17 - Body size in African Middle Pleistocene Homo  pp. 319-346

Body size in African Middle Pleistocene <i>Homo</i>

By Steven E. Churchill, Lee R. Berger, Adam Hartstone-Rose and B. Headman Zondo

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It is generally accepted that archaic humans of the African later Early and early Middle Pleistocene constituted the source population for anatomically modern humans. Due to limited fossil and archaeological records, however, relatively little is known about the morphology, behaviour and ecology of these presumed ancestors of modern humans. Fragmentary fossils (variously attributed to Homo heidelbergensis, H. rhodesiensis and H. helmei) from across Africa suggest that these archaic humans were both taller and more massive than their extant modern human descendants in this region, and perhaps had a body shape that was stockier and less ‘nilotic’ than seen among extant sub-Saharan Africans. Fragmentary fossils attributed to Homo sapiens, on the other hand, appear to represent individuals closer in body size to the means of recent sub-Saharan Africans. Since body size and shape are critical to the ecology, energetics and thermoregulatory adaptations of early humans, these differences in morphology may signal important adaptive changes at the time of the origins of modern humans. Comparative analyses of femoral and orbital dimensions support the claim that Middle Pleistocene Africans were of greater body size (both stature and mass) and had greater mass/stature ratios than modern Africans, and support the claim that early African H. sapiens were of smaller body size than their Middle Pleistocene ancestors.

Aiello, L. C.Wells, C. K.2002Energetics and the evolution of the genus Annual Review of Anthropology31323
Aiello, L. C.Wood, B. A.1994Cranial variables as predictors of hominine body massAmerican Journal of Physical Anthropology95409
Aiello, L. C.Wood, B.Key, C.1999Morphological and taxonomic affinities of the Olduvai ulna (OH 36)American Journal of Physical Anthropology10989
Allbrook, D.1961The estimation of stature in British and East African MalesJournal of Forensic Medicine815
Anderson, E.1984Martin, P. S.Klein, R. G.Quaternary Extinctions: a Prehistoric RevolutionTucson, AZUniversity of Arizona Press
Arsuaga, J.-L.Lorenzo, C.Carretero, J.-M.1999A complete human pelvis from the Middle Pleistocene of SpainNature399255
Auerbach, B. M.Ruff, C. B.2004Human body mass estimation: a comparison of ‘morphometric’ and ‘mechanical’ methodsAmerican Journal of Physical Anthropology125331
Berger, L. R.Parkington, J. E.1995Brief communication: a new Pleistocene hominid-bearing locality at Hoedjiespunt, South AfricaAmerican Journal of Physical Anthropology98601
Brauer, G.Leakey, R. E.1986A new archaic cranium from Eliye Springs, West Turkana, KenyaZeitschrift für Morphologie und Anthropologie76245
Brauer, G.Leakey, R. E.1986The ES-11693 cranium from Eliye Springs, west Turkana, Kenya (Hominidae)Journal of Human Evolution15289
Brauer, G.Yokoyama, Y.Falgueres, C.1997Modern human origins backdatedNature386337
Brink, J. S.1993Postcranial evidence for the evolution of the black wildebeest, : an exploratory studyPalaeontologica Africana3061
Brook, B. W.Bowman, D. M. J. S.2004The uncertain blitzkrieg of Pleistocene megafaunaJournal of Biogeography31517
Brooks, A. S.Yellen, J. E.Nevell, L.2005Hovers, E.Kuhn, S.Transitions Before the Transition: Evolution and Stability in the Middle Paleolithic and Middle Stone AgeNew YorkKluwer
Brown, J. H.Maurer, B. A.1986Body size, ecological dominance and Cope’s ruleNature324248
Calder, W. A.1984Size, Function, and Life HistoryCambridge, MAHarvard University Press
Cameron, N.Tanner, J. M.Whitehouse, R. H.1982A longitudinal analysis of the growth of limb segments in adolescenceAnnals of Human Biology9211
Churchill, S. E.1993Peterkin, G. L.Bricker, H. M.Mellars, P. A.Hunting and Animal Exploitation in the Later Palaeolithic and Mesolithic of EuropeAmerican Anthropological Association Archaeological Paper
Churchill, S. E.2002Of assegais and bayonets: reconstructing prehistoric spear useEvolutionary Anthropology11185
Churchill, S. E.2006Havarti, K.Harrison, T.Neanderthals Revisited: New Approaches and PerspectivesNew YorkSpringer
Churchill, S. E.Berger, L. R.Parkington, J.2000A Middle Pleistocene human tibia from Hoedjiespunt, Western Cape, South AfricaSouth African Journal of Science96367
Churchill, S. E.Pearson, O. M.Grine, F. E.1996Morphological affinities of the proximal ulna from Klasies River Main Site: archaic or modern?Journal of Human Evolution31213
Churchill, S. E.Rhodes, J. A.2006How strong were the Neandertals? Leverage and muscularity at the shoulder and elbow in Mousterian foragersPeriodicum Biologorum108457
Clark, J. D.Beyene, Y.WoldeGabriel, G.2003Stratigraphic, chronological and behavioral contexts of Pleistocene from Middle Awash, EthiopiaNature423747
Clark, J. D.De Heinzelin, J.Schick, K. D.1994African : old radiometric ages and young Oldowan assemblages in the Middle Awash Valley, EthiopiaScience2641907
Clarke, R. J.1985Delson, E.Ancestors: the Hard EvidenceNew YorkAlan R. Liss, Inc
Cohen, P.1996Fitting a face to NgalobaJournal of Human Evolution30373
Conroy, G. C.Jolly, C. J.Cramer, D.1978Newly discovered fossil hominid skull from the Afar depression, EthiopiaNature27667
Conroy, G. C.Pickford, M.Senut, B.1993Diamonds in the desert: the discovery of Evolutionary Anthropology246
Conroy, G. C.Weber, G. W.Seidler, H.2000Endocranial capacity of the Bodo cranium determined from three-dimensional computed tomographyAmerican Journal of Physical Anthropology113111
Day, M. H.1969Omo human skeletal remainsNature2221135
Day, M. H.Leakey, M. D.Magori, C.1980A new hominid fossil skull (L.H.18) from the Ngaloba Beds, Laetoli, northern TanzaniaNature28455
Day, M. H.Leakey, R. E. F.1974New evidence of the genus Homo from East Rudolf, Kenya (III)American Journal of Physical Anthropology41367
Day, M. H.Twist, M. H. C.Ward, S.1991Les vestiges post-craniens d’Omo I (Kibish)L’Anthropologie95595
De Villiers, H.1973Human skeletal remains from border cave, Ingwavuma District, KwaZuluAnnals of the Transvaal Museum28229
Deino, A.McBrearty, S.200240Ar/(39)Ar dating of the Kapthurin Formation, Baringo, KenyaJournal of Human Evolution42185
deMenocal, P. B.2004African climate change and faunal evolution during the Plio- PleistoceneEarth and Planetary Science Letters2203
Dreyer, T. F.1935A hominid skull from Florisbad, Orange Free State, with a note on the endocranial cast by C.U. Ariens KappersProceedings of the Amsterdam Academy of Sciences38119
Eisenberg, J. F.1990Damuth, J.MacFadden, B. J.Body Size in Mammalian Paleobiology: Estimation and Biological ImplicationsCambridgeCambridge University Press
Estévez, J.2004Vanishing carnivores: what can the disappearance of large carnivores tell us about the Neanderthal world?International Journal of Osteoarchaeology14190
Feldesman, M. R.Fountain, R. L.1996Race’ specificity and the femur/stature ratioAmerican Journal of Physical Anthropology100207
Feldesman, M. R.Kleckner, J. G.Lundy, J. K.1990The femur/stature ratio and estimates of stature in mid- and late-Pleistocene fossil hominidsAmerican Journal of Physical Anthropology83359
Feldesman, M. R.Lundy, J. K.1988Stature estimates for some African Plio- Pleistocene fossil hominidsJournal of Human Evolution17583
Fisher, R.McBrearty, S.2002The comparative morphology of hominin postcranial remains from the Kapthurin Formation, Baringo District, KenyaAmerican Journal of Physical Anthropology11770
Goudie, A. S.1999Slack, P.Environments and Historical ChangeOxfordOxford University Press
Graves, R. R.Lupo, A. C.McCarthy, R. C.2010Just how strapping was KNM-WT 15000?Journal of Human Evolution59542
Grine, F. E.Jungers, W. L.Tobias, P. V.1995Fossil Homo femur from Berg Aukas, Northern NamibiaAmerican Journal of Physical Anthropology97151
Grine, F. E.Pearson, O. M.Klein, R. G.1998Additional human fossils from Klasies River Mouth, South AfricaJournal of Human Evolution3595
Grun, R.Brink, J. S.Spooner, N. A.1996Direct dating of Florisbad hominidNature382500
Hartwig-Scherer, S.1993Body weight prediction in early fossil hominids: towards a taxon-‘independent’ approachAmerican Journal of Physical Anthropology9217
Hiernaux, J.1968La Diversité Humaine en Afrique Subsaharienne: Recherches BiologiquesBruxellesInstitut de Sociologie, Université Libre de Bruxelles
Holliday, T. W.Ruff, C. B.1997Ecogeographical patterning and stature prediction in fossil hominids: comment on M. R. Feldesman and R. L. FountainAmerican Journal of Physical Anthropology1996100
Hostetler, S. W.Clark, P. U.2000Tropical climate at the last glacial maximum inferred from glacier mass-balance modelingScience2901747
Howell, D. C.2001www.uvm.edu/~dhowell/StatPages
Howell, F. C.1978Maglio, V.Cooke, H. B. S.Evolution of African MammalsCambridge, MA
Howells, W. W.1973Cranial Variation in ManCambridge, MAPeabody Museum
Hublin, J. J.1991
Hublin, J. J.1992Recent human evolution in Northern AfricaPhilosophical Transactions of the Royal Society of London337185
Humphrey, L. T.1998Growth patterns in the modern human skeletonAmerican Journal of Physical Anthropology10557
Johnson, C. N.2002Determinants of loss of mammal species during the Late Quaternary ‘megafauna’ extinctions: life history and ecology, but not body sizeProceedings of the Royal Society of London2692221
Kappelman, J.1996Evolution of body mass and relative brain size in fossil hominidsJournal of Human Evolution30243
Kappelman, J.1997They might be giantsNature387126
Katzmarzyk, P. T.Leonard, W. R.1998Climatic influences on human body size and proportions: ecological adaptations and secular trendsAmerican Journal of Physical Anthropology106483
Klein, R. G.1984The Large Mammals of Southern Africa: Late Pliocene to RecentRotterdamBalkema
Klein, R. G.1994Southern Africa before the Iron AgeEnglewood Cliffs, NJPrentice Hall
Leonard, J. A.Vilà, C.Fox-Dobbs, K.2007Megafaunal extinctions and the disappearance of a specialized wolf ecomorphCurrent Biology171146
Lieberman, D. E.Devlin, M. J.Pearson, O. M.2001Articular area responses to mechanical loading: effects of exercise, age, and skeletal locationAmerican Journal of Physical Anthropology11626677
Martin, R.1928Lehrbuch der AnthropologieJenaVerlag von Gustav Fischer
McBrearty, S.Brooks, A. S.2000The revolution that wasn’t: a new interpretation of the origin of modern human behaviorJournal of Human Evolution39453
McDougall, I.Brown, F. H.Fleagle, J. G.2005Stratigraphic placement and age of modern humans from Kibish, EthiopiaNature433733
McHenry, H. M.1992Body size and proportions in early hominidsAmerican Journal of Physical Anthropology87407
McHenry, H. M.Coffing, K.2000Australopithecus to : transformations in body and mindAnnual Review of Anthropology29125
McNab, B. K.1983Energetics, body size, and the limits to endothermyJournal of Zoology1991
McNab, B. K.1990Damuth, J.MacFadden, B. J.Body Size in Mammalian Paleobiology: Estimation and Biological ImplicationsCambridgeCambridge University Press
Morris, A. G.1992Brauer, GSmith, F. H.Continuity or Replacement. Controversies in EvolutionRotterdamBalkema
Owen-Smith, N.1987Pleistocene extinctions: the pivotal role of megaherbivoresPaleobiology13351
Pearson, O. M.2000Postcranial remains and the origin of modern humansEvolutionary Anthropology9229
Pearson, O. M.Grine, F. E.1996Morphology of the Border Cave hominid ulna and humerusSouth African Journal of Science92231
Pearson, O. M.Grine, F. E.1997Re-analysis of the hominid radii from Cave of Hearths and Klasies river mouth, South AfricaJournal of Human Evolution32577
Pearson, O.Jungers, W.Grine, F.2001The reliability of estimates of hominin body mass derived from bi-iliac breadth and statureAmerican Journal of Physical Anthropology32118
Peters, J.Gautier, A.Brink, J. S.1994Late Quaternary extinction of ungulates in sub-Saharan Africa: a reductionist’s approachJournal of Archaeological Science2117
Pfeiffer, S.1998Klasies River Mouth post-cranial comparisonsJournal of Human Evolution34A17
Pfeiffer, S.Zehr, M. K.1996A morphological and histological study of the human humerus from Border CaveJournal of Human Evolution3149
Pontzer, H.Rolian, C.Rightmire, G. P.2010Locomotor anatomy and biomechanics of the Dmanisi homininsJournal of Human Evolution58492
Potts, R.Behrensmeyer, A. K.Deino, A.2004Small mid-Pleistocene hominin associated with east African Acheulean technologyScience30575
Pycraft, W. P.1928Pycraft, W. P.Elliot Smith, G.Yearsley, M. et al.Rhodesian Man and Associated RemainsLondonBritish Museum
Reynolds, S. C.2007Mammalian body size changes and Plio-Pleistocene environmental shifts: implications for understanding hominin evolution in eastern and southern AfricaJournal of Human Evolution53528
Rightmire, G. P.1978Florisbad and human population succession in Southern AfricaAmerican Journal of Physical Anthropology48475
Rightmire, G. P.1996The human cranium from Bodo, Ethiopia: evidence for speciation in the Middle Pleistocene?Journal of Human Evolution3121
Rightmire, G. P.1998Human evolution in the Middle Pleistocene: the role of Evolutionary Anthropology6218
Rightmire, G. P.Deacon, H. J.1991Comparative studies of Late Pleistocene human remains from Klasies River Mouth, South AfricaJournal of Human Evolution20131
Roberts, D. F.Bainbridge, D. R.1963Nilotic physiqueAmerican Journal of Physical Anthropology21341
Rosenberg, K. R.Lu, Z.Ruff, C. B.2006Body size, body proportions, and encephalization in a Middle Pleistocene archaic human from northern ChinaProceedings of the National Academy of Science1033552
Ruff, C. B.1991Climate and body shape in hominid evolutionJournal of Human Evolution2181
Ruff, C. B.1993Climatic adaptation and hominid evolution: the thermoregulatory imperativeEvolutionary Anthropology253
Ruff, C. B.1994Morphological adaptation to climate in modern and fossil hominidsYearbook of Physical Anthropology3765
Ruff, C. B.2000Body mass prediction from skeletal frame size in elite athletesAmerican Journal of Physical Anthropology113507
Ruff, C. B.2007Body size prediction from juvenile skeletal remainsAmerican Journal of Physical Anthropology133698
Ruff, C. B.2010Body size and body shape in early hominins: implications of the Gona pelvisJournal of Human Evolution58166
Ruff, C. B.Trinkaus, E.Holliday, T. W.1997Body mass and encephalization in Pleistocene Nature387173
Ruff, C.Walker, A. C.1993Walker, A. C.Leakey, R. E.The Nariokotome SkeletonCambridgeHarvard University Press
Schmitt, D. O.Churchill, S. E.Hylander, W. L.2003Experimental evidence concerning spear use in Neandertals and early modern humansJournal of Archaeological Science30103
Shea, J. J.2006The origins of lithic projectile point technology: evidence from Africa, the Levant, and EuropeJournal of Archaeological Science33823
Simpson, S. W.Quade, J.Levin, N. E.2008Science3221089
Singer, R.Wymer, J. J.1982The Middle Stone Age at Klasies River Mouth in South AfricaChicagoUniversity of Chicago Press
Smith, R. J.1996Biology and body size in human evolution: statistical inference misappliedCurrent Anthropology37451
Solan, M.Day, M. H.1992The Baringo (Kapthurin) ulnaJournal of Human Evolution22307
Spoor, F.Leakey, M. G.Gathogo, P. N.2007Implications of new early fossils from Ileret, east of Lake Turkana, KenyaNature448688
Steele, D. G.McKern, T. W.1969A method for assessment of maximum long bone length and living stature from fragmentary long bonesAmerican Journal of Physical Anthropology31215
Steudel-Numbers, K. L.Tilkens, M. J.2004The effect of lower limb length on the energetic cost of locomotion: implications for fossil homininsJournal of Human Evolution4795
Steudel-Numbers, K.Weaver, T. D.2006Froude number corrections in anthropological studiesAmerican Journal of Physical Anthropology13127
Steyn, M.İşcan, M. Y.1999Osteometric variation in the humerus: sexual dimorphism in South AfricansForensic Science International10677
Tobias, P. V.1971Human skeletal remains from the Cave of Hearths, Makapansgat, Northern TransvaalAmerican Journal of Physical Anthropology34335
Trinkaus, E.1993A note on the KNM-ER 999 hominid femurJournal of Human Evolution24493
Trinkaus, E.1993Femoral neck-shaft angles of the Qafzeh-Skhul early modern humans, and activity levels among immature Near Eastern Paleolithic hominidsJournal of Human Evolution25393
Trinkaus, E.Churchill, S. E.Ruff, C. B.1994Postcranial robusticity in . II: Humeral bilateral asymmetry and bone plasticityAmerican Journal of Physical Anthropology931
Trinkaus, E.Ruff, C. B.Conroy, G. C.1999The anomalous archaic Homo femur from Berg Aukas, Namibia: a biomechanical assessmentAmerican Journal of Physical Anthropology110379
Trotter, M.Gleser, G. C.1952Estimation of stature from long bones of American whites and NegroesAmerican Journal of Physical Anthropology10463
Weaver, T. D.Steudel-Numbers, K.2005Does climate or mobility explain the differences in body proportions between Neandertals and their Upper Paleolithic successors?Evolutionary Anthropology14218
White, T. D.Asfaw, B.DeGusta, D.2003Pleistocene from Middle Awash, EthiopiaNature423742