Earliest complete hominin fifth metatarsal-implications for the evolution of the lateral column of the foot

American Journal of Physical Anthropology

Volumn 140, Issue 3, 2009, Pages 532-545

  Zipfel, Bernhard ^a   DeSilva, Jeremy M ^b   Kidd, Robert S ^a,c  

^a UNIVERSITY OF THE WITWATERSRAND   (South Africa)

^b Boston University   (United States)

^c UNIVERSITY OF WESTERN SYDNEY   (Australia)

Author keywords

Bipedalism; Fossil metatarsal; Hominins; Sterkfontein

Indexed keywords

APE; ARTICLE; BIOMECHANICS; CALCANEUS; COMPARATIVE ANATOMY; EVOLUTION; FOOT; FOSSIL HOMINID; HEAD; HUMAN; METATARSAL BONE; MORPHOLOGICAL TRAIT; NONHUMAN; PLESIOMORPHY; SOUTH AFRICA; TARSAL BONE;

ANATOMY, COMPARATIVE; ANIMALS; EVOLUTION; FEMALE; FOOT; FOSSILS; HOMINIDAE; HUMANS; LOCOMOTION; MALE; METATARSAL BONES; MULTIVARIATE ANALYSIS;

EID: 70349911502     PISSN: 00029483     EISSN: 10968644     Source Type: Journal    
DOI: 10.1002/ajpa.21103     Document Type: Article

References (131)

1. Aiello L, Dean C. 1990. An introduction to human evolutionary anatomy. London: Academic Press.
2. Albrecht G. 1980a. Multivariate analysis and the study of form, with special reference to canonical variates analysis. Am Zool 20:679-693.
3. Albrecht G. 1980b. Weighted versus unweighted canonical variate analysis in morphometrics. Am Zool 20:820.
4. Albrecht G. 1992. Assessing the affinities of fossils using canonical variates and generalized distances. J Hum Evol 7:49-69.
5. Antón SC. 2003. Natural history of Homo erectus. Yearb Phys Anthropol 122:126-170.
6. Archibald JD, Lovejoy CO, Heiple KG. 1972. Implications of relative robusticity in the Olduvai metatarsus. Am J Phys Anthropol 37:93-96.
7. Bennett MR, Harris MRJ, Richmond BG, Braun DR, Mbua E, Kiura P, Olago D, Kibunjia M, Omuombo C, Behrensmeyer AK, Huddart D, Gonzalez S. 2009. Early hominin foot morphology based on 1.5-million-year-old footprints from Ileret, Kenya. Science 323:1197-1201.
8. Berger LR, Lacruz R, de Ruiter DJ. 2002. Brief communication: revised age estimates of Australopithecus-bearing deposits at Sterkfontein, South Africa. Am J Phys Anthropol 119:192-197.
9. Berillon G. 1999. Geometric pattern of the hominoid hallucal tarsometatarsal complex. Quantifying the degree of hallux abduction in early hominids. C R Acad Sci Ser IIa Earth Planet Sci 328:627-633.
10. Berillon G. 2003. Assessing the longitudinal structure of the early hominin foot: a two-dimensional architecture analysis. Hum Evol 18:113-122.
11. Blackith RE, Reyment RA. 1971. Multivariate morphometrics. London: Academic Press.
12. Blackwood CB, Yuen TJ, Sangeorzan BJ, Ledoux WR. 2005. The midtarsal joint locking mechanism. Foot Ankle Int 26:1074-1080.
13. Bojsen-Møller F. 1979. Calcaneocuboid joint and stability of the longitudinal arch of the foot at high and low gear push off. J Anat 129:165-176. (Pubitemid 9256024)
14. Bojsen-Møller F, Lamoreux L. 1979. Significance of free dorsiflexion of the toes in walking. Acta Orthop Scand 50:471-479. (Pubitemid 10233027)
15. Brunet M, Guy F, Pilbeam D, Mackaye HT, Likius A, Ahounta D, Beauvilain A, Blondel C, Bocherens H, Boisserie J-R, de Bonis L, Coppens Y, Dejax J, Denys C, Duringer P, Eisenmann V, Fanone G, Fronty P, Geraads D, Lehmann T, Lihoreau F, Louchart A, Mahamat A, Merceron G, Mouchelin G, Otero O, Campomanes PP, Ponce De Leon M, Rage J-C, Sapanet M, Schuster M, Sudre J, Tassy P, Valentin X, Vignaud P, Viriot L, Zazzo A, Zollikofer C. 2002. A new hominid from Upper Miocene of Chad, Central Africa. Nature 418:145-151. (Pubitemid 34773762)
16. Bryant FB, Yarnold PR. 2001. Principal-components analysis and exploratory and confirmatory factor analysis. In: Grimm LG, Yarnold PR, editors. Reading and understanding multivariate statistics. Washington, DC: American Psychological Association. p 99-108.
17. Cavanaugh PR, Rodgers MM, Iiboshi A. 1987. Pressure distributions under symptom-free feet during barefoot study. Foot Ankle 7:262-276. (Pubitemid 17062912)
18. Chen W, Tang F, Ju C. 2001. Stress distribution of the foot during mid-stance to push-off in barefoot gait: a 3-D finite element analysis. Clin Biomech (Bristol, Avon) 16:614-620. (Pubitemid 32722139)
19. Clarke RJ. 2006. A deeper understanding of the stratigraphy of Sterkfontein hominid site. Trans R Soc South Afr 61:111-120.
20. Clarke RJ, Tobias PV. 1995. Sterkfontein member 2 foot bones of the oldest South African hominid. Science 269:521-524.
21. D'Août K, Aerts P, De Clercq D, De Meester K, Van Elsacker L. 2002. Segment and joint angles of hind limb during bipedal and quadrupedal walking of the bonobo (Pan paniscus). Am J Phys Anthropol 119:37-51. (Pubitemid 34976774)
22. Day MH, Leakey REP. 1974. New evidence of the genus Homo from East Rudolf, Kenya III. Am J Phys Anthropol 41:367-380.
23. Day MH, Napier JR. 1964. Hominid fossils from Bed I, Olduvai Gorge, Tanganyika: fossil foot bones. Nature 201:969-970.
24. DeLoison Y. 2003. Anatomie des fossils de pieds des hominidés D'Afrique du Sud dates entre 2,4 et 3,5 millions dánnées. Interprétaton quant à leur mode de locomotion. Biometr Hum Anthropol 21:189-230.
25. Delson E. 1988. Chronology of South African australopiths site units. In: Grine FE, editor. Evolutionary history of the "robust" australopithecines. New York: Aldine de Gruyter. p 317-325.
26. DeSilva JM. 2008. Vertical climbing adaptations in the ape ankle and midfoot. Implications for locomotion in Miocene catarrhines and Plio-Pleistocene hominins, Ph.D. thesis, University of Michigan.
27. DeSilva JM. Revisiting the "midtarsal break." Am J Phys Anthropol (submitted).
28. DeSilva JM, MacLatchy LM. 2008. Revisiting the midtarsal break. Am J Phys Anthropol [Suppl] 46:89.
29. Duncan AS, Kappelman J, Shapiro LJ. 1994. Metatarsophalangeal joint function and positional behavior in Australopithecus afarensis. Am J Phys Anthropol 93:67-82. (Pubitemid 24020099)
30. Elftman H, Manter J. 1935a. The evolution of the human foot, with especial reference to the joints. J Anat 70:56-67.
31. Elftman H, Manter J. 1935b. Chimpanzee and human feet in bipedal walking. Am J Phys Anthropol 20:69-79.
32. Eliot J, Jungers WL. 2000. Fifth metatarsal morphology does not predict presence or absence of fibularis tertius muscle in hominids. J Hum Evol 38:333-342.
33. Feibel CS, Agnew A, Latimer B, Demas M, Marshall F, Waane S, Schmid P. 1996. The Laetoli hominid footprints - a preliminary report on the conservation and scientific restudy. Evol Anthropol 4:149-154. (Pubitemid 126089035)
34. Gebo DL. 1992. Plantigrady and foot adaptation in African apes: implications for hominin origins. Am J Phys Anthropol 89:29-58.
35. Gebo DL, Schwartz GT. 2006. Foot bones from Omo: implications for hominid evolution. Am J Phys Anthropol 129:499-511.
36. Godfrey L, Sutherland M, Paine R, Williams F, Boy D, Vuillaume- Randriamanantena M. 1995. Limb joint surface areas and their ratios in Malagasy lemurs and other mammals. Am J Phys Anthropol 97:11-36.
37. 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 63:164.
38. Grausz HM, Leakey REF, Walker AC, Ward CV. 1988. Associated cranial and postcranial bones of Australopithecus boisei. In: Grine FE, editor. Evolutionary history of the ''robust'' australopithecines. New York: Aldine de Gruyter. p 127-132.
39. Griffin NL, Richmond BG. 2005. Cross-sectional geometry of the human forefoot. Bone 37:253-260.
40. Haile-Selassie Y. 2001. Late Miocene hominids from the Middle Awash, Ethiopia. Nature 412:178-181. (Pubitemid 32674712)
41. Harcourt-Smith W, Aiello LC. 2004. Fossils, feet and the evolution of human bipedal locomotion. J Anat 204:403-416. (Pubitemid 38786689)
42. Harcourt-Smith WEH. 2002. Form and function in the hominoid tarsal skeleton, Ph.D. thesis, University College London, London.
43. Harcourt-Smith WEH, Aiello LC. 1999. An investigation into the degree of hallux abduction of OH 8. Am J Phys Anthropol [Suppl] 28:145.
44. Harcourt-Smith WEH, Hilton C. 2005. Did Australopithecus afarensis make the Laetoli footprint trail? New insights into an old problem. Am J Phys Anthropol [Suppl] 40:112.
45. Häusler M. 2002. New insights into the locomotion of Australopithecus africanus based on the pelvis. Evol Anthropol 11 (S1):53-57. (Pubitemid 37239401)
46. Häusler M, Berger LR. 2001. StW 441/465: a new fragmentary ilium of a small-bodied Australopithecus africanus from Sterkfontein, South Africa. J Hum Evol 40:411-417. (Pubitemid 32511856)
47. Hayafune N, Hayafune Y, Jacob H. 1999. Pressure and force distribution characteristics under the normal foot during the push-off phase in gait. Foot 9:88-92. (Pubitemid 29358832)
48. Hicks JH. 1954. The mechanics of the foot. II. The plantar aponeurosis and the arch. J Anat 88:25-31.
49. Hills A, Hennig E, McDonald M, Bar-Or O. 2001. Plantar pressure differences between obese and non-obese adults: a biomechanical analysis. Int J Obes 25:1674-1679. (Pubitemid 33052079)
50. Hunt KD. 1994. The evolution of human bipedality: ecology and functional morphology. J Hum Evol 26:183-202. (Pubitemid 124013669)
51. Jolicoeur P. 1963. The multivariate generalisation of the allometry equation. Biometrics 19:497-499.
52. Jungers WL. 1988. Relative joint size and hominid locomotor adaptations with implications for the evolution of hominid bipedalism. J Hum Evol 17:247-265.
53. Kanatli U, Yetkin H, Bolukbasi S. 2003. Evaluation of the transverse metatarsal arch of the foot with gait analysis. Arch Orthop Trauma Surg 123:148-150. (Pubitemid 36747760)
54. Kibii JM, Clarke RJ. 2003. A reconstruction of the StW 431 Australopithecus pelvis based on newly discovered fragments. South Afr J Sci 99:225-226.
55. Kidd R. 1995. An investigation into the patterns of morphological variation in the proximal tarsus of selected human groups, apes and fossils: a morphometric analysis, Ph.D. thesis, University of Western Australia.
56. Kidd R. 1998. The past is the key to the present: thoughts on the origins of human foot structure, function and dysfunction as seen from the fossil record. Foot 8:75-84. (Pubitemid 28339704)
57. Kidd R. 1999. Evolution of the hindfoot: a model of adaptation with evidence from the fossil record. J Am Podiatr Med Assoc 89:2-17.
58. Kidd R, 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.
59. Kidd RS, O'Higgins P, Oxnard CE. 1996. The OH8 foot: a reappraisal of the functional morphology of the hindfoot utilizing a multivariate analysis. J Hum Evol 31:269-291. (Pubitemid 27115540)
60. Kuman K, Clarke RJ. 2000. Stratigraphy, artifact industries and hominid associations for Sterkfontein, Member 5. J Hum Evol 38:827-847. (Pubitemid 30341924)
61. Lamy P. 1986. The settlement of the longitudinal plantar arch of some African Plio-Pleistocene hominins: a morphological study. J Hum Evol 15:31-46. (Pubitemid 16082354)
62. Langdon JH, Bruckner J, Baker HH. 1991. Pedal mechanics and bipedalism in early hominins. In: Coppens Y, Senut B, editors. Origine(s) de la Bipédie chez les Homininés. Paris: Centre National de la Recherche Scientifique. p 59-167.
63. Largey A, Bonnel F, Canovas F, Subsol G, Chemouny S, Banegas F. 2007. Three-dimensional analysis of the intrinsic anatomy of the metatarsal bones. J Foot Ankle Surg 46:434-441. (Pubitemid 350016950)
64. Latimer B. 1991. Locomotion adaptations in Australopithecus afarensis: the issue of arboreality. In: Coppens Y, Senut B, editors. Origine(s) de la Bipédie chez les Homininés. Paris: Centre National de la Recherche Scientifique. p 169-176.
65. 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. (Pubitemid 19076958)
66. Latimer B, Lovejoy CO. 1990a. Metatarsophalangeal joints of Australopithecus afarensis. Am J Phys Anthropol 83:13-23.
67. Latimer B, Lovejoy CO. 1990b. Hallucial tarsometatarsal joint in Australopithecus afarensis. Am J Phys Anthropol 82:125-133. (Pubitemid 20229884)
68. Latimer B, Lovejoy CO, Johanson DC, Coppens Y. 1982. Hominid tarsal, metatarsal, and phalangeal bones recovered from the Hadar formation: 1974-77 collections. Am J Phys Anthropol 57:701-719. (Pubitemid 12089989)
69. Latimer B, Ohman JC, Lovejoy CO. 1987. Talocrural joint in African hominoids: implications for Australopithecus afarensis. Am J Phys Anthropol 74:155-175. (Pubitemid 18046101)
70. Lauder GV. 1995. On the inference of function from structure. In: Thomason JJ, editor. Functional morphology in vertebrate paleontology. Cambridge: Cambridge University Press. p 1-18.
71. Leakey L, Tobias P, Napier J. 1964. A new species of the genus Homo from Olduvai Gorge. Nature 202:5-7.
72. Leakey MD, Hay RL. 1979. Pliocene footprints in the Laetoli beds at Laetoli, northern Tanzania. Nature 278:317-323.
73. Lebatard AE, Bourlés DL, Duringer P, Jolivet M, Braucher R, Carcaillet J, Schuster M, Arnaud N, Monié P, Lihoreau F, Likius A, Mackaye HT, Vignaud P, Brunet M. 2008. Cosmogenic nuclide dating of Sahelanthropus tchadensis and Australopithecus bahrelghazali: Mio-Pliocene hominids from Chad. Proc Natl Acad Sci USA 105:3226-3231. (Pubitemid 351723535)
74. Ledoux W, Hillstrom H. 2002. The distributed plantar vertical forces of the neutrally aligned pes planus feet. Gait Posture 15:1-9. (Pubitemid 34059246)
75. Lewis OJ. 1980. The joints of the evolving foot. Part III. The fossil evidence. J Anat 131:275-298.
76. Lewis OJ. 1989. Functional morphology of the evolving hand and foot. Oxford: Clarendon Press.
77. Lordkipanidze D, Jashashville T, Vekua A, Ponce de Léon MS, Zollikofer CPE, Rightmire GP, 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 449:305-310. (Pubitemid 47443479)
78. Lovejoy CO. 1974. The gait of australopithecines. Yearb Phys Anthropol 17:147-161.
79. Lovejoy CO. 2005a. The natural history of gait and posture. Part 1. Spine and pelvis. Gait Posture 21:95-112.
80. Lovejoy CO. 2005b. The natural history of gait and posture. Part 2. Hip and thigh. Gait Posture 21:113-124.
81. Lovejoy CO. 2007. The natural history of gait and posture. Part 3. The knee. Gait Posture 25:325-341.
82. Luger EJ, Nissan M, Karpf A, Steinberg S, Dekel S. 1999. Patterns of weight distribution under the metatarsal heads. J Bone Joint Surg (Br) 81:199-202. (Pubitemid 29327052)
83. Manter JT. 1946. Distribution of compression forces in the joints of the human foot. Anat Rec 3:313-321.
84. Marchi D. 2005. The cross-sectional geometry of the hand and foot bones of the Hominoidea and its relationship to locomotor behavior. J Hum Evol 49:743-761. (Pubitemid 41824067)
85. McHenry H, Jones AL. 2006. Hallucial convergence in early hominins. J Hum Evol 50:534-539.
86. McHenry HM. 1994. Behavioral ecological implications of early hominid body size. J Hum Evol 27:77-87.
87. McKee JK, Thackeray JF, Berger LR. 1995. Faunal assemblage seriation of southern African Pliocene and Pleistocene fossil deposits. Am J Phys Anthropol 106:235-250.
88. Meldrum DJ. 2002. Midfoot flexibility and the evolution of bipedalism. Am J Phys Anthropol [Suppl] 34:111-112.
89. Meldrum DJ, Wunderlich RE. 1998. Midfoot flexibility in ape foot dynamics, early hominid footprints and bipedalism. Am J Phys Anthropol [Suppl] 26:161.
90. Moggi-Cecchi J, Grine FE, Tobias PV. 2006. Early hominid dental remains from Members 4 and 5 of the Sterkfontein Formation (1966-1996 excavations): catalogue, individual associations, morphological descriptions and initial metrical analysis. J Hum Evol 50:239-328. (Pubitemid 43291727)
91. Morton DJ. 1922. Evolution of human foot. Am J Phys Anthropol 5:305-325.
92. Patake SM, Mysorekar VR. 1977. Diaphysial nutrient foramina in human metatacarpals and metatarsals. J Anat 124:299-304.
93. Pickering TR, Clarke RJ, Moggi-Cecchi J. 2004. Role of carnivores in the accumulation of the Sterkfontein Member 4 hominid assemblage: a taphanomic reassessment of the complete hominid fossil sample (1936-1999). Am J Phys Anthropol 125: 1-15. (Pubitemid 39095831)
94. Reed KE, Kitching JM, Grine FE, Jungers WL, Sokoloff L. 1993. Proximal femur of Australopithecus africanus from Member 4, Makapansgat, South Africa. Am J Phys Anthropol 92:1-15. (Pubitemid 23240711)
95. Reyment RA, Blackwith RE, Campell NA. 1984. Multivariate morphometrics. London: Academic Press.
96. Richmond BG, Jungers WL. 2008. Orrorin tugenensis femoral morphology and the evolution of hominin bipedalism. Science 319:1662-1665.
97. Robinson JT. 1972. Early hominid posture and locomotion. Chicago: Chicago University Press.
98. Sanders WJ. 1998. Comparative morphometric study of the australopithecine vertebral series Stw-H8/H41. J Hum Evol 34: 249-302. (Pubitemid 28325237)
99. Sarmiento E. 1991. Functional and phylogenetic implications of the differences in the pedal skeleton of australopithecines. Am J Phys Anthropol [Suppl] 12:157-158.
100. Scott SH, Winter DA. 1993. Biomechanical model of the human foot: kinematics during the stance phase of walking. J Biomech 26:1091-1104.
101. Senut B, Pickford M, Gommery D, Mein P, Cheboi K, Coppens Y. 2001. First hominid from the Miocene (Lukeino formation, Kenya). C R Acad Sci Ser IIA Earth Planet Sci 332:137-144.
102. Singh I. 1960. Variations in the metatarsal bones. J Anat 94: 345-350.
103. Stern JT, Susman RL. 1983. The locomotor anatomy of Australopithecus afarensis. Am J Phys Anthropol 60:279-317.
104. 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, editors. Origine(s) de la Bipédie chez les Homininés. Paris: Centre National de la Recherche Scientifique. p 99-111.
105. Strait DS, Grine FE. 2004. Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa. J Hum Evol 47:399-452.
106. Susman RL. 1983. Evolution of the human foot: evidence from the Plio-Pleistocene hominids. Foot Ankle 3:365-376.
107. Susman RL. 1988. New postcranial remains from Swartkrans and their bearing on the functional morphology and behavior of Paranthropus robustus. In: Grine FE, editor. Evolutionary history of the "robust" australopithecines. Hawthorne: Aldine de Gruyter. p 149-172.
108. Susman RL. 2004. Hominid postcranial remains from Swartkrans. In: Brain CK, editor. Swartkrans: a cave's chronicle of early man. Pretoria: Transvaal Museum Northern Flagship Institution. p 118-136.
109. Susman RL. 2008. Brief communication: evidence bearing on the status of Homo habilis at Olduvai Gorge. Am J Phys Anthropol 137:356-361.
110. Susman RL, Brain TM. 1988. New first metatarsal (SKX 5017) from Swartkrans and the gait of Paranthropus robustus. Am J Phys Anthropol 79:451-454.
111. Susman RL, de Ruiter DJ. 2004. New hominin first metatarsal (SK 1813) from Swartkrans. J Hum Evol 47:171-181.
112. Susman RL, Stern JT. 1982. Functional morphology of Homo habilis. Science 217:931-934.
113. Susman RL, Stern JT. 1991. Locomotor behavior of early hominids: epistemology and fossil evidence. In: Coppens Y, Senut B, editors. Origine(s) de la Bipédie chez les Homininés. Paris: Centre National de la Recherche Scientifique. p 121-131.
114. Susman RL, Stern JT, Jungers WL. 1984. Arboreality and bipedality in the Hadar hominids. Folia Primatol 43:113-156.
115. Susman RL, Stern JT, Jungers WL. 1985. Locomotor adaptations in the Hadar hominids. In: Delson E, editor. Ancestors: the hard evidence. New York: Alan R. Liss. p 184-192.
116. Swartz SM. 1989. The functional morphology of weight-bearing: limb joint surface area allometry in anthropoid primates. J Zool 218:441-460.
117. Tobias PV. 1998. History of the discovery of a fossilised Little Foot at Sterkfontein, South Africa, and the light that it sheds on the origins of hominin bipedalism. Mitt Berliner Ges Anthropol Ethnol Urgeschichte 19:47-56.
118. Touissant M. Macho GA, Tobias PV, Partridge TC, Hughes AR. 2003. The third partial skeleton of a late Pliocene hominin (StW 431) from Sterkfontein, South Africa. South Afr J Sci 99:215-223.
119. Tuttle RH. 1985. Ape footprints and Laetoli impressions: a response to the SUNY claims. In: Tobias PV, editor. Hominid evolution: past, present and future. New York: Alan R. Liss. p 129-133.
120. Vereecke E, D'Aoûte KD, Clercq DD, Elsaker LV, Aerts P. 2003. Dynamic plantar pressure distribution during terrestrial locomotion of bonobos (Pan paniscus). Am J Phys Anthropol 120: 373-383. (Pubitemid 36323854)
121. Vrba ES. 1985. Early hominids in southern Africa: updated observations on chronological and ecological background. In: Tobias PV, editor. Hominid evolution: past, present and future. New York: Alan R. Liss. p 195-200.
122. 39Ar dating of the Denen Dora member of the Hadar formation. Geology 22:6-10.
123. Ward CV. 2002. Interpreting the posture and locomotion of Australopithecus afarensis: where do we stand? Yearb Phys Anthropol 45:185-215.
124. Weishaupt D, Treiber K, Jacob HAC, Kundert H, Hodler J, Marincek B, Zaneti M. 2002. MR imaging of the forefoot under weight-bearing conditions: position-related changes of the neurovascular bundles and the metatarsal heads in asymptomatic volunteers. J Magn Reson Imaging 16:75-84. (Pubitemid 34700845)
125. White TD. 1980. Evolutionary implications of Pliocene hominid footprints. Science 208:175-176.
126. White TD, Suwa G. 1987. Hominid footprints at Laetoli: facts and interpretations. Am J Phys Anthropol 72:485-514. (Pubitemid 17062891)
127. Wood BA. 1974. Olduvai Bed I postcranial fossils: a reassessment. J Hum Evol 3:373-378.
128. Wunderlich RE. 1999. Pedal form and plantar pressure distribution in anthropoid primates, Ph.D. thesis, State University of New York at Stony Brook.
129. Zipfel B. 2004. Morphological variation in the metatarsal bones of selected recent and pre-pastoral humans from South Africa, Ph.D. thesis, University of the Witwatersrand.
130. Zipfel B, Kidd R. 2006. Hominin first metatarsals (SKX 5017 and SK 1813) from Swartkrans: a morphometric analysis. HOMO J Comp Hum Biol 57:117-131.
131. 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 434:754-759. (Pubitemid 44739517)