The Evolution of the Primate Hand pp 485-514 | Cite as
The Hands of Miocene Hominoids
Abstract
Hands of extant hominoids are highly derived compared with those of non-hominoid catarrhines. The evolution of the ape hand started from an appendage very well suited for powerful pollical-assisted grasping that supplied a balancing function in response to the loss of tail as seen in the early Miocene Proconsul (or Ekembo). Nacholapithecus from the early Middle Miocene of Africa had hands that were adapted for a primitive pollical-assisted grasping function, but absolutely large and more powerful. This might be an initial change toward forelimb-dominated positional behaviors while retaining the generally plesiomorphic postcrania. Pierolapithecus from the late Middle Miocene of Europe is clearly derived and may represent an ‘intermediate’ form evolving toward the extant great ape-like suspension/orthogrady in a mosaic way: enhanced ulnar deviation and midcarpal supination, yet lacking complete radial-side loading of the wrist and specializations for a hook-like grip. Late Miocene Eurasian great apes exhibit varying degrees (or modes) of suspension and orthograde adaptations. However, the hand anatomy that invokes such behaviors is not always morphologically identical across these apes, suggesting independent specialization for those behaviors. A reduction of the pollex in relation to the fingers, a trait common in extant apes, is not observed in any of these species. This may suggest that above-branch quadrupedalism/climbing with power grasping was an indispensable locomotor behavior, even in these apes. It is unclear in the fossil record when and where extant ape-like hands evolved.
Keywords
Late Miocene Ulnar Deviation Proximal Surface Polynomial Curve Fitting Midcarpal JointNotes
Acknowledgments
We thank the editors for inviting us to contribute to this volume and providing us with thoughtful comments on an early version of the manuscript. We are grateful to the National Museums of Kenya for permission to study original specimens under their care. We thank Ashley Hammond, Jay Kelley, Tracy Kivell, Salvador Moyà-Solà, Lorenzo Rook, and Randy Susman for providing us original fossil photographs. This work is supported by grants from the JSPS Grant-in-Aid (#22255006, 25257408) to M.N., from the Spanish Ministerio de Economía y Competitividad (CGL2014-54373-P) and National Science Foundation (NSF-BCS 1316947) to S.A., from the American Association of Physical Anthropologists (Professional Development Program) to S.A., and from the Natural Sciences and Engineering Research Council of Canada, the National Geographic Society, and the University of Toronto to D.R.B.
References
- Alba DM (2012) Fossil apes from the Vallès-Penedès Basin. Evol Anthropol 21:254–269PubMedCrossRefGoogle Scholar
- Alba DM, Almécija S, Moyà-Solà S (2010) Locomotor inferences in Pierolapithecus and Hispanopithecus: reply to Deane and Begun (2008). J Hum Evol 59:143–149PubMedCrossRefGoogle Scholar
- Alba DM, Almécija S, Casanovas-Vilar I, Méndez JM, Moyà-Solà S (2012) A partial skeleton of the fossil great ape Hispanopithecus laietanus from Can Feu and the mosaic evolution of crown-hominoid positional behaviors. PLoS One 7:e39617PubMedPubMedCentralCrossRefGoogle Scholar
- Allen KL, McCrossin ML (2007) Functional morphology of the Kenyapithecus hand from Maboko Island (Kenya). Am J Phys Anthropol 132(Suppl 44):62Google Scholar
- 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 behaviors. Proc R Soc B 274:2375–2384PubMedPubMedCentralCrossRefGoogle Scholar
- Almécija S, Alba DM, Moyà-Solà S (2009) Pierolapithecus and the functional morphology of Miocene ape hand phalanges: Paleobiological and evolutionary implications. J Hum Evol 57:284–297PubMedCrossRefGoogle Scholar
- Almécija S, Alba DM, Moyà-Solà S (2012) The thumb of Miocene apes: new insights from Castell de Barberà (Catalonia, Spain). Am J Phys Anthropol 148:436–450PubMedCrossRefGoogle Scholar
- Almécija S, Shrewsbury M, Rook L, Moyà-Solà S (2014) The morphology of Oreopithecus bambolii pollical distal phalanx. Am J Phys Anthropol 153:582–597PubMedCrossRefGoogle Scholar
- Almécija S, Smaers JB, Jungers WL (2015) The evolution of human and ape hand proportions. Nat Commun 6:7717Google Scholar
- Beard KC, Teaford MF, Walker A (1986) New wrist bones of Proconsul africanus and P. nyanzae from Rusinga Island, Kenya. Folia Primatol 47:97–118PubMedCrossRefGoogle Scholar
- Begun DR (1988) Catarrhine phalanges from the late Miocene (Vallesian) of Rudabánya, Hungary. J Hum Evol 17:413–438CrossRefGoogle Scholar
- Begun DR (1992) Miocene fossil hominids and the chimp-human clade. Science 257:1929–1933PubMedCrossRefGoogle Scholar
- Begun DR (1993) New catarrhine phalanges from Rudabánya (Northeastern Hungary) and the problem of parallelism and convergence in hominoid postcranial morphology. J Hum Evol 24:373–402CrossRefGoogle Scholar
- Begun DR (1994) Relations among the great apes and humans: new interpretations based on the fossil great ape Dryopithecus. Yearb Phys Anthropol 37:11–63Google Scholar
- Begun DR (2002) European hominoids. In: Hartwig W (ed) The primate fossil record. Cambridge University Press, Cambridge, pp 231–253Google Scholar
- 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 Publishers, New York, pp 9–33CrossRefGoogle Scholar
- Begun DR (2007) Fossil record of Miocene hominoids. In: Henke W, Tattersall I (eds) Handbook of paleoanthropology. Springer, Berlin, pp 921–977CrossRefGoogle Scholar
- Begun DR (2009) Dryopithecins, Darwin, de Bonis, and the European origin of the African apes and human clade. Geodiversitas 31:789–816CrossRefGoogle Scholar
- Begun DR (2010) Miocene hominids and the origins of the African apes and humans. Annu Rev Anthropol 39:67–84CrossRefGoogle Scholar
- Begun DR, Kivell TL (2011) Knuckle-walking in Sivapithecus? The combined effects of homology and homoplasy with possible implications for pongine dispersals. J Hum Evol 60:158–170PubMedCrossRefGoogle Scholar
- Begun DR, Nargolwalla MC, Kordos L (2012) European Miocene hominids and the origin of the African ape and human clade. Evol Anthropol 21:10–23PubMedCrossRefGoogle Scholar
- Begun DR, Kordos L (2011) New postcrania of Rudapithecus hungaricus from Rudabánya (Hungary). Am J Phys Anthropol 144(Suppl 52):86 (abstract)Google Scholar
- 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 26:89–165CrossRefGoogle Scholar
- Begun DR, Ward CV (2005) Comment on “Pierolapithecus catalaunicus, a new Middle Miocene great ape from Spain”. Science 308:203cCrossRefGoogle Scholar
- Böhme M, Aziz HA, Prieto J, Bachtadse V, Schweigert G (2011) Bio-magnetostratigraphy and environment of the oldest Eurasian hominoid from the Early Miocene of Engelswies (Germany). J Hum Evol 61:332–339PubMedCrossRefGoogle Scholar
- De Bonis L, Koufos GD (1994) Our ancestors’ ancestor: Ouranopithecus is a Greek link in human ancestory. Evol Anthropol 3:75–83CrossRefGoogle Scholar
- De Bonis L, Koufos GD (2014) First discovery of postcranial bones of Ouranopithecus macedoniensis (Primates, Hominoidea) from the late Miocene of Macedonia (Greece). J Hum Evol 74:21–36PubMedCrossRefGoogle Scholar
- Cartmill M, Milton K (1977) The lorisiform wrist joint and the evolution of “brachiating” adaptations in the Hominoidea. Am J Phys Anthropol 47:249–272PubMedCrossRefGoogle Scholar
- Casanovas-Vilar I, Alba DM, Garcés M, Robles JM, Moyà-Solà S (2011) Updated chronology for the Miocene hominoid radiation in Western Eurasia. Proc Natl Acad Sci U S A 108:5554–5559PubMedPubMedCentralCrossRefGoogle Scholar
- Corruccini RS, Ciochon RL, McHenry HM (1975) Osteometric shape relationships in the wrist joint of some anthropoids. Folia Primatol 24:250–274PubMedCrossRefGoogle Scholar
- Cote SM (2004) Origins of the African hominoids: an assessment of the palaeobiogeographical evidence. C R Palevol 3:323–340CrossRefGoogle Scholar
- 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 55:691–701PubMedCrossRefGoogle Scholar
- DeSilva JM, Morgan ME, Barry JC, Pilbeam D (2010) A hominoid distal tibia from the Miocene of Pakistan. J Hum Evol 58:147–154PubMedCrossRefGoogle Scholar
- de Terra H (1956) New approach to the problem of man's origin. Science 124:1282–1285PubMedCrossRefGoogle Scholar
- Erikson GE (1963) Brachiation in New World monkeys and in anthropoid apes. Symp Zool Soc Lond 10:135–164Google Scholar
- Ersoy A, Kelley J, Andrews P, Alpagut B (2008) Hominoid phalanges from the middle Miocene site of Paşalar, Turkey. J Hum Evol 54:518–529PubMedCrossRefGoogle Scholar
- Godinot M, Beard KC (1991) Fossil primate hands: a review and an evolutionary inquiry emphasizing early forms. Hum Evol 6:307–354CrossRefGoogle Scholar
- Hamrick MW, Meldrum DJ, Simons EL (1995) Anthropoid phalanges from the Oligocene of Egypt. J Hum Evol 28:121–145CrossRefGoogle Scholar
- Harrison T (1986) A reassessment of the phylogenetic relationships of Oreopithecus bambolii Gervais. J Hum Evol 15:541–583CrossRefGoogle Scholar
- 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. CNRS, Paris, pp 235–244Google Scholar
- Harrison T (2010) Dendropithecoidea, Proconsuloidea and Hominoidea (Catarrhini, Primates). In: Werdelin L, Sanders WJ (eds) Cenozoic mammals of Africa. University of California Press, Berkeley, pp 429–469CrossRefGoogle Scholar
- Heizmann EPJ, Begun DR (2001) The oldest European hominoid. J Hum Evol 41:463–481PubMedCrossRefGoogle Scholar
- Inouye SE (1994) Ontogeny of knuckle-walking hand postures in African apes. J Hum Evol 26:459–485CrossRefGoogle Scholar
- Ishida H, Kunimatsu T, Nakatsukasa M, Nakano Y (1999) New hominoid genus from the middle Miocene of Nachola, Kenya. Anthropol Sci 107:189–191CrossRefGoogle Scholar
- Ishida H, Kunimatsu Y, Takano T, Nakano Y, Nakatsukasa M (2004) Nacholapithecus skeleton from the Middle Miocene of Kenya. J Hum Evol 46:67–101CrossRefGoogle Scholar
- Jenkins FA Jr (1981) Wrist rotation in primates: a critical adaptation for brachiators. Symp Zool Soc Lond 48:429–451Google Scholar
- Jenkins FA Jr, Fleagle JG (1975) Knuckle-walking and the functional anatomy of the wrists in living apes. In: Tuttle RH (ed) Primate functional morphology and evolution. Mouton, The Hague, pp 213–231Google Scholar
- Jouffroy FK, Godinot M, Nakano Y (1991) Biometrical characteristics of primate hands. Hum Evol 6:269–306CrossRefGoogle Scholar
- Jungers WL (1987) Body size and morphometric affinities of the appendicular skeleton in Oreopithecus bambolii (IGF11778). J Hum Evol 16:445–456CrossRefGoogle Scholar
- Kappelman J, Richmond BG, Seiffert ER, Maga AM, Ryan TM (2003) Hominoidea (Primates). In: Fortelius M, Kappelman J, Sen S, Bernor RL (eds) Geology and paleontology of the Miocene Sinap Formation, Turkey. Columbia University Press, New York, pp 90–124Google Scholar
- Kelley J (1997) Paleobiological and phylogenetic significance of life history in Miocene hominoids. In: Begun DR, Ward CV, Rose MD (eds) Function, phylogeny, and fossils: Miocene hominoid evolution and adaptations. Plenum Press, New York, pp 173–208CrossRefGoogle Scholar
- Kelley J (2001) Phylogeny and sexually dimorphic characters: canine reduction in Ouranopithecus. In: Koufos GD, Andrews P, de Bonis L (eds) Phylogeny of the Neogene hominoid primates of Eurasia. Cambridge University Press, Cambridge, pp 269–283CrossRefGoogle Scholar
- Kelley J (1988) A new large species of Sivapithecus from the Siwaliks of Pakistan. J Hum Evol 17:305–324CrossRefGoogle Scholar
- Kelley J (2002) The hominoid radiation in Asia. In: Hartwig WC (ed) The primate fossil record. Cambridge University Press, Cambridge, pp 369–384Google Scholar
- Kelley J (2005) Twenty-five years contemplating Sivapithecus taxonomy. In: Lieberman DE, Smith RH, Kelley L (eds) Interpreting the past: essays on human, primate, and mammal evolution. Brill, Boston, pp 123–143Google Scholar
- Kelley J, Andrew P, Alpagut B (2008) A new hominoid species from the middle Miocene site of Paşalar, Turkey. J Hum Evol 54:455–479PubMedCrossRefGoogle Scholar
- Kivell TL, Begun DR (2009) New primate carpal bones from Rudabánya (late Miocene, Hungary): taxonomic and functional implications. J Hum Evol 57:697–709PubMedCrossRefGoogle Scholar
- Kunimatsu Y, Nakatsukasa M, Sawada Y, Sakai T, Hyodo M, Hyodo H, Itaya T, Nakaya H, Saegusa H, Mazurier A, Saneyoshi M, Tsujikawa H, Yamamoto A, Mbua E (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 U S A 104:19220–19225PubMedPubMedCentralCrossRefGoogle Scholar
- Leakey M, Walker A (1997) Afropithecus: function and phylogeny. In: Begun DR, Ward CV, Rose MD (eds) Function, phylogeny, and fossils: Miocene hominoid evolution and adaptation. Plenum Press, New York, pp 225–239CrossRefGoogle Scholar
- Leakey RE, Leakey MG, Walker AC (1988) Morphology of Afropithecus turkanensis from Kenya. Am J Phys Anthropol 76:289–307PubMedCrossRefGoogle Scholar
- Le Minor JM (1988) The ventral metacarpo- and metatarsophalangeal sesamoid bones: comparative and evolutionary aspects. Gebenbaurs Morphol Jahrb 134:693–731Google Scholar
- Lewis OJ (1969) The hominoid wrist joint. Am J Phys Anthropol 30:251–268PubMedCrossRefGoogle Scholar
- Lewis OJ (1972) Osteological features characterizing the wrists of monkeys and apes, with a reconsideration of this region in Dryopithecus (Proconsul) africanus. Am J Phys Anthropol 36:45–58PubMedCrossRefGoogle Scholar
- Lewis OJ (1989) Functional morphology of the evolving hand and foot. Clarendon Press, OxfordGoogle Scholar
- Lovejoy CO, Simpson SW, White TD, Asfaw B, Suwa G (2009a) Careful climbing in the Miocene: the forelimbs of Ardipithecus ramidus and humans are primitive. Science 326:70e71–70e78Google Scholar
- Lovejoy CO, Suwa G, Simpson SW, Matternes JH, White TD (2009b) The great divides: Ardipithecus ramidus reveals the postcrania of our last common ancestors with African apes. Science 326:100–106PubMedGoogle Scholar
- Madar SI, Rose MD, Kelley J, MacLatchy L, Pilbeam D (2002) New Sivapithecus postcranial specimens from the Siwaliks of Pakistan. J Hum Evol 42:705–752PubMedCrossRefGoogle Scholar
- McCrossin ML (1994) The phylogenetic relationships, adaptations, and ecology of Kenyapithecus. Ph.D. dissertation, University of California, BerkeleyGoogle Scholar
- McCrossin ML, Benefit BR (1994) Maboko Island and the evolutionary history of Old World monkeys and apes. In: Corruccini RS, Ciochon RL (eds) Integrative paths to the past: paleoanthropological advances in honor of F. Clark Howell. Prentice Hall, Englewood Cliffs, pp 95–122Google Scholar
- McCrossin ML, Benefit BR, Gitau SN, Palmer AK, Blue KT (1998) Fossil evidence for the origins of terrestriality among Old World higher primates. In: Strasser E, Fleagle J, Rosenberger A, McHenry H (eds) Primate locomotion. Plenum Press, New York, pp 353–396CrossRefGoogle Scholar
- McHenry HM, Corruccini RS (1983) The wrist of Proconsul africanus and the origin of hominoid postcranial adaptations. In: Ciochon RL, Corruccini RS (eds) New interpretations of ape and human ancestry. Plenum Press, New York, pp 353–366CrossRefGoogle Scholar
- McNulty KP, Begun DR, Kelley J, Manthi FK, Mbua EN (2015) A systematic revision of Proconsul with the description of a new genus of early Miocene hominoid. J Hum Evol 84:42–61Google Scholar
- Meldrum DJ, Pan Y (1988) Manual proximal phalanx of Laccopithecus robustus from the Latest Miocene site of Lufeng. J Hum Evol 17:719–731CrossRefGoogle Scholar
- Morbeck ME (1975) Dryopithecus africanus forelimb. J Hum Evol 4:39–46CrossRefGoogle Scholar
- Moyà-Solà S, Köhler M (1996) A Dryopithecus skeleton and the origins of great-ape locomotion. Nature 379:156–159PubMedCrossRefGoogle Scholar
- Moyà-Solà S, Köhler M, Rook L (1999) Evidence of hominid-like precision grip capability in the hand of the Miocene ape Oreopithecus. Proc Natl Acad Sci U S A 96:313–317PubMedPubMedCentralCrossRefGoogle Scholar
- Moyà-Solà S, Köhler M, Alba DM, Casanovas-Vilar I, Galindo J (2004) Pierolapithecus catalaunicus, a new Middle Miocene great ape from Spain. Science 306:1339–1344PubMedCrossRefGoogle Scholar
- Moyà-Solà S, Köhler M, Alba DM (2005a) Response to comment on “Pierolapithecus catalaunicus, a new Middle Miocene great ape from Spain”. Science 308:203dCrossRefGoogle Scholar
- Moyà-Solà S, Köhler M, Rook L (2005b) The Oreopithecus thumb: a strange case in hominoid evolution. J Hum Evol 49:395–404PubMedCrossRefGoogle Scholar
- Moyà-Solà S, Köhler M, Alba DM, Casanovas-Vilar I, Galindo J, Robles JM, Cabrera L, Garcés M, Almécija S, Beamud E (2009) First partial face and upper dentition of the Middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallès-Penedès Basin, Catalonia, NE Spain): taxonomic and phylogenetic implications. Am J Phys Anthropol 139:126–145PubMedCrossRefGoogle Scholar
- Nakatsukasa M, Kunimatsu Y (2009) Nacholapithecus and its importance for understanding hominoid evolution. Evol Anthropol 18:103–119CrossRefGoogle Scholar
- Nakatsukasa M, Kunimatsu Y, Nakano Y, Takano T, Ishida H (2003) Comparative and functional anatomy of phalanges in Nacholapithecus kerioi, a Middle Miocene hominoid from northern Kenya. Primates 160:469–482Google Scholar
- Napier JR (1960) Studies of the hands of living primates. Proc Zool Soc Lond 134:647–657CrossRefGoogle Scholar
- Napier J (1993) Hands [revised by Russell H. Tuttle]. Princeton University Press, PrincetonGoogle Scholar
- Napier JR, Davis PR (1959) The fore-limb skeleton and associated remains of Proconsul africanus. Fossil mammals of Africa, No. 16. British Museum (Natural History), LondonGoogle Scholar
- O’Connor BL (1975) The functional morphology of the cercopithecoid wrist and inferior radioulnar joints, and their bearing on some problems in the evolution of the Hominoidea. Am J Phys Anthropol 43:113–122PubMedCrossRefGoogle Scholar
- Ogihara N, Almécija S, Nakatsukasa M, Nakano Y, Kikuchi Y, Kunimatsu Y, Makishima H, Shimizu D, Takano T, Tsujikawa H, Kagaya M, Ishida H, (2016) Carpal bones of Nacholapithecus kerioi, a middle Miocene hominoid from Northern Kenya. Am J Phys Anthropol 160:469–482Google Scholar
- Patel BA, Susman RL, Rossie JB, Hill A (2009) Terrestrial adaptations in the hands of Equatorius africanus revisited. J Hum Evol 57:763–772PubMedCrossRefGoogle Scholar
- Pilbeam DR, Rose MD, Badgley C, Lipschutz B (1980) Miocene hominoids from Pakistan. Postilla 181:1–94Google Scholar
- Pina M, Alba DM, Almécija S, Fortuny J, Moyà-Solà S (2012) Brief communication: Paleobiological inferences on the locomotor repertoire of extinct hominoids based on femoral neck cortical thickness: the fossil great ape Hispanopithecus laietanus as a test-case study. Am J Phys Anthropol 149:142–148PubMedCrossRefGoogle Scholar
- Preuschoft H (1973) Body posture and locomotion in some East African Miocene Dryopithecinae. In: Day M (ed) Human evolution. Taylor and Francis, London, pp 13–46Google Scholar
- Rafferty KL (1990) The functional and phylogenetic significance of the carpometacarpal joint of the thumb in anthropoid primates. M.A. thesis, New York UniversityGoogle Scholar
- Rafferty KL, Walker A, Ruff CB, Rose MD, Andrews PJ (1995) Postcranial estimates of body weight in Proconsul, with a note on a distal tibia of P. major from Napak, Uganda. Am J Phys Anthropol 97:391–402Google Scholar
- Richmond BG (2006) Functional morphology of the midcarpal joint in knuckle-walkers and terrestrial quadrupeds. In: Ishida H, Tuttle R, Pickford M, Ogihara N, Nakatsukasa M (eds) Human origins and environmental backgrounds. Springer, New York, pp 105–122CrossRefGoogle Scholar
- Richmond BG, Jungers WL (2008) Orrorin tugenensis femoral morphology and the evolution of hominin bipedalism. Science 319:1662–1665PubMedCrossRefGoogle Scholar
- Richmond BG, Strait DR (2000) Evidence that humans evolved from a knuckle-walking ancestor. Nature 404:382–385PubMedCrossRefGoogle Scholar
- Richmond BG, Whalen M (2001) Forelimb function, bone curvature and phylogeny of Sivapithecus. In: de Bonis L, Koufos GD, Andrews P (eds) African and Eurasian Miocene Hominoids and the origins of the Hominidae. Cambridge University Press, Cambridge, pp 326–348Google Scholar
- Richmond BG, Begun DR, Strait DS (2001) Origin of human bipedalism: the knuckle-walking hypothesis revisited. Yearb Phys Anthropol 44:70–105CrossRefGoogle Scholar
- Rose MD (1984) Hominoid postcranial specimens from the Middle Miocene Chinji Formation, Pakistan. J Hum Evol 13:503–516CrossRefGoogle Scholar
- Rose MD (1986) Further hominoid postcranial specimens from the Late Miocene Nagri Formation of Pakistan. J Hum Evol 15:333–367CrossRefGoogle Scholar
- Rose MD (1988) Functional anatomy of the cheiridia. In: Schwartz JH (ed) Orangutan biology. Oxford University Press, New York, pp 299–310Google Scholar
- Rose MD (1992) Kinematics of the trapezium-1st metacarpal joint in extant anthropoids and Miocene hominoids. J Hum Evol 22:255–266CrossRefGoogle Scholar
- Rose MD (1993) Locomotor anatomy of Miocene hominoids. In: Gebo DL (ed) Postcranial adaptation in nonhuman primates. Northern Illinois University Press, DeKalb, pp 252–272Google Scholar
- Rose MD (1994) Quadrupedalism in some Miocene catarrhines. J Hum Evol 26:387–411CrossRefGoogle Scholar
- Rose MD, Nakano Y, Ishida H (1996) Kenyapithecus postcranial specimens from Nachola, Kenya. Afr Stud Monogr Suppl 24:3–56Google Scholar
- Ruff CB (2003) Long bone articular and diaphyseal structure in Old World monkeys and apes. II: Estimation of body mass. Am J Phys Anthropol 120:16–37PubMedCrossRefGoogle Scholar
- Ruff CB, Walker A, Teaford MF (1989) Body mass, sexual dimorphism and femoral proportions of Proconsul from Rusinga and Mfangano Islands, Kenya. J Hum Evol 18:515–536Google Scholar
- Sarmiento EE (1987) The phylogenetic position of Oreopithecus and its significance in the origin of the Hominidae. Am Mus Novit 2881:1–44Google Scholar
- Sarmiento EE (1988) Anatomy of the hominoid wrist joint: its evolutionary and functional implications. Int J Primatol 9:281–345CrossRefGoogle Scholar
- Schmitt D (1994) Forelimb mechanics as a function of substrate type during quadrupedalism in two anthropoid primates. J Hum Evol 26:441–457CrossRefGoogle Scholar
- Schmitt D (2003) Insights into the evolution of human bipedalism from experimental studies of humans and other primates. J Exp Biol 206:1437–1448PubMedCrossRefGoogle Scholar
- Schön MA, Ziemer LK (1973) Wrist mechanism and locomotor behavior of Dryopithecus (Proconsul) africanus. Folia Primatol 20:1–11PubMedCrossRefGoogle Scholar
- Sherwood RJ, Ward S, Hill A, Duren DL, Brown B, Downs W (2002) Preliminary description of the Equatorius africanus partial skeleton (KNM-TH 28860) from Kipsaramon, Tugen Hills, Baringo District, Kenya. J Hum Evol 42:63–73PubMedCrossRefGoogle Scholar
- Spoor CF, Sondaar PY, Hussain ST (1991) A hominoid hamate and first metacarpal from the Late Miocene Nagri Formation of Pakistan. J Hum Evol 21:413–424CrossRefGoogle Scholar
- Straus WL (1963) The classification of Oreopithecus. In: Washburn SL (ed) Classification and human evolution. Aldine Publishing Co., Chicago, pp 146–177Google Scholar
- Straus WL Jr (1940) The posture of the great ape hand in locomotion and its phylogenetic implications. Am J Phys Anthropol 27:199–207CrossRefGoogle Scholar
- Susman RL (1979) Comparative and functional morphology of hominoid fingers. Am J Phys Anthropol 50:215–236PubMedCrossRefGoogle Scholar
- Susman RL (2004) Oreopithecus bambolii: an unlikely case of hominidlike grip capability in a Miocene ape. J Hum Evol 46:105–117PubMedCrossRefGoogle Scholar
- Susman RL (2005) Oreopithecus: still apelike after all these years. J Hum Evol 49:405–411PubMedCrossRefGoogle Scholar
- 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 448:921–924PubMedCrossRefGoogle Scholar
- Szalay FS, Delson E (1979) Evolutionary history of the primates. Academic Press, New YorkGoogle Scholar
- Tallman M, Almécija S, Reber SL, Alba DM, Moyà-Solà S (2013) The distal tibia of Hispanopithecus laietanus: more evidence for mosaic evolution in Miocene apes. J Hum Evol 64:319–327PubMedCrossRefGoogle Scholar
- Tuttle RH (1967) Knuckle-walking and the evolution of hominoid hands. Am J Phys Anthropol 26:171–206CrossRefGoogle Scholar
- Tuttle RH (1975) Parallelism, brachiation, and hominoid phylogeny. In: Luckett WP, Szalay FS (eds) Phylogeny of the primates. Plenum Press, New York, pp 447–480CrossRefGoogle Scholar
- Walker A (1997) Proconsul: function and phylogeny. In: Begun DR, Ward CV, Rose MD (eds) Function, phylogeny, and fossils: Miocene hominoid evolution and adaptations. Plenum Press, New York, pp 209–224CrossRefGoogle Scholar
- Walker AC, Pickford M (1983) New postcranial fossils of Proconsul africanus and Proconsul nyanzae. In: Ciochon RL, Corruccini RS (eds) New interpretations of ape and human ancestry. Plenum Press, New York, pp 325–351CrossRefGoogle Scholar
- Ward CV (1998) Afropithecus, Proconsul, and the primitive hominoid skeleton. In: Strasser E, Fleagle J, Rosenberger A, McHenry H (eds) Primate locomotion: recent advances. Plenum Press, New York, pp 337–352CrossRefGoogle Scholar
- Ward CV (2007) Postcranial and locomotor adaptations of hominoids. In: Henke W, Tattersall I (eds) Handbook of paleoanthropology. Springer Verlag, Berlin, pp 1011–1030CrossRefGoogle Scholar
- Ward S, Brown B, Hill A, Kelley J, Downs W (1999) Equatorius: a new hominoid genus from the middle Miocene of Kenya. Science 285:1382–1386PubMedCrossRefGoogle Scholar
- Wu R, Xu Q, Lu Q (1986) Relationship between Lufeng Sivapithecus and Ramapithecus and their phylogenetic position. Acta Anthropol Sinica 5:1–30Google Scholar