY E. D. G., ONYANGO B., LAMELA-LOPEZ R., FORREST F. L., HE H., LANE T. P., FROUIN M., NOMADE S., WILSON E. P., BARTILOL S. K., ROTICH N. K. & POTTS R. 2023. - Expanded geographic distribution and dietary strategies of the earliest Oldowan hominins and Paranthropus. Science 379 (6632): 561- 566. https://doi.org/10.1126/science.abo7452
- POBINER B. L. 2020. - The zooarchaeology and paleoecology of early hominin scavenging.Evolutionary Anthropology: Issues, News, and Reviews 29 (2): 68-82. https://doi.org/10.1002/evan.21824
- PONTZER H., ROLIAN C., RIGHTMIRE G. P., JASHASHVILI T., PONCE DE LEON M. S., LORDKIPANIDZE D. & ZOLLIKOFER C. 2010. - Locomotor anatomy and biomechanics of the Dmanisi hominins. Journal of Human Evolution 58 (6): 492-504. https:// doi.org/10.1016/j.jhevol.2010.03.006
- PRAT S., BRUGAL J.-P., ROCHE H. & TEXIER P.-J. 2003. - Nouvelles decouvertes de dents d'hominides dans le membre Kaitio de la formation de Nachukui (1,65-1,9 Ma), Ouest du lac Turkana (Kenya). Comptes Rendus Palevol 2 (8): 685-693. https://doi. org/10.1016/j.crpv.2003.07.001
- PRAT S., BRUGAL J.-P., TIERCELIN J.-J., BARRAT J.-A., BOHN M., DELAGNES A., HARMAND S., KIMEU K., KIBUNJIA M., TEXIER P.-J. & ROCHE H. 2005. - First occurrence of early Homo in the Nachukui Formation (WestTurkana, Kenya) at 2.3-2.4 Myr.Journal of Human Evolution 49 (2): 230-240.https://doi.org/10.1016/j. jhevol.2005.03.009
- PROST J. H. 1965. - A Definitional System for the Classification of Primate Locomotion. American Anthropologist 67 (5): 1198- 1214. https://doi.org/10.1525/aa.1965.67.5.02a00060
- RAVENTOS-IZARD G., POTAU J. M., CASADO A., PASTOR J. F. & ARIAS-MARTORELL J. 2023. - The morphofunctional implications of the glenoid labrum of the glenohumeral joint in hominoids. American Journal of Biological Anthropology181 (2): 195-205. https://doi.org/10.1002/ajpa.24729
- RICHMOND B. G., GREEN D. J., LAGUE M. R., CHIRCHIR H., BEHRENSMEYER A. K., BOBE R., BAMFORD M. K., GRIFFIN N. L., GUNZ P., MBUA E., MERRITT S. R., POBINER B., KIURA P., KIBUNJIA M., HARRIS J. W. K. & BRAUN D. R. 2020. - The upper limb of Paranthropus boisei from Ileret, Kenya. Journal of Human Evolution 141: 102727. https://doi.org/10.1016/j. jhevol.2019.102727
- RIGHTMIRE G. P. 2001. - Patterns of hominid evolution and dispersal in the Middle Pleistocene. Quaternary International 75 (1): 77-84. https://doi.org/10.1016/S1040- 6182(00)00079-3
- ROACH N. T. & RICHMOND B. G. 2014. - Clavicle length, throwing performance and the reconstruction of the Homo erectus shoulder. Journal of Human Evolution 80: 107-113. https://doi. org/10.1016/j.jhevol.2014.09.004
- ROACH N. T. & RICHMOND B. G. 2015. - Humeral torsion does not dictate shoulder position, but does influence throwing speed. Journal of Human Evolution 85: 206-211. https://doi. org/10.1016/j.jhevol.2015.05.011
- ROACH N. T., LIEBERMAN D. E., GILL T. J., PALMER W. E. & GILL T. J. 2012. - The effect of humeral torsion on rotational range of motion in the shoulder and throwing performance: Humeral torsion and range of motion in the shoulder. Journal of Anatomy 220 (3): 293-301. https://doi.org/10.1111/j.1469- 7580.2011.01464.x
- ROACH N. T., VENKADESAN M., RAINBOW M. J. & LIEBERMAN D. E. 2013. - Elastic energy storage in the shoulder and the evolution of high-speed throwing in Homo. Nature 498 (7455): 483-486. https://doi.org/10.1038/nature12267
- ROSE M. D. 1988. - Another look at the anthropoid elbow. Journal of Human Evolution 17 (1-2): 193-224. https://doi. org/10.1016/0047-2484(88)90054-1
- RUFF C., HOLT B. & TRINKAUS E. 2006. - Who's afraid of the big bad Wolff?: "Wolff's law" and bone functional adaptation. American Journal of Physical Anthropology 129 (4): 484-498. https://doi.org/10.1002/ajpa.20371
- RYAN T. M., CARLSON K. J., GORDON A. D., JABLONSKI N., SHAW C. N. & STOCK J. T. 2018. - Human-like hip joint loading in Australopithecus africanus and Paranthropus robustus. Journal of Human Evolution 121: 12-24.https://doi.org/10.1016/j. jhevol.2018.03.008
- SCHNEIDER C. A., RASBAND W. S. & ELICEIRI K. W. 2012. - NIH Image to ImageJ: 25 years of image analysis. Nature Methods 9 (7): 671-675. https://doi.org/10.1038/nmeth.2089
- SELBY M. S. & LOVEJOY C. O. 2017. - Evolution of the hominoid scapula and its implications for earliest hominid locomotion: Selby and Lovejoy. American Journal of Physical Anthropology 162 (4): 682-700. https://doi.org/10.1002/ajpa.23158
- SEMAW S., RENNE P., HARRIS J. W. K., FEIBEL, CRAIG S, BERNOR R. L., FESSEHA N. & MOWBRAY K. 1997. - 2.5-million-year-old stone tools from Gona, Ethiopia.Nature 385: 333-336. https://doi.org/10.1038/385333a0
- SENUT B. 1981. - L'humerus et ses articulations chez les Hominides plio-pleistocenes. CNRS Editions (Cahiers de paleontologie), Paris, 141 p.
- SHEETS H. D., KIM K. & MITCHELL C. E. 2004. - A combined landmark and outline-based approach to ontogenetic shape change in the Ordovician trilobite Triarthrus becki, in ELEWA A. M. T. (ed.), Morphometrics. Springer, Berlin, Heidelberg: 67-82. https://doi.org/10.1007/978-3-662- 08865-4_6
- SKINNER M. M., STEPHENS N. B., TSEGAI Z. J., FOOTE A. C., NGUYEN N. H., GROSS T., PAHR D. H., HUBLIN J.-J. & KIVELL T. L. 2015. - Human-like hand use in Australopithecus africanus. Science 347 (6220): 395-399.https://doi.org/10.1126/ science.1261735
- SPONHEIMER M., PASSEY B. H., DE RUITER D. J., GUATELLI-STEINBERG D., CERLING T. E. & LEE-THORP J. 2006. - Isotopic Evidence for Dietary Variability in the Early Hominin Paranthropus robustus. Science 314 (5801): 980-982. https:// doi.org/10.1126/science.1133827
- SQUYRES N. & DELEON V. B. 2015. - Clavicular curvature and locomotion in anthropoid primates: A 3D geometric morphometric analysis.American Journal of Physical Anthropology 158 (2): 257-268. https://doi.org/10.1002/ajpa.22785
- STAMMERS R. C., CARUANA M. V. & HERRIES A. I. R. 2018. - The first bone tools from Kromdraai and stone tools from Drimolen, and the place of bone tools in the South African Earlier Stone Age. Quaternary International 495: 87-101. https://doi. org/10.1016/j.quaint.2018.04.026
- STERN J. T. & SUSMAN R. L. 1983. - The locomotor anatomy of Australopithecus afarensis. American Journal of Physical Anthropology 60 (3): 279-317. https://doi.org/10.1002/ ajpa.1330600302
- SUSMAN R. L. & BRAIN T. M. 1988. - New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus. American Journal of Physical Anthropology 77 (1): 7-15. https:// doi.org/10.1002/ajpa.1330770103
- SUSMAN R. L., DE RUITER D. & BRAIN C. K. 2001. - Recently identified postcranial remains of Paranthropus and Early Homo from Swartkrans Cave, South Africa. Journal of Human Evolution 41 (6): 607-629. https://doi.org/10.1006/jhev.2001.0510
- TAYLOR C., MASAO F., NJAU J. K., SONGITA A. V. & HLUSKO L. J. 2024. - OH 89: A newly described ~1.8-million-year-old hominid clavicle from Olduvai Gorge.Peer Community Journal 4. https://doi.org/10.1101/2023.02.02.526656
- THOMPSON N. E., RUBINSTEIN D., & LARSON S. G. 2018. - Great ape thorax and shoulder configuration - An adaptation for arboreality or knuckle walking? Journal of Human Evolution 125: 15-26. https://doi.org/10.1016/j.jhevol.2018.09.005
- THRUN M. C. 2021. - Distance-based clustering challenges for unbiased benchmarking studies. Scientific Reports 11: 18988. https://doi.org/10.1038/s41598-021-98126-1
- TROYANSKAYA O., CANTOR M., SHERLOCK G., BROWN P., HASTIE T., TIBSHIRANI R., BOTSTEIN D. & ALTMAN R. B. 2001. - Missing value estimation methods for DNA microarrays. Bioinformatics 17 (6): 520-525. https://doi.org/10.1093/bioinformatics/17.6.520
- UNGAR P. S., GRINE F. E. & TEAFORD M. F. 2008. - Dental Microwear and Diet of the Plio-Pleistocene Hominin Paranthropus boisei. PLoS ONE 3 (5): 10. https://doi.org/10.1371/ journal.pone.0002044
- UNGAR P. S., GRINE F. E. & TEAFORD M. F. 2006. - Diet in Early Homo: a review of the evidence and a new model of adaptive versatility. Annual Review of Anthropology 35 (1): 209-228. https://doi.org/10.1146/annurev.anthro.35.081705.123153
- VANDERMEERSH B. 1995. - The Postcranial Remains of the Regourdou 1 Neandertal: the shoulder and arm remains. Journal of Human Evolution 28 (5): 439-476. https://doi.org/10.1006/ jhev.1995.1034
- VILLMOARE B., KIMBEL W. H., SEYOUM C., CAMPISANO C. J., DIMAGGIO E. N., ROWAN J., BRAUN D. R., ARROWSMITH J. R. & REED K. E. 2015. - Early Homo at 2.8 Ma from Ledi-Geraru, Afar, Ethiopia. Science 347 (6228): 1352-1355. https://doi. org/10.1126/science.aaa1343
- VOISIN J.-L. 2000. - L'epaule des hominides : Aspects architecturaux et fonctionnels, references particulieres a la clavicule. These de doctorat, Museum national d'histoire naturelle, Paris, 442 p.
- VOISIN J.-L. 2006. - Clavicle, a neglected bone: morphology and relation to arm movements and shoulder architecture in primates. The Anatomical Record Part A: Discoveries in Molecular, Cellular, and Evolutionary Biology 288A (9): 944-953. https:// doi.org/10.1002/ar.a.20354
- VOISIN J.-L., ROPARS M. & THOMAZEAU H. 2014. - The human acromion viewed from an evolutionary perspective.Orthopaedics & Traumatology: Surgery & Research 100 (8): S355-S360.https:// doi.org/10.1016/j.otsr.2014.09.011
- WALLACE I. J., MIDDLETON K. M., LUBLINSKY S., KELLY S. A., JUDEX S., GARLAND JR. T. & DEMES B. 2010. - Functional significance of genetic variation underlying limb bone diaphyseal structure.American Journal of Physical Anthropology 143 (1): 21-30. https://doi.org/10.1002/ajpa.21286
- WALKER A. & LEAKEY R. E. (eds) 1993. - The Nariokotome Homo erectus skeleton. Springer-Verlag, Berlin, 457 p.
- WANG W. J. & CROMPTON R. H. 2004. - The role of loadcarrying in the evolution of modern body proportions. Journal of Anatomy 204 (5): 417-430. https://doi.org/10.1111/j.0021- 8782.2004.00295.x
- WARD C. V. 2015. - Postcranial and locomotor adaptations of Hominoids, in HENKE W. & TATTERSALL I. (eds), Handbook of Paleoanthropology. Springer, Heidelberg, Berlin: 1363-1386.
- WARD J. H. 1963. - Hierarchical Grouping to Optimize an Objective Function. Journal of the American Statistical Association 58 (301): 236-244. https://doi.org/10.1080/01621459.1963.10500845
- WHITE T. D. & FOLKENS P. A. 2005. - Thorax: Sternum and Ribs, in WHITE T. D. & FOLKENS P. A. (eds), The Human Bone Manual. Ch. 10. Academic Press, Burlington, MA: 153-176.
- WOOD B. & CONSTANTINO P. 2007. - Paranthropus boisei: Fifty years of evidence and analysis.American Journal of Physical Anthropology 134 (S45): 106-132.https://doi.org/10.1002/ajpa.20732
- WOOD B. & STRAIT D. 2004. - Patterns of resource use in early Homo and Paranthropus. Journal of Human Evolution 46 (2): 119-162. https://doi.org/10.1016/j.jhevol.2003.11.004
- WYNN J. G., ALEMSEGED Z., BOBE R., GRINE F. E., NEGASH E. W. & SPONHEIMER M. 2020. - Isotopic evidence for the timing of the dietary shift toward C4 foods in eastern African Paranthropus. Proceedings of the National Academy of Sciences 117 (36): 21978- 21984. https://doi.org/10.1073/pnas.2006221117
- YEGIAN A. K., TUCKER Y., GILLINOV S. & LIEBERMAN D. E. 2021. - Shorter distal forelimbs benefit bipedal walking and running mechanics: implications for hominin forelimb evolution. American Journal of Physical Anthropology 175 (3): 589-598. https://doi.org/10.1002/ajpa.24274
- YOUNG N. M., CAPELLINI T. D., ROACH N. T. & ALEMSEGED Z. 2015. - Fossil hominin shoulders support an African ape-like last common ancestor of humans and chimpanzees. Proceedings of the National Academy of Sciences 112 (38): 11829-11834. https://doi.org/10.1073/pnas.1511220112
- ZYLSTRA M. 2000. - Functional morphology of the hominoid forelimb, implications for knuckle-walking and the origin of hominid bipedalism Thesis. PhD thesis, University of Toronto, 231 p.