Miocene hominids and the origins of the African apes and humans

Annual Review of Anthropology

Volumn 39, Issue , 2010, Pages 67-84

  Begun, David R ^a  

^a UNIVERSITY OF TORONTO   (Canada)

Author keywords

Dryopithecus; fossil apes; Griphopithecus; hominid origins; hominines; paleobiogeography

Indexed keywords

ANCESTRY; BRAIN; FOSSIL RECORD; HOMINID; MIOCENE; MORPHOLOGY; PALEOBIOGEOGRAPHY; PALEOBIOLOGY; SEXUAL DIMORPHISM;

AFRICA; EUROPE;

DRYOPITHECUS; GRIPHOPITHECUS; HOMINIDAE; PAN;

EID: 77957991162     PISSN: 00846570     EISSN: None     Source Type: Book Series    
DOI: 10.1146/annurev.anthro.012809.105047     Document Type: Article

References (139)

1. Agustí. 2007. The biotic environments of the late Miocene hominids. See Henke & Tattersall 2008, pp. 979-1009
2. Agustí J, Sanz de Siria A, Garcés M. 2003. Explaining the end of the hominoid experiment in Europe. J. Hum. Evol. 45:145-53
3. Almécija S, Alba DM, Moyà-Solà S, Köhler M. 2007. Orang-like manual adaptations in the fossil hominoid Hispanopithecus laietanus: first steps towards great ape suspensory behaviours. Proc. R. Soc. Ser. B 274:2375-84
4. Andrews P, Martin L. 1987. Cladistic relationships of extant and fossil hominoids. J. Hum. Evol. 16:101-18
5. Ashley-Montagu MF. 1935. The premaxilla in the primates. Q. Rev. Biol. 10:32-59; 181-208
6. Beard KC, Teaford MF, Walker A. 1986. New wrist bones of Proconsul africanus and Proconsul nyanzae from Rusinga Island, Kenya. Folia Primatol. 47:97-118
7. Begun DR. 1992a. Miocene fossil hominids and the chimp-human clade. Science 257:1929-33
8. Begun DR. 1992b. Phyletic diversity and locomotion in primitive European hominids. Am. J. Phys. Anthropol. 87:311-40
9. Begun DR. 1994. Relations among the great apes and humans: new interpretations based on the fossil great ape Dryopithecus. Yearb. Phys. Anthropol. 37:11-63
10. Begun DR. 1996. Events in European hominoid evolution. Europal 10:16-20
11. Begun DR. 1999. Hominid family values: morphological and molecular data on relations among the great apes and humans. In The Mentalities of Gorillas and Orangutans: Comparative Perspectives, ed. ST Parker, RW Mitchell, HL Miles, pp. 2-42. Cambridge, UK: Cambridge Univ. Press
12. Begun DR. 2001. African and Eurasian Miocene hominoids and the origins of the Hominidae. In Hominoid Evolution and Environmental Change in the Neogene of Europe. Vol. 2. Phylogeny of the Neogene Hominoid Primates of Eurasia, ed. L de Bonis, G Koufos, P Andrews, pp. 231-53. Cambridge, UK: Cambridge Univ. Press
13. Begun DR. 2002. European hominoids. In The Primate Fossil Record, ed. W Hartwig, pp. 339-68. Cambridge: Cambridge Univ. Press
14. Begun DR. 2003. Planet of the Apes. Sci. Am. 289:74-83
15. Begun DR. 2004. Knuckle-walking and the origin of human bipedalism. In From Biped to Strider: The Emergence of Modern Human Walking, ed. DJ Meldrum, CE Hilton, pp. 9-33. New York: Kluwer
16. Begun DR. 2005. Sivapithecus is east and Dryopithecus is west, and never the twain shall meet. Anthropol. Sci. 113:53-64
17. Begun DR. 2007. Fossil record of Miocene hominoids. See Henke & Tattersall 2007, pp. 921-77. Berlin: Springer
18. Begun DR. 2009. Dryopithecins, Darwin, de Bonis, and the European origin of the African apes and human clade. Geodiversitas 31:789-816
19. Begun DR, Geraads D, Güleç E. 2003a. The Çandir hominoid locality: Implications for the timing and pattern of hominoid dispersal events. Cour. Forschung. Inst. Senckenberg. 240:251-65
20. Begun DR, Güleç E. 1998. Restoration of the Type and Palate of Ankarapithecus meteai: Taxonomic, Phylogenetic, and Functional Implications. Am. J. Phys. Anthropol. 105:279-314
21. Begun DR, Güleç E, Geraads D. 2003b. Dispersal patterns of Eurasian hominoids: Implications from Turkey. Deinsea 10:23-39
22. Begun DR, Kordos L. 1997. Phyletic affinities and functional convergence in Dryopithecus and other Miocene and living hominids. See Begun et al. 1997, pp. 291-316
23. Begun DR, Kordos L. 2004. Cranial evidence of the evolution of intelligence in fossil apes. In The Evolution of Thought: Evolutionary Origins of Great Ape Intelligence, ed. AE Russon, DR Begun, pp. 260-79. Cambridge: Cambridge Univ. Press
24. Begun DR, Nargolwalla M, Kordos L. 2008. Revision of the Dryopithecinae: Phylogenetic and paleobiogeographic implications. Am J. Phys. Anthropol. S46:66
25. Begun DR, Nargolwalla MC. 2004. Late Miocene hominid biogeography: some recent perspectives. Evol. Anthropol. 13:234-38
26. Begun DR, Teaford MF, Walker A. 1994. Comparative and functional anatomy of Proconsul phalanges from the Kaswanga primate site, Rusinga Island, Kenya. J. Hum. Evol. 26:89-165
27. Begun DR, Ward CV, Deane AS, Kivell TL, Nargolwalla MC, Taylor ND. 2006. Stem hominine or hominid? The phylogeny and functional anatomy of Pierolapithecus. Am. J. Phys. Anthropol. S42:63
28. Begun DR, Ward CV, Rose MD, eds. 1997. Function, Phylogeny and Fossils: Miocene Hominoid Evolution and Adaptations. New York: Plenum
29. Benefit BR, McCrossin ML. 1995. Miocene Hominoids and hominid origins. Annu. Rev. Anthropol. 24:237-56
30. Bernor RL. 2007. New apes fill the gap. Proc. Natl. Acad. Sci. USA 104:19661-62
31. Bernor RL, Rook L. 2003. Palaeozoogeography of the Rudabanya fauna. Palaeontolograph. Ital. 89 (2002):21-25
32. Bonis L de. 1983. Phyletic relationships of Miocene hominoids and higher primate classification. See Corruccini & Ciochon 1983, pp. 625-49
33. Bonis L de, Bouvrain G, Geraads D, Koufos G. 1990. New hominid skull material form the late Miocene of Macedonia in Northern Greece. Nature 345:712-14
34. Bonis L de, Bouvrain G, Geraads D, Melentis J. 1974. Première découverte d'un primate hominöide dans le Miocène supérieur de Macédoine (Grèce). C. R. Acad. Sci. Paris Ser. D 278:3063-66
35. Bonis L de, Bouvrain G, Koufos G, Malentis J. 1986. Succession and dating of the late Miocene primates of Macedonia. See Else & Lee 1986, pp. 107-14
36. Bonis L de, Koufos G. 1993. The face and mandible of Ouranopithecus macedoniensis: description of new specimens and comparisons. J. Hum. Evol. 24:469-91 (Pubitemid 23394689)
37. Bonis L de, Koufos G. 1994. Our ancestors' ancestor: Ouranopithecus is a Greek link in human ancestry. Evol. Anthropol. 3:75-83
38. Bonis L de, Koufos G. 1997. The phylogenetic and functional implications of Ouranopithecus macedoniensis. See Begun et al. 1997, pp. 317-26
39. Burchak-Abramovitsch NO, Gabashvili EG. 1950. Discovery of a fossil anthropoid in Georgia (in Russian). Priroda, Moscow 9:70-72
40. Cave AJE. 1949. Notes on the nasal fossa of a young Chimpanzee. Proc. Zool. Soc. London 119:61-63
41. Cave AJE. 1961. The frontal sinus of the gorilla. Proc. Zool. Soc. London 136:359-73
42. Cave AJE, Haines RW. 1940. The paranasal sinues of the anthropoid apes. J. Anat. 74:493-523
43. Chaimanee Y, Jolly D, Benammi M, Tafforeau P, Duzer D, et al. 2003. A middle Miocene hominoid from Thailand and orangutan origins. Nature 422:61-65
44. Chaimanee Y, Suteethorn V, Pratueng J, Vidthayanon C, Marandat B, Jaeger J-J. 2004. A new orang-utan relative from the Late Miocene of Thailand. Nature 427:439-41
45. Corruccini RL, Ciochon RS, eds. 1983. New Interpretations of Ape and Human Ancestry. New York: Plenum
46. Darwin C. 1871. The Descent of Man. London: Murray
47. Deane AS. 2007. Inferring dietary behavior from Miocene hominoids: a high resolution morphometric approach to incisal crown curvature. PhD thesis. Univ. Toronto, Toronto. 338 pp.
48. Deane AS, Begun DR. 2008. Broken fingers: retesting locomotor hypotheses for fossil hominoids using fragmentary proximal phalanges and high-resolution polynomial curve fitting (HR-PCF). J. Hum. Evol. 55:691-701
49. Else JC, Lee PC, eds. 1986. Primate Evolution. New York: Cambridge Univ. Press
50. Fortelius M, Werdelin L, Andrews P, Bernor RL, Gentry A, et al. 1996. Provinciality, diversity, turnover, and paleoecology in land mammal faunas of the later Miocene of Western Eurasia. In The Evolution of Western Eurasian Neogene Mammal Faunas, ed. RL Bernor, V Fahlbusch, H-W Mittmann, pp. 415-48. New York: Columbia Univ. Press
51. Goodman M, Porter CA, Czelusniak J, Page SL, Schneider H, et al. 1998. Toward a phylogenetic classification of primates based on DNA evidence complemented by fossil evidence. Mol. Phylogen. Evol. 9:585-98
52. Greenfield LO. 1974. Taxonomic reassessment of two Ramapithecus specimens. Folia Primatol. 22:97-115
53. Greenfield LO. 1980. A late divergence hypothesis. Am. J. Phys. Anthropol. 52:351-65
54. Güleç E, Sevim A, Pehlevan C, Kaya F. 2007. A new great ape from the late Miocene of Turkey. Anthropol. Sci. 115:153-58
55. Harrison T. 2010. Apes among the tangled branches of human origins. Science 327:532-34
56. Hartwig W, ed. 2002. The Primate Fossil Record. Cambridge: Cambridge Univ. Press
57. Heizmann E, Begun DR. 2001. The oldest European hominoid. J. Hum. Evol. 41:465-81
58. Henke W, Tattersall I, eds. 2007. Handbook of Palaeoanthropology. Vol. 2: Primate Evolution and Human Origins. Berlin: Springer
59. Hill A, Ward S. 1988. Origin of the Hominidae: the record of African large hominoid evolution between 14 my and 4 my. Yearb. Phys. Anthropol. 31:48-83
60. Hopwood AT. 1933. Miocene primates from Kenya. Zool. J. Linnean Soc. London 38:437-64
61. Ishida H, Kunimatsu Y, Takano T, Nakano Y, Nakatsukasa M. 2004. Nacholapithecus skeleton from the middle Miocene of Kenya. J. Hum. Evol. 46:1-35
62. Ishida H, Nakatsukasa M, Kunimatsu Y. 2000. Erection of a new genus and species: Nacholapithecus kerioi for a middle Miocene hominoid from Nachola area, northern Kenya. Anthropol. Sci. 108:92
63. Jablonski NG. 2002. Fossil Old World monkeys: the late Neogene. See Hartwig 2002, pp. 255-99
64. Kappelman J, Kelley J, Pilbeam D, Sheikh KA, Ward S, et al. 1991. The earliest occurrence of Sivapithecus from the middle Miocene Chinji Formation of Pakistan. J. Hum. Evol. 21:61-73
65. Kay RF, Ungar PS. 1997. Dental evidence for diet in some Miocene catarrhines with comments on the effects of phylogeny on the interpretation of adaptation. See Begun et al. 1997, pp. 131-51
66. Kelley J. 2002. The hominoid radiation in Asia. See Hartwig 2002, pp. 369-84
67. Kelley J, Andrews P, Alpagut B. 2008. A new hominoid species from the middle Miocene site of Pasalar, Turkey. J. Hum. Evol. 54:455-79 (Pubitemid 351448143)
68. Kordos L, Begun DR. 2001a. A new cranium of Dryopithecus from Rudabánya, Hungary. J. Hum. Evol. 41:689-700
69. Kordos L, Begun DR. 2001b. Primates from Rudabánya: allocation of specimens to individuals, sex and age categories. J. Hum. Evol. 40:17-39
70. Koufos GD. 2003. Late Miocene mammal events and biostratigraphy in the Eastern Mediterranean. Deinsea 10:343-71
71. Koufos GD. 2007. Potential hominoid ancestors for Hominidae. In Handbook of Palaeoanthropology. Vol. 3: Phylogeny of Hominids, ed. W Henke, I Tattersall, pp. 1347-77. Berlin: Springer
72. Kretzoi M. 1969. Geschichte der Primaten und der Hominisation. Symp. Biol. Hung. 9:3-11
73. Krogman WM. 1930a. Studies in the growth change in the skull and face of Anthropoids. II Ectocranial and endocranial suture closure in the anthropoids and apes. Am. J. Anat. 46:315-53
74. Krogman WM. 1930b. Studies in the growth change in the skull and face of Anthropoids. V. Growth changes in the skull and face of the ourang-utan. Am. J. Anat. 47:343-65
75. Kunimatsu Y, Nakatsukasa M, Sawada Y, Sakai T, Hyodo M, et al. 2007. A new Late Miocene great ape from Kenya and its implications for the origins of African great apes and humans. Proc. Natl. Acad. Sci. USA 104:19220-25
76. Lartet E. 1856. Note sur un grand singe fossile qui se rattache au groupe des singes supérieurs. C. R. Acad. Sci. 43:219-23
77. Leakey LSB. 1962. A new Lower Pliocene fossil primate from Kenya. Ann. Mag. Nat. Hist. 13:689-96
78. Leakey M, Walker A. 1997. Afropithecus: function and phylogeny. See Begun et al. 1997, pp. 225-39
79. Leakey REF, Leakey MG, Walker AC. 1988. Morphology of Afropithecus turkanesis from Kenya. Am. J. Phys. Anthropol. 76:289-307
80. Lovejoy CO. 2009. Reexamining human origins in light of Ardipithecus ramidus. Science 326:74 e1-8
81. Lovejoy CO, Latimer B, Suwa G, Asfaw B, White TD. 2009a. Combining prehension and propulsion: the foot of Ardipithecus ramidus. Science 326:72 e1-8
82. Lovejoy CO, Simpson SW, White TD, Asfaw B, Suwa G. 2009b. Careful climbing in the Miocene: The forelimbs of Ardipithecus ramidus and humans are primitive. Science 326:70 e1-8
83. 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 326:73-106
84. Lovejoy CO, Suwa G, Spurlock L, Asfaw B, White TD. 2009d. The pelvis and femur of Ardipithecus ramidus: the emergence of upright walking. Science 326:71 e1-6
85. Madar SI, Rose MD, Kelley J, MacLatchy L, Pilbeam D. 2002. New Sivapithecus postcranial specimens from the Siwaliks of Pakistan. J. Hum. Evol. 42:705-52
86. Martin L, Andrews P. 1993. Renaissance of Europe's ape. Nature 365:494
87. Moyà-Solà M, Köhler M. 1995. New partial cranium of Dryopithecus Lartet, 1863 (Hominoidea, Primates) from the upper Miocene of Can Llobateres, Barcelona, Spain. J. Hum Evol. 29:101-39
88. Moyà-Solà S, Alba DM, Almecija S, Casanovas-Vilar I, Köhler M, et al. 2009a. A unique middle Miocene European hominoid and the origins of the great ape and human clade. Proc. Natl. Acad. Sci. USA 106:9601-6
89. Moyà-Solà S, Köhler M. 1996. A Dryopithecus skeleton and the origins of great ape locomotion. Nature 379:156-59
90. Moyà-Solà S, Köhler M, Alba DM, Casanovas-Vilar I, Galindo J. 2004. Pierolapithecus catalaunicus, a new middle Miocene great ape from Spain. Science 306:1339-44
91. Moyà-Solà S, Köhler M, Alba DM, Casanovas-Vilar I, Galindo J, et al. 2009b. First partial face and upper dentition of the middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallès-Penèdes Basin, Catalonia, NE Spain): taxonomic and phylogenetic implications. Am. J. Phys. Anthropol. 139:126-45
92. Nakatsukasa M, Yamanaka A, Kunimatsu Y, Shimizu D, Ishida H. 1998. A newly discovered Kenyapithecus skeleton and its implications for the evolution of positional behavior in Miocene East African hominoids. J. Hum. Evol. 34:659-64
93. Nakatsukasa M, Tsujikawa H, Shimizu D, Takano T, Kunimatsu Y, et al. 2003. Definitive evidence for tail loss in Nacholapithecus, an East African Miocene hominoid. J. Hum. Evol. 45:179-86
94. Nargolwalla MC. 2009. Euasian middle and late Miocene hominoid paleobiogeography and the geographic origins of the Homininae. PhD thesis. Univ. Toronto, Toronto. 259 pp.
95. Nargolwalla MC, Hutchison MP, Begun DR. 2006. Middle and late Miocene terrestrial vertebrate localities and paleoenvironments in the Pannonian Basin. Beitr. Paläontolog. 30:319-32
96. Pickford M. 1986. The geochronology of Miocene higher primate faunas of East Africa. See Else & Lee 1986, pp. 21-33
97. Pickford M, Morales J. 1994. Biostratigraphy and palaeobiogeography of East Africa and the Iberian peninsula. Palaeogeog. Palaeoclimatol. Palaeoecol. 112:297-322
98. Pickford M, Sawada Y, Tayama R, Matsuda Y-K, Itaya T, et al. 2006. Refinement of the age of the Middle Miocene Fort Teman Beds, Western Kenya, and its implications for Old World biochronology. C. R. Geosci. 338:545-55
99. Pilbeam D, Young N. 2004. Hominoid evolution: synthesizing disparate data. C. R. Palevol. 3:305-21
100. Pilbeam DR. 1982. New hominoid skull material from the Miocene of Pakistan. Nature 295:232-34
101. Pilbeam DR. 1997. Research on Miocene hominoids and hominid origins: the last three decades. See Begun et al. 1997, pp. 13-28
102. Pilbeam DR, Simons EL. 1971. Humerus of Dryopithecus from Saint Gaudens, France. Nature 229:406-7
103. Richmond BG, Begun DR, Strait DS. 2001. Origin of human bipedalism: the knuckle-walking hypothesis reconsidered. Yearb. Phys. Anthropol. 44:70-105
104. Rook L, Bernor RL. 2004. Recent advances on multidisciplinary research at Rudabánya, late Miocene (MN 9), Hungary: a compendium. Ancestry of the African ape/human clade? Palaeontolograph. Ital. 89:30-31
105. Rose MD. 1986. Further hominoid postcranial specimens from the Late Miocene Nagri Formation of Pakistan. J. Hum. Evol. 15:333-67
106. Rose MD. 1988. Another look at the anthropoid elbow. J. Hum. Evol. 17:193-224
107. Rose MD. 1989. New postcranial specimens of catarrhines from the Middle Miocene Chinji Formation, Pakistan: description and a discussion of proximal humeral functional morphology in anthropoids. J. Hum. Evol. 18:131-62
108. Rose MD. 1997. Functional and phylogenetic features of the forelimb in Miocene hominoids. See Begun et al. 1997, pp. 79-100
109. Rose MD, Nakano Y, Ishida H. 1996. Kenyapithecus postcranial specimens from Nachola, Kenya. Afr. Study Monog. 24:3-56
110. Rossie JB. 2005. Anatomy of the nasal cavity and paranasal sinuses in Aegyptopithecus and Early Miocene African catarrhines. Am. J. Phys. Anthropol. 126:250-67
111. Schultz AH. 1926. Fetal growth of man and other primates. Q. Rev. Biol. 1:465-521
112. Schultz AH. 1936. Characters common to higher primates and characters specific to man. Q. Rev. Biol. 11:425-55
113. Schwartz JH. 1997. Lufengpithecus and hominoid phylogeny. Problems in delineating and evaluating phylogenetically relevant characters. See Begun et al. 1997, pp. 363-88
114. Shea BT. 1985. On aspects of skull form in African apes and orangutans, with implications for hominoid evolution. Am. J. Phys. Anthropol. 68:329-42
115. Shea BT. 1988. Phylogeny and skull form in the hominoid primates. In Orang-Utan Biology, ed. J Schwartz, pp. 233-45. New York: Oxford Univ. Press
116. Simons EL, Pilbeam DR. 1965. Preliminary revision of the Dryopithecinae (Pongidae, Anthropoidea). Folia Primatol. 3:81-152
117. Solounias N, Plavcan JM, Quade J, Witmer L. 1999. The paleoecology of the Pikermian Biome and the Savanna myth. In The Evolution of Neogene Terrestrial Ecosystems in Europe, ed. J Agusti, L Rook, P Andrews, pp. 436-53. Cambridge: Cambridge Univ. Press
118. Spassov N. 2009. Bulgaria, national museum of natural history. Soc. Vert. Paleontol. News Bull. 197:12
119. Spassov N, Geraads D. 2008. The latest prehumans of Europe: discovery of a late Miocene hominoid in Bulgaria of about 7 Ma. Bul. Acad. Sci. 2:2-4
120. Stewart C-B, Disotell TR. 1998. Primate evolution-in and out of Africa. Curr. Biol. 8:582-88
121. 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
122. Suwa G, Asfaw B, Kono RT, Kubo D, Lovejoy CO, White TD. 2009a. The Ardipithecus ramidus skull and its implications for hominid origins. Science 326:68-87
123. Suwa G, Kono RT, Katoh S, Asfaw B, Beyene Y. 2007. A new species of great ape from the late Miocene epoch in Ethiopia. Nature 448:921-24
124. Suwa G, Kono RT, Simpson SW, Asfaw B, Lovejoy CO, White TD. 2009b. Paleobiological implications of the Ardipithecus ramidus dentition. Science 326:69-99
125. Szalay F, Delson E. 1979. Evolutionary History of the Primates. New York: Academic
126. Ungar PS. 1996. Dental microwear of European Miocene catarrhines: evidence for diets and tooth use. J. Hum. Evol. 31:335-66
127. Ungar PS, Kay RF. 1995. The dietary adaptations of European Miocene catarrhines. Proc. Natl. Acad. Sci. USA 92:5479-81
128. Ward CS, Brown B. 1986. The facial skeleton of Sivapithecus indicus. In Comparative Primate Biology, ed. DR Swindler, J Erwin, pp. 413-52. New York: Alan R. Liss
129. Ward CV. 1993. Torso morphology and locomotion in Proconsul nyanzae. Am. J. Phys. Anthropol. 92:291-328
130. Ward CV. 1997a. Functional anatomy and phyletic implications of the hominoid trunk and hindlimb. See Begun et al. 1997, pp. 101-30
131. Ward CV. 2007. Postcranial and locomotor adaptations of hominoids. See Henke & Tattersall 2007, pp. 1011-30
132. Ward CV, Walker AC, Teaford MF. 1991. Proconsul did not have a tail. J. Hum. Evol. 21:215-20
133. Ward S. 1997b. The taxonomy and phylogenetic relationships of Sivapithecus revisited. See Begun et al. 1997, pp. 269-90
134. Ward S, Brown B, Hill A, Kelley J, Downs W. 1999. Equatorius: a new hominoid genus from the middle Miocene of Kenya. Science 285:1382-86
135. Ward SC, Kimbel WH. 1983. Subnasal alveolar morphology and the systemic postion of Sivapithecus. Am. J. Phys. Anthropol. 61:157-71
136. Ward SC, Pilbeam DR. 1983. Maxillofacial morphology of Miocene Hominoids from Africa and Indo-Pakistan. See Corruccini & Ciochon 1983, pp. 211-38
137. White TD. 2002. Earliest hominids. See Hartwig 2002, pp. 407-17
138. White TD, Asfaw B, Beyene Y, Haile-Selassie Y, Lovejoy CO, et al. 2009. Ardipithecus ramidus and the paleobiology of early hominids. Science 326:64-86
139. Wood B, Richmond BG. 2000. Human evolution: taxonomy and paleobiology. J. Anat. 196:19-60