Index 271. Dryopithecus, taxonomic identification and re-classification, 15, 18 22

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1 INDEX accessory processes, in Proconsul lumbar vertebrae, adaptive radiation, and vertical climbing in primates, adduction, and gluteus maximus muscle, 138 adipose tissue, adolescence, Aegyptopithecus, 22 Africa extant primate distribution, 179, 180(fig.), 192, 193, Miocene environmental change, Miocene hominid distribution, 181(fig.) phytochores, 178(fig.), 180(fig.), 181(fig.), 193 sub-saharan faunal resemblance to Eurasia, Afropithecus, 25 and phytochores, 181(fig.) Afropithecus turkanensis, taxonomic identification and re-classification, 20, 24 Aka Aiteputh Formation composition of, habitat, 215 K-Ar ages, 81(fig.), 85(table) as part of Neogene System, 71, 72 taxonomic identification from, 24 See also Samburu Hills alpha taxonomy, , 253 alternating field (AF) demagnetization, Andrews, Peter, 18 anestrous period in hominoids, angle of retroglenoid tubercle, and Otavipithecus namibiensis, 34(table) angulation of odontoid, Australopithecus antiquus seu afarensis, 36 ankle displacement in primates, 100, 101(fig.), 102 Anthropoidea/anthropoids allometric studies, 18 alpha taxonomy, dietary models, initial description and early studies, models of positional behavior, paleoenvironmental models, re-classification and phylogenic interpretations, apes, African distribution affected by humidity, 192 distribution and seasonal environments, 179 and eurytopy, 179 genome, 249 habitats, 179, 180(fig.), 192, 193, 213(fig.) and humans, joint function and morphology in, and knuckle-walking (see knuckle-walking) links to Miocene ancestors, 16, 25 and phytochores, (figs.), 193 social structure, 236(fig.), and stenotopy, 179 and territoriality, 179, 182(fig.) See also apes, great; bonobo; chimpanzee; gorilla apes, Asian joint function and morphology, wrist mobility, 118, 120 See also gibbons; orangutan apes, great female-philopatric groups, 238 female transfer, 237 interbirth intervals, male tolerance of copulation by other group males, natal grouping of males, 237 nonmatrilineal groups, 238 pair bonds, superior vertebro-articular process angle, 38(fig.) See also apes, African; apes, Asian arborealism and Ardipithecus kadabba, 202 in australopithecines, 150, 201 and Cercopithecoidea, 46 and Dendropithecus, 18 and divergence of human lineage from ape lineage, 232 energetic costs of suspensory locomotion, 152,

2 268 Index arborealism (cont.) in generalized bipeds, and Mabokopithecus harrisoni, 216 and Nacholapithecus, 103, and Orrorin tugenensis, and Otavipithecus namibiensis, 34 physiological constraints, and Pliopithecus, 18 as pre-adaptation to bipedalism, and wrist mobility, See also bipedalism, arboreal origin of; climbing Arctic ice cap, and East Side Story, 187 Ardipithecus, 202 Astaracian appearance of taxa, 63 faunal turnover and open country mammalian taxa, Ateles geoffroyi. See spider monkey Australopithecinae/australopithecines locomotor behavior, mental map capacity, 186 taxonomy, 251(table), 253 Australopithecus and arboreal origins of bipedalism, computer simulation as predictor of locomotion, hamate, 114(fig.), 117(fig.) joint range of motion measurements, kinematics of bipedal locomotion, Australopithecus afarensis and bipedal locomotion, 37, 150 cervical vertebrae, and environment, 201 femur, 203(fig.) and Laetoli hominid, locomotion and limb length, 165 studies on sacrum, Australopithecus antiquus seu afarensis, baboon (Papio) first studies of locomotion, 149 hand postures, 106(fig.) joint range of motion measurements, Baringo Basin, taxonomic identification from, 24 basal metabolic rate, 171 Basmajian, John V., 3 behavioral changes, and hominid evolution and diet, 49 50, , 239, and threat displays, See also social structure of hominoids bending-extension movements, 33, 37 bent-hip, bent-knee gait, 152, 157, 167 Berg Aukas, taxonomic identification from, 34 biostratigraphy, mammalian, and faunal change in sub-saharan Africa, faunal resemblance between sub-saharan Africa and Eurasia, bipedal energetics compared to quadrupedal energetics, 152, 158, , 167, computer models, and humans, 165, 167, 171 and macaques, , recommendations for further research, 150 bipedal locomotion and cineradiography, 150 computer simulation, fast bipedal motion, 139 and kinematics, 150 as mechanism of evolutionary divergence, 196 and morphophysiology, 157 obligate, 149 origins of (see bipedalism, arboreal origin of; bipedalism, origins of) and trained primates, (see also macaque, Japanese) See also specific extant and fossil species bipedal standing in rats. See rat bipedalism, arboreal origin of, in Ardipithicus, 202 in australopithecines, in Orrorin, , 203(fig.), 204(fig.) physiological constraints in arboreal activities, in Preanthropus, in Sahelanthropus, bipedalism, origins of, and adipose tissue, and adolescence, and dentition, and diet (see diet) and East Side Story, geographic scenarios, 183(fig.), and humidity, and instep evolution, and mental maps, 186

3 Index 269 and Miocene vegetation, open country hypothesis, and paleoenvironments, , , 188(fig.), and phytocores, 178(fig.), 180(fig.), 181(fig.), 193 and posture and body size, 193 and precipitable water, 188(fig.) prehominid evolution, and ranges, 184 savanna hypothesis, , , 205 and social structure of primate groups, and stages of terrestrialization, 264(fig.) and stenotypy vs. eurytopy, , 185(fig.) terrestrialization hypothesis, , and territoriality, 182(fig.) and threat displays, See also bipedalism, arboreal origin of Bishop, Walter, 19 body proportions and size and adolescent growth spurt, and hominid evolution, 193 and interbirth intervals, in Nacholapithecus, 8 and terrestriality, 51 53, 55 in Victoriapithecus macinnesi, 51 53, 55 bones. See joints; limb length; vertebral column; and specific bones bonobo (Pan paniscus) filiation, 17 social structure, See also apes, African; Pan brachiation. See arborealism Bukwa II, taxonomic identification from, 19 Buluk, taxonomic identification from, 24 buttocks. See gluteal muscle C5, in hominoid and nonhominoid primates, capitate extension, 108 interspecific differences in breadth, 112 measurements, 109(fig.), 115(fig.), 116(fig.) photographs of primate, 118(fig.) range of motion, 115 shape variables in primates, 117(fig.), 117(table) capuchin (Cebus sp.) limb movement, 100, 101, 102 limb position, 99, 100 locomotor energetics, 158 vertical climbing, 5, , 99(fig.), 101(fig.), 102(table) carbon dioxide measurements (macaques), carpal anatomy cluster diagram, 52(fig.) resemblance between African apes and humans, 116 See also midcarpal joint Catarrhini/catarrhines midcarpal joint, 105, , 24, 25 and terrestriality, 55, 105, 108 Ceboidea/ceboids carpal measurements, 52(fig.) humeral measurements, 47(fig.), 49(fig.) radial measurements, 51(fig.) ulnar measurements, 48(fig.), 50(fig.) and Victoriapithecus, 46 Cebus. See capuchin Cenozoic faunal change, geological map of volcanic rocks, 73(fig.) See also Miocene Central Pattern Generator (CPG), 169 Centre for Prehistory and Palaentology, 21 cephalic swinging movements, and Plio-Pleistocene hominids, 37 Cercopithecinae/cercopithecines diet, dentition, and terrestriality, and Victoriapithecus, Cercopithecoidea/cercopithecoids, in African habitats, 213(fig.) and arborealism, 46 carpal measurements, 52(fig.) diet and dentition, 46, 49 divergence of clad, earliest remains of, 53 and folivory, 46 humeral measurements, 47(fig.), 49(fig.) hypothesis on evolution, 19 radial measurements, 51(fig.) and terrestriality, 48 ulnar measurements, 48(fig.), 50(fig.) cervical vertebrae in chimpanzee, 35 lordosis, 35, 36, 37 in Miocene hominoids, 34 in Plio-Pleistocene hominids, characteristic remanent magnetization (ChRM), 86 Cheboi, Kiptalam, 7

4 270 Index Chief Kerio, 8 chimpanzee (Pan troglodytes) cervical lordosis, 35 culture, joint range of motion measurements, limb movement, limb position, 99, 100 locomotion and pressure distribution of feet, 149 locomotor energetics, 158 possible ancestors of, 19 20, 25 social structure, thoracic-lumbar kyphosis, 35 vertical climbing, , (figs.) See also apes, African; bonobo; Pan ChRM. See characteristic remanent magnetization cineradiography, and bipedal locomotion, 150 cladistics, climbing, , (figs.) and adolescent growth spurt, and Australopithecus antiquus seu afarensis, 37 digitigrade climbing, and divergence of human lineage from ape lineage, 232 and gluteus maximus muscle, human foot contact on trees, 194(fig.) and kinematics, 97 and limb function, 102(table), and limb position, 99(fig.), modeling in extinct species, 37, 103 and paleoenvironment, 195 vertical climbing hypothesis, 5, 205 See also arborealism cluster analysis of faunas from sub-saharan Africa and Eurasia, Colobinae/colobines, 213(fig.) and Victoriapithecus, 48 computer simulation of bipedal locomotion, calculation of locomotion, and innervation of skeletal muscle, 169(fig.) integration diagram at alpha motoneuron, 170(fig.) locomotion diagram, 171(fig.) musculoskeletal model, 168(fig.), nervous model, neuromusculoskeletal model, 170(fig.) predictive simulation, 171(fig.), 172 continental change, and East Side Story, 187 cooperant bipedalism, cortical thickness in femur, 127(fig.), 128(fig.), CPG. See Central Pattern Generator crouching, and gluteus maximus muscle, , culture, evolution of, curvatures, spinal. See lordosis daily travel range, 184 Dendropithecus and arborealism, , 22, 25 See also Limnopithecus dentition and bipedalism, in cercopithecoids, 46, and dietary adaptations, 46, 49 50, in Kenyapithecus, 216, 261 in Nacholapithecus, 8, 261, 262 in Otavipithecus namibiensis, 191 in Samburupithecus, 8,262 in Simiolus leakeyorum, 217 in Victoriapithecus, 46, 49, 217 diet and bipedalism, and cercopithecoids, 46, changes and evolution of hominids, , 239 dietary models, dispersed food hypothesis, faunivores, folivory, 39, 46, 217 frugivory, 46, as mechanism of evolutionary divergence, sclerocarp feeders, 216, 217 size of food patches, 239 and terrestriality, 49 50, 55 See also dentition; foraging; specific species digitigrade locomotion, 106, 108, Dionysopithecus, 25 dispersed food hypothesis, DNA, difference between chimp and human, 249 Dolichopithecus, 48 dorsal musculature, in Proconsul, 32 Dryopithecinae, 16, 22 Dryopithecus africanus, as ancestor of Pan troglodytes, 20

5 Index 271 Dryopithecus, taxonomic identification and re-classification, 15, East Side Story (ESS) and bipedal locomotion, and hominid origins, and Orrorin tugenensis, 225 Eastern Rift Valley, 7 ecological niches and Cercopithecoidae, and Hominidae, and Paranthropinae, 252 and Victoriapithecus macinnesi, See also diet elbow displacement in primates, 101 electromyography (EMG), 1, 3, 149 in humans, 136, , 140(fig.), 229(fig.) in Japanese macaque, , 231(fig.) in spider monkeys, , 230(fig.) energetics. See bipedal energetics; locomotor energetics Eppelshiem femur, 15 Equatorius africanus, taxonomic identification and re-classification, 24, Erythrocebus patas. See patas monkey ESS. See East Side Story estrus sex ratio hypothesis, Ethiopia, taxonomic identification from, 35 eurytopy, 181, , 185(fig.) evolution of bipedalism. See bipedalism, origins of; specific bones, muscles, and joints exercise, changes resulting from. See macaque, Japanese; rat Far East Side Story, 186, fauna and associations with Orrorin, change in Late Cenozoic Eurasia and sub-saharan Africa, first and last appearance, formulas for calculating resemblance, 60 and half-life, of Namurungule, 61 from Neogene sub-saharan Africa, 62(table), 65(fig.), 66(fig.) turnover and paleoenvironmental change, faunivores, feeding strategies. See diet female primates. See primates, female femur Eppelshiem femur, 15 human bone morphology, 131 of Lucy, 203(fig.) of Orrorin tugenensis, (figs.) of rat, Ferembach, Denise, 16, 17 fifth cervical vertebra, flexibility, and vertebral column, 41. See also joints folivory, 39, 46, 217 footprints human, 200(fig.) and Laetoli hominid, foraging dispersed food hypothesis, ecology in primates, terrestrial, 54 See also diet force detection pole, introduction of, 5 forelimbs forelimb hauling/propulsion, 102, 103 humerus (cercopithecines and ceboids), 49(fig.) radius (cercopithecines and ceboids), 51(fig.) ulna (cercopithecines and ceboids), 50(fig.) in vertical climbing, See also digitigrade locomotion; knuckle-walking; palmigrade walking forest habitat, and Victoriapithecus macinnesi, See also arborealism Fort Ternan alpha taxonomy, 211 fossil distribution, 215 habitat, 215 Miocene anthropoids, taxonomic identification from, 20, 21 fossil primate collection, Nairobi, 21 frugivory, 46, function, muscle, defined, 136 gait. See locomotion, modes of; posture and gait galagos, jumping behavior, 5 gap-crossing strategies, 5 generalized biped, and arborealism, heart rate, 226(fig.), 231(table) lineage divergence, 232 muscle EMGs, 229(fig.), 230(fig.), 231(fig.) and musculature, oxygen consumption, 227(fig.) relative metabolic rate, 227(fig.), 228(table) genome, ape and human, 249 geography of hominid origins, geologic maps Cenzoic volcanic rocks, 73(fig.) Kenya Rift, 73(fig.) Nachola-Samburu hills, 73(fig.), 76(fig.) Tugen Hills, 73(fig.)

6 272 Index geological background of Miocene hominoids, of Nacholapithecus, of Orrorin tugenensis, 82 83, 92 94, 204 of Sahelanthropus tchadensis, 205 of Samburupithecus, 80 82, See also specific formations geomagnetic polarity time scale Lukeino Formation, 90(fig.) Samburu Hills, 88(fig.) See also paleomagnetic data gibbons (Hylobates sp.) bipedalism experiments, 4 hand postures, 106(fig.) joint range of motion measurements, limb movement, , 149 vertical climbing, 5, , 99(fig.), 101(fig.), 102(table) See also apes, Asian; Hylobates gluteal muscle, action, advantages of hypertrophy in hominids, EMG recordings during walking, 140(fig.) evolution of, function, and hypertrophy in hominids, morphology, origin in Homo, 138(fig.) placement in Gorilla, 137(fig.) placement in Homo, 137(fig.) and quiescence, gluteus maximus muscle. See gluteal muscle gluteus medius, 136 gluteus minimus, 136 gluteus superficialis, 137, 139 gorilla (Gorilla), 25 capitate, 115(fig.), 116(fig.), 117(fig.), 118(fig.) Dryopithecus major as ancestor of, 20 gluteal muscle placement, 137(fig.) gluteus superficialis, 139 hand postures, 106(fig.) ischial callosities, 136(fig.) joint range of motion measurements, lineage separation and chimpanzees, 20 and phytocores, (figs.) social structure, See also apes, African Gunther, Michael, 5 Hadar, taxonomic identification from, 35 half-life concept, hamate, 109(fig.), 112, 114(fig.), 117(fig.), 117(table), 118(fig.) hand postures of primates, 106(fig.). See also digitigrade walking; knuckle-walking; palmigrade walking; wrist joint Hayama, S., 3, 5 heart rate, 226(fig.), , 231(fig.) heat balance, of bipedal locomotion in patas monkeys, 158, 159 hind limb in vertical climbing, hind limb of bipedally trained animals. See macaque, Japanese; rat femur hip displacement, 101(fig.), 102 extension, 138, stabilization, Hipparion, and Namurungule Formation, 89 Hiwegi, taxonomic identification from, 22 home ranges, 184, 185(fig.) Hominidae/hominids ability to exploit new environments, advantages of buttocks hypertrophy, and climbing, Kanapoi, 200 Laetoli, locomotion, 131, 150, 157, 165 morphology and energy efficiency, 167 origins and evolution, 16, 179, , , 193, 225 storage of adipose tissue, taxonomy, , 251(fig.) vertebral column, See also bipedalism, origins of Homininae/hominins and knuckle-walking, taxonomy, 251(table), Hominoidea/hominoids dispersal, distribution, 189(fig.) evolution, 59 69, 192 evolution of social structure, evolution of vertebral column, extinction, fossils, 77(fig.) geological background, history of taxonomic identification of hominid species, and locomotor cost, 164 lumbar vertebrae, 31 Miocene anthropoid studies, and origin of bipedalism, and stenotypy vs. eurytopy, 185(fig.) and terrestriality, 45 55

7 Index 273 vertebral column, See also biostratigraphy, mammalian; bipedalism, origins of; social structure of hominoids Homo, ancestral type, 18 Homo ergaster, 150 Homo sapiens cervical lordosis, 35, 36, 37 computer simulations of bipedal locomotion, EMGs of limb muscles, 136, , 140(fig.), 229(fig.) and eurytopy, 181 evolution of instep, evolution of social structure, genome, 249 gluteal muscle, joint range of motion measurements, knuckle-walking ancestor, lumbar lordosis, 35, 39 occupations and relative metabolic rate, 228(fig.) and permanent bipedal locomotion, 34 posture, 34, 135, 139 relationships with extant apes, superior vertebro-articular process angle, 38(fig.), 39 thoracic kyphosis, 35 See also locomotion, human Hopwood, A. Tindell, 15 humerus (cercopithecines and ceboids), 49(fig.) humidity, and hominoid evolution, 192 hunting behavior, and gluteus maximus muscle, 143, 145 hurdler, photo of, 142(fig.) Hylobates, 97, 106(fig.) joint range of motion measurements, taxonomy, 251(table) See also gibbons Hylobatidae/hylobatid apes, 16, 17, 20, 21, 22 ichnology, Ikeda, Jiro, 1, 2 iliac crest, and attachment of m. erector spinae in apes, iliac tuberosity, and attachment of m. erector spinae in Proconsul, 32 Imanishi, Kinji, 6 innervation of skeletal muscle, 169(fig.) instep evolution, musculoskeletal morphology, 168 interbirth intervals of great apes, ischial tuberosities, 135 Ishida, Hidemi academic training, 3 career appointments, 3, 4, 6, 8 doctoral thesis, 3 postdoctoral career, 3 9 research interests, 3 6, 8 students, 2(table) Ishida, S., 6, 7 Itani, Jun ichiro, 3, 6 joints ankle displacement, 100, 101(fig.), 102 elbow displacement, 101 hip displacement, 101(fig.), 102 knee displacement, 101(fig.), 102 midcarpal joint, morphology and function, sacrum and sacro-iliac joint, shoulder displacement, 100, 101(fig.) and vertical climbing patterns, wrist joint, 107, 108, , 111(fig.), , 118, 120 See also knuckle-walking Jouffroy, Francoise K., 5 K-Ar ages of Aka Aiteputh Formation, 81(fig.), 83 84, 85(table) of Kabarnet Trachyte, of Kaparaina Basalt, of Kongia Formation, 81(fig.) of Lukeino Formation, 90(fig.) of Namurungule Formation yielding Samburupithecus, 81(fig.) of Neogene formation at Nachola-Samburu Hills, 77(fig.) Kabarnet Trachyte composition of, 82 correlation with Lukeino Formation, 91(fig.) K-Ar ages, Kalepithecus, 21, Kalodirr, taxonomic identification from, 24 Kaloma, taxonomic identification from, 24 Kanapoi hominid, 200 Kaparaina Basalt, K-Ar ages, Kapcheberek Member, composition of, 82 83

8 274 Index Karamoja District, taxonomic identification from, 19 Kaswanga Primate Site, 8 Kawai, Masao, 6 Kenya geologic map of the Rift, 73(fig.) geology, 6 8 joint expedition with Japan, 8 volcanism of the Rift, 75 See also Baringo Basin; Buluk; Fort Ternan; Hiwegi; Kalodirr; Kaloma; Kipsaramon; Koru; Lake Victoria; Lothidok Hills; Maboko; Majiwa; Mfangano Island; Muruyur Formation; Nachola; Namurungule; Ngorora Formation; Nyakach; Ombo; Rusinga Island; Samburu Hills; Songhor; Tugen Hills; Winam Gulf Kenyapithecus dentition, 216, 261 as a sclerocarp feeder, , kinematics/kinetics and bipedal locomotion, 150 and human musculoskeletal morphology, 168 and performing macaques, 164 position of markers for experiments, 98(fig.) vertical climbing in primates, 97 Kipsaramon, taxonomic identification from, 24 Kirimun Formation, 6, 7 knee displacement in primates, 101(fig.), 102 knuckle-walking, 35, , , 205 and capitates and hamates, (figs.) and divergence of human lineage from ape lineage, 232 and hand postures, 106(fig.) and joint range of motion, 107, , 111(fig.), 118, 120 as locomotion unique to gorillas and chimpanzees, 106 and lunate articular surface, 116(fig.) metacarpophalangeal joint postures, 107 and the midcarpal joint, , 113(fig.) and morphometric assessment, morphometric measurement techniques, and scaphoid central facet, 115(fig.) and triquetral facet angle, 114(fig.) and wrist extension hypothesis, 108 Kogolepithecus, Kondo, Shiro, 3, 4 Kongia Formation composition and distribution of, K-Ar ages, 81(fig.) as part of Neogene System, 71, 72 Koru, 15, 16 Koyaguchi, T, 7 Laetoli faunal associations, 201 Laetoli hominid, Lake Victoria, taxonomic identification from, 16 language, development of, 254 Lartet, Edouard, 9, 15 Leakey, Louis, 16, 18, 20, 21 Leakey, Mary, 20 Leakey, Richard, 6 LeGros Clark, Wilfrid, 16 limb function and vertical climbing in primates, 102(table), limb length and adolescent growth spurt, and bipedal locomotion, 157 and increase in range size, 184 limb position, and vertical climbing in primates, , 99(fig.) limbs, front. See forelimbs limbs, hind. See gluteal muscle; hind limb in vertical climbing; macaque, Japanese; rat femur Limnopithecus posture and gait, , 21 23, 25 lithofacies, and Lukeino Formation, 90(fig.) locomotion and Australopithecinae, , 253 energetics of (see locomotor energetics) evolution of, 131, 151 (see also bipedalism, arboreal origin of; bipedalism, origins of ) and Hominidae, and Homininae, and Homo sapiens (see locomotion, human) modes of (see locomotion, modes of) and Paranthropinae, 252 prehominid, See also posture and gait locomotion, human and adolescent growth spurt, vs. ape, 232 calculation of, EMGs of limb muscles, 136, , 140(fig.), 229(fig.) and heart rate, 226(fig.) musculoskeletal model, nervous model,

9 Index 275 neuromusculoskeletal model, 170(fig.) and oxygen measurement, 227(fig.) and relative metabolic rate, 227(fig.) stick diagram of, 171(fig.) locomotion, modes of bent-hip, bent-knee gait, 152, 157, 167 and bone morphology, 131 cost of bipedalism vs. quadrupedal locomotion, 152, 158, , 167, cost of quadrupedal vs. suspensory locomotion, 152, 158 digitigrade walking, 106, 108, 193 fast bipedal motion, 139 and gluteal muscle, palmigrade walking, 106, 193, 199 rat locomotion, 124(fig.), ungulate locomotion, 186 See also arborealism; bipedalism; climbing; knuckle-walking; posture and gait; specific species locomotor energetics and bipedalism, , bipedalism vs. quadrupedal locomotion, 152, 158, , 167, carbon dioxide concentration, 161(fig.), 163(fig.) carbon dioxide production rates, 163(fig.) computer models, and foraging ecology in primates, formula for, 159 in hominoids, and human musculoskeletal morphology, 168 and increase in range size, 184 in macaques, , in nonhuman primates, physiological load, quadrupedal vs. suspensory locomotion, 152, 158 respiratory quotient and energy cost, and terrain, lordosis cervical, 35, 36, 37 in chimpanzees, 35 in humans, 35, 36, 37, 39 lumbar, 35, 39, 151, 158 in performing macaques, 151, 158 loris, positional behavior studies, 5 Lothidok Hills, taxonomic identification from, 16 Lucy femur, 203(fig.) studies on sacrum, 41 See also Australopithecus afarensis Lukeino Formation correlation with Kabarnet Trachyte, 91(fig.) K-Ar ages, 90(fig.) lithofacies, 90(fig.) paleomagnetic data, 90(fig.) lumbar lordosis in humans, 35, 39 in performing macaques, 151, 158 lumbar vertebrae in humans, 35 in Miocene hominoids, 31 34, 32(table), 41(fig.) in Plio-Pleistocene hominids, 39, 41(fig.) in Proconsul, transverse processes of Hominoidea, 33(fig.) in Ugandapithecus, See also lumbar lordosis m. erector spinae, m. extensor caudae lateralis, m. intertransversarius, m. longissimus, Maboko alpha taxonomy, 211 fossil distribution, habitat, 214 Miocene anthropoids, taxonomic identification from, 16, 19, 22 24, and Victoriapithecus macinnesi, Mabokopithecus diet, , Macaca fascicularis, 54 Macaca nemestrina, 54 macaque, Japanese (Macaca fuscata) acquisition of lumbar lordosis, 151 bipedally trained, 5, , EMGs of limb muscles, energetics of locomotion, , heart rate during locomotion, limb position and movement, respiratory quotient, vertical climbing, 5, , 99(fig.), 101(fig.), 102(table) MacInnes, Donald, 16 magnetostratigraphy. See paleomagnetic data/magnetostratigraphic ages Majiwa, taxonomic identification from, 24 male primates. See primates, male mammalian biostratigraphy. See biostratigraphy, mammalian mammalian sites, Neogene sub-saharan Africa, 64(fig.)

10 276 Index mammalian taxa, from Neogene sub-saharan Africa, 65(fig.), 66(fig.) mandibular fossae, Otavipithecus namibiensis, 34 mass-specific carbon dioxide production, in macaque locomotion, 161, 161(fig.) mass-specific locomotor cost formula for primates, 159 in macaque locomotion, mass-specific oxygen consumption, in macaque locomotion, 161 Matano, Shozo, 4 Matsuda, T., 7 mental map, 186 Mesopithecus pentelici, 48 metabolism, and bipedal locomotion, 164. See also locomotor energetics metacarpophalangeal joint postures, knuckle-walking, 107 Mfangano Island, taxonomic identification from, 22, 23 Micropithecus 19, 21, 23 25, 211 midcarpal joint, breadth in primates, 113(fig.) broad midcarpal joint as adaptation to stability, 119 and capitates and hamates, 117(fig.), 117(table), 118(fig.) comparisons between humans, African apes, and Asian Apes, and lunate articular surface, 116(fig.) metacarpophalangeal joint postures, 107 morphometric assessment, morphometric measurement techniques, and scaphoid central facet, 115(fig.) and triquetral facet angle, 114(fig.) and wrist extension hypothesis, 108 and wrist joint range of motion, 111(fig.) Miocene ages and geological backgrounds of hominoids, desert biomes, faunal change in sub-saharan Africa, history of taxonomic identification of hominid species, paleoenvironments, 176(fig.) phytochores, primate distribution biases, 212 tree distribution, 183(fig.) See also paleoenvironment; specific formations and fossil species Mitsushio, H., 7 monkeys, African. See baboon; patas monkey monkeys, Asian. See Macaca fascicularis; Macaca nemestrina; macaque, Japanese monkeys, New World. See capuchin; spider monkey Moroto, taxonomic identification from, 19, 20 Morotopithecus, See also Afropithecus turkanensis Morotopithecus bishopi, 20 morphology and energy efficiency in hominids, 167 and locomotion in fossil hominids, 131 macaque studies and morphophysiology, of the midcarpal joint in knuckle-walkers and terrestrial quadrupeds, See also body proportions and size; joints; muscles; posture and gait; vertebral column; specific muscles, bones, and joints morphometrics assessment in knuckle-walkers and terrestrial quadrupeds, measurement techniques in knuckle-walkers and terrestrial quadrupeds, Mount Elgon, taxonomic identification from, 19 Muruyur Formation alpha taxonomy, 211 habitat, 215 Miocene anthropoids from, taxonomic identification from, 24 muscle action, defined, 136 muscles, 135 attachment and cortical thickness in femur, 131 attachment in hominoid spine, contractile forces and bone morphology, 130 EMGs of human limb muscles, 136, , 140(fig.), 229(fig.) energy consumption model, 171 and generalized biped, and model of human locomotion, 168(fig.), morphology and the origin of bipedalism, myological studies, 3 Nachola alpha taxonomy, 211 geologic map, 73(fig.), 76(fig.) geology of, habitat, 215 K-Ar and magnetostratigraphic ages, 77(fig.) location of, 73(fig.)

11 Index 277 and Miocene anthropoids, and Neogene System, 71, 72, 77(fig.) taxonomic identification from, 24 See also Samburu Hills Nacholapithecus age of, 78(fig.), 79(fig.), 89, 93 and arborealism, 103, dentition, 261, 262 geological background, Nacholapithecus kerioi, 7, 8, 24, 211 Nakano, Yoshihiko, 7 Nakaya, H., 7 Namib Desert and vegetation during the Miocene, Namibia. See Berg Aukas; Otavi Mountains Namurungule Formation age samples, composition of, 74, distribution of, 74 habitat, 215 and Hipparion, 89 magnetostratigraphy of, and paleoenvironments, as part of Neogene System, 71, 72 and Samburupithecus, 81(fig.) taxonomic identification from, 24 See also Samburu Hills Napak, taxonomic identification from, 19, 20 natal grouping of male apes, 237 natural remanent magnetization (NRM), Neogene. See also Miocene Neogene formation at Nachola-Samburu Hills, 77(fig.) Neogene sub-saharan Africa faunal sets, 62(table), 65(fig.), 66(fig.) mammalian sites, 64(fig.) mammalian taxa, 66(fig.) Neogene System, divisions of, 71 neontological primatology, neuro-musculoskeletal model of human locomotion, 170(fig.) Ngorora Formation, taxonomic identification from, 24 Niemitz, Carsten, 5 nonlocomotor behaviors, and gluteus maximus muscle, nonmatrilineal groups, and great apes, 238 North Side Story, NRM. See natural remanent magnetization Nyakach alpha taxonomy, 211 taxonomic identification from, 24 Nyanzapithecus, 23 25, 211 obligate bipedal locomotion, 149 occupations, human, and relative metabolic rate, 228(fig.) Okada, Morihiko, 4 Ombo, taxonomic identification from, 24 open country hypothesis, See also savanna hypothesis open country taxa, Miocene, operational sex ratio, 241 orangutan (Pongo) hand postures, 106(fig.) joint range of motion measurements, social structure, Oreopithecidae, 25 Oreopithecus, taxonomic identification and re-classification, 21, 23, 211 Orrorin age of, and faunal associations, and phytocores, 182 Orrorin tugenensis age of, 94 arboreal adaptations, as early bipedal hominid, 71, 204 and East Side Story, 225 femora of, (figs.) geological background of, 82 83, 204 orthograde posture, and Homo sapiens, 34 Otavi Mountains, taxonomic identification from, 34 Otavipithecus, Otavipithecus namibiensis, 34 angle of retroglenoid tubercle, 34(table) and arborealism, 34 dentition, 191 jaw, 191(fig.) oxygen measurement of locomotion in humans, 227(fig.) in macaques, pair bonds, and great apes, paleoecology and hominid bipedalism, of proto-hominids and the mental map, 186 of Victoriapithecus macinnesi, See also diet; paleoenvironment

12 278 Index paleoenvironment in African from Early Miocene to present, 176(fig.) and Australopithecus, 201 and faunal turnover, and hominid evolution, 6, 59 68, , , and Hominidae niches, and neontological primatology, and Orrorin tugenensis, 203(fig.) and Paranthropinae niches, 252 and Paranthropus, 201 and Sahelanthropus tchadensis, 205 and social structure of primates, and Victoriapithecus macinnesi, paleomagnetic data/magnetostratigraphic ages, 86 Lukeino Formation, 90(fig.) Nachola, 77(fig.) Namurungule Formation, Samburu Hills, 77(fig.), 87(fig.), 88(fig.) palmigrade walking, 106, 193, 199 Pan and dispersed food hypothesis, female dispersal, female-philopatric groups, 240 gluteus superficialis, 139 male-philopatric social structure, and phytocores, (figs.) time of separation from gorilla lineages, 20 See also bonobo; chimpanzee Pan-Homo, last common ancestor, 105 Pan paniscus. See bonobo Pan troglodytes. See chimpanzee Panidae, taxonomy, 251(table) Papio anubis. See baboon Paranthropinae, taxonomy, 251(table), 252 Paranthropus, 201 Parapapio lothagamensis, 48 patas monkey (Erythrocebus patas) hand postures, 106(fig.) joint range of motion measurements, locomotor energetics, pelvis rotation of, stabilization of, phylogenetic position, and social structure of primates, phytochores, 178(fig.), 180(fig.), 181(fig.), , 193 Pickford, Martin, 7 Pilbeam, D. R., 18, 20 plantigrade climbing, Platydontopithecus, 25 Plio-Pleistocene hominids, lower cervical vertebrae, lumbar vertebrae, 39 sacrum and sacro-iliac joint, upper cervical vertebrae, Pliocene taxa, appearance of, 63 Pliopithecus, taxonomic identification and re-classification, 15, 17, 18, 21. See also Limnopithecus Pongidae/pongids, taxonomic identification and re-classification, 21, 22, 25, 251(table) Pongo. See orangutan posterior gluteal line, 137 postional behavior, models of, posture and gait of Australopithecinae, , 253 of Australopithecus antiquus seu afarensis, 37 of bipedally trained animals (see macaque, Japanese; rat) of chimpanzees and gorillas, 193 computer simulations, and gluteus maximus muscle, 139 and hominid evolution, 193 of humans, 34, 135, 139, of Limnopithecus, 17 of Nacholapithecus, of Orrorin tugenensis, 71, 204 simulations of pathological gaits, 172 of Victoriapithecus macinnesi, 216 See also arborealism; knuckle-walking; locomotion, modes of Praeanthropous africanus, 24, predation among primates, Preuschoft, Holger, 5 primates and adaptive radiation, , 135 angular displacement of joints, 101(fig.) climbing (see climbing) distribution of extant species in East Africa, (figs.), distribution of extinct species in East Africa, 181(fig.) evolution of social structure, (see also social structure of hominoids) evolution of terrestriality, 105 (see also terrestriality) female primates (see primates, female) gluteus superficialis of nonhuman primates, 137 locomotor energetics of nonhuman primates, locomotor patterns, , (see also locomotion, modes of )

13 Index 279 male primates (see primates, male) musculoskeletal systems, 135 (see also specific bones and muscles) origins of terrestriality, and phytocores, (figs.) and predation, superior vertebro-articular process angle of nonhominoid primates, 38(fig.) trained primates (see macaque, Japanese) See also apes, African; apes, Asian; specific species primates, female estrus sex ratio hypothesis, female dispersal, female-philopatric groups, 235, 238, 240 female transfer, 237 interbirth intervals, See also social structure of hominoids primates, male male coalitions, male-philopatric social structure, natal grouping of male apes, 237 toleration of copulation by other males, See also social structure of hominoids primatology, neontological, principal component plots of humerus, ceboids and cercopithecines, 47(fig.) Proconsul distribution, 181(fig.) vertebral column, Proconsulidae, taxonomic identification and re-classification, 15 19, Procynocephalus, 48 pronograde semiterrestrial locomotion, 158 Propliopithecus, 16 propulsive plantarflexion, 103 Proterozoic Mozambique Belt, 72 quadrupedism vs. bipedal locomotion, 152, 158, , 167, effects of gravity on, 5 energetics, 152, 158, , and propulsive plantarflexion, 103 rat experiments, , vs. suspensory locomotion, 152, 158 quadrupeds, terrestrial. See knuckle-walking; terrestriality quiescence, and gluteus maximus muscle, radio-scaphoid articulation, extension-limiting role of, 115 radius (cercopithecines and ceboids), 51(fig.) Ramapithecus, taxonomic identification and re-classification, 19, 21 Rangwapithecus 21, 22, 23, 25, 211 rat, applications to fossil studies, bipedal standing exercises, changes in femur (see rat femur) modes of locomotion, 124(fig.), running exercises, rat femur, bone density, 126, 129(table) bone mass and load, changes in response to environment, compact bone thickness, 126(fig.), 127 compressive load, 127 cortical thickness, 127(fig.), 128(fig.), cross-sectional morphology, 125(fig.), 129(table) relative metabolic rate (RMR) during human locomotion, 227(fig.) during human occupations, 228(fig.) respiratory quotient (RQ), retroglenoid tubercle, 36 rib-cage, 32 Rift System, East African, and evolution of hominoids, 71 riverine delta habitat and Victoriapithecus macinnesi, RMR. See relative metabolic rate Rormuch Sills, composition of, 82 RQ. See respiratory quotient running and gluteus maximus muscle, 139, rat experiments, Rusinga Island, taxonomic identification from, 16, 17, 22, 23 sacro-iliac joint, sacrum, Sahara, and vegetation during Miocene, Sahelanthropus, 190 Sahelanthropus tchadensis, Samburu Hills paleomagnetic data, 87(fig.), 88(fig.) taxonomic identification from, 24 See also Nachola

14 280 Index Samburupithecus age of, dentition, 262 geological background, 80 82, Samburupithecus kiptalami, 7,24 sarumawashi, 151 savanna hypothesis, , , 205 savannitization, 68 sclerocarp feeders, 216, 217 Sebei District, taxonomic identification from, 19 sensory information, diagram of integration of, 170(fig.) sex ratio estrus hypothesis, operational, 241 socionomic, 242 shoulder displacement in primates, 100, 101(fig.) Simiolus diet and dentition, , 23, 24, 25, 211 Simons, Elwyn, 17, 18 Simpson s formula, Sivapithecus, taxonomic identification and re-classification, 16, 18, 19, 23 skeletal muscle, innervation of, 169(fig.) skeletal specializations, and primates, 135. See also specific bones and joints social structure of hominoids, , 236(fig.) African great apes, dispersed food hypothesis, and environmental change, 242(fig.) estrus sex ratio hypothesis, evolution of early human social structure, socionomic sex ratio, 242 Songhor, taxonomic identification from, 16, 22 South Africa, taxonomic identification from, 35 South Side Story and hominid origins, spider monkey (Ateles geoffroyi) EMGs of limb muscles, , 230(fig.) heart rate during locomotion, , 231(fig.) limb position and movement, locomotor energetics, vertical climbing, 5, , 99(fig.), 101(fig.), 102(table) spinous processes, cervical, in Plio-Pleistocene hominids, 37 Sprague Dawley rat. See rat standing posture, and gluteus maximus muscle, 139 stenotypy and apes, 179 vs. eurytopy in African hominoids, 185(fig.) as factor in hominid evolution, stooping, and gluteus maximus muscle, human, stratigraphy Namurungule Formation, Neogene formation at Nachola-Samburu Hills, 77(fig.) See also biostratigraphy, mammalian; paleomagnetic data/magnetostratigraphic ages superior articular facet, Australopithecus antiquus seu afarensis, 35 superior vertebro-articular process angle, 38(fig.), 39 suspensory locomotion. See arborealism Swartkrans, taxonomic identification from, 35 taxonomy, of Australopithecinae, 251(table) of Hominidae, , 251(table) of Homininae, of Hominoidea, 251(table) of Hylobatidae, 251(table) of Panidae, 251(table) of Paranthropinae, 252 of Pongidae, 251(table) See also specific regions and species taxonomy, alpha of Australopithecinae, 253 of Miocene Anthropoidea, temporary base camp hypothesis, terrain, and locomotor costs, terrestriality in African apes, body size as causal factor in development of, and diet, 49 50, 55 of fossil apes, hypothesis, 260, and joint adaptations, origins of in higher primates, 45 55, 105 and paleoenvironment, and skeletal measurements, stages of, 264(fig.) terrestrial catarrhines, 55, 105, 108 ungulate adaptations, 186 and Victoriapithecus, and wrist postures, 106 See also bipedal locomotion; knuckle-walking territoriality and evolution of African apes, 179 and origin of bipedalism, 184

15 Index 281 thermal demagnetization, Theropithecus oswaldi, and terrestriality, 48 thoracic kyphosis, in humans, 35 thoracic-lumbar kyphosis, in chimapanzees, 35 threat display, and bipedalism, Tirr Tirr Formation composition and distribution of, 75 as part of Neogene System, 71, 72 tool-using, development of, 254 trait aquisition, Ceboidea, 48 transverse processes lumbar vertebrae of Hominoidea, 33(fig.) in Otavipithecus namibiensis, 34 in Plio-Pleistocene hominids, 37 triquetral angle in primates, 114(fig.) trunk, rotation of, and gluteus maximus muscle, Tugen Hills geologic map, 73(fig.) taxonomic identification from, 24 Turkanapithecus, 24, 25 Turolian faunas, 60, 61, 68 Tuttle, Russell, 3 Uganda. See Bukwa II; Karamoja District; Moroto; Mount Elgon; Napak; Sebei District Ugandapithecus and flexibility, 41 lumbar vertebrae, Ugandapithecus major, 20. See also Proconsul ulna (cercopithecines and ceboids), 50(fig.) Van Couvering, John, 22 Van Couvering, Judith, 22 vertebral body ventral crest, Australopithecus antiquus seu afarensis, 36 vertebral column cervical vertebrae, differences between apes and humans, evolution, flexibility in primates, 41 lordosis (see lordosis) lumbar accessory processes, lumbar transverse processes, lumbar vertebrae, 31 34, 32(table), 33(fig.), 39, 41(fig.) Miocene hominoids, Plio-Pleistocene hominids, sacrum and sacro-iliac joint, trunk characteristics, Victoriapithecidae, 45 Victoriapithecus dentition, 46, 49, 217 disputed ancestors of, 19 as sclerocarp feeder, 217 sister taxa of, , and terrestriality, Victoriapithecus macinnesi body size, 51 53, 52(table) diet, 49 50, 55 ecological niche, fossil distribution, 53 and paleoenvironment, postcranial features, 46 principal component analyses of humerus, 46 48, 47(fig.) principal component analyses of ulna, 46 48, 48(fig.) as a semiterrestrial quadruped, 216 skeletal measurements, , 211 type site, 23 volcanism, 67, 73(fig.), 75 West Side Story, Winam Gulf, taxonomic identification from, 16 wrist joint and knuckle-walking, 107, 108, 118, 120 and limited extension, range of motion in primates, , 111(fig.) and suspensory motion, 118, 120 Xenopithecus, 16, 23, 24 Xenopithecus koruensis, 15 Yerkes Regional Primate Research Center, 3 Yonin gumi, 4 zygapophyses, 39, 40(fig.)

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