Paleontology and Geology of Laetoli: Human Evolution in Context (eBook)

Paleontology and Geology of Laetoli: Human Evolution in Context 3
Preface 7
Contents 9
Contributors 11
Chapter 1: Introduction: The Laetoli Hominins and Associated Fauna 15
Introduction 15
The Laetoli Fauna 16
Paleoecology 25
References 27
Chapter 2: Rodents 29
Introduction 29
Material and Methods 30
Systematics 30
Discussion and Conclusion 60
Appendix 2.1 List of new material attributed to Pedetes laetoliensis from the Upper Laetolil Beds (ULB) 64
Appendix 2.2 List of specimens attributed to Xerus janenschi from the Upper Laetolil Beds (ULB) and the Upper Ndolanya Beds (UNB) 65
Appendix 2.3 List of specimens attributed to Saccostomus major from the Upper Laetolil Beds (ULB) and S.cf. major from th 65
Appendix 2.4 List of specimens attributed to Heterocephalus quenstedti from the Upper Laetolil Beds (ULB) 66
References 66
Chapter 3: The Lower Third Premolar of Serengetilagus praecapensis (Mammalia: Lagomorpha: Leporidae) from Laetoli, Tanzania 68
Introduction 68
Stratigraphic Context 70
Materials and Methods 71
Results 73
Occlusal Morphology 73
Measurements 74
Comparisons with Serengetilagus from Other Localities 76
Conclusions 77
Appendix 3.1 Specimens of Serengetilagus praecapensis from Laetoli, Tanzania, included in this study 78
References 78
Chapter 4: Macroscelidea 80
Introduction 80
Methods 81
Systematic Paleontology 81
Discussion 82
Conclusions 86
Appendix 4.1 Species and provenance of comparative specimens of extant Rhynchocyon 86
References 87
Chapter 5: Galagidae (Lorisoidea, Primates) 88
Introduction 88
Systematics 89
Description of EP 1064/03 90
Morphology of Laetolia sadimanensis 90
Taxonomic and Phylogenetic Relationships 92
Conclusions 92
References 93
Chapter 6: Cercopithecids (Cercopithecidae, Primates) 95
Introduction 95
Material and Methods 96
Systematics and Description 97
Parapapio Jones, 1937 97
Distribution and Taxonomy 97
Parapapio ado (Hopwood, 1936) 98
Description of Cranio-Dental Morphology 102
Comparisons 113
Papionini gen. et sp. indet. 117
Description of Cranio-Dental Morphology 117
Rhinocolobus Leakey, 1982 118
Distribution and Taxonomy 119
cf. Rhinocolobus sp. 119
Description of Cranio-Dental Morphology 121
Comparisons 127
Cercopithecoides Mollett, 1947 129
Taxonomy and Distribution 130
Cercopithecoides sp. 130
Description of Cranio-Dental Morphology 130
Comparisons 133
Postcranial Remains of Fossil Cercopithecids from Laetoli 134
Postcranials Attributed to Parapapio 135
Postcranials Attributed to the Large Papionin 140
Postcranials Attributed to cf. Rhinocolobus sp. 141
Postcranials of Cercopithecoides sp. 143
Unassigned Cercopithecid Postcranials 145
Temporal and Spatial Distribution of Fossil Cercopithecids at Laetoli 146
Conclusions 148
References 149
Chapter 7: Hominins from the Upper Laetolil and Upper Ndolanya Beds, Laetoli 152
Introduction 152
Material 154
Provenance 155
New Hominins from the Upper Laetolil Beds 159
EP 2400/00 159
Location and Stratigraphic Provenance 159
Preservation 159
Morphology 160
Comparisons 161
EP 162/00 164
Location and Stratigraphic Provenance 164
Preservation 164
Morphology 165
Comparisons 165
MB.Ma. 8294 166
Location and Stratigraphic Provenance 166
Preservation 166
Morphology 166
Comparisons 167
L.H. 29 167
Location and Stratigraphic Provenance 167
Preservation 167
Morphology 168
Comparisons 169
LAET 79-5447 170
Location and Stratigraphic Provenance 170
Preservation 170
Morphology 170
Comparisons 171
New Hominins from the Upper Ndolanya Beds 172
EP 1500/01 172
Location and Stratigraphic Provenance 172
Preservation 174
Morphology 174
Comparisons 178
EP 1000/98 179
Location and Stratigraphic Provenance 179
Preservation 179
Morphology 180
Comparisons and Functional Implications 182
LAET 75-3817 185
Location and Stratigraphic Provenance 185
Preservation 186
Description 186
Comparisons 187
Discussion 187
Conclusions 193
References 194
Chapter 8: Carnivora 200
Introduction 200
Material and Methods 201
Systematic Paleontology 201
Genus Nyctereutes Temminck, 1838 209
?Nyctereutes barryi sp. nov. (Fig. 8.1) 209
Synonymy: cf. Canis brevirostris Barry, 1987 209
Genus ?Canis 211
cf. Canis sp. A (Fig. 8.3) 211
cf. Canis sp. B (Fig. 8.5) 213
Genus Otocyon Müller, 1836 213
aff. Otocyon sp. (Fig. 8.6) 213
Genus Prepoecilogale Petter, 1987 215
Prepoecilogale bolti (Cooke, 1985) (Fig. 8.8) 215
Prepoecilogale sp. 215
Genus Mellivora Storr, 1780 215
Mellivora sp. 215
Tribe Aonyxini Sokolov, 1973 216
Aonyxini gen. et sp. nov. (Fig. 8.9) 216
Mustelidae indet. 216
Genus Viverra Linnaeus, 1758 217
Viverra leakeyi Petter, 1963 217
Genus Genetta Cuvier, 1817 217
Genetta sp. (Fig. 8.10) 217
aff. Viverridae 217
Genus Herpestes Illiger, 1811 218
Herpestes palaeoserengetensis Dietrich, 1942 (Fig. 8.12) 218
Herpestes cf. H. palaeoserengetensis 219
Herpestes ichneumon Linnaeus, 1758 220
Genus Galerella Gray, 1865 220
Galerella sp. 220
Genus Helogale Gray, 1862 222
Helogale palaeogracilis (Dietrich, 1942) 222
Helogale cf. H. palaeogracilis 224
Genus Mungos Geoffroy St.-Hilaire and Cuvier, 1795 224
Mungos dietrichi Petter, 1963 (Fig. 8.16) 224
Mungos sp. nov.? (Fig. 8.16) 225
Genus Crocuta Kaup, 1828 226
Crocuta dietrichi Petter and Howell, 1989 226
Genus Parahyaena (Hendey, 1974) 227
Parahyaena howelli Werdelin, 2003 (Fig. 8.18) 227
Genus Ikelohyaena Werdelin and Solounias, 1991 229
Ikelohyaena cf. I. abronia Hendey, 1974 229
Genus Lycyaenops Kretzoi, 1938 230
Lycyaenops cf. L. silberbergi (Broom in Broom and Schepers, 1946) (Fig. 8.19) 230
Genus ?Pachycrocuta Kretzoi, 1938 230
?Pachycrocuta sp. (Fig. 8.20) 230
Genus Proteles Geoffroy St.-Hilaire, 1824 230
aff. Proteles sp. (Fig. 8.21) 231
Genus Dinofelis Zdansky, 1924 231
Dinofelis petteri Werdelin and Lewis, 2001 231
Genus Homotherium Fabrini, 1890 232
Homotherium sp. (Fig. 8.22) 232
Genus Panthera Oken, 1816 232
Panthera sp. aff. P. leo (Linnaeus, 1758) (Fig. 8.23) 232
Panthera sp. cf. P. pardus (Linnaeus, 1758) (Fig. 8.25) 234
Genus Acinonyx Browokes, 1828 235
Acinonyx sp. (Fig. 8.27) 235
Genus Caracal Gray, 1843 or Genus Leptailurus Severtzow, 1858 237
Caracal sp. or Leptailurus sp. (Fig. 8.28) 237
Genus Felis Linnaeus, 1758 237
Felis sp. (Fig. 8.29) 238
Discussion 238
Canidae 238
Mustelidae 239
Viverridae 239
Herpestidae 239
Hyaenidae 239
Felidae 239
Summary 240
Note 241
References 241
Chapter 9: Proboscidea 244
Introduction 244
Abbreviations 245
Systematic Paleontology 245
Discussion 268
References 269
Chapter 10: Orycteropodidae 274
Introduction 274
Material 275
Description 275
Cranio-Dental Specimens 275
Postcranials 277
Comparisons 283
Conclusions 284
References 284
Chapter 11: Rhinocerotidae 286
Introduction 286
Materials and Methods 286
Institutional Abbreviations 286
Material 287
Stratigraphy 287
Mesowear 287
Systematic Paleontology 288
Synonymy 288
Material 289
Lower Laetolil Beds (NMT) 289
Upper Laetolil Beds (NMT) 289
Probably Upper Laetolil Beds (KNM) 290
Remarks and Comparisons 290
Crania 291
Permanent Upper Dentition 291
Upper Deciduous Dentition 292
Mandible and Lower Dentition 292
Material 292
Upper Ndolanya Beds (NMT) 292
Remarks and Comparisons 292
Material 293
Upper Ndolanya Beds (NMT) 293
Remarks and Comparisons 293
Restricted Synonymy 293
Material 293
Upper Laetolil Beds (NMT) 293
Remarks and Comparisons 293
Cranium 293
Material 294
Lower Laetolil Beds (NMT) 294
Upper Laetolil Beds (NMT) 294
Probably Upper Laetolil Beds (KNM) 294
Upper Ndolanya Beds (NMT) 294
Ngaloba Beds (NMT) 294
Remarks and Comparisons 294
Permanent Dentition 294
Postcrania 295
Discussion 295
Taxonomic History of the Laetoli Rhinoceroses 295
Relationships of the Plio-Pleistocene Dicerotini 296
Early Evolution of Dicerotini 298
Paleoecology of the Laetoli Rhinoceroses 300
Conclusions 302
References 303
Chapter 12: Equidae 306
Introduction 306
Abbreviations, Definitions and Methods 308
Abbreviations in Text 309
Measurement Table Abbreviations 309
General Analyses 309
Abbreviations for Laetoli Horizons 310
Metric Procedures 310
Statistical Analysis 310
Metrical Results 310
Maxillary P2 310
Calcanea 318
Astragali 318
Metacarpal III 318
Metatarsal III 320
First Phalanges III 322
Summary of Metrical Results 324
Description of the Material 324
Maxillary Teeth 324
Upper Laetolil Beds 324
Upper Ndolanya Beds 326
Mandibular Cheek Teeth 328
Lower Laetolil Beds 328
Upper Laetolil Beds 328
Upper Ndolanya Beds 329
Postcrania 332
1PH III – Cranial and Caudal Comparisons 333
Later Material 333
Equus from Emboremony 1 and 2 (Ngaloba Beds) 333
Systematics 333
Discussion 334
References 335
Chapter 13: Suidae 338
Introduction 338
Materials and Methods 338
Systematic Paleontology 339
Discussion 344
References 348
Chapter 14: Giraffidae 349
Introduction 349
Cranial Specimens and Taxonomic Attribution of Giraffids 350
Lower Laetolil Beds 350
Upper Laetolil Beds 351
Upper Laetolil Dento-Gnathic Specimens 351
Giraffa aff. jumae 352
Giraffa stillei 353
Sivatherium maurusium 358
Upper Laetolil Beds Postcranial Specimens 358
Giraffa aff. jumae 359
Giraffa stillei 361
Sivatherium maurusium 362
Upper Ndolanya Beds 363
Upper Ndolanya Beds Dento-Dnathic Specimens 364
Upper Ndolanya Beds Postcranial Specimens 365
Giraffa aff. jumae 365
Giraffa aff. stillei 365
Additional Specimen from the Upper Ndolanya Beds 366
Discussion 366
Lower Laetolil Beds 366
Giraffa aff. stillei 366
Sivatherium 366
Giraffidae indet. 367
Upper Laetolil Beds 367
Giraffa aff. jumae 368
Giraffa stillei 368
Upper Ndolanya Beds 369
Giraffa aff. jumae 369
Giraffa aff. stillei and Giraffa aff. pygmaea 369
Sivatherium aff. maurusium 369
Diversity of Giraffids at Laetoli 370
References 371
Chapter 15: Bovidae 373
Introduction 373
Systematics: Bovidae of the Upper Unit of the Laetolil Beds 375
Material 376
Description 376
Discussion 376
Other Species and Occurrences 377
Occurrence and Range 377
Generic Diagnosis 377
Remarks 377
Holotype 378
Type Locality 378
Material 378
Diagnosis 378
Description 378
Discussion 378
Material 379
Description 379
Discussion 379
Holotype 379
Type Locality 379
Material 379
Diagnosis 381
Description 381
Discussion 382
Material 383
Description 383
Discussion 383
Material 384
Description 384
Generic Diagnosis 385
Remarks 385
Syntypes 385
Type Locality 385
Horizon and Range 385
Material 385
Diagnosis 385
Discussion 385
Generic Diagnosis 385
Remarks 385
Holotype 386
Name 386
Material 386
Diagnosis 387
Description 387
Discussion 388
Diagnosis 389
Remarks 389
Lectotype 389
Type Locality 389
Horizon and Range 389
Material 389
Diagnosis 390
Description 390
Discussion 390
Material 390
Description 390
Discussion 390
Lectotype 391
Type Locality 391
Horizon and Range 391
Material 391
Diagnosis 392
Description 392
Discussion 393
Other Genera 393
Material 393
Description 393
Discussion 393
Short Diagnosis 394
Remarks 394
Material 394
Description 395
Discussion 396
Diagnosis 398
Remarks 398
Lectotype 398
Type Locality 398
Horizon and Range 398
Material 398
Diagnosis 398
Description 398
Discussion 399
Other Species 399
Occurrences and Range 399
Diagnosis 399
Remarks 400
Holotype 400
Type Locality 400
Horizon and Range 400
Material 400
Diagnosis 401
Description 401
Discussion 401
Material 402
Description 403
Discussion 403
Material 405
Description 405
Systematics: Bovidae of the Lower Unit of the Laetolil Beds 405
Material 405
Material 405
Description 405
Material 405
Material 405
Material 405
Description 406
Material 406
Description 406
Material 407
Description 407
Material 407
Description 407
Discussion 407
Material 407
Description 407
Conclusions 407
Systematics: Bovidae of the Upper Ndolanya Beds 408
Material 408
Description 408
Discussion 408
Bovini sp. or spp. indet.Material 408
Description 409
Discussion 409
Material 410
Description 410
Discussion 410
Description 411
Discussion 411
Material 411
Description 411
Discussion 411
Material 411
Description 412
Other Species 412
Short Diagnosis 412
Remarks 412
Material 412
Diagnosis 412
Description 412
Discussion 413
Type Locality 413
Material 413
Description 413
Material 413
Description 414
Discussion 414
Other Species 414
Occurrence and Range 414
Diagnosis 414
Holotypes 414
Horizon and Range 414
Material 414
Diagnosis 414
Description 414
Discussion 415
The Position of Parestigorgon gadjingeri 415
Material 415
Description 416
Discussion 416
Material 416
Diagnosis 416
Description 416
Discussion 416
Holotype 417
Type Locality 417
Horizon and Range 417
Name 417
Material 417
Diagnosis 417
Description 419
Discussion 420
Conclusions 422
Appendix 423
A. Upper Laetolil Beds 423
Locality 1 423
Laetoli Locality 1NW 425
Laetoli Locality 2 425
Laetoli Locality 3 429
Laetoli Locality 4 430
Laetoli Locality 5 431
Laetoli Locality 6 432
Laetoli Locality 7 434
Laetoli Loc. 7E 437
Laetoli Locality 8 437
Laetoli Locality 9 439
Laetoli Locality 9 S 442
Laetoli Locality 10 444
Laetoli Locality 10E 446
Laetoli Locality 10NE 450
Laetoli Locality 10W 450
Laetoli Locality 11 453
Laetoli Locality 12 455
Laetoli Locality 12E 455
Laetoli Locality 13 456
Laetoli Locality 13 “Snake Gully” 457
Laetoli Locality 13E 458
Laetoli Locality 15 458
Laetoli Locality 16 459
Laetoli Locality 17 460
Laetoli Locality 19 460
Laetoli Locality 20 461
Laetoli Locality 21 461
Laetoli Locality 22 462
Laetoli Locality 22E 463
Laetoli Locality 24 463
B. Upper Ndolanya Beds 464
Laetoli Locality 1 464
Laetoli Locality 7E 464
Laetoli Locality 9 466
Laetoli Locality 14 466
Laetoli Locality 15 466
Laetoli Locality 18 467
Laetoli Locality 22E 469
Laetoli Locality 22S 469
Silal Artum 470
C. Lower Laetolil Beds 471
Kakesio 471
Esere 472
Emboremony 472
D. Ngaloba Beds 472
Emboremony 472
References 473
Chapter 16: Amphibia and Squamata 476
Introduction 476
Systematic Account 476
Discussion and Conclusions 485
References 486
Chapter 17: Tortoises (Chelonii, Testudinidae) 488
Introduction 488
Material 489
Systematics and Description 493
Comments 499
Comments 502
Paleobiology and Paleoecology 503
Age Structure and Mortality in Stigmochelys brachygularis 503
Fossil Eggs 506
Paleoecological Considerations 507
Other Fossil Tortoises from Africa 507
Conclusions 509
References 510
Chapter 18: Aves 513
Introduction 513
Material and Methods 513
Systematic Paleontology 514
Discussion 536
Appendix: List of the Modern Comparative Osteological Specimens Examined 539
References 540
Chapter 19: Beetles (Insecta: Coleoptera) 542
Introduction 542
Tenebrionidae (W. Schawaller) 542
Tentyriini Species a (? Genus Tentyria) 543
? Tentyriini Species B (? Genus) 545
? Tentyriini Species C (? Genus) 545
Molurini Species a (? Genus Arturium) 546
Biology 546
Scarabaeidae (F.-T. Krell) 546
Dynastinae: Oryctini/Pentodontini 547
Melolonthinae: Schizonychini, Species a 548
Ichnofossils 549
Biology 553
Conclusions 554
References 554
Chapter 20: Lepidoptera, Insecta 556
Introduction 556
Lepidoptera Phylogeny and Classification, with Special Reference to Saturniidae 556
Fossil History of Lepidoptera 557
Description 558
Taxonomy 559
Paleobiology 559
References 560
Chapter 21: Trace Fossils Interpreted in Relation to the Extant Termite Fauna at Laetoli, Tanzania 562
Introduction 562
Exposures at Locality 9 562
Fossil Termite Hives 563
Other Fossilized Structures 564
Exposures at Locality 10W 565
Fossil Termite Nests 565
Flattened Ovoids 565
Probable Macrotermes Nests 567
Shafted Chambers 568
Inverted Pear-Shape Chambers 568
Vertical Shafts 568
Anastomosing Passages and Root Casts 568
Kakesio 3 569
Discussion of Preservation 569
Modern Termites at Laetoli 569
Hodotermes mossambicus 569
Macrotermes herus 569
Odontotermes montanus 570
Discussion 570
References 571
Chapter 22: Gastropoda 573
Introduction 573
Taxonomy and Ecology 574
Systematics 574
Interpretation of the Paleoecology Using the Gastropod Assemblage 586
Sampling Issues 586
Overall Character of the Pliocene Fauna 586
Paleoenvironmental Interpretation 588
Lower Laetolil Beds 589
Upper Laetolil Beds 590
Upper Ndolanya Beds 591
Note 591
References 592
Index 594