ThisPageIntentionallyLeftBlank C O N T E N T S Tables&Illustrations ix APrimateDay 55 Preface xvii PrimateDiets 55 Locomotion 57 SocialLife 59 Individuals,Groups,andCommunities 62 ONE Adaptation,Evolution, WhyPrimatesLiveinGroups 63 ProtectionfromPredators • ImprovedAccessto andSystematics Food • AccesstoMates • Assistancein Adaptation 1 ProtectingandRearingOffspring • Evolution 1 Phylogeny • PolyspecificGroups SpeciesandSpeciation • Patternsof PrimateLifeHistories 67 Evolution • Phylogeny LifeHistoryDiversity TaxonomyandSystematics 5 PrimateCommunities 72 Bibliography 9 Bibliography 73 TWO FOUR ThePrimateBody Prosimians Size 11 Strepsirhines 82 CranialAnatomy 12 MalagasyStrepsirhines 85 BonesoftheSkull • TeethandChewing • Cheirogaleids • Lemurids • Lepilemurids(or TongueandTaste • MusclesofFacialExpression Megaladapids) • Indriids • Daubentoniids TheBrainandSenses 19 SubfossilMalagasyProsimians 104 TheBrain • CranialBloodSupply • Palaeopropithecines,orSlothLemurs • Olfaction • Vision • Hearing SubfossilLemurids • SubfossilLepilemurids TheTrunkandLimbs 27 AdaptiveRadiationofMalagasyPrimates 109 AxialSkeleton • UpperLimb • Lower GalagosandLorises 110 Limb • LimbProportions Galagids • Lorisids SoftTissues 36 AdaptiveRadiationofGalagosandLorises 116 DigestiveSystem • ReproductiveSystem PhyleticRelationshipsofStrepsirhines 117 GrowthandDevelopment 39 Tarsiers 118 Bibliography 43 PhyleticRelationshipsofTarsiers 120 Bibliography 122 THREE PrimateLives FIVE PrimateHabitats 47 NewWorldAnthropoids ForestHabitats • PrimatesinTropical AnatomyofHigherPrimates 133 Ecosystems Platyrrhines 136 LandUse 52 Pitheciines • Callicebinae • Aotines • ActivityPatterns 52 Atelines • Cebinae • Callitrichines v vi CONTENTS AdaptiveRadiationofPlatyrrhines 171 AnatomicalCorrelatesofSocial PhyleticRelationshipsofPlatyrrhines 173 Organization 306 Bibliography 174 AdaptationandPhylogeny 308 Bibliography 309 SIX OldWorldMonkeys TEN CatarrhineAnatomy 185 TheFossilRecord Cercopithecoids 185 GeologicalTime 316 Cercopithecines • Colobines Paleomagnetism • PlateTectonicsand AdaptiveRadiationofOldWorldMonkeys 220 ContinentalDrift • Paleoclimate PhyleticRelationshipsofOldWorld FossilsandFossilization 320 Monkeys 221 Paleoenvironments 323 Bibliography 218 ReconstructingBehavior 324 Paleobiogeography 325 Bibliography 326 SEVEN ApesandHumans Hominoids 235 ELEVEN Hylobatids • Pongids • SizeandEvolutionof PrimateOrigins theAfricanApes • Hominids Archontans—PrimatesandOther AdaptiveRadiationofHominoids 256 Mammals 330 PhyleticRelationshipsofHominoids 258 TreeShrews • FlyingLemurs • Bats Bibliography 259 Plesiadapiforms 332 Purgatorius • Plesiadapids • Carpolestids • Saxonellids • Paromomyids • Micromomyids • Microsyopids • EIGHT Palaechthonids • Picrodontids • Picromomyids PrimateCommunities AdaptiveRadiationofPlesiadapiforms 342 PrimateBiogeography 267 PlesiadapiformsandPrimates 344 EcologyandBiogeography 270 ThePhylogeneticOriginsofPrimatesamong ComparingPrimateCommunities 272 theArchonta 345 Bibliography 282 TheAdaptiveOriginofPrimates 346 Bibliography 347 NINE PrimateAdaptations TWELVE EffectsofSize 283 FossilProsimians SizeandDiet • SizeandLocomotion • Size TheFirstModernPrimates 354 andLifeHistory • SizeandEcology TheOriginofProsimians 356 AdaptationstoDiet 291 DentalAdaptations • OtherOralAdaptations • Adapoids 356 DigestiveTractAdaptations • Dietand Notharctines • Cercamoniines Ranging • DietandSocialGroups (Protoadapines) • Adapines • Sivaladapines LocomotorAdaptations 297 AreAdapoidsStrepsirhines? 369 ArborealQuadrupeds • FossilLorisesandGalagos 371 TerrestrialQuadrupeds • Leapers • Omomyoids 372 SuspensoryPrimates • Bipeds • Omomyids • Anaptomorphines • LocomotorCompromises • Locomotion, Omomyines • Microchoerids • Asian Posture,andEcology Omomyoids • AfricanOmomyoids • Tarsiids CONTENTS vii Omomyoids,Tarsiers,andHaplorhines 383 TheEvolutionofLivingHominoids 480 AdaptiveRadiationsofEoceneProsimians 385 Bibliography 483 PhyleticRelationshipsofAdapidsand Omomyids 387 Bibliography 388 SIXTEEN FossilOldWorldMonkeys Victoriapithecids:TheEarliestOldWorld THIRTEEN Monkeys 491 EarlyAnthropoids FossilCercopithecids 495 EoceneAnthropoidsfromAsia 397 FossilCercopithecines • FossilColobines EoceneandOligoceneAnthropoidsfromAfrica SummaryofFossilCercopithecoids 504 andArabia 399 Bibliography 506 FossilPrimatesfromFayum,Egypt • Parapithecids • Propliopithecids • Oligopithecids OtherNorthAfricanandArabianEarly SEVENTEEN Anthropoids 415 Hominids,theBipedalPrimates Oman • Algeria • Tunisia Australopithecines 511 EarlyAnthropoidAdaptations 416 Ardipithecus • Australopithecus • PhyleticRelationshipsofEarly Paranthropus Anthropoids 417 AustralopithecineAdaptationsandHominid ProsimianOriginsofAnthropoids 418 Origins 523 SolvingAnthropoidOrigins 421 PhyleticRelationshipsofEarlyHominids 529 Bibliography 421 EarlyHomo 530 Homohabilis • Homorudolfensis • Homoerectus andHomoergaster LateHomo 535 FOURTEEN Homoheidelbergensis • Homoneanderthalensis • FossilPlatyrrhines Homosapiens ThePlatyrrhineFossilRecord 427 HumanPhylogeny 539 TheEarliestPlatyrrhines 430 HumansasanAdaptiveRadiation 542 ThePatagonianPlatyrrhines 431 Bibliography 542 AMoreModernCommunity 436 PleistocenePlatyrrhines 441 CaribbeanPrimates 441 SummaryofFossilPlatyrrhines 443 EIGHTEEN PatternsinPrimateEvolution PlatyrrhineOrigins 444 Bibliography 447 PrimateAdaptiveRadiations 551 BodySizeChanges • DietaryDiversity • LocomotorDiversity PatternsinPrimatePhylogeny 556 FIFTEEN PrimateEvolutionattheSpeciesLevel 558 FossilApes MosaicEvolution 561 LatestOligocenetoMiddleMioceneApesfrom PrimateExtinctions 563 Africa 453 ClimaticChanges • Competition • Predation AdaptiveRadiationofProconsulids 454 • LimitingPrimateExtinctions PhyleticRelationshipsofProconsulidsand Bibliography 567 OtherAfricanMioceneApes 465 Glossary 571 EurasianFossilApes 467 ClassificationofOrderPrimates 577 Pliopithecids • Oreopithecus • Pongidae Index 581 ThisPageIntentionallyLeftBlank T A B L E S & I L L U S T R A T I O N S Tables 6.8 FamilyCercopithecidae,Subfamily COLOBINAE,odd-nosedmonkeys 218 1.1 Aclassificationofthetuftedcapuchin 7.1 FamilyHYLOBATIDAE 239 monkey 5 7.2 FamilyPONGIDAE 243 2.1 Skeletalproportions 35 7.3 FamilyHOMINIDAE 255 4.1 FamilyCheirogaleidae 84 11.1 SuperfamilyINCERTAESEDIS 336 4.2 FamilyLemuridae 89 11.2 SuperfamilyPLESIADAPOIDEA 336 4.3 FamilyMegaladapidae 94 11.3 SuperfamilyMICROSYOPOIDEA 341 4.4 FamilyIndriidae 96 11.4 SuperfamilyINCERTAESEDIS 421 4.5 FamilyDaubentoniidae 99 12.1 FamilyIncertaesedis 356 4.6 FamilyGalagidae 109 12.2 FamilyNotharctidae,Subfamily 4.7 FamilyLorisidae 111 NOTHARCTINAE 360 4.8 FamilyTarsiidae 115 12.3 FamilyNotharctidae,Subfamily 5.1 SubfamilyPithecinae 140 CERCAMONIINAE 364 5.2 SubfamilyCallicebinae 147 12.4 FamilyIncertaesedis 365 5.3 SubfamilyAotinae 148 12.5 FamilyAdapidae,Subfamily 5.4 FamilyAtelinae 149 ADAPINAE 367 5.5 FamilyCebinae 157 12.6 FamilySivaladapidae,Subfamily 5.6 SubfamilyCallitrichinae 161 SIVALADAPINAE 369 6.1 FamilyCercopithecidae,Subfamily 12.7 SuperfamilyLORISOIDEA 371 CERCOPITHECINAE,macaques 192 12.8 FamilyOmomyidae,Subfamily 6.2 FamilyCercopithecidae,Subfamily ANAPTOMORPHINAE 377 CERCOPITHECINAE,mangabeys 194 12.9 FamilyOmomyoidea,Subfamily 6.3 FamilyCercopithecidae,Subfamily OMOMYINAE 380 CERCOPITHECINAE,baboonsand 12.10 SuperfamilyOmomyoidea,Family geladas 196 MICROCHOERIDAE 381 6.4 FamilyCercopithecidae,Subfamily 12.11 SuperfamilyOmomyoidea,Subfamily CERCOPITHECINAE,mandrillsand Incertaesedis 382 drills 199 12.12 FamilyTARSIIDAE 382 6.5 FamilyCercopithecidae,Subfamily 13.1 SuperfamilyIncertaesedis 398 CERCOPITHECINAE,guenons 201 13.2 SuperfamilyParapithecoidea 404 6.6 FamilyCercopithecidae,Subfamily 13.3 SuperfamilyPropliopithecoidea 411 Colobinae,Africancolobusmonkeys 207 14.1 InfraorderPLATYRRHINI 428 6.7 FamilyCercopithecidae,Subfamily 15.1 EARLYANDMIDDLEMIOCENE COLOBINAE,langursandleaf APES 456 monkeys 211 15.2 FamilyPliopithecidae 468 ix x TABLES&ILLUSTRATIONS 15.3 FamilyOREOPITHECIDAE 472 2.24 Skeleteonofbaboonfoot 35 15.4 FamilyPONGIDAE 475 2.25 Footskeletonandsoleofsixprimates 35 16.1 FamilyVictoriapithecidae,Subfamily 2.26 Orangutandigestivesystem 37 Victoriapithecinae 492 2.27 Gorillareproductiveorgans 38 16.2 FamilyCercopithecidae,Subfamily 2.28 Primatefetalmembranes 40 CERCOPITHECINAE 496 2.29 Timingofprimatelifehistoryevents 41 16.3 FamilyCercopithecidae,Subfamily 2.30 Humanandnonprimatemammalgrowth COLOBINAE 501 curves 42 17.1 FamilyHOMINIDAE 512 2.31 Primategrowthcurves 43 2.32 Primatedentaleruptionsequences 43 Illustrations CHAPTER3 3.1 Primategeographicdistribution 48 CHAPTER1 3.2 Primatehabitats 49 1.1 Sharedspecializationsandancestral 3.3 Rainforestmicrohabitats 50 features 4 3.4 Primatelanduse 53 1.2 Aprimateclassification 6 3.5 Benefitsandcostsofdiurnalityand 1.3 Twoapproachestoapeandhuman nocturnality 54 classification 9 3.6 Primateactivityhistograms 56 3.7 Spatialdistributionoffoodresources 56 CHAPTER2 3.8 Primatelocomotorbehaviors 58 2.1 Mouselemurandgorilla,thesmallestand 3.9 Primatefeedingpostures 59 largestlivingprimates 12 3.10 Primatesocialgroups 60 2.2 Mammaliansizeranges 13 3.11 Primatelifehistorydiversity 69 2.3 Thehumanskull 14 3.12 Modelofmammalianlifehistory 70 2.4 Capuchinandlemurskulls 15 3.13 Primatelifehistoryparameters 71 2.5 Siamangdentalformula 16 3.14 Parentalcareandinfantgrowth 72 2.6 Primitiveprimatedentition 16 2.7 Primateskullandchewingmuscles 17 2.8 Primatetongues 18 CHAPTER4 2.9 Primatemusclesoffacialexpression 18 4.1 Geographicdistributionofextant 2.10 Primatebrains 20 prosimians 81 2.11 Functionalareasofthehumanbrain 21 4.2 Gradisticandphyleticdivisionof 2.12 Primatecranialbloodsupply 22 primates 82 2.13 Primatenasalregions 23 4.3 Malagasystrepsirhinedentitions 83 2.14 Primateandcatnoses 24 4.4 Strepsirhineskeletalfeatures 84 2.15 Primateandraccoonorbits 25 4.5 Strepsirhineskulls 84 2.16 Primateearstructure 26 4.6 ForesttypesofMadagascar 85 2.17 Primateexternalears 27 4.7 Sixcheirogaleidgenera 87 2.18 Primatetympanicregions 28 4.8 Predationofmouselemurbyowl 88 2.19 Spidermonkeyskeleton 29 4.9 Dietofforestheightpreferenceforfive 2.20 Baboonskeletonandlimbmusculature 30 sympatricMalagasyprosimians 89 2.21 Terminologyforanatomical 4.10 Threelemuridspecies 92 orientation 31 4.11 Gentlebamboolemurs 95 2.22 Skeletonofababoonhand 32 4.12 Threesympatricbamboolemurs 96 2.23 Handskeletonandpalmofsix 4.13 Sportivelemurs 97 primates 32 4.14 Sifaka,indris,andwoollylemurs 99 TABLES&ILLUSTRATIONS xi 4.15 Indriidskulls 101 5.22 White-lippedtamarinandsaddle-back 4.16 Indriskeleton 102 tamarin 166 4.17 Aye-aye 103 5.23 Liontamarin 169 4.18 ReconstructedMalagasynaturalsetting(ca. 5.24 Marmosetandtamarinlower 8,000–1,000B.P.) 105 dentition 170 4.19 Palaeopropithecusingensskeleton 107 5.25 Marmosetfaces 171 4.20 TheadaptivediversityofMalagasy 5.26 Pygmymarmoset 171 prosimians 109 5.27 Adaptivediversityofplatyrrhines 172 4.21 FivesympatriclorisoidsfromGabon 111 5.28 Platyrrhinephylogeny 174 4.22 Dietandforestheightpreferenceforfive sympatricgalagosandlorisesfrom CHAPTER6 Gabon 112 6.1 Anatomicalfeaturesofcercopithecoidsand 4.23 Slowlorisandslenderloris 116 hominoids 187 4.24 Phylogenyofstrepsirhines 117 6.2 Geographicdistributionof 4.25 Tarsiusskull,dentitionandskeleton 119 cercopithecoids 188 4.26 Spectraltarsier 121 6.3 Anatomicalfeaturesofcolobinesand cercopithecines 189 CHAPTER5 6.4 Skullsofcercopithecinesand 5.1 Anatomicalfeaturesofanthropoidsand colobines 190 prosimians 134 6.5 Crab-eatingmacaqueandpig-tailed 5.2 Anthropoidcranialanddental macaque 191 features 135 6.6 Diet,forestheightpreference,and 5.3 Geographicdistributionofextantand locomotorbehaviorofsixsympatric extinctplatyrrhines 137 Malaysiancatarrhines 192 5.4 Platyrrhineandcatarrhinecranial 6.7 Mangabeysandcercopithecoidmolecular characteristics 137 phylogeny 194 5.5 Platyrrhinedentitionsadaptedtodifferent 6.8 Savannahbaboon 196 foodtypes 138 6.9 Baboonskeleton 197 5.6 Platyrrhineskulls 138 6.10 Geladas 198 5.7 Twopithecineprimates 141 6.11 Mandrill 199 5.8 SevensympatricSuriname 6.12 Guenonfaces 202 platyrrhines 142 6.13 Crownedguenon,greaterspot-nosed 5.9 Diet,forestheightpreferences,and guenon,moustachedguenon 203 locomotorbehaviorforsevensympatric 6.14 Dietandforestheightpreferencesoffive Surinameplatyrrhines 143 Gabonmonkeys 204 5.10 Beardedsakiskeleton 145 6.15 DeBrazza’smonkeys 205 5.11 Titimonkey 146 6.16 Vervetmonkeys 206 5.12 Nightmonkey 148 6.17 Redcolobusandblack-and-white 5.13 Redhowlingmonkey 151 colobus 208 5.14 Woollymonkey 153 6.18 GeographicdistributionofAsianleaf 5.15 Blackspidermonkey 154 monkeys 212 5.16 Woollyspidermonkey 156 6.19 Purple-facedmonkeyandHanuman 5.17 Tuftedcapuchinandsquirrel langur 213 monkey 158 6.20 Spectacledlangurandbandedleaf 5.18 Squirrelmonkeyskeleton 159 monkey 215 5.19 Unusualfeaturesofcallitrichines 162 6.21 Locomotorandanatomicaldifferences 5.20 Goeldi’smonkey 164 betweenPresbytismelalophosand 5.21 Tamarinfaces 165 Trachypithecusobscura 216 xii TABLES&ILLUSTRATIONS 6.22 Proboscismonkey 218 8.6 Distributionofprimategroupsizeinfour 6.23 Goldenmonkey 219 majorgeographicalareas 273 6.24 AdaptiveradiationofOldWorld 8.7 Ecologicaldistributionofspeciesintwo monkeys 220 MalagasyandtwoAfricanprimate 6.25 MorphologicalphylogenyofOldWorld communities 274 monkeys 221 8.8 Ecologicaldistributionofspeciesintwo 6.26 MolecularphylogeniesofOldWorld AsianandtwoSouthAmericanprimate monkeys 222 communities 275 8.9 Diurnalandnocturnalprimate CHAPTER7 communities,Ranomafana, 7.1 Geographicdistributionofextant Madagascar 276 apes 236 8.10 Diurnalandnocturnalprimate 7.2 Characteristicskeletalfeaturesofextant communities,Marazolaza,Madagascar 277 apes 237 8.11 Diurnalandnocturnalprimate 7.3 Geographicdistributionandfacial communities,TaiForest,IvoryCoast 278 characteristicsofextantgibbons 238 8.12 DiurnalprimatesofKualaLompat, 7.4 Lowerjawsofasiamangandan Malaysa 279 orangutan 239 8.13 PrimatecommunityofRaleighvallen- 7.5 Gibbonskeleton 240 VoltsbergNatureReserve,Surinam 280 7.6 White-handedgibbonand siamang 241 7.7 Locomotorbehaviorandfeedingpostures CHAPTER9 oftheMalayansiamang 242 9.1 Mathematicalrelationshipoflinear,areal, 7.8 Orangutan 244 andvolumetricdimensions 284 7.9 Mountaingorilla 246 9.2 Femoraofagorillaandapygmy 7.10 Gorillaskeleton 247 marmoset 285 7.11 Chimpanzee 249 9.3 Growthallometry,intraspecificallometry, 7.12 Bonobochimpanzee 252 interspecificallometry 286 7.13 SkullallometryofAfricanapes 253 9.4 Correlationofprimatedietaryhabitsand 7.14 Human 254 bodysize 287 7.15 Humanskeleton 254 9.5 Correlationofprimatelocomotorbehavior 7.16 Femalelifespanofhumansandother andbodysize 289 vertebrates 257 9.6 Locomotorconsequencesof 7.17 Adaptiveradiationofliving bodysize 289 hominoids 258 9.7 Posturalconsequencesofbodysize 290 7.18 Phyleticrelationshipsamong 9.8 Dailyfoodintakeandbodysize 291 hominoids 258 9.9 Primateanatomicaladaptationsto diet 293 CHAPTER8 9.10 Foodpropertiesanddentalblades 294 8.1 Distributionoflivingprimatesinfour 9.11 Anatomicalfeaturesofaprimatearboreal biogeographicalareas 268 quadruped 298 8.2 Relationshipofprimatespeciesandtropical 9.12 Anatomicalfeaturesofaprimateterrestrial rainforestarea 269 quadruped 300 8.3 Relationshipofprimatespeciesandannual 9.13 Anatomicalfeaturesofaprimate rainfall 270 leaper 301 8.4 Distributionofprimatebodysizesinfour 9.14 Anatomicalfeaturesofasuspensory biogeographicalareas 271 primate 302 8.5 Distributionofprimateactivitypatternsand 9.15 Anatomicalfeaturesofabipedal dietinfourmajorgeographicalareas 272 primate 303
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