Abstract
Bipedalism is a highly specialized and unusual form of primate locomotion that is found today only in modern humans. The majority of extinct taxa within the Hominini were bipedal, but the degree to which they were bipedal remains the subject of considerable debate. The significant discoveries of fossil hominin that remains in the last 40 years have resulted in this debate becoming increasingly focused on how bipedal certain fossil taxa were, rather than on the overall process. Although the early hominin fossil record remains poor, evidence points to at least two distinct adaptive shifts. First, there was a shift to habitual bipedalism, as typified by certain members of Australopithecus, but possibly including earlier genera such as Ardipithecus and Orrorin. Such taxa were bipedal, but also retained a number of significant adaptations to arboreal climbing. The second shift was to fully obligate bipedalism and coincides with the emergence of the genus Homo. By the Early Pleistocene, certain members of Homo had acquired a postcranial skeleton indicating fully humanlike striding bipedalism. The final part of this chapter reviews why bipedalism was selected for. There have been many theoretical explanations, and the most robust remain those linked to the emergence of more varied habitats. Such an environmental shift would have involved strong selection for new behavioral strategies most likely linked to the efficient procurement of food.
Access this chapter
Tax calculation will be finalised at checkout
Purchases are for personal use only
References
Aiello LC, Dean MC (1990) An introduction to human evolutionary anatomy. Academic, London
Aiello LC, Wood B, Key C, Wood C (1998) Laser scanning and palaeoanthropology: an example from Olduvai Gorge, Tanzania. In: Strasser E, Fleagle J, Rosenberger A, McHenry H (eds) Primate locomotion: recent advances. Plenum, New York, pp 223–236
Alemseged Z, Spoor F, Kimbel WH, Bobé R, Geraads D, Reed D, Wynn JG (2006) A juvenile early hominin skeleton from Dikika, Ethiopia. Nature 443:296–301
Almécija S, Tallman M, Alba D, Pina M, Moyà-Solà S, Jungers WL (2013) The femur of Orrorin tugenensis exhibits morphometric affinities with both Miocene apes and later hominins. Nat Commun 4:2888. doi:10.1038/ncomms3888
Avis V (1962) Brachiation: the crucial issue for man’s ancestry. Southwestern J Anthropol 18:119–148
Bacon A-M (1997) Presence of an obturator groove in some Cercopithecoids and small-sized Platyrrhines. Comptes Rendus de l’Academie des Sciences–Serie III, Sciences de la Vie 320(5):421–425
Bartholomew GA, Birdsell JB (1953) Ecology and the protohominids. Am Anthrop 55:481–498
Begun DR (2004) Knuckle-walking and the origin of human bipedalism. In: Meldrum DJ, Hilton CE (eds) From biped to strider: the emergence of modern human walking, running and resource transport. Kluwer Academic/Plenum, New York
Behrensmeyer AK, Reed KE (2013) Reconstructing the habitats of Australopithecus: paleoenvironments, site taphonomy and faunas. In: Reed KE, Fleagle JG, Leakey REF (eds) The paleobiology of Australopithecus. Springer, Dordrecht, pp 41–59
Bennett MR, Harris JWK, Richmond BG, Braun DR, Mbua E, Kiura P, Olago D, Kibunjia M, Omuomb C, Behrensmeyer AK, Huddart D, Gonzalez S (2009) Early Hominin foot morphology based on 1.5-million-year-old footprints from Ileret, Kenya. Science 323:1174–1201
Berger LR, Tobias PV (1996) A chimpanzee-like tibia from Sterkfontein, South Africa and its implications for the interpretation of bipedalism in Australopithecus africanus. J Hum Evol 30:343–348
Berger LR, de Ruiter DJ, Churchill SE, Schmid P, Carlsonm KJ, Dirks PHGM, Kibii JM (2010) Australopithecus sediba: a new species of Homo-like Australopith from South Africa. Science 328:195–204
Berillon G (1999) Geometric pattern of the hominoid hallucal tarsometatarsal complex. Quantifying the degree of hallux abduction in early hominids. Comptes-Rendus de l’Académie des Sciences, Paris Série IIa 328:627–633
Berillon G (2000) Le pied des hominoïdes Miocènes et des hominidés fossiles. Architecture, locomotion, évolution. CNRS Editions, Paris
Berillon G (2003) Assessing the longitudinal structure of the early hominid foot: a two-dimensional architecture analysis. Hum Evol 18(3–4):113–122
Bramble DM, Lieberman DE (2004) Endurance running and the evolution of Homo. Nature 432:345–352
Broom R, Schepers GWH (1946) The South-African ape-men: the Australopithecineae. Transvaal Museum Memoir no 2, Pretoria
Brunet M, Guy F, Pilbeam D, Mackaye HT, Likius A, Ahounta D et al (2002) A new hominid from Upper Miocene of Chad, Central Africa. Nature 418:145–151
Cant JGH (1987) Positional behavior of female Bornean orangutans Pongo pygmaeus. Am J Primatol 12:71–90
Carrier DR (1984) The energetic paradox of human running and hominid evolution. Curr Anthropol 25:483–495
Chaplin G, Jablonski NG, Cable NT (1994) Physiology, thermoregulation and bipedalism. J Hum Evol 27:497–510
Clarke RJ (1998) First ever discovery of an associated skull and skeleton of an ape-man. S Afr J Sci 94(10):460–463
Clarke RJ (1999) Discovery of complete arm and hand of the 3.3 million-year-old Australopithecus skeleton from Sterkfontein. S Afr J Sci 95:477–480
Clarke RJ (2002) Newly revealed information on Sterkfontein member 2 Australopithecus skeleton. S Afr J Sci 98:523–526
Clarke RJ (2013) Australopithecus from Sterkfontein Caves, South Africa. In: Reed KE, Fleagle JG, Leakey REF (eds) The paleobiology of Australopithecus. Springer, Dordrecht, pp 105–123
Clarke RJ, Tobias PV (1995) Sterkfontein member 2 foot bones of the oldest South African hominid. Science 269:521–524
Crompton RH, Li Y, Thorpe SK, Wang WJ, Savage R, Payne R, Carey TC, Aerts P, Van Elsacker L, Hofstetter A, Gunther MM, D’Aout K, De Clerq D (2003) The biomechanical evolution of erect bipedality. Cour Forsch-Inst Senckenberg 243:115–126
Dainton M (2001) Did our ancestors knuckle-walk? Nature 410:325–326
Dart RA (1925) Australopithecus africanus: the ape-man of South Africa. Nature 115:195–197
Darwin C (1871) The descent of man and selection in relation to sex. Random House, New York
Davis PR (1964) Hominid fossils from Bed I, Olduvai Gorge, Tanganyika. A tibia and fibula. Nature 201:967–968
Day MH (1969) Femoral fragment of a robust australopithecine from Olduvai Gorge, Tanzania. Nature 232:383–387
Day MH, Molleson TI (1976) The puzzle from JK2: a femur and a tibial fragment from (OH 34) from Olduvai Gorge, Tanzania. J Hum Evol 5:455–465
Day MH, Napier JR (1964) Fossils from Bed I, Olduvai Gorge, Tanganyika. Fossil foot bones. Nature 201:969–970
Day MH, Wickens EH (1980) Laetoli Pliocene hominid footprints and bipedalism. Nature 286:385–387
Day MH, Wood BA (1968) Functional affinities of the Olduvai Hominid 8 talus. Man 3:440–455
Deloison Y (1999) L’Homme ne descend pas d’un Primate arboricole ! une évidence méconnue. Biom Hum et Anthropol 17:147–150
DeSilva JM, Throckmorton ZJ (2010) Lucy’s flat feet: the relationship between the ankle and rearfoot arching in early Hominins. PLoS One 5(12):e14432. doi:10.1371/journal.pone.0014432
DeSilva JM, Holt KG, Churchill SE, Carlson KJ, Walker CS, and Berger LR (2013) The Lower Limb and Mechanics of Walking in Australopithecus sediba. Science 340:6129. DOI: 10.1126/science.1232999
Dobson SD (2005) Are the differences between Stw 431 (Australopithecus africanus) and AL 288–1 (A. afarensis) significant? J Hum Evol 49:143–154
Domínguez-Rodrigo M, Pickering TR, Baquedano E, Mabulla A, Mark DF et al (2013) First partial skeleton of a 1.34-million-year-old Paranthropus boisei from Bed II, Olduvai Gorge, Tanzania. PLoS One 8(12):e80347. doi:10.1371/journal.pone.0080347
Doran D, Hunt KD (1995) Comparative locomotor behavior of chimpanzees and bonobos: species and habitat differences. In: Wrangham RW, McGrew WC, de Waal F, Heltne PG (eds) Chimpanzee cultures. Harvard University Press, Cambridge, pp 93–108
Drake R, Curtis GH (1987) K-Ar geochronology of the Laetoli fossil localities. In: Leakey MD, Harris JM (eds) Laetoli: a Pliocene site in Northern Tanzania. Clarendon, Oxford, pp 48–52
Drapeau MSM, Harmon EH (2013) Metatarsal torsion in monkeys, apes, humans and australopiths. J Hum Evol 64:93–108
Duncan AS, Kappelman J, Shapiro LJ (1994) Metatarsophalangeal joint function and positional behaviour in Australopithecus afarensis. Am J Phys Anthropol 93:67–81
Dunsworth HM, Walker A (2002) Early genus Homo. In: Hartwig WC (ed) The primate fossil record. Cambridge University Press, Cambridge, pp 419–435
Elftman H, Manter J (1935) Chimpanzee and human feet in bipedal walking. Am J Phys Anthropol 20:69–79
Fleagle JG (1999) Primate adaptation and evolution, 2nd edn. Academic, New York
Fleagle JG, Stern JT, Jungers WL, Susman RL, Vangor AK, Wells JP (1981) Climbing: a biomechanical link with brachiation and bipedalism. Symp Zool Soc Lond 48:359–373
Foley RA, Elton S (1998) Time and energy: the ecological context for the evolution of bipedalism. In: Strasser E, Fleagle J, Rosenberger A, McHenry H (eds) Primate locomotion: recent advances. Plenum, New York, pp 419–433
Gagneux P, Varki A (2000) Genetic differences between humans and great apes. Mol Phylogenet Evol 18(1):2–13
Galik K, Senut B, Pickford M, Gommery D, Treil J, Kuperavage AJ, Eckhardt RB (2004) External and internal morphology of the BAR 1002’00 Orrorin tugenensis femur. Science 305:1450–1453
Gebo DL (1992) Plantigrady and foot adaptation in African apes: implications for hominid origins. Am J Phys Anthropol 89:29–58
Gebo DL (1996) Climbing, brachiation, and terrestrial quadrupedalism: historical precursors of hominid bipedalism. Am J Phys Anthropol 101:55–92
Gomberg DN, Latimer B (1984) Observations on the transverse tarsal joint of A. afarensis, and some comments on the interpretation of behaviour from morphology. Am J Phys Anthropol Suppl 63(2):164
Gregory WK (1916) Studies on the evolution of the primates. Bull Am Mus Nat Hist 35:239–355
Gregory WK (1927) Two views on the origin of man. Science 65:601–605
Gregory WK (1928) The upright posture of man: review of its origin and evolution. Proc Am Philos Soc 67:339–376
Haeusler MF (2001) New insights into the locomotion of Australopithecus africanus: implications of the partial skeleton of Stw 431 (Sterkfontein, South Africa). Ph D thesis, University of Zurich
Haeusler MF, McHenry HM (2004) Body proportions of Homo habilis reviewed. J Hum Evol 46:433–465
Haile-Selassie Y (2001) Late Miocene hominids from the Middle Awash, Ethiopia. Nature 412:178–181
Haile-Selassie Y, Suwa G, White TD (2004) Late Miocene teeth from Middle Awash, Ethiopia, and early hominid dental development. Science 303:1503–1505
Haile-Selassie Y, Latimer B, Alene M, Deino AL, Gilbert L, Melillo SM, Saylor BZ, Scott GR, Lovejoy CO (2010) An early Australopithecus postcranium from Woranso-Mille, Ethiopia. Proc Natl Acad Sci 107:12121–12126
Haile-Selassie Y, Saylor BZ, Deino A, Levin NF, Alene M, Latimer BM (2012) A new hominin foot from Ethiopia shows multiple Pliocene bipedal adaptations. Nature 483:565–570
Harcourt-Smith WEH (2002) Form and function in the hominoid tarsal skeleton. Ph D thesis, University College London, London
Harcourt-Smith WEH, Aiello LC (1999) An investigation into the degree of hallux abduction of OH 8. Am J Phys Anthropol Suppl 28:145
Harcourt-Smith WEH, Aiello LC (2004) Fossils, feet and the evolution of bipedal locomotion. J Anat 204:403–416
Harcourt-Smith WEH, Hilton C (2005) Did Australopithecus afarensis make the Laetoli footprint trail? New insights into an old problem. Am J Phys Anthropol 126(S40):112
Harcourt-Smith WEH, O’Higgins P, Aiello LC (2002) From Lucy to Littlefoot: a three dimensional analysis of Plio-Pleistocene hominin tarsal remains. Am J Phys Anthrop Suppl 34:82
Harcourt-Smith WEH, O’Higgins P, Aiello LC (2003) 3D morphometrics and the evolution of bipedality. Am J Phys Anthropol Suppl 36:109
Harrison T (1987) A reassessment of the phylogenetic relationships of Oreopithecus bambolii Gervais. J Hum Evol 15:541–583
Harrison T (1991) The implications of Oreopithecus bambolii for the origins of bipedalism. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 233–244
Harrison T (2010) Apes among the tangled branches of human origins. Science 327:532–534
Hartwig-Scherer S, Martin RD (1991) Was ‘Lucy’ more human than the ‘child’? Observations on early hominid postcranial skeletons. J Hum Evol 21:439–449
Hay RL, Leakey MD (1982) The fossil footprints of Laetoli. Sci Am 246:50–57
Herries AIR, Pickering R, Adams JW, Curnoe D, Warr G, Latham AG, Shaw J (2013) A multi-disciplinary perspective on the age of Australopithecus in Southern Africa. In: Reed KE, Fleagle JG, Leakey REF (eds) The paleobiology of Australopithecus. Springer, Dordrecht, pp 21–39
Holliday TW, Dugan J (2003) The utility of the lateral meniscal notch in distinguishing taxa of early hominins. Am J Phys Anthropol Suppl 36:116
Hunt KD (1990) Implications of chimpanzee positional behaviour in the Hominoidea. Am J Phys Anthropol 81:242
Hunt KD (1994) The evolution of human bipedality: ecology and functional morphology. J Hum Evol 26(3):183–203
Hunt KD (1996) The postural feeding hypothesis: an ecological model for the origin of bipedalism. S Afr J Sci 9:77–90
Hürzeler J (1968) Questions et réflexions sur l’histoire des anthropomorphes. Verh Naturf Ges Basel 69:1–47
Jablonski NG, Chaplin G (1993) Origin of habitual terrestrial bipedalism in the ancestor of the Hominidae. J Hum Evol 24:259–280
Johanson DC, Lovejoy CO, Burnstein AH, Heiple KG (1976) Functional implications of the Afar knee joint. Am J Phys Anthropol 44:188
Johanson DC, Masao FT, Eck GG, White TD, Walter RC, Kimbel WH et al (1987) New partial skeleton of Homo habilis from Olduvai Gorge, Tanzania. Nature 327:205–209
Jolly CJ (1970) The seed-eaters: a new model of hominid differentiation based on a baboon analogy. Man 5(1):5–26
Jungers WL (1982) Lucy’s limbs: skeletal allometry and locomotion in Australopithecus afarensis. Nature 297:676–678
Jungers WL, Stern JT (1983) Body proportions, skeletal allometry and locomotion in the Hadar hominids: a reply to Wolpoff. J Hum Evol 12:673–684
Jungers WL, Harcourt-Smith WEH, Wunderlich RE, Tocheri MW, Larson SG, Due RA, Djubiantono T, Morwood MJ (2009) The foot of Homo floresiensis. Nature 459:81–84
Keith A (1903) The extent to which the posterior segments of the body have been transmuted and suppressed in the evolution of man and allied primates. J Anat Physiol 37:18–40
Keith A (1923) Man’s posture: its evolution and disorders. Br Med J 1:451–454, 499–502, 545–548, 587–590, 624–626, 669–672
Kelly RE (2001) Tripedal knuckle-walking: a proposal for the evolution of human locomotion and handedness. J Theor Biol 213:333–358
Kibii JM, Clarke RJ (2003) A reconstruction of the Stw 431 Australopithecus pelvis based on newly discovered fragments. S Afr J Sci 99:225–226
Kibii JM, Churchill SE, Schmid P, Carlson KJ, Reed ND, de Ruiter DJ, Berger LR (2011) A partial pelvis of Australopithecus sediba. Science 333:1407–1411
Kidd RS, Oxnard C (2005) Little foot and big thoughts: a re-evaluation of the Stw573 foot from Sterkfontein, South Africa. HOMO J Comp Hum Biol 55:189–212
Kidd RS, O’Higgins P, Oxnard CE (1996) The OH8 foot: a reappraisal of the hindfoot utilizing a multivariate analysis. J Hum Evol 31:269–291
Kimbel WH, Suwa G, Asfaw B, Rak Y, White TD (2014) Ardipithecus ramidus and the evolution of the human cranial base. Proc Natl Acad Sci 111(3):948–953
Kivell TL, Kibii JM, Churchill SE, Schmid P, Berger LR (2011) Australopithecus sediba hand demonstrates mosaic evolution of locomotor and manipulative abilities. Science 333:1411–1417
Kivell TL, Schmitt D (2009) Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor. Proc Natl Acad Sci 106:14241–14246
Köhler M, Moyà-Solà S (1997) Ape-like or hominid-like? The positional behavior of Oreopithecus bambolii reconsidered. Proc Natl Acad Sci 94:11747–11750
Korey KA (1990) Deconstructing reconstruction: the OH 62 humerofemoral index. Am J Phys Anthropol 83:25–33
Kummer B (1965) Die Biomechanik der aufrechten Haltung. Mitt Naturf Ges Bern N F 22:239–259
Larson SG, Jungers WL, Morwood MJ, Sutikna T, Jatmiko EWS, Due RA, Djubiantono T (2007) Homo floresiensis and the evolution of the hominin shoulder. J Hum Evol 53:718–731
Latimer B (1991) Locomotor adaptations in Australopithecus afarensis: the issue of arboreality. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 169–176
Latimer B, Lovejoy CO (1982) Hominid tarsal, metatarsal, and phalangeal bones recovered from the Hadar formation: 1974–77 collections. Am. J. Phys. Anthropol. 57:701–719
Latimer B, Lovejoy CO (1989) The calcaneus of Australopithecus afarensis and its implications for the evolution of bipedality. Am J Phys Anthropol 78:369–386
Latimer B, Lovejoy CO (1990a) Hallucial tarsometatarsal joint in Australopithecus afarensis. Am J Phys Anthropol 82:125–133
Latimer B, Lovejoy CO (1990b) Metatarsophalangeal joints of Australopithecus afarensis. Am J Phys Anthropol 83:13–23
Latimer B, Ohman JC, Lovejoy CO (1987) Talocrural joint in African hominoids: implications for Australopithecus afarensis. Am J Phys Anthropol 74:155–175
Leakey MD, Harris JM (1987) Laetoli: a Pliocene site in Northern Tanzania. Clarendon, Oxford
Leakey MD, Hay RL (1979) Pliocene footprints in the Laetoli Beds at Laetoli, northern Tanzania. Nature 278:317–323
Leakey LSB, Tobias PV, Napier JR (1964) A new species of the genus Homo from Olduvai Gorge. Nature 202:7–9
Leakey REF, Leakey MG, Behrensmeyer AK (1978) The hominid catalogue. In: Leakey MG, Leakey RE (eds) Koobi Fora research project, volume 1: the fossil hominids and an introduction to their context, 1968–1974. Clarendon, Oxford
Leakey MG, Fiebel CS, McDougall I, Walker AC (1995) New four-million-year-old hominid species from Kanapoi and Allia Bay, Kenya. Nature 376:565–571
Leakey MG, Feibel CS, McDougall I, Ward C, Walker A (1998) New specimens and confirmation of an early age for Australopithecus anamensis. Nature 393:62–66
Lewis OJ (1980) The joints of the evolving foot. Part III. The fossil evidence. J Anat 131(2):275–298
Lewis OJ (1989) Functional morphology of the evolving hand and foot. Clarendon, Oxford
Lordkipanidze D, Jashashvili T, Vekua V, Ponce de León MS, Zollikofer CPE, Rightmire GP, Pontzer H, Ferring R, Oms O, Tappen M, Bukhsianidze M, Agustí J, Kahlke R, Kiladze G, Martinez-Navarro B, Mouskhelishvili A, Nioradze M, Rook L (2007) Postcranial evidence from early Homo from Dmanisi, Georgia. Nature 449:305–310
Lorenzo C, Arsuaga J-L, Carretero J-M (1999) Hand and foot remains from the Gran Dolina Early Pleistocene site (Sierra de Atapuerca, Spain). J Hum Evol 37:501–522
Lovejoy CO (1973) The gait of Australopithecines. Yearbook Phys Anthropol 17:147–161
Lovejoy CO (1975) Biomechanical perspectives on the lower limb of early hominids. In: Tuttle RH (ed) Primate Morphology and Evolution. Mouton, The Hague, pp 291–326
Lovejoy CO (1978) A biomechanical review of the locomotor diversity of early hominids. In: Jolly CJ (ed) Early hominids of Africa. St Martin’s Press, New York, pp 403–429
Lovejoy CO (1981) The origin of man. Science 211:344–350
Lovejoy CO (1988) Evolution of human walking. Sci Am 259:82–89
Lovejoy CO, Heiple KG, Burnstein AH (1973) The gait of Australopithecus africanus. Am J Phys Anthropol 38:757–780
Lovejoy CO, Heiple KG, Meindl RS (2001) Palaeoanthropology: did our ancestors knuckle-walk? Nature 410:325–326
Lovejoy CO, Meindl RS, Ohman JC, Heiple KG, White TD (2002) The Maka femur and its bearing on the antiquity of human walking: applying contemporary concepts of morphogenesis to the human fossil record. Am J Phys Anthropol 119(2):97–133
Lovejoy CO, Latimer B, Suwa G, Asfaw B, White TD (2009a) Combining prehension and propulsion: the foot of Ardipithecus ramidus. Science 326:72e1–72e8
Lovejoy CO, Simpson SW, White TD, Asfaw D, Suwa G (2009b) Careful climbing in the Miocene: the forelimbs of Ardipithecus ramidus and humans are primitive. Science 326:70e1–70e8
Lovejoy CO, Suwa G, Simpson SW, Matternes JH, White TD (2009c) The great divides: Ardipithecus ramidus reveals the postcrania of our last common ancestors with African apes. Science 326:100–106
Lovejoy CO, Suwa G, Spurlock L, Asfaw B, White TD (2009d) The pelvis and femur of Ardipithecus ramidus: the emergence of upright walking. Science 326:71e1–71e6
Marzke MW (1983) Joint function and grips of the Australopithecus afarensis hand, with special reference to the region of the capitate. J Hum Evol 12:197–211
McHenry HM (1986) The first bipeds: a comparison of the A. afarensis and A. africanus postcranium and implications for the evolution of bipedalism. J Hum Evol 15:177–191
McHenry HM (1991) First steps? Analyses of the postcranium of early hominids. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 133–141
McHenry HM (2002) Introduction to the fossil record of human ancestry. In: Hartwig WC (ed) The primate fossil record. Cambridge University Press, Cambridge, pp 401–405
McHenry HM, Berger LR (1998a) Body proportions of Australopithecus afarensis and A. africanus and the origin of the genus Homo. J Hum Evol 35:1–22
McHenry HM, Berger LR (1998b) Limb lengths in Australopithecus and the origin of the genus Homo. S Afr J Sci 94:447–450
McHenry HM, Brown CC (2008) Side steps: the erratic pattern of hominin postcranial change through time. J Hum Evol 55:639–651
McHenry HM, Jones AL (2006) Hallucial convergence in early hominids. J Hum Evol 50:534–539
Morton DJ (1924) Evolution of the human foot II. Am J Phys Anthropol 7:1–52
Morton DJ (1935) The human foot. Its evolution, physiology and functional disorders. Columbia University Press, New York
Moyà-Solà S, Köhler M (1996) A Dryopithecus skeleton and the origins of great-ape locomotion. Nature 379:156–159
Napier JR (1964) The evolution of bipedal walking in the hominids. Archives de Biologie (Liège) 75:673–708
Napier JR (1967) The antiquity of human walking. Sci Am 216(4):56–66
Napier JR, Davis PR (1959) The forelimb skeleton and associated remains of Proconsul africanus. Fossil Mammals Afr 16:1–69
O’Higgins P, Elton S (2007) Walking in trees. Science 316:1292–1294
Ohman JC, Krochta TJ, Lovejoy CO, Mensforth RP, Latimer B (1997) Cortical bone distribution in the femoral neck of hominoids: implications for the locomotion of Australopithecus afarensis. Am J Phys Anthropol 104:117–132
Page SL, Goodman M (2001) Catarrhine phylogeny: noncoding DNA evidence for a diphyletic origin of the mangabeys and a for a human-chimpanzee clade. Mol Phylogenet Evol 18(1):14–25
Partridge TC, Shaw J, Heslop D, Clarke RJ (1999) The new hominid skeleton from Sterkfontein, South Africa: age and preliminary assessment. J Quat Sci 14(4):293–298
Partridge TC, Granger DE, Caffee MW, Clarke RJ (2003) Lower Pliocene hominid remains from Sterkfontein. Science 300:607–612
Pauwels F (1980) Biomechanics of the locomotor apparatus: contributions on the functional anatomy of the locomotor apparatus. Springer, Berlin
Pickford M, Senut B (2001) The geological and faunal context of Late Miocene hominid remains from Lukeino, Kenya. Comptes Rendus de l’Académie des Sci IIA Earth Planetary Sci 332(2):145–152
Pickford M, Senut B, Gommery D, Treil J (2002) Bipedalism in Orrorin tugenensis revealed by its femora. Comptes Rendus Palevol 1(4):191–255
Plavcan JM, Lockwood CA, Kimbel WH, Lague MR, Harmon EH (2005) Sexual dimorphism in Australopithecus afarensis revisited: how strong is the case for a human-like pattern of dimorphism? J Hum Evol 48:313–320
Pontzer H, Rolian C, Rightmire GP, Jashashvili T, Ponce de León MS, Lordkipanidze D, Zollikofer CP (2010) Locomotor anatomy and biomechanics of the Dmanisi hominins. J Hum Evol 58:492–504
Potts R (1998) Environmental hypotheses of hominin evolution. Yearbook Phys Anthropol 41:93–136
Proctor DJ, Broadfield D, Proctor K (2008) Quantitative three-dimensional shape analysis of the proximal hallucial metatarsal articular surface in Homo, Pan, Gorilla, and Hylobates. Am J Phys Anthropol 135:216–224
Prost JH (1980) Origin of bipedalism. Am J Phys Anthropol 52:175–189
Rafferty KL (1998) Structural design of the femoral neck in primates. J Hum Evol 34:361–383
Raichlen D, Gordon A, Harcourt-Smith WEH, Foster A, Randall Hass W Jr (2010) Human-like biomechanics are preserved in the Laetoli footprints. PLoS One 5(3):e9769. doi:10.1371/journal.pone.0009769
Raichlen DA, Armstrong H, Lieberman DE (2011) Calcaneus length determines running economy: implications for endurance running performance in modern humans and Neanderthals. J Hum Evol 60:299–308
Reed KE (1997) Early hominid evolution and ecological change through the African Plio-Pleistocene. J Hum Evol 32:289–322
Reno PL, Meindl RS, McCollum MA, Lovejoy CO (2003) Sexual dimorphism in Australopithecus afarensis was similar to that of modern humans. Proc Natl Acad Sci 100:9404–9409
Richmond BG, Jungers WL (2008) Orrorin tugenensis femoral morphology and the evolution of bipedalism. Science 319:1662–1665
Richmond BG, Strait DS (2000) Evidence that humans evolved from a knuckle-walking ancestor. Nature 404:382–385
Richmond BG, Begun DR, Strait DS (2001) Origin of human bipedalism: the knuckle-walking hypothesis revisited. Yearbook Phys Anthropol 44(Suppl 33):71–105
Robbins LM (1987) Hominid footprints from site G. In: Leakey MD, Harris JM (eds) Laetoli: a Pliocene site in Northern Tanzania. Clarendon, Oxford, pp 497–502
Robinson JT (1972) Early hominid posture and locomotion. University of Chicago Press, Chicago
Robinson JT (1978) Evidence for locomotor differences between gracile and robust early hominids from South Africa. In: Jolly C (ed) Early hominids of Africa. Duckworth, London, pp 441–457
Rodman PS, McHenry HM (1980) Bioenergetics and the origin of hominid bipedalism. Am J Phys Anthropol 52:103–106
Rook L, Bondioli L, Köhler M, Moyà-Solà S, Macchiarelli R (1999) Oreopithecus was a bipedal ape after all: evidence from the iliac cancellous architecture. Proc Natl Acad Sci 96:8795–8799
Rose MD (1984) A hominine hip bone, KNM-ER 3228 from East Lake Turkana, Kenya. Am J Phys Anthropol 63:371–378
Rose MD (1991) The process of bipedalization in hominids. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 37–48
Ruff C (2009) Relative limb strength and locomotion in Homo habilis. Am J Phys Anthropol 138:90–100
Ruff CB, Walker AC (1993) Body size and body shape. In: Walker AC, Leakey RE (eds) The Nariokotome Homo erectus skeleton. Harvard Press, Cambridge, MA, pp 234–265
Russo GA, Kirk EC (2013) Foramen magnum position in bipedal mammals. J Hum Evol 65:656–670
Russo GA, Shapiro LJ (2013) Reevaluation of the lumbosacral region of Oreopithecus bambolii. J Hum Evol 65:253–265
Sarmiento EE (1994) Terrestrial traits in the hands and feet of gorillas. Am Mus Novit 3091:1–56
Sarmiento EE (1998) Generalized quadrupeds, committed bipeds and the shift to open habitats: an evolutionary model of hominid divergence. Am Mus Novit 3250:1–78
Sarmiento EE (2000) The Os navicular of humans, great apes, OH 8, Hadar, and Oreopithecus: function, phylogeny, and multivariate analyses. Am Mus Novit 3288:2–38
Sarmiento EE (2010) Comment on the paleobiology and classification of Ardipithecus ramidus. Science 328:1105b
Sawada Y, Pickford M, Senut B, Itaya T, Hyodo M, Miura T, Kashine C, Chujo T, Fujii H (2002) The age of Orrorin tugenensis, an early hominid from the Tugen Hills, Kenya. C. R. Paleovol 1:293–303
Schmid P (1983) Eine Rekonstruktion des Skelettes von AL 288–1 (Hadar) und deren Konsequenzen. Folia Primatologica 40:283–306
Schmid P (1991) The trunk of the australopithecines. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 225–234
Schmid P (2004) Functional interpretations of the Laetoli footprints. In: Meldrum DJ, Hilton CE (eds) From biped to strider: the emergence of modern human walking, running and resource transport. Kluwer Academic/Plenum, New York
Schrago CG, Voloch CM (2013) The precision of the hominid timescale estimated by relaxed clock methods. J Evol Biol n/a–n/a. doi:10.1111/jeb.12076
Schultz AH (1930) The skeleton of the trunk and limbs of higher primates. Hum Biol 2:303–438
Senut B (1981a) Humeral outlines in some hominoid primates and the Plio-Pleistocene hominids. Am J Phys Anthropol 56:275–283
Senut B (1981b) L’humerus et ses articulations chez les hominides Plio-Pleistocenes. Cahiers de Paleontologie (Paleoanthropologie). Editions du CNRS, Paris
Senut B (1989) Climbing as a crucial preadaptation to for human bipedalism. Ossa 14:35–44
Senut B, Tardieu C (1985) Functional aspects of Plio-Pleistocene hominid limb bones: implications for taxonomy and phylogeny. In: Delson E (ed) Ancestors: the hard evidence. Alan R. Liss, New York, pp 193–201
Senut B, Pickford M, Gommery D, Mein P, Cheboi K, Coppens Y (2001) First hominid from the Miocene (Lukeino formation, Kenya). Comptes Rendus de l’Académie des Sci IIA Earth Planetary Sci 332(2):137–144
Shipman P (1986) Scavenging or hunting in early hominids: theoretical framework and tests. Am Anthropol 88:27–43
Sinclair ARE, Leakey MD, Norton-Griffiths M (1986) Migration and hominid bipedalism. Nature 324:307–308
Spoor F, Wood BA, Zonneveld F (1994) Implications of early hominid labyrinth morphology for evolution of human bipedal locomotion. Nature 369:645–648
Springer MS, Meredith RW, Gatesy J, Emerling CA, Park J, Rabosky DL, Stadler T, Steiner C, Ryder OA, Janečka JE, Fisher CA, Murphy WJ (2012) Macroevolutionary dynamics and historical biogeography of primate diversification inferred from a species supermatrix. PLoS One 7(11):e49521. doi:10.1371/journal.pone.0049521
Steiper ME, Seiffert ER (2012) Evidence for a convergent slowdown in primate molecular rates and its implications for the timing of early primate evolution. Proc Natl Acad Sci 109:6006–6012
Stern JT (1975) Before bipedality. Yearbook Phys Anthropol 19:59–68
Stern JT (2000) Climbing to the top: a personal memoir of Australopithecus afarensis. Evol Anthropol 9(3):113–133
Stern JT, Susman RL (1983) The locomotor anatomy of Australopithecus afarensis. Am J Phys Anthropol 60:279–317
Stern JT, Susman RL (1991) “Total morphologocal pattern” versus the “magic trait”: conflicting approaches to the study of early hominid bipedalism. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 99–111
Steudel KL (1994) Locomotor energetics and hominid evolution. Evol Anthropol 3:42–48
Straus WL (1949) The riddle of man’s ancestry. Quart Rev Biol 24:200–223
Straus WL (1957) Oreopithecus bambolii. Science 126:345–346
Straus WL (1962) Fossil evidence of the evolution of the erect, bipedal posture. Clin Orthop 25:9–19
Stringer C, Andrews P (2005) The complete world of human evolution. Thames and Hudson, London
Susman RL (1989) New hominid fossils from the Swartkrans formation (1979–1986 excavations): postcranial specimens. Am J Phys Anthropol 79:451–474
Susman RL, Brain TM (1988) New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus. Am J Phys Anthropol 77:7–15
Susman RL, Creel N (1979) Functional and morphological affinities of the subadult hand (OH 7) from Olduvai Gorge. Am J Phys Anthropol 51:211–332
Susman RL, de Ruiter DJ (2004) New hominin first metatarsal (SK 1813) from Swartkrans. J Hum Evol 47:171–181
Susman RL, Stern JT (1982) Functional morphology of Homo habilis. Science 217:931–934
Susman RL, Stern JT, Jungers WL (1984) Arboreality and bipedality in the Hadar hominids. Folia Primatol 43:283–306
Susman RL, Stern JT, Jungers WL (1985) Locomotor adaptations in the Hadar hominids. In: Delson E (ed) Ancestors: the hard evidence. Alan R Liss, New York, pp 184–192
Suwa G, Asfaw B, Kono RT, Kubo D, Lovejoy CO, White TD (2009) The Ardipithecus ramidus skull and its implications for hominid origins. Science 325:68e1–68e7
Taieb M, Johanson DC, Coppens Y, Bonnefille R, Kalb J (1974) Découverte d’hominidés dans le séries Plio-Pleistocénes d’Hadar (Bassin de l’Awash; Afar, Ethiopia). C R Acad Sci Paris 279:735–738
Tardieu C (1981) Morpho-functional analysis of the articular surfaces of the knee-joint in primates. In: AB Chiarelli and RS Corruccini (eds) Primate Evolutionary Biology. Berlin: Springer-Verlag, pp. 68–80
Tardieu C (1983) L’articulation du Genou. Analyse Morpho-Fonctionnelle Chez les Primates. Application aux Hominidés Fossiles. Cah. Paléoanthrop. Paris: C.N.R.S., pp. 1–108
Tardieu C (1986) Evolution of the knee menisci in primates. In: J Else and J Lee (eds) Primate Evolution. Cambridge, Cambridge University Press, pp. 183–190
Tardieu C (1993) L’angle bicondylaire du fémur est-il homologue chez l’homme et les primates non humains? Réponse ontogénétique. Bull Mém Soc Anthropol. Paris 5:159–168
Tardieu C (1994) Development of the femoral diaphysis in humans: Functional and evolutionary significance. Folia Primatol. 63:53–58
Tardieu C, Trinkaus E (1994) Early ontogeny of the human femoral bicondylar angle. Am J Phys Anthropol 95:183–195
Thackeray F, Gommery D, Braga J (2002) Australopithecine postcrania (Sts 14) from the Sterkfontein Caves, South Africa: the skeleton of ‘Mrs Ples’? S Afr J Sci 98:211–212
Thorpe SK, Crompton RH (2005) The locomotor ecology of wild orangutans (Pongo pygmaeus abelii) in the gunung leuser ecosystem, Sumatra, Indonesia: a multivariate analysis using log-linear modelling. Am J Phys Anthropol 127:58–78
Thorpe SK, Holder RL, Crompton RH (2007) Origin of human bipedalism as an adaptation for locomotion on flexible branches. Science 316:1328–1331
Tocheri MW, Orr CM, Larson SG, Sutikna T, Jatmiko T, Saptomo EW, Due RA, Djubiantono T, Morwood MJ, Jungers WL (2007) The primitive wrist of Homo floresiensis and its implications for hominin evolution. Science 317:1743–1745
Trinkaus E (1983) Functional aspects of Neanderthal pedal remains. Foot Ankle 3(6):377–390
Tuttle RH (1968) Propulsive and prehensile capabilities in the hands and feet of the great apes: a preliminary report. In: VIIIth congress of anthropological and ethnological sciences, Tokyo, pp 31–32
Tuttle RH (1970) Postural, propulsive, and prehensile capabilities in the cheiridia of chimpanzees and other great apes. In: Bourne GH (ed) The chimpanzee, vol 2. Karger, Basel/New York, pp 167–253
Tuttle RH (1974) Darwin’s apes, dental apes, and the descent of man: normal science in evolutionary anthropology. Curr Anthropol 15:389–398
Tuttle RH (1975) Knuckle-walking and knuckle-walkers: a commentary on some recent perspectives in hominoid evolution. In: Tuttle RH (ed) Primate functional morphology and evolution. Mouton, The Hague, pp 203–212
Tuttle RH (1981) Evolution of hominid bipedalism and prehensile capabilities. Philos Trans R Soc Lond B 292:89–94
Tuttle RH (1987) Kinesiological inferences and evolutionary implications from Laetoli bipedal trails G-1, G-2/3 and A. In: Leakey MD, Harris JM (eds) Laetoli: a Pliocene site in Northern Tanzania. Clarendon, Oxford, pp 497–502
Tuttle RH, Basmajian JV (1974) Electromyography of forearm musculature in Gorilla and problems related to knuckle-walking. In: Jenkins FA (ed) Primate locomotion. Academic, New York, pp 293–348
Tuttle RH, Webb D, Weidl E, Baksh M (1990) Further progress on the Laetoli trails. J Archaeol Sci 17:347–362
Tuttle RH, Webb DM, Tuttle NI (1991) Laetoli footprint trails and the evolution of hominid bipedalism. In: Coppens Y, Senut B (eds) Origine(s) de la bipédie chez les hominidés. Editions du CNRS, Paris, pp 187–198
Van Couvering JA (2000) The Pliocene. In: Delson E, Tattersall I, Van Couvering JA, Brooks AS (eds) Encyclopedia of human evolution and prehistory, 2nd edn. Garland, New York, pp 574–576
Videan EN, McGrew WC (2002) Bipedality in chimpanzee (Pan troglodytes) and bonobo (Pan paniscus): testing hypotheses on the evolution of bipedalism. Am J Phys Anthropol 118:184–190
Wallace IJ, Demes B, Jungers WL, Alvers M, Su A (2008) The bipedalism of the Dmanisi hominins: Pigeon-toed early Homo? Am J Phys Anthropol 136:375–378
Wang W, Crompton RH, Carey TS, Gunther MM, Li Y, Savage R, Sellers WI (2004) Comparison of inverse-dynamics musculo-skeletal models of AL 288–1 Australopithecus afarensis and KNM-WT 15000 Homo ergaster to modern humans, with implications for the evolution of bipedalism. J Hum Evol 47:453–478
Ward CV (2002) Interpreting the posture and locomotion of Australopithecus afarensis: where do we stand? Yearbook Phys Anthropol Suppl 35:185–215
Ward CV, Walker A, Leakey MG (1999) The new species Australopithecus anamensis. Evol Anthropol 7:197–205
Ward CV, Leakey MG, Walker A (2001) Morphology of Australopithecus anamensis from Kanapoi and Allia Bay, Kenya. J Hum Evol 41:255–368
Ward CV, Kimbel WH, Johanson DC (2011) Complete fourth metatarsal and arches in the foot of Australopithecus afarensis. Science 331:750–753
Washburn SL (1960) Tools and human evolution. Sci Am 203:63–75
Washburn SL (1967) Behavior and the origin of man. Proc R Anthropol Inst 3:21–27
Webb NM, Harcourt-Smith WEH, Pontzer H (2013) An analysis of the Ardipithecus ramidus pelvis reconstruction using 3D geometric morphometric techniques. Am J Phys Anthropol S56:287
Wheeler PE (1984) The evolution of bipedality and loss of functional body hair in hominids. J Hum Evol 13:91–98
Wheeler PE (1988) Stand tall and stay cool. New Scientist 118(1613):62–65
Wheeler PE (1991) The influence of bipedalism on the energy and water budgets of early hominids. J Hum Evol 21:117–136
Wheeler PE (1993) The influence of stature and body form on hominid energy and water budgets; a comparison of Australopithecus and early Homo physiques. J Hum Evol 24:13–28
Wheeler PE (1994) The foraging times of bipedal and quadrupedal hominids in open equatorial environments (a reply to Chaplin, Jablonski & Cable, 1994). J Hum Evol 27:511–517
White TD (2002) Earliest hominids. In: Hartwig WC (ed) The primate fossil record. Cambridge University Press, Cambridge, pp 407–417
White TD, Suwa G (1987) Hominid footprints at Laetoli: facts and interpretations. Am J Phys Anthropol 72:485–514
White TD, Suwa G, Asfaw B (1994) Australopithecus ramidus, a new species of early hominid from Aramis, Ethiopia. Nature 371:306–312
White TD, Suwa G, Asfaw B (1995) Australopithecus ramidus, a new species of early hominid from Aramis, Ethiopia: a corrigendum. Nature 375:88
White TD, Asfaw B, Beyene Y, Haile-Selassie Y, Lovejoy CO, Suwa G, WoldeGabriel G (2009) Ardipithecus ramidus and the paleobiology of early hominids. Science 326:75–85
White TD, Suwa G, Lovejoy CO (2010) Response to comment on the paleobiology and classification of Ardipithecus ramidus. Science 328:1105
Williams SA (2010) Morphological integration and the evolution of knuckle-walking. J Hum Evol 58:432–440
Wood BA (1974) A Homo talus from East Rudolf, Kenya. J Anat 117:203–204
Wood BA (1993) Four legs good, two legs better. Nature 363:587–588
Wood BA (2002) Hominid revelations from Chad. Nature 418:133–135
Wood Jones F (1916) Arboreal man. Edward Arnold, London
Wood Jones F (1929) Man’s place among the mammals. Arnold, London
Wood BA, Collard M (1999) The human genus. Science 284:65–71
Wood BA, Harrison T (2011) The evolutionary context of the first hominins. Nature 470:347–352
Wrangham RW (1980) Bipedal locomotion as a feeding adaptation in Gelada Baboons, and its implications for hominid evolution. J Hum Evol 9:329–331
Zipfel B, DeSilva JM, Kidd RS, Carlson KJ, Churchill SE, Berger LR (2011) The foot and ankle of Australopithecus sediba. Science 333:1417–1420
Zollikofer CPE, Ponce de León MS, Lieberman DE, Guy F, Pilbeam D, Likius A, Mackaye HT, Vignaud P, Brunet M (2005) Virtual cranial reconstruction of Sahelanthropus tchadensis. Nature 434:755–759
Author information
Authors and Affiliations
Corresponding author
Editor information
Editors and Affiliations
Rights and permissions
Copyright information
© 2015 Springer-Verlag Berlin Heidelberg
About this entry
Cite this entry
Harcourt-Smith, W.E.H. (2015). Origin of Bipedal Locomotion. In: Henke, W., Tattersall, I. (eds) Handbook of Paleoanthropology. Springer, Berlin, Heidelberg. https://doi.org/10.1007/978-3-642-39979-4_48
Download citation
DOI: https://doi.org/10.1007/978-3-642-39979-4_48
Published:
Publisher Name: Springer, Berlin, Heidelberg
Print ISBN: 978-3-642-39978-7
Online ISBN: 978-3-642-39979-4
eBook Packages: Biomedical and Life SciencesReference Module Biomedical and Life Sciences