Leakey, L. B. New fossil skull from Olduvai. Nature 184, 491–493 (1959).
Leakey, L. B., Tobias, P. V. & Napier, J. R. A new species of the genus Homo from the Olduvai Gorge. Nature 202, 7–9 (1964).
Tocheri, M. W., Orr, C. M., Jacofsky, M. C. & Marzke, M. W. The evolutionary history of the hominin hand since the last common ancestor of Pan and Homo. J. Anat. 212, 544–562 (2008).
Plummer, T. W. et al. Expanded geographic distribution and dietary strategies of the earliest Oldowan hominins and Paranthropus. Science 379, 561–566 (2023).
Wood, B. A. & Richmond, B. G. Human evolution: taxonomy and paleobiology. J. Anat. 196, 19–60 (2000).
Panger, M. A., Brooks, A. S., Richmond, B. G. & Wood, B. Older than the Oldowan? Rethinking the emergence of hominin tool use. Evol. Anthropol. 11, 235–245 (2002).
Tocheri, M. W. et al. The primitive wrist of Homo floresiensis and its implications for hominin evolution. Science 317, 1743–1745 (2007).
Susman, R. L. Hand of Paranthropus robustus from Member 1, Swartkrans: fossil evidence for tool behavior. Science 240, 781–784 (1988).
Susman, R. L. New hominid fossils from the Swartkrans formation (1979–1986 excavations): postcranial specimens. Am. J. Phys. Anthropol. 79, 451–474 (1989).
Susman, R. L. Fossil evidence for early hominid tool use. Science 265, 1570–1573 (1994).
Susman, R. L. Response: thumbs, tools, and early humans. Science 268, 589 (1995).
Trinkaus, E. & Long, J. C. Species attribution of the Swartkrans Member 1 first metacarpals: SK 84 and SKX 5020. Am. J. Phys. Anthropol. 83, 419–424 (1990).
Grine, F. E., Mongle, C. S., Fleagle, J. G. & Hammond, A. S. The taxonomic attribution of African hominin postcrania from the Miocene through the Pleistocene: associations and assumptions. J. Hum. Evol. 173, 103255 (2022).
Moyà-Solà, S., Kohler, M., Alba, D. M. & Almécija, S. Taxonomic attribution of the Olduvai hominid 7 manual remains and the functional interpretation of hand morphology in robust australopithecines. Folia Primatol. 79, 215–250 (2008).
Hatala, K. G. et al. Footprint evidence for locomotor diversity and shared habitats among early Pleistocene hominins. Science 386, 1004–1010 (2024).
Brown, F. H. & Feibel, C. S. Revision of lithostratigraphic nomenclature in the Koobi Fora region, Kenya. J. Geol. Soc. 143, 297–310 (1986).
Leakey, M. G. et al. New fossils from Koobi Fora in northern Kenya confirm taxonomic diversity in early Homo. Nature 488, 201–204 (2012).
Neubauer, S. et al. Reconstruction, endocranial form and taxonomic affinity of the early Homo calvaria KNM-ER 42700. J. Hum. Evol. 121, 25–39 (2018).
Wood, B. A. Hominid Cranial Remains. Koobi Fora Research Project Vol. 4 (Oxford Univ. Press, 1991).
Martin, L. B., Boyde, A. & Grine, F. E. Enamel structure in primates: a review of scanning electron microscope studies. Scanning Microsc. 2, 1503–1526 (1988).
Mongle, C. S., Strait, D. S. & Grine, F. E. Expanded character sampling underscores phylogenetic stability of Ardipithecus ramidus as a basal hominin. J. Hum. Evol. 131, 28–39 (2019).
Mongle, C. S., Strait, D. S. & Grine, F. E. An updated analysis of hominin phylogeny with an emphasis on re-evaluating the phylogenetic relationships of Australopithecus sediba. J. Hum. Evol. 175, 103311 (2023).
Lockey, A. L., Alemseged, Z., Hublin, J.-J. & Skinner, M. M. Maxillary molar enamel thickness of Plio-Pleistocene hominins. J. Hum. Evol. 142, 102731 (2020).
Skinner, M. M., Alemseged, Z., Gaunitz, C. & Hublin, J.-J. Enamel thickness trends in Plio-Pleistocene hominin mandibular molars. J. Hum. Evol. 85, 35–45 (2015).
Tobias, P. V. The Cranium and Maxillary Dentition of Australopithecus (Zinjanthropus) Boisei. Vol. 2, Olduai Gorge (Cambridge Univ. Press, 1967).
Rak, Y. Y. The Australopithecine Face (Academic, 1983).
Strait, D. S., Grine, F. E. & Moniz, M. A. A reappraisal of early hominid phylogeny. J. Hum. Evol. 32, 17–82 (1997).
Wood, B. A. & Engleman, C. A. Analysis of the dental morphology of Plio-Pleistocene hominids. V. Maxillary postcanine tooth morphology. J. Anat. 161, 1–35 (1988).
Kimbel, W. H., White, T. D. & Johanson, D. C. Cranial morphology of Australopithecus afarensis: a comparative study based on a composite reconstruction of the adult skull. Am. J. Phys. Anthropol. 64, 337–388 (1984).
Almécija, S., Moyà-Solà, S. & Alba, D. M. Early origin for human-like precision grasping: a comparative study of pollical distal phalanges in fossil hominins. PLoS ONE 5, e11727 (2010).
Richmond, B. G. et al. The upper limb of Paranthropus boisei from Ileret, Kenya. J. Hum. Evol. 141, 102727 (2020).
Susman, R. L. Comparative and functional morphology of hominoid fingers. Am. J. Phys. Anthropol. 50, 215–236 (1979).
Bucchi, A., Luengo, J., Del Bove, A. & Lorenzo, C. Insertion sites in manual proximal phalanges of African apes and modern humans. Am. J. Phys. Anthropol. 173, 556–567 (2020).
Syeda, S. M., Tsegai, Z. J., Cazenave, M., Skinner, M. M. & Kivell, T. L. Cortical bone architecture of hominid intermediate phalanges reveals functional signals of locomotion and manipulation. Am. J. Biol. Anthropol. 184, e24902 (2024).
Syeda, S. M., Tsegai, Z. J., Cazenave, M., Skinner, M. M. & Kivell, T. L. Cortical bone distribution of the proximal phalanges in great apes: implications for reconstructing manual behaviours. J. Anat. 243, 707–728 (2023).
Susman, R. L. & Stern, J. T. Functional morphology of Homo habilis. Science 217, 931–934 (1982).
Domínguez-Rodrigo, M. et al. First partial skeleton of a 1.34-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania. PLoS ONE 8, e80347 (2013).
Lague, M. R. et al. Humeral anatomy of the KNM-ER 47000 upper limb skeleton from Ileret, Kenya: implications for taxonomic identification. J. Hum. Evol. 126, 24–38 (2019).
Baab, K. L., Copes, L. E., Ward, D. L., Wells, N. & Grine, F. E. Using modern human cortical bone distribution to test the systemic robusticity hypothesis. J. Hum. Evol. 119, 64–82 (2018).
Latimer, B. & Lovejoy, C. O. Metatarsophalangeal joints of Australopithecus afarensis. Am. J. Phys. Anthropol. 83, 13–23 (1990).
Fernández, P. J. et al. Evolution and function of the hominin forefoot. Proc. Natl Acad. Sci. USA 131, 8746–8751 (2018).
Trinkaus, E. & Patel, B. A. An Early Pleistocene human pedal phalanx from Swartkrans, SKX 16699, and the antiquity of the human lateral forefoot. C. R. Palevol 15, 978–987 (2016).
Venkadesan, M. et al. Stiffness of the human foot and evolution of the transverse arch. Nature 579, 97–100 (2020).
DeSilva, J., McNutt, E., Benoit, J. & Zipfel, B. One small step: a review of Plio-Pleistocene hominin foot evolution. Am. J. Phys. Anthropol. 168, 63–140 (2019).
Susman, R. L. & Brain, T. M. New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus. Am. J. Phys. Anthropol. 77, 7–15 (1988).
Venkataraman, V. V., Kraft, T. S. & Dominy, N. J. Tree climbing and human evolution. Proc. Natl Acad. Sci. USA 110, 1237–1242 (2013).
Harmand, S. et al. 3.3-million-year-old stone tools from Lomekwi 3, West Turkana, Kenya. Nature 521, 310–315 (2015).
Rein, T. R. The correspondence between proximal phalanx morphology and locomotion: implications for inferring the locomotor behavior of fossil catarrhines. Am. J. Phys. Anthropol. 146, 435–445 (2011).
Neufuss, J., Robbins, M. M., Baeumer, J., Humle, T. & Kivell, T. L. Gait characteristics of vertical climbing in mountain gorillas and chimpanzees. J. Zool. 306, 129–138 (2018).
Ruff, C. B. & Wood, B. A. The estimation and evolution of hominin body mass. Evol. Anthropol. 32, 223–237 (2023).
Smith, R. J. & Jungers, W. L. Body mass in comparative primatology. J. Hum. Evol. 32, 523–559 (1997).
Marzke, M. in Stone Knapping: The Necessary Conditions for a Uniquely Hominin Behaviour (eds Roux V. & Brill, B.) 243–255 (McDonald Institute Monograph, 2006).
Neufuss, J., Robbins, M. M., Baeumer, J., Humle, T. & Kivell, T. L. Manual skills for food processing by mountain gorillas (Gorilla beringei beringei) in Bwindi Impenetrable National Park, Uganda. Biol. J. Linn. Soc. 127, 543–562 (2019).
Sponheimer, M., Daegling, D. J., Ungar, P. S., Bobe, R. & Paine, O. C. C. Problems with Paranthropus. Quat. Int. 650, 40–51 (2023).
Wood, B. A. Olduvai Bed I post-cranial fossils: a reassessment. J. Hum. Evol. 3, 373–378 (1974).
Kivell, T. L. Evidence in hand: recent discoveries and the early evolution of human manual manipulation. Philos. Trans. R. Soc. Lond. B Biol. Sci. 370, 20150105 (2015).
Marzke, M. W. et al. Comparative 3D quantitative analyses of trapeziometacarpal joint surface curvatures among living catarrhines and fossil hominins. Am. J. Phys. Anthropol. 141, 38–51 (2010).
Lorenzo, C., Arsuaga, J. L. & Carretero, J. M. Hand and foot remains from the Gran Dolina Early Pleistocene site (Sierra de Atapuerca, Spain). J. Hum. Evol. 37, 501–522 (1999).
Ward, C. V., Tocheri, M. W., Plavcan, J. M., Brown, F. H. & Manthi, F. K. Early Pleistocene third metacarpal from Kenya and the evolution of modern human-like hand morphology. Proc. Natl Acad. Sci. USA 111, 121–124 (2014).
O’Brien, K., Hebdon, N. & Faith, J. T. Paleoecological evidence for environmental specialization in Paranthropus boisei compared to early Homo. J. Hum. Evol. 177, 103325 (2023).
du Plessis, A., le Roux, S. G. & Guelpa, A. The CT scanner facility at Stellenbosch University: an open access X-ray computed tomography laboratory. Nucl. Instrum. Methods Phys. Res. B 384, 42–49 (2016).
Robinson, J. T. Dentition of the Australopithecinae Vol. Memoir No. 9 (Transvaal Museum, 1956).
Grine, F. E., Martin, L. B. in Evolutionary History of the ‘Robust’ Australopithecines (ed. Grine, F. E.) 3–42 (Aldine de Gruyter, 1988).
Stern, J. T. Jr, Jungers, W. L. & Susman, R. L. Quantifying phalangeal curvature: an empirical comparison of alternative methods. Am. J. Phys. Anthropol. 97, 1–10 (1995).
Domínguez-Rodrigo, M. et al. Earliest modern human-like hand bone from a new >1.84-million-year-old site at Olduvai in Tanzania. Nat. Commun. 6, 7987 (2015).
Tocheri, M. W. et al. Functional capabilities of modern and fossil hominid hands: three-dimensional analysis of trapezia. Am. J. Phys. Anthropol. 122, 101–112 (2003).
Tocheri, M. W., Razdan, A., Williams, R. C. & Marzke, M. W. A 3D quantitative comparison of trapezium and trapezoid relative articular and nonarticular surface areas in modern humans and great apes. J. Hum. Evol. 49, 570–586 (2005).
Schlager, S. in Statistical Shape and Deformation Analysis 217–256 (Elsevier, 2017).
Gunz, P., Mitteroecker, P. & Bookstein, F. L. in Developments in Primatology: Progress and Prospects (eds Zheng, G. et al.) 73–98 (Kluwer Academic–Plenum, 2006).
Gunz, P. & Mitteroecker, P. Semilandmarks: a method for quantifying curves and surfaces. Hystrix 24, 103–109 (2013).
Wickham, H. Ggplot2 (Springer International, 2016).
Adams, D. C. & Otárola-Castillo, E. geomorph: an rpackage for the collection and analysis of geometric morphometric shape data. Methods Ecol. Evol. 4, 393–399 (2013).
Baken, E. K., Collyer, M. L., Kaliontzopoulou, A. & Adams, D. C. geomorph v4.0 and gmShiny: enhanced analytics and a new graphical interface for a comprehensive morphometric experience. Methods Ecol. Evol. 12, 2355–2363 (2021).
Orr, C. M. et al. New wrist bones of Homo floresiensis from Liang Bua (Flores, Indonesia). J. Hum. Evol. 64, 109–129 (2013).
Mongle, C. S., Wallace, I. J. & Grine, F. E. Cross-sectional structural variation relative to midshaft along hominine diaphyses. I. The forelimb. Am. J. Phys. Anthropol. 158, 386–397 (2015).
Doube, M. BoneJ: free and extensible bone image analysis in ImageJ. Bone 47, 1076–1079 (2010).
Harcourt-Smith, W. E. H. et al. The foot of Homo naledi. Nat. Commun. 6, 8432 (2015).
Desilva, J. M., McNutt, E., Zipfel, B., Ward, C. V. & Kimbel, W. H. Associated Australopithecus afarensis second and third metatarsals (A.L. 333-133) from Hadar, Ethiopia. J. Hum. Evol. 146, 102848 (2020).
Susman, R. L., Stern, J. T. & Jungers, W. L. Arboreality and bipedality in the Hadar hominids. Folia Primatol. 43, 113–156 (1984).
Goodenberger, K. E. et al. Functional morphology of the hallucal metatarsal with implications for inferring grasping ability in extinct primates. Am. J. Phys. Anthropol. 156, 327–348 (2015).
Pontzer, H. et al. Locomotor anatomy and biomechanics of the Dmanisi hominins. J. Hum. Evol. 58, 492–504 (2010).
DeSilva, J. M., Proctor, D. J. & Zipfel, B. A complete second metatarsal (StW 89) from Sterkfontein Member 4, South Africa. J. Hum. Evol. 63, 487–496 (2012).
Drapeau, M. S. M. & Harmon, E. H. Metatarsal torsion in monkeys, apes, humans and australopiths. J. Hum. Evol. 64, 93–108 (2013).
Maddison, W. P. & Maddison, D. R. Mesquite: a modular system for evolutionary analysis. Mesquite https://www.mesquiteproject.org (2003).
Prang, T. C. Custom MATLAB script for metatarsal torsion calculation. Figshare https://doi.org/10.6084/m9.figshare.29991643 (2025).