The Human Career: Human Biological and Cultural Origins [3rd ed.] 9780226439655

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The human Career

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The human Career Human Biological and Cultural Origins

Third ediTion

Richard G. Klein

T h e u n I V e r S I T Y O F C h I C a G O P r e S S • C h i cago an d L o n do n

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Richard G. Klein is professor of anthropology and human biology at Stanford University. His books include Ice Age Hunters of the Ukraine and (with Kathryn Cruz-Uribe) he Analysis of Animal Bones from Archeological Sites, both published by the University of Chicago Press.

The university of Chicago Press, Chicago 60637 The university of Chicago Press, Ltd., London © 1989, 1999, 2009 by The university of Chicago all rights reserved. Published 2009 Printed in the united States of america 18 17 16 15 14 13 12 11 10 09

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ISBn-13: 978-0-226-43965-5 (cloth) (cloth) ISBn-10: 0-226-43965-8

Library of Congress Cataloging-in-Publication Data Klein, richard G. The human career : human biological and cultural origins / richard G. Klein. —3rd ed. p. cm. Includes bibliographical references and index. ISBn-13: 978-0-226-43965-5 (cloth: alk. paper) ISBn-10: 0-226-43965-8 (cloth: alk. paper) 1. human beings—Origin. 2. Fossil hominids. 3. human evolution. I. Title. Gn281.K55 2009 599.93' 8—dc22 2008029270

a The paper used in this publication meets the minimum requirements of the american national Standard for Information Sciences—Permanence of Paper for Printed Library materials, anSI Z39.48-1992.

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ConTenTs

List of Illustrations vii List of Tables xvii Preface to the Third edition xix Preface to the Second edition xxiii Preface to the First edition xxvii 1

Evolution, Classification, and Nomenclature 1

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The Geologic Time Frame 19

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The Primate Background 65

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The Australopiths and Homo habilis 131

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Evolution of the Genus Homo 279

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The Neanderthals and Their Contemporaries 435

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Anatomically Modern Humans 615

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Synopsis: Anatomy, Behavior, and Modern Human Origins 725 references 753 reference Index 935 Site Index 000 Subject Index 000

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1.1. he process of speciation as visualized by gradualists and by advocates of punctu-

ated equilibrium 1.2. A currently popular phylogeny of the hominins 1.3. A popular hierarchical classiication of the living Primates down to the family

level 1.4. A cladogram illustrating the evolutionary relationships among humans, chim-

panzees, gorillas, orangutans, gibbons, rhesus monkeys, and spider monkeys 2.1. he geologic timescale and the proposed ages of some important evolutionary

events 2.2. Stages in the evolution of Mimomys, Arvicola, and Microtus 2.3. Middle and late Pleistocene vole stratigraphy of central and southeastern Europe 2.4. Shed mature antlers of red deer from Mosbach, Germany, and Ilford, England 2.5. he average number of plates on third molars in successive species of

Mammuthus 2.6. Time ranges of the Elephantinae in Africa 2.7. Time ranges of fossil suids in Africa ater 10 Ma 2.8. Time ranges of the later Miocene to Holocene equids in Africa 2.9. Time ranges covered by various numerical dating methods 2.10. A proposed curve for calibrating radiocarbon ages to calendar (solar) years 2.11. Global paleomagnetic stratigraphy for the past 5 million years 2.12. Global temperature change over the past 70 million years 2.13. he δ oxygen-18 record for the past 900,000 years 2.14. he maximum extent of Quaternary glaciation and associated sea-level change 3.1. Reconstructed skull of Australopithecus africanus 3.2. Right upper and lower dentitions of various primates 3.3. Reconstructed skeleton of Australopithecus afarensis 3.4. Diagrammatic section through the right ear region of a therian mammal 3.5. Geographic distribution of the living nonhuman primates 3.6. Nostril orientation in New World and Old World monkeys

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3.7. Cephalic views of the rib cage and right shoulder girdle of an adult macaque

monkey and of an adult human 3.8. Lower molars of Propliopithecus and Colobus 3.9. Occlusal relation between the upper canine, lower canine, and lower third pre-

molar in a nonhuman catarrhine primate and a human being 3.10. Quadrupedal postures in an Old World monkey, a human, a gorilla, and a chim-

panzee 3.11. Skulls of male and female adult mandrills 3.12. Skulls of a Malagasy lemur and a New World monkey 3.13. Biomolecular estimates for the divergence times between great apes, Old World

monkeys, New World Monkeys, and strepsirrhines 3.14. Changing positions of the continents from the late Triassic to the middle Eocene 3.15. Two interpretations of early primate phylogeny 3.16. Skulls, foot bones, and reconstructions of Paleocene and Eocene primates 3.17. Eocene and Oligocene localities with possible or probable higher primates fossils 3.18. Mandibles of Apidium phiomense, Propliopithecus chirobates, and P. zeuxis 3.19. Facial and lateral views of the skull of Propliopithecus zeuxis 3.20. Anterior and distal views of the distal let humerus in various monkeys and in

Propliopithecus zeuxis 3.21. Temporal distribution of fossil hominoid genera 3.22. Relative positions of Africa and Eurasia in the early and middle Miocene 3.23. Approximate locations of the main African Miocene fossil hominoid sites 3.24. Reconstructed skull and skeleton of Proconsul heseloni 3.25. Fossil hominoid localities dating between about 16–15 and 8–7 Ma in relation to

the historic distribution of the chimpanzees and the gorilla 3.26. Reconstructed skeleton of Pliopithecus vindobonensis 3.27. Skulls of Sivapithecus indicus, Pongo pygmaeus, and Pan troglodytes 3.28. A provisional phylogeny of the Primates 4.1. Approximate locations of African Plio-Pleistocene fossil sites 4.2. Facial skeleton and endocast of Australopithecus africanus from Taung 4.3. he position of the skull relative to the spinal column in a gorilla and a modern

human 4.4. Ages of some important South African hominin fossil sites, inferred from mam-

malian species 4.5. Dating of the earliest artifact industries in Africa and Eurasia 4.6. Time spans of the most commonly recognized hominin species between 4.4 and

1.0 million years ago 4.7. hree stages in the evolution of the Swartkrans australopith cave 4.8. he location of Olduvai Gorge on the Serengeti Plain 4.9. Schematic stratigraphy of Olduvai Gorge 4.10. Schematic stratigraphic column for the Laetoli area 4.11. Distribution of fossiliferous geological formations in the Lake Turkana Basin 4.12. Schematic stratigraphy of the Koobi Fora Formation and correlation to the Shun-

gura Formation 4.13. he approximate numbers of hominin fossils by member within the principal

formations of the Omo Group

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4.14. Correlation of the Plio-Pleistocene successions in the Late Turkana Basin 4.15. Schematic stratigraphy of the Hadar Formation 4.16. he skull of Sahelanthropus tchadensis before reconstruction 4.17. Posterior view of the Orrorin tugensis proximal femur 4.18. Insertion of the masseter and temporalis muscles on a modern human skull 4.19. Mandibular dentitions of a female chimpanzee, Ardipithecus ramidus, and Aus-

tralopithecus afarensis 4.20. Upper canines and lower premolars of Dryopithecus, male and female chimpan-

zees, Ardipithecus ramidus, and Australopithecus afarensis 4.21. Lower deciduous anterior premolars of Dryopithecus, pygmy chimpanzee, com-

mon chimpanzee, Ardipithecus ramidus, Australopithecus afarensis, Au. africanus, Paranthropus robustus, P. boisei, and living humans 4.22. Schematic views of the skull of Australopithecus afarensis 4.23. Basal views of the skull of Pan troglodytes, Australopithecus afarensis, A. africanus,

A. robustus, and modern Homo sapiens 4.24. Occlusal views of mandibles of Australopithecus anamensis and A. afarensis 4.25. Lingual views of mandibles of Australopithecus anamensis and A. afarensis 4.26. Proximal tibiae of a chimpanzee, a living human, and Australopithecus anamensis 4.27. Lateral and anterior views of cranium AL 444-2, Australopithecus afarensis from

Hadar 4.28. Facial and occipital views of Pan troglodytes, Australopithecus afarensis, A. africa-

nus, Paranthropus robustus, P. boisei, and Homo habilis 4.29. Palates of a chimpanzee, various australopiths, and a modern human 4.30. Upper canine morphology of various australopiths, the chimpanzee, and modern

humans 4.31. Cross sections of the mandibular body below the fourth premolar in various

australopiths and in Homo habilis 4.32. Lower third premolar morphology in the chimpanzee, in Australopithecus afaren-

sis, and in modern humans 4.33. he superior margin of the mandibular ramus in Australopithecus afarensis,

Gorilla gorilla, Paranthropus robustus, Pan troglodytes, and modern Homo sapiens 4.34. Occipital views of cranial venous sinus systems typical for modern humans and

most other hominoids and for Australopithecus afarensis, Paranthropus boisei, and P. robustus 4.35. humb metacarpals of a modern human, Paranthropus robustus, Australopithecus

afarensis, and a chimpanzee 4.36. Lower limb and knee joint of a modern human, Australopithecus afarensis, and a

chimpanzee 4.37. Skulls and pelvises of a chimpanzee, Australopithecus afarensis, and a living human 4.38. Facial views of KNM-WT 40000 (Kenyanthropus platyops) and KNM-ER 1470

(Homo rudolfensis) 4.39. Basicranial lexion in gracile australopiths, robust australopiths, Homo habilis,

and modern H. sapiens 4.40. Superior and lateral views of the skull of Australopithecus garhi 4.41. Maxillae of Australopithecus garhi and Paranthropus boisei 4.42. Relative limb bone proportions in Pan troglodytes, Australopithecus afarensis,

possible Australopithecus garhi, Homo ergaster, and Homo sapiens

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4.43. Skulls of Paranthropus aethiopicus and P. boisei 4.44. A mandible of Paranthropus robustus articulated with a reconstructed skull of

P. boisei 4.45. A reconstructed male skull of Paranthropus robustus 4.46. Reconstructed skulls of Homo habilis from Koobi Fora 4.47. Palatal view of maxilla AL 666-1 (Homo cf. habilis) from Hadar 4.48. Cladograms showing the evolutionary relationships of the australopiths and

Homo suggested by endocranial capacity and by cheek tooth area 4.49. A working phylogeny of the hominins 4.50. he basic types of stone artifacts found at Oldowan and Acheulean sites in

Africa 4.51. A range of typical Oldowan stone tools and their conventional typological

designations 4.52. Oldowan artifacts from sites EG10 and EG12, East Gona, Ethiopia 4.53. he relation between handedness and lake form 4.54. A partial loor plan of site DK 1 at Olduvai Gorge 4.55. Fossilized hyena feces, a cut-marked bone, and various tooth-marked bones 4.56. he abundance of large mammals at the FLK Zinj site, Olduvai Gorge Bed I 5.1. he phylogeny of the genus Homo 5.2. he occupied world roughly 500,000 years ago 5.3. Principal fossiliferous localities in Java 5.4. he main Chinese sites with fossils of primitive Homo 5.5. Approximate ages of the main sites with fossils of Homo ergaster, H. erectus, early

H. neanderthalensis, and early H. sapiens 5.6. Tentative correlation of global marine oxygen-isotope stages and key European

human fossil sites 5.7. Franz Weidenreich’s restorations of Homo erectus skulls from Java and China 5.8. Skulls of Javan Homo erectus and a robust modern person 5.9. Front and side views of Ngandong skull XI and Sangiran Skull 17 5.10. Browridge form in Indonesian and Chinese Homo erectus 5.11. Mandibles of Indonesian Homo erectus and a robust modern person 5.12. Skull KNM-ER 3733 (Homo ergaster) from Koobi Fora, East Turkana 5.13. Skull of KNM-WT 15000 (Homo ergaster) from Nariokotome III, West Turkana 5.14. Reconstructed skeletons of Homo ergaster and Australopithecus afarensis 5.15. Fossils of Homo ergaster from Swartkrans Cave 5.16. Fossil skullcap from the Daka Member of the Bouri Formation, Middle Awash

Valley 5.17. Skullcap of Olduvai Hominid 9 from Upper Bed II, Olduvai Gorge 5.18. Skullcap of archaic Homo from the Narmada Valley, India 5.19. Fossil skull from Swanscombe, England 5.20. Rear views of skulls of Homo erectus, early H. sapiens, modern H. sapiens, early

H. neanderthalensis, and classic H. neanderthalensis 5.21. Skulls 4, 5, and 6 from the Sima de los Huesos, Atapuerca 5.22. Facial view of skull 5 from the Sima de los Huesos, Atapuerca 5.23. Mandibles AT 505 and AT 605 from the Sima de los Huesos, Atapuerca

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5.24. Midsagittal sections through the occipital bones of Homo erectus, early H. nean-

derthalensis, and later H. neanderthalensis 5.25. Fossil skulls from Steinheim (Germany), Arago (France), and Petralona (Greece) 5.26. Fossil human mandible from Montmaurin, France 5.27. he principal fossil skull from Bodo, Middle Awash 5.28. Fossil skull from deposits of seasonal Lake Ndutu 5.29. Fossil skull from Kabwe, Zambia 5.30. Fossil skullcap from Elandsfontein, South Africa 5.31. Skull 1 from Jebel Irhoud, Morocco 5.32. Fossil skull from Singa, Sudan 5.33. Fossil skull from Florisbad, South Africa 5.34. Fossil skull from Dali, China 5.35. Fossil skull from Jinniushan, China 5.36. Fossil skull from Maba, China 5.37. Probable human dispersal routes from Africa superimposed on the locations of

early archaeological or human fossil sites in Africa and Eurasia 5.38. A schematic section through the Dmanisi site, Georgia 5.39. Skull 2700 and mandible 2735 from Dmanisi, Georgia 5.40. Mandible KNM-ER 922 from East Turkana, Kenya, and mandible D211 from

Dmanisi, Georgia 5.41. Skulls D2280 and D2282 from Dmanisi, Georgia 5.42. Skull 3444 and mandible 3900 from Dmanisi, Georgia 5.43. he approximate locations of European sites with early human fossils or artifacts 5.44. Reconstructed fossil skullcap from Ceprano, Italy 5.45. A juvenile maxilla from the Gran Dolina, Atapuerca 5.46. Bifaces from site TK, Bed II, Olduvai Gorge 5.47. he approximate locations of the main African Acheulean sites 5.48. Earlier and later Acheulean bifaces from the Bouri Formation, Middle Awash 5.49. Earlier and later Acheulean hand axes from Sterkfontein Cave and Kathu Pan 5.50. Earlier and later Acheulean cleavers from Sterkfontein Cave and Elandsfontein 5.51. Later Acheulean artifacts from Elandsfontein Cutting 10 5.52. Artifacts associated with Homo erectus at Zhoukoudian Locality 1 5.53. Locations of the main excavated early Paleolithic sites in western Asia 5.54. Acheulean bifaces from Torralba, Spain 5.55. Late Acheulean artifacts from southern England 5.56. Tayacian artifacts from La Caune de l’Arago, France 5.57. Bone and artifact concentrations at Bilzingsleben, Germany 5.58. Floor plan of an early Middle Paleolithic layer at Le Lazaret Cave, France 5.59. A laked fragment of elephant bone from Bilzingsleben, Germany 5.60. Wooden throwing spears from Schöningen, Germany 5.61. An elephant tibia shat fragment with scored lines from Bilzingsleben, Germany 5.62. A putative igurine from Berkehat Ram, Golan Heights, and the “Venus of

Lespugue” 5.63. Proportionate representation of diferent large mammals at the Elandsfontein

Acheulean site

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5.64. Proportionate representation of bovid species at the Elandsfontein Acheulean

site 5.65. Proportionate representation of carnivore species at the Elandsfontein Acheulean

site 5.66. A schematic section through the Sima de los Huesos, Atapuerca 6.1. he European and west Asian realm of the Neanderthals 6.2. Approximate locations of European and north African Middle Paleolithic sites 6.3. Skullcap of Neanderthal 1 from Feldhofer Cave 6.4. Approximate locations of southwest Asian Middle and Upper Paleolithic sites 6.5. Derived features on Neanderthal skulls from La Ferrassie Cave (France) and

Amud Cave (Israel) 6.6. Neanderthal skull from La Quina Cave (France) and the non-Neanderthal skull

from Broken Hill (Zambia) 6.7. Adult male Neanderthal and modern human skeletons 6.8 Labyrinthine inner ear structure in pygmy chimpanzees, modern humans, and

the La Ferrassie 1 Neanderthal 6.9. Shoveling in Neanderthal upper front teeth from Krapina, Croatia 6.10. he dentition of Shanidar Neanderthal skull 1, illustrating the retromolar space

and labial wear 6.11. Taurodont upper cheek teeth from Krapina Cave, Croatia 6.12. Superior margin of the mandibular ramus in a recent human, near-modern

humans, and Neanderthals 6.13. Coniguration of the mandibular foramen in a modern human from Vogelherd,

Germany, and a Neanderthal from La Chapelle-aux-Saints, France 6.14. Femurs of a Neanderthal and of a modern person 6.15. he axillary border of the scapula in Neanderthals and modern humans 6.16. Hand skeletons of a Neanderthal and an early-modern human 6.17. he shat/neck angle on proximal femurs of a Neanderthal and a modern Eur-

american male 6.18. Reconstructed physiques of a Neanderthal and of an early-modern European 6.19. Superior view of the reconstructed Neanderthal pelvis from Kebara Cave, Israel 6.20. Approximate locations of key north African Aterian and Mousterian/Middle

Stone Age (MSA) sites 6.21. Approximate locations of east African Middle and Later Stone Age sites 6.22. Approximate locations of the main MSA sites in southern Africa 6.23. A fossil skull from Jebel Irhoud, Morocco, and a Neanderthal skull from Spy

Cave, Belgium 6.24. Facial and lateral views of the near-modern human skull from Ngaloba, Tanzania 6.25. Near-modern human skull and mandible from Dar es Soltan 2, Morocco 6.26. he principal adult skull from Herto, Ethiopia 6.27. Skulls 1 and 2 from the Omo-Kibish Formation, Ethiopia 6.28. Occlusal and buccal views of mandibles 41815 and 16424 from Klasies River Main,

South Africa 6.29. he reconstructed face and skull of a Neanderthal associated with Châtelperro-

nian artifacts at Saint Césaire, France 6.30. Skull 9 from Qafzeh Cave, Israel

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6.31. Partial reconstitution of a lat, discoidal core by reitting of lakes from Mouste-

rian Site C, Maastricht-Belvédère, the Netherlands 6.32. Stages in the manufacture of a classic Levallois core 6.33. Two variants of the Levallois technique for producing elongated lakes or blades 6.34. he technique that Quina Mousterians oten used to “slice” lakes from an elon-

gated core 6.35. Basic Mousterian and MSA stone-tool types, as deined by François Bordes 6.36. Reduction of a simple convex sidescraper to other sidescraper types 6.37. Convergent sidescrapers from the Mousterian site of Biache-Saint-Vaast, France 6.38. Utilized Levalloiso-Mousterian artifacts from Kebara Cave, Israel 6.39. Schematic section through the deposits of Combe-Grenal Cave, France 6.40. Correlation between the archaeological sequences at Le Moustier and Combe-

Grenal, France 6.41. Paleolithic cultural stratigraphy and chronology of the Levant 6.42. Aterian artifacts from Algeria 6.43. Howieson’s Poort artifacts from Nelson Bay Cave, South Africa 6.44. Barbed bone points from Katanda, Democratic Republic of the Congo 6.45. he putative Mousterian lute from Divje Babe Cave 1, Slovenia 6.46. Proposed tick shell beads and an incised red ocher fragment from Blombos Cave,

South Africa 6.47. Floor plan and section through “ind level II” at Ariendorf, Germany 6.48. Plan of Molodova I, level 4, Ukraine 6.49. Approximate locations of South African sites that illuminate Middle and Later

Stone Age coastal ecology 6.50. he minimum numbers of eland, Cape bufalo, and bushpig in the Stone Age

levels of Klasies River Cave 1, Nelson Bay Cave, Die Kelders Cave 1, and Byneskranskop Cave 1, South Africa 6.51. Mortality proiles of eland and bufalo in the MSA layers of Klasies River Cave 1 6.52. Distal humerus breadth in known-age fur seals and in fossil or subfossil fur seal

samples from Stone Age and brown hyena sites on the coast of southern Africa 6.53. he maximum length of Cape turban shell opercula in Stone Age sites on the

south coast of South Africa 6.54. he maximum length of granite limpet shells in Stone Age sites on the west coast

of South Africa 6.55. Breadths of angulate tortoise distal humeri in Stone Age sites on the south coast

of South Africa 6.56. Breadths of angulate tortoise distal humeri in Stone Age sites on the west coast of

South Africa 6.57. Face of Shanidar 1, showing the crushed outer margin of the let orbit 6.58. Chronological arrangement of late Pleistocene cultural units and fossil human

types in Africa and Eurasia 6.59. he distribution of the Aurignacian, Châtelperronian, Uluzzian, and Szeletian/

Jerzmanowician Industries in Europe and western Asia roughly 37 ka 6.60. Stone and bone artifacts from the Châtelperronian layers of the Grotte du Renne,

France 6.61. Plan and reconstruction of the Châtelperronian hut emplacement in level XI of

the Grotte du Renne, France

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6.62. Two proposed routes for basal Aurignacian dispersal across Europe 6.63. Skulls of individual 1 from Shanidar Cave, Iraq, and of individual 3 from

Předmostí, Czech Republic 6.64. Bifacial leaf-shaped points from the upper deposits of Szeleta Cave, Hungary 6.65. Body proportions in the early-modern or near-modern Last Interglacial occu-

pants of Israel and the Last Glacial Neanderthals of Europe 6.66. Artifacts that characterize diferent variants of the Early Upper Paleolithic in

southwestern Asia 7.1. Reconstructed skulls of the Cro-Magnon 1 early-modern European and the La

Chapelle-aux-Saints Neanderthal 7.2. Approximate locations of major western and central European Upper Paleolithic

sites 7.3. Skull of an anatomically modern child from Kostenki XV, Russia 7.4. Skull of an anatomically modern adolescent from the Pataud Rockshelter, France 7.5. A phylogram based on complete mtDNA sequences from ity-three geographi-

cally dispersed individuals 7.6. Hypothetical routes of modern human dispersal from Africa 7.7. Typical Upper Paleolithic tool types 7.8. Typical Aurignacian split-base bone points, “pendants,” and chipped stone

artifacts 7.9. Gravettian stone artifacts 7.10. Solutrean artifacts from La Riera Cave, Spain 7.11. Magdalenian bone and stone artifacts from El Juyo Cave, Spain 7.12. Climate and cultural stratigraphy ater 186,000 years ago 7.13. Abundance of mammal species in the Magdalenian layers of El Juyo Cave Al-

tamira Cave, Spain 7.14. he chronology of major late Paleolithic technological innovations in Eurasia 7.15. Approximate locations of Siberian Mousterian and Upper Paleolithic sites 7.16. Structural ruins at the Mezin Upper Paleolithic site, Ukraine 7.17. Structural ruins at the Mezhirich Upper Paleolithic site, Ukraine 7.18. Structural ruins at the Pushkari 1 Upper Paleolithic site, Ukraine 7.19. Grooved antler artifacts from the Korolevo I Siberian Upper Paleolithic site 7.20. Use of the spear-thrower and a decorated example from southern France 7.21. Bone artifacts from the Later Stone Age layers of Nelson Bay Cave, South Africa 7.22. Approximate locations of key decorated caves in France and Spain and of caves in

Germany with some of the oldest known ivory igurines 7.23. Mammoth ivory “lion-man” from an early Aurignacian layer at Holhenstein-

Stadel, Germany 7.24. he early Aurignacian “Venus” from Galgenberg Hill, near Krems, Austria 7.25. Approximate locations of the Upper Paleolithic Gravettian sites with female

igurines or engravings 7.26. Mammoth ivory “Venus” igurines from Kostenki 1, layer 1, Russia 7.27. Plan of a burial pit dated to roughly 30 ka at Kostenki XV on the Don River,

Russia 7.28. Approximate locations of major Paleolithic sites in eastern Europe 7.29. Human igurines from the Mal’ta Upper Paleolithic site, Siberia

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7.30. Structural remnants at the Mal’ta Upper Paleolithic site, Siberia 7.31. Grave with a fragmentary child’s skeleton at the Mal’ta Upper Paleolithic site,

Siberia 7.32. Northeastern Siberia and northwestern North America at the height of the Last

Glaciation 7.33. he main in situ occurrences of Clovis Paleo-Indian artifacts and of like-aged

archaeological sites in the Americas 7.34. Map of Sunda and Sahul at the height of the Last Glaciation 8.1. A working phylogeny of the hominins from 4.5 million years ago to the present 8.2. Artifact types that characterize major Stone Age culture-stratigraphic units 8.3. Hominin Encephalization and Megadontia Quotients versus time

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lisT of Tables

1.1. A classiication of living people at twenty-two potential levels in the Linnaean

hierarchy 2.1. Basic parameters of some isotopes used in numerical dating 3.1. A provisional classiication of the Primates to the level of the suborder and to the

level of the family within the suborder Anthropoidea 3.2. Subdivision of the Hominoidea, with special reference to the Hominidae 4.1. Sedimentary units with hominoid fossils, artifacts, or both in the Tugen Hills 4.2. Hominin fossils from Lothagam 4.3. Hominin taxa and artifact traditions in the Koobi Fora Formation 4.4. Hominin taxa in the Lower Ormo River Basin 4.5. Hominin taxa and artifact traditions in the Nachukui Formation 4.6. Hominin fossil localities in the Middle Awash region 4.7. he earliest putative hominins 4.8. African Pliocene sites with large fossil samples and no hominins 4.9. he principal craniums and mandibles from which the evolutionary relationships

of the australopiths and early Homo must be inferred 4.10. Body, brain, and dental size estimates for extant Pan troglodytes, Homo sapiens,

and various fossil hominins 4.11. Sites with fossils of Homo that antedate 1.9–1.8 million years ago 4.12. he main sites assigned to the Oldowan Industrial Complex 5.1. he main Javan Homo erectus sites 5.2. he main Chinese Homo erectus sites 5.3. African sites with fossils of Homo ergaster 5.4. African sites with fossils of early Homo sapiens 5.5. European sites with fossils of early Homo neanderthalensis 5.6. Endocranial capacity in Homo ergaster and later fossil representatives of Homo 5.7. he minimum numbers of postcranial elements recovered from the Sima de los

Huesos, Atapuerca, through 1995 5.8. Characters that fossils from the Sima de los Huesos, Atapuerca, share with fossils

of Homo erectus, the classic Neanderthals, and H. sapiens

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5.9. Sites with fossils that could document late Homo erectus or H. heidelbergensis in

China 5.10. European sites that have provided only doubtful evidence for human presence

before 600,000 years ago 5.11. he main sites that document human presence in Europe before Global Isotope

Stage 5 5.12. Early west Asian archaeological and human fossil sites 5.13. Key African archaeological sites where Homo ergaster, H. heidelbergensis, or early

H. sapiens probably made the artifacts 5.14. he principal east and central Asian early archaeological sites 6.1. European sites that have produced important Neanderthal fossils since 1920 6.2. he African contemporaries of the Neanderthals 6.3. African sites where numeric dating indicates the Middle Stone Age (MSA) began

before 250–200 ka 6.4. he principal west and central Asian Mousterian sites that have provided Nean-

derthal or near-modern human remains 6.5. he principal stratiied MSA sites in southern Africa 6.6. East African sites that were occupied in the gap between 60 and 30 ka when

southern and northern Africa seem to have been largely abandoned 6.7. Mousterian/MSA sites with wooden artifacts 6.8. Mousterian/MSA sites with proposed art or personal ornaments 6.9. Prominent open-air Mousterian and MSA sites in Europe, western Asia, and

Africa 6.10. Mousterian/MSA sites that may preserve remains of structures 6.11. Mousterian/MSA sites that have provided remains of possible or probable food

plants 6.12. South African sites that illuminate Middle and Later Stone Age coastal foraging 6.13. North African, European, and west Asian coastal or near-coastal sites to which

Mousterian/MSA people brought shells of edible species 7.1. Non-European sites, in order of discovery, that have provided especially informa-

tive fossils of early fully modern humans 7.2 he principal African sites that have provided modern or near-modern human

fossils older than 50 ka 7.3. Sites with Neanderthal bones that have provided mitochondrial DNA 7.4. African sites with ostrich eggshell beads that antedate 30 ka 7.5. Late Paleolithic sites with remains of plants that people probably introduced

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PrefaCe To The Third ediTion

Paleoanthropology continues to prosper, and the broad pattern of human evolution is becoming ever clearer, even if many subsidiary issues remain to be settled. his book summarizes the overall pattern as a series of ive phases: (1) an australopithecine (or australopith) phase from 4.5 Ma (million years ago) or before until roughly 2.5 Ma; (2) a phase initiated by the appearance of the genus Homo and the irst archaeological sites about 2.5 Ma; (3) a phase marked by the emergence of Homo ergaster (or early African Homo erectus) 1.8–1.7 Ma and the irst dispersal of humans from Africa; (4) a phase initiated by an abrupt increase in brain size to near the modern average roughly 700–600 ka (thousands of years ago), the concomitant appearance of more sophisticated stone artifacts in Africa, and a subsequent Out-of-Africa event that produced the irst permanent settlement of Europe; and (5) a phase beginning about 50 ka when modern humans spread from Africa to swamp or replace nonmodern Eurasians. he phase scheme is a heuristic device rooted in chronological change, but the fossil record itself is patently more complex. Ater 3 Ma, it involved repeated branching events that brought forth contemporaneous hominin (“human”) species. By 50 ka there were at least three nonoverlapping species, Homo sapiens in Africa, Homo neanderthalensis in Eurasia, and Homo erectus in the Far East. It was of course Homo sapiens that spread from Africa 50 ka to replace the others. his book retains the fundamental structure of its predecessor, but each chapter has been substantially revised and updated. he irst chapter outlines current understanding of the evolutionary process. he second addresses the geologic time frame for human evolution, with emphasis on numeric dating methods and on climatic change as a major driving (natural selective) factor. It devotes roughly equal space to the radiocarbon, radiopotassium, and luminescence dating methods, although the reliability of the luminescence methods is oten questionable. heir main

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strength is their applicability to materials and time ranges that neither radiocarbon nor radiopotassium cover. he third chapter summarizes primate evolution before the emergence of hominins roughly 7 Ma. It is the only chapter whose length has been reduced from prior editions. he reason is that as the prehominin primate fossil record has grown, so have professional disagreements on how to interpret it, and it has become increasingly diicult to synthesize succinctly and to relate directly to human evolution. Chapters 4–7 present the ive basic phases of human evolution listed above, together with the branching phylogenetic tree that crosscuts them. hese chapters are longer than their counterparts in the previous edition, mainly because they incorporate the implications of important new discoveries. he new indings include fossils dated between 7 and 5 Ma that may document the oldest hominin species; fossils dated to 2.5 Ma that shed fresh light on the australopithecine roots of Homo; sites that may imply people reached northern China as much as 1.6 Ma; other sites that suggest people sporadically penetrated southern Europe between 1.1 Ma and 800 ka and northern Europe by perhaps 700 ka; new dates and additional fossils to document the evolution of Homo neanderthalensis in Europe between 500 and 70 ka; additional dates and fossils that illuminate the evolution of Homo sapiens in Africa over the same interval; and inally, the on-going retrieval of Neanderthal DNA, which ever more strongly underscores the divergence of the Neanderthals from modern humans. Like all overviews of human evolution, the one presented here is a narrative, but it is constrained by evidence, and the inclusion of fresh evidence and analyses means that it approximates the actual course of human evolution more closely than its predecessors. Chapter 8 summarizes the main points in chapters 4–7. It is titled “Conclusion,” but it is actually a synopsis or abstract that could be read irst as well as last. Not all the most recent indings are equally persuasive, and in some cases, including for example, the settlement of northern China by 1.6 Ma and of northern Europe by 700 ka, they require conirmation from additional discoveries. his follows from the realization that the fossil and archaeological records are inherently noisy and that multiple wellsubstantiated observations are required to identify a valid signal. My own belief, repeatedly expressed in the text, is that a irst occurrence should be treated as a possible accident and even a second should be regarded as a possible coincidence. Only repeated, independent, mutually consistent occurrences can document a reliable pattern. his philosophy explains what some may ind surprising—I have addressed the “hobbit” (Homo loresiensis) only in passing, even though it may one day bear out its frequent billing as one of the two or three most important fossil inds of the past hundred years.

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he recurring shortage of reliable, mutually supportive observations largely explains what lay observers and science writers oten see as the contentious nature of paleoanthropology, in which practitioners stake out contrary positions that seem irresolvable. Some issues may be truly irresolvable, but many others require only fresh observations to reduce or eliminate ambiguity, and the problem is that new observations often appear slowly. To see real progress, we must oten then be willing to wait years or even decades. he Neanderthals provide a case in point. As recently as twenty years ago, some prominent specialists argued that the Neanderthals were directly ancestral to modern Europeans or that we could never know their evolutionary relationships, for their bones told us only about their behavior. Today, thanks to fresh fossil inds and dates, to multiple, consistent analyses of the genes of living humans, and perhaps above all, to the spectacular recovery of DNA from Neanderthal bones, almost all specialists agree that the Neanderthals represent an evolutionary dead end. Many authorities who formerly placed the Neanderthals on the line to modern Europeans now argue only that they interbred with modern human invaders. However, genes increasingly deny even this fallback position, and the bottom line is that the Neanderthal controversy, which had raged for over a century, is now essentially concluded. his allows us to concentrate on other questions like the reason for their demise. Specialists disagree sharply on this issue, but as laid out in this book, genes again have the power to narrow the alternatives. Like the previous editions, this book difers from most commercial texts on human evolution in the extensive integration of the fossil and archaeological records and in the amount of supporting detail for both. he quantity of information in the previous editions makes me suspect that they were used more as references than as texts. In an attempt to enhance the value of this version as a readable text, I have removed much of the detailed information to tables, which I believe makes for smoother reading and easier access. I’ve also removed all bibliographic citations to separate paragraphs that immediately follow each topic within each chapter so that students can treat the citations as optional. An alternative scheme, which I used in the second edition, was to replace conventional in-text author-date citations with numbered references, but professionals disliked having to turn to the bibliography to see who was being cited. I hope both students and professionals will welcome the present system. he literature on human evolution is mushrooming, and this edition contains about 1,700 more references than its predecessor. he new references appeared mostly in the last ten years, and they represent only those whose titles or abstracts suggested that I needed to read them. hey comprise only a fraction of the articles and books I might have read, and I apologize to anyone who produced an important source that I inadvertently overlooked.

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Finally, I’m grateful to readers who pointed out errors or omissions in the previous editions, and I hope they ind I have made appropriate corrections. I owe a special debt to Kathryn Cruz-Uribe, who illustrated most of the fossils and artifacts in the earlier editions and who produced ity new high-quality illustrations for this one. I thank David DeGusta, Teresa Steele and Tim Weaver for thoughtful comments on parts of the present text.

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PrefaCe To The seCond ediTion

Paleoanthropology—the parent discipline for human paleontology and paleolithic archeology—has never been more vigorous or productive. In the ten years since the irst edition of this book was published, paleoanthropologists have discovered human fossils that antedate 4 million years ago and that show ever more clearly the ape origins of the human family; they have found new fossils and archeological sites that imply a complex history of branching events within the genus Homo; and they have uncovered a wealth of new fossil, archeological, and genetic evidence that the last shared ancestor of all living humans existed in Africa no more than 200,000 years ago. he purpose of this edition is to show how the new discoveries and analyses supplement previous ones to reveal the basic course of human evolution. Two conclusions are especially clear. he irst is that the australopithecines and other very early people who lived between 5 and 1.8 million years ago were intermediate in appearance and behavior between apes and unquestionable humans and that the irst “true” humans, who appeared 1.8 to 1.7 million years ago, were morphologically and behaviorally intermediate between the australopithecines and living people. An ironic corollary is that the human family now provides one of the most persuasive fossil cases for the occurrence of macroevolution. he second major conclusion is that fresh fossil, archeological, genetic and geochronological indings conirm earlier ones, suggesting that modern humans originated in Africa and later replaced other kinds of people in Eurasia. he irst edition espoused this “Out-of-Africa” scenario only cautiously, and it retained an earlier view that human evolution comprised a series of grades or stages, from the relatively apelike australopithecines through Homo habilis, Homo erectus, and “early” (or “archaic”) Homo sapiens to modern Homo sapiens. xxiii

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he richer fossil, archeological, and genetic records that bolster “Out of Africa” now indicate that the stage system must be discarded in favor of a branching scheme. hus the text argues that ater an initial human dispersal from Africa by 1 million years ago, at least three geographically distinct human lineages emerged. hese culminated in three separate species: Homo sapiens in Africa, Homo neanderthalensis in Europe, and Homo erectus in eastern Asia. Homo sapiens then spread from Africa beginning perhaps 50,000 years ago to extinguish or swamp its archaic Eurasian contemporaries. he spread was prompted by the development of the uniquely modern ability to innovate and to manipulate culture in adaptation. his ability may have followed on a neural transformation or on social and technological changes among Africans who already had modern brains. Whichever alternative is favored, the fossil, archeological, and genetic data now show that African H. sapiens largely or wholly replaced European H. neanderthalensis. he situation in eastern Asia is more obscure, because the fossil, archeological, and geochronological data are much sparser, but fresh data will probably show that H. sapiens replaced H. erectus in the same way. he change from a synthesis centered on stages to one based on branches required a new chapter titled “Evolution of the Genus Homo” in place of the irst edition’s separate chapters titled “Homo erectus” and “Early Homo sapiens.” he new chapter stresses the fossil and archeological evidence for multiple, contemporaneous species within fossil Homo, but it also acknowledges the continuing meagerness of the record, particularly before the advent of unequivocal Homo sapiens and Homo neanderthalensis by 130,000 years ago. To ensure that readers appreciate just what the evidence is (and what it is not), the new chapter illustrates many of the key fossils and artifacts. Most of the remaining chapters retain their original titles, but every one has been updated and expanded, and they all have many new illustrations. he total number of igures for the book has been increased from 151 to 223, and I could not have produced them without the willing, skilled assistance of my archeological colleague and friend Kathryn Cruz-Uribe. Even a cursory examination of the following pages will reveal her enviable ability to render informative, tasteful line drawings of complex fossils and artifacts. I used the computer programs Adobe Streamline and Macromedia Freehand to enhance her drawings with labels for key landmarks or features. Like the irst edition, this one difers from most commercial texts in emphasizing not only the broad pattern of human evolution but also the fossil and archeological data behind it. he book includes far more fossil and archeological detail than standard introductions to human evolution, and it is intended to be as much a sourcebook as a text. Nonetheless, the irst edition was oten used as a text, and with this in mind I

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have tried to make the second edition more accessible to undergraduate students. he most signiicant change is the addition of an introductory chapter that outlines how species evolve and how evolutionary biologists (including paleoanthropologists) classify and name them. My own students have oten complained about the standard academic author-date referencing system used in the irst edition, because it obstructs text low and intelligibility, especially when many citations are chained together. In this edition, I have therefore adopted a system in which numbers in parentheses in the text refer to numbered items in a summary reference list. A secondary beneit of this system is that it shortens the text by more than ive thousand words. he reference list is alphabetized, so works by a particular author are easy to locate. here is also a separate “reference index” that lists the text pages on which individual numbered works are cited. I used the Endnote computer program to ensure that the numbers cited in the text matched the numbered works in the reference list. he reference list includes more than 2,500 separate items, yet this is only a tiny fraction of the literature on human evolution. It is in fact only a small fraction of the items that have appeared since the irst edition was published, and it may neglect some signiicant discovery or idea in a reference I have missed. I hope that anyone who sees a glaring omission will notify me so I will not overlook it in the future. In preparing the second edition, I have beneited from the research and writing of innumerable colleagues, but I particularly want to thank James Bischof, Frank Brown, Kathryn Cruz-Uribe, Robert Franciscus, Clark Howell, and Henry McHenry for critical comments on portions of the text. I am also grateful to the many wonderful students at the University of Chicago and Stanford University who endured the lectures and text drats that underlie both editions.

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PrefaCe To The firsT ediTion

Supericially, the study of human evolution seems remarkably abstruse and impractical. Yet, each year the popular press covers major inds, large prime-time audiences watch televised documentaries, and thousands of students enroll in pertinent university courses. Australopithecus and Olduvai Gorge feature in nationally syndicated cartoons, and some discoverers of important fossils are better known than their counterparts in more practical ields such as physics and medicine. Clearly, in spite of its apparent irrelevance to everyday afairs, the origin of the human species is intensely interesting to most modern humans, who are fascinated by the growing number of fossils, artifacts, and related facts that scientists have amassed. hey want to know what these data tell us about the appearance and behavior of our remote ancestors. his book is a summary of what I think the data say. here are many perspectives on how the data should be interpreted, and this book, of necessity, relects just one. In writing it, I have tried to steer a middle course between what I see as two extreme approaches— one in which the data are simply a springboard for stimulating speculation about what might have happened in the past and another in which they are meaningless except to test and eliminate all but one competing explanation of what really happened. he diiculty with the irst perspective is that it emphasizes imagination over validity. he diiculty with the second, whose roots lie in a perception of how the physical sciences have advanced, is that it assumes an unrealistic degree of control over data quantity and quality. In fact, substantial control is unusual in human evolutionary studies, where carefully planned experiments are rare and most data are obtained through excavations and ield surveys whose success oten depends more on chance than on design. Under these circumstances, I think that the physical sciences provide a less suitable role model than the judicial system, in which oten limited evidence is

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weighed to determine which of two or more competing explanations or interpretations seems most reasonable. In most instances the possible alternatives are not pared unequivocally to one, but one is selected because it seems more justiiable, given the evidence on hand. Of course, in human evolutionary studies, as in the judicial system, both laypeople and specialists may disagree about what is reasonable or justiiable and also about the soundness of the supporting evidence. All too oten the evidence is incomplete, ambiguous, or even contradictory, and it cannot be used to bolster any particular theory or explanation very strongly. In this book I have tried hard to present the major competing opinions on prominent unresolved issues, and whenever possible I have explained why I think one view is more reasonable than others. More oten than I would like, I have had to say that a irmer choice will require more data. I know that not everyone will agree with the positions I have taken or even with my decision to abstain on some matters. However, I think that such diferences of opinion are unavoidable, given the imperfect nature of the evidence; and the point is that this book is inevitably just one of many possible summaries of what we know about human evolution. Its success will depend on the extent to which the readers, experts, and laypeople alike think that the presentation and argumentation are sensible. Philosophical approaches aside, there are several possible ways to organize the evidence for human evolution. he way I have chosen is perhaps the most conventional, focusing on a series of chronologically successive stages—beginning with the earliest Primates, dating from perhaps 80 million years ago, and ending with the emergence of anatomically modern people within the past 200,000 years. he presentation does depart from the norm, however, in that I have given roughly equal weight to the fossil record and to the accompanying archeological evidence over the 2.5 million year interval for which this is available. Most summaries focus largely on the fossils or on the archeology, thereby forgoing one of the major points that I have sought to make—namely, that the human form and human behavior have evolved together and that neither can be fully understood or appreciated without a full understanding of the other. At the same time, however, it remains true that the fossils are far easier to arrange into a set of chronologically successive, interrelatable units, and, since the fossil record is also far longer than the archeological one, I have relied on the fossils to deine the chronologically successive stages that structure the text. his is not to say that there are no problems in deining the fossil units, but these pale beside the diiculties in deining and interrelating corresponding archeological categories. he diference stems from our much weaker understanding of the mechanisms underlying artifactual (cultural) change and diferentiation.

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A second way in which this survey difers from many others is that it includes more information on speciic sites, fossils, artifacts, and so forth—in short, it is more detailed, with more concern for the factual evidence that underlies our understanding of human origins. It is a formal rendering of the lectures I give in an upper-level undergraduate course at the University of Chicago, and it has been written with upperlevel undergraduates, graduate students, and professionals in mind. In my experience, the audience for whom it is primarily intended will already have a basic understanding of how evolution occurs (through natural selection, mutation, gene low, and gene drit), and therefore this latter topic is not explicitly addressed. However, at least some members of the audience will lack essential background information on skeletal anatomy, zoological classiication and nomenclature, stone tool typology and technology, and especially the geologic time frame for human evolution, and so these subjects are covered. In general I think the book is too detailed to be a central text in lower-level courses, especially ones that also deal with modern human variation, genetics, and the like; but I hope it will ind use there as one of the sources the instructor consults or recommends to those students who are especially curious about the fossil and archeological records. In keeping with the comparatively technical orientation of the book, I have employed an in-text citation system that is common in professional scientiic publications. I rejected the usual textbook system of grouping sources at the end of each major section or chapter because I felt the target audience for the book would prefer to know precisely where to look for further information on a particular topic. I also wanted to give credit directly where it was due. I rejected a system of linking references to numbers because I thought the risk of serious error would be too great when so many references are involved. he large number of references was unavoidable, given the broad theme, but I have tried to keep the list manageable by stressing recent sources that can serve as guides to older ones, and I have also excluded many non-English primary sources in favor of secondary English ones with their own extensive bibliographies of important non-English publications. No synthesis of human evolution would be successful without good illustrations, but these can be very expensive and time consuming to produce. As a result, even many commercially produced texts are underillustrated. I have attempted to compensate for the limitations of time and expense that were important here by adapting many illustrations from published sources, which are gratefully acknowledged. hanks mainly to the eforts of Kathryn Cruz-Uribe, most of the illustrations have been substantially modiied to support pertinent points in the text and to provide stylistic consistency. In addition, whenever possible, I have labeled

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important features directly on fossils, artifacts, site plans, and so forth, and I have attempted to make the captions freestanding supplements to the text, to emulate the useful sidebars that are common in commercially produced texts. My goal was to make the illustrations especially helpful to those with little or no prior knowledge of skeletal anatomy, stone artifacts, stratigraphies, and such. It was not easy to choose a title for the book, because the most obvious ones, such as “Human Origins” and “Human Evolution,” have been used many times before. he inal choice—he Human Career—is the name of a graduate course I took at the University of Chicago in 1962, in which F. Clark Howell introduced me to the concept of human evolutionary studies as an amalgam of human paleontology and paleolithic archeology. Howell’s alternative name for the subject matter, “paleoanthropology,” would do equally well—though it too has been used before and has oten been applied to human paleontology alone rather than to the broader paleontological/archeological ield that Howell had in mind. In both the title and the text, I intend the vernacular term human (and its complement, people) to refer to all members of the zoological family Hominidae, as conventionally deined, and not simply to living humans. My own research on human evolution has focused mostly on behavioral (archeological) evidence from middle and late Quaternary sites in southern Africa and parts of Europe, and my acquaintance with the remainder of the record comes mainly from published sources. In synthesizing these, I have tried hard to make the text as accurate, comprehensive, and up-to-date as possible, and I have been greatly helped by comments and criticisms from Peter Andrews, Kathryn Cruz-Uribe, Janette Deacon, Leslie Freeman, Fred Grine, Clark Howell, Philip Rightmire, Chris Stringer, Russell Tuttle, and Tom Volman. I hope they ind that I have employed their suggestions productively and that I have not introduced any new errors in the process. Inevitably, however, some defects remain, and I would be grateful to hear from anyone who inds a speciic problem or who has suggestions on how the interpretations or overall organization can be improved.

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eVOLuTIOn, CLaSSIFICaTIOn, anD nOmenCLaTure

1

his chapter introduces basic evolutionary terms and concepts that are essential if we are to synthesize sites, fossils, artifacts, dates, and other facts into a coherent overview of human evolution. he discussion draws on muc h more extensive treatments of process and principle in works listed immediately below. GENERAL SOURCES: Cain 1960; Dawkins 1987; Dobzhansky 1962; Eldredge 1991, 1995; Gould 2002; Levinton 1988; Maynard Smith 1989; Mayr 1942, 1963, 2001; Ridley 1986, 1993; Simpson 1953, 1961; Stanley 1979, 1981

The Biological Species he species is the least arbitrary and most fundamental evolutionary unit, and it must be understood before any consideration of evolution, even one focused tightly on a single species like Homo sapiens. Evolutionary biologists deine a species as a group (or population) of organisms that look more or less alike and that interbreed to produce fertile ofspring. his deinition is oten called the biological species concept. In practice, individuals are usually assigned to a species based on their appearance, but it is their membership in the same procreative unit that validates (or invalidates) the assignment. hus, no matter how detailed the resemblances between two groups of organisms, if individuals cannot exchange genes between groups, the two populations must be assigned to diferent species. Species ordinarily comprise lesser units known as breeding populations (sometimes called demes or Mendelian populations) in which most individuals ind their mates. For example, the American black bear species (Ursus americanus) ranges across much of North America, but black bears in Alaska and central Mexico are unlikely to mate, and they are thus reasonably assigned to separate breeding populations. he maximum

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breeding population is always the species, since no members may ind mates outside it. Mountain chains, deserts, large water bodies, or other barriers between breeding populations may prevent them from interbreeding, but as long as interbreeding is possible and fertile ofspring would result, the breeding populations still belong to the same species. Breeding populations tend to luctuate in size, membership, and total number over relatively short periods, which means they are harder to delineate than species. However, as discussed below, they are central to evolution, since they are potential species in the making. he modern deinition of the species, founded in interbreeding capacity, is not the only conceivable one, and it is worthwhile to consider why it has replaced a deinition that was popular before evolution became widely accepted in the second half of the nineteenth century. his earlier concept deined each species according to the characters of a type specimen, usually the irst one to be found or described. Specimens that were not identical to the type specimen were allowed in the same species, but they were regarded as “sports” or deviants, and their anomalous features did not contribute to the species description. he typological deinition of the species was problematic even at the time it was most popular. From the beginning, its arbitrariness was apparent, and it loundered when additional specimens turned out to typify the species better than the original type. It was also diicult to decide just how diferent two specimens had to be before they could be assigned to diferent species. Typological practice tended to exaggerate small differences, and the result was an excessive number of species. Most important, however, the typological deinition literally ixed species in time. Species were immutable, for if they could change then there could be no type specimen, only an endless series of deviants. he typological deinition was therefore incompatible with organic evolution, and the infraspeciic (within-species) variation it explicitly suppressed was fundamental to Darwin’s novel idea of how species evolved, via natural selection of the ittest (discussed below). Only the modern biological species deinition is consistent with the idea of evolution, but ironically, only the typological deinition is directly applicable to the fossil record. he disjunct arises partly because interbreeding cannot be observed among fossils, and paleontologists must rely strictly on morphological features. In practice, with the biological species concept in mind, they commonly use skeletal variability within living species to evaluate whether diferences among fossils are likely to mean interbreeding could not occur. In this case the fossils are assigned to diferent species. However, the degree of skeletal variability varies from species to species, which means there can be no universal standard. Sometimes, as in the case of the African guenon monkeys (Cercopithecus), even members of diferent species are diicult or impossible

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to distinguish from skeletons alone. In other cases, for example, modern Homo sapiens, substantial skeletal variation occurs within a single unquestioned biological species. Paleontologists face the additional diiculty that their specimens are usually not complete skeletons. Isolated bones or fragments of bones are much more common, and this helps to explain why fossil species are oten so hotly debated. Finally, there is the problem that the identiication of fossil species inevitably requires dividing the evolutionary continuum into arbitrary units. If the record is complete enough, some dividing lines will fall between succeeding generations that could have interbred and that therefore should not be assigned to diferent species. his problem seems insuperable, except for the observation that whereas evolution is continuous, the fossil record oten is not. As discussed below, the fossil history of most species suggests long periods of morphological stability or stasis, followed either by extinction or by brief bursts of rapid change during which new species emerge. In this circumstance, the chances of inding fossils from populations in the process of speciation are slim, and the frequent absence of such fossils therefore poses no theoretical problem for the biological species concept. heir absence means that most fossil species, therefore, have discrete beginning and endpoints that probably correspond at least broadly to the reproductive boundaries of living species.

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Natural Selection Nonspecialists oten think that Charles Darwin (1809–1882) devised the concept of evolution in he Origin of Species (published in 1859). In fact, others had proposed the idea long before, and Darwin’s monumental contribution was the discovery of a natural mechanism to explain how evolution had occurred. Before Darwin, the notion that species had evolved was hardly less mystical than the more common traditional idea that they had been created supernaturally. Darwin’s great discovery was the principle of natural selection, which he synthesized from his own natural history observations and from the writings of other eminent nineteenth-century thinkers, perhaps above all homas Malthus (1766–1834) on population growth. As framed by Darwin and still employed by evolutionary biologists, natural selection is grounded in three observations: (1) that individuals within a breeding population vary in morphology and behavior, (2) that ofspring tend to inherit features of morphology and behavior from their parents, and (3) that not all individuals contribute equal numbers of ofspring to the next generation. If we assume that diferential capacity to survive and reproduce is linked to morphology and behavior, then any novel traits that enhance survival and reproduction will tend to increase in frequency from one generation to the next. Such traits are said to be adaptive or in

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the parlance of evolutionary biology, to make their bearers more it, and natural selection can be deined as the sum of environmental forces or pressures that determine itness. Fitness is deined totally, if somewhat tautologically, as an individual’s ability to produce ofspring who themselves survive and reproduce. As Darwin noted, natural selection is exempliied by the well-known process that people have used to produce breeds of domestic animals. In this case the selective force is human preference or desire, and an individual animal’s itness (ability to survive and reproduce) depends on the extent to which it exhibits desired characteristics. New breeds can be produced relatively quickly, even within historical memory, if selection is stringent enough. Darwin recognized that far more time would be required to transform one species into another or to produce two species from one, and he probably would not have proposed natural selection as the underlying mechanism if geologists had not simultaneously demonstrated the great antiquity of the earth. Darwin followed contemporary geological research closely, and he was especially inluenced by the work of Charles Lyell (1797–1875), oten known as the founder of modern geology and the formulator of the uniformitarian principle that guides it. In its simplest form, uniformitarianism is the notion that “the present is the key to the past.” Less cryptically, it is the idea that erosion, sedimentation, volcanism, crustal movements, and other geologic processes that can be observed today have operated throughout earth history and are suicient to explain the geologic record. From a uniformitarian perspective, for example, crustal uplit can explain the occurrence of shelly marine limestone far above modern sea level. To accept this, an observer must accept only that uplit and other geologic processes that produce only small-scale efects over the short term can produce much greater large-scale efects over eons. In essence, Darwin’s idea that natural selection explains the origin of species simply extended the uniformitarian principle to the biological realm.

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Genetics and Evolution Darwin believed that natural selection operated on individuals, and itness in the classical Darwinian sense applies exclusively to individuals. But modern evolutionary biologists oten think of selection as operating on particular traits. he diference is profound, because it relects an understanding of how traits are inherited—in short, of genetics—that was totally lacking in Darwin’s day. Strictly speaking, the fundamental nature of inheritance had been established, but the discoverer, the Austrian priest Gregor Mendel (1822–1884), published his indings in an obscure journal and his pioneering research was recognized only in 1900, long ater both he and Darwin had died.

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When Darwin wrote he Origin of Species, it was generally believed that ofspring blended the traits they inherited from their parents and they then passed on a blend of blends to their own ofspring. his view of inheritance presented an immediate problem for the idea of natural selection, for it followed that a selectively advantageous novelty (or mutation) could not become ixed in a population. Instead, it would become diluted or lost through progressive blending with the much larger number of less advantageous alternatives. In essence, Mendel showed that inheritance was not a blending process but, instead, involved discrete units or particles, inherited half from one parent and half from the other. he particles, which we now call genes, retain their integrity from generation to generation, even though their expression may vary depending on the presence of other genes or on particular environmental conditions. hus an adaptive or advantageous novelty will not necessarily be diluted out of existence but could spread if it confers greater itness. Today a particular gene may be deined by its position on a chromosome (its locus) or by its singular chemical structure. Evolutionary divergence may be investigated directly at the gene level by comparing genetic material, deoxyribonucleic acid (DNA), or its proximate products (proteins) among related organisms. For those interested in the origin of species, however, it is useful to deine a gene simply as a Mendelian particle or a discrete unit of inheritance. he complete set of genes an individual possesses is known as his or her genotype. he genotype interacts with the environment to produce the phenotype, which is the sum of observable morphological and behavioral traits. he total complement of genes that characterizes a species or one of its subunits (a breeding population) is known as its gene pool, and this can be described by the frequencies of diferent genes (or gene variants) it contains. Evolution can then be deined as change in gene frequencies through time, which is completely consistent with Darwin’s deinition of evolution as “descent with modiication.” Four forces are commonly said to drive changes in gene frequencies and thus to drive evolution. he most important is natural selection, formulated essentially in Darwin’s terms. he others are mutation, gene low, and random gene drit. Mutation refers to a spontaneous, chance change in the chemical structure of a gene. Mutation is thought to be unusual (generally afecting fewer than one in 100,000 genes), and in complex organisms like people, any mutations that do occur are much more likely to be harmful than helpful. his means that relative to natural selection, mutation has much less potential to alter gene frequencies. Mutation is important primarily as the ultimate source of all genetic variability, the raw material on which natural selection acts. However, if mutation were somehow to cease, natural selection could continue to operate indeinitely on the genetic variability that already exists. his is particularly true since the

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variability in most species includes genes whose efects are largely suppressed or masked by those of others. In changed circumstances, natural selection could reduce the frequency of the masking genes, resulting in signiicant, even profound morphological change totally in the absence of mutation. Gene low is simply the exchange of genes between populations. By deinition, the populations must belong to the same species, else gene low would be impossible. In a population that is newly formed from two others, the new gene frequencies will be intermediate, and in the absence of natural selection, their precise values will depend on the relative sizes of the two original populations. Whatever the exact outcome, however, in a strictly technical sense evolution has occurred. Random gene drit is change in gene frequencies due strictly to chance. Intuition alone predicts that random drit is most likely to occur in small populations and to afect genes that are infrequent to begin with and this has been borne out empirically. In a special instance called the founder principle or efect, chance alone is likely to produce frequency diferences between a large parent population and a small emigrant population that splintered from it. he founder principle is oten used to explain some of the genetic diferences between Native Americans and the east Asian ancestral population from which they derive. Genetics has illuminated the evolutionary process in ways that Darwin could never imagine, but its application to the fossil record is not straightforward. his is because fossils contain no genetic information, with the exception of some relatively recent fossils like those of the Neanderthals discussed in chapters 6 and 7. Fossils reveal only the phenotype, which may change independently of the genotype and vice versa. his is illustrated most clearly by the occurrence of genes that that have no obvious function and that are therefore presumed to be immune to natural selection. Such genes are said to be “neutral,” and unlike genes whose form is maintained by selection, they oten accumulate changes (mutations) at a relatively rapid and more or less constant rate. he degree of similarity in neutral genes among breeding populations or species may reveal their evolutionary relationships independently of the fossil record. However, the relatively constant rate of change that neutral genes exhibit contrasts sharply with the irregular rate of morphological change that appears to characterize fossil species in deep time. As discussed below, evolution in the fossil record oten appears to have been episodic, meaning it was marked by short, rapid bursts of phenotypic change that punctuated long intervals of little or no change. he genetic basis for this remains unclear. Conceivably, the rapid bursts oten relect the wholesale reformulation of genotypes under natural selection, but it is at least equally likely that the morphological shits stem from changes in only one or a few genes that profoundly altered the mode and tempo

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of individual development. For the moment, the bottom line is that the relation between genetic change and the origin of species in deep time is poorly understood, and it’s become an active area of research.

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Speciation Many evolutionary biologists distinguish between microevolution, in which genetic change is relatively minor and occurs within separate breeding populations that continue to comprise a single species, and macroevolution, in which genetic change is far more dramatic and ultimately produces separate species. In the case of macroevolution, the separate species could be an ancestor and a descendant many generations apart or they could be contemporaneous, divergent descendants of what was once a single species. Faced with a relatively sparse database and long timespans, paleontologists generally focus on macroevolution, and they must therefore be especially concerned with speciation, the process by which species form. Darwin believed that natural selection caused species to change more or less continuously and that the slow accumulation of gradual change eventually produced new species. He was not particularly concerned with how one species could evolve into two or more, but, Darwin’s preeminent twentieth-century successor, Ernst Mayr (1904–2005), advanced the concept of geographic speciation, or adaptive radiation, to address this. According to this model, if a species expands into diverse environments, as many oten have, or if environmental change physically separates populations of a single species, adaptation to local circumstances (natural selection) will cause the populations to diverge genetically. he divergence will tend to increase through time, particularly if isolation by distance or natural barriers impedes gene low. Later, some previously isolated populations may come back into contact where there are niches (ecological opportunities) for each, but any hybrids they produce will be less well-adapted to either niche than nonhybrids. Selection will thus tend to prune away the hybrids, and the trend to genetic divergence will continue. Given time, the degree of divergence can preclude interbreeding, and the separate populations would then be species in the full biological sense. For present purposes, the key point is that following on Darwin, the adaptive radiation model assumes anagenesis, or gradual change along separate branches of the evolutionary tree. he fundamental concept involved is now oten called phyletic gradualism. Phyletic gradualism is plausible in perhaps every respect but one—it fails to explain the common mode of evolutionary change in the fossil record. he typical pattern there is for a species to appear relatively abruptly, to change very little during the course of its existence. And then to disappear relatively quickly. In Darwin’s time, one could argue,

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as Darwin did, that the pattern relects large gaps in the record, but the record is now far more complete and the pattern persists. Many paleontologists have therefore abandoned gradualism in favor of what Eldredge and Gould (1972) have termed punctuated equilibrium. he key idea behind punctuated equilibrium is that true evolutionary innovations appear and spread suddenly and infrequently. It is at these points of abrupt change, oten sparked by major climatic or environmental shits, that new species tend to arise. Major climatic shits not only open up fresh ecological opportunities, they also extinguish existing species, clearing the ecological playing ield for new ones. Viewed from the present, the fossil record appears to show a sudden alteration ater a period of constancy, a species-spawning event captured in a lash of geologic time, which punctuates an otherwise prolonged period of evolutionary stasis. In other words, stability is the norm, while speciation is the rarer but essential exception. he emphasis in punctuated equilibrium is strongly on branching or cladogenesis (the formation of branches or clades) as an event, and anagenesis is largely denied (or replaced by stasis). Figure 1.1 shows how the punctuationalist model, unlike gradualism, postulates signiicant evolutionary change only at branching time and not along branches. In the punctuated equilibrium model, new species are thought to arise mostly in small isolated populations where genetic changes (mutations) are especially likely to take hold and become dominant. In large populations or in small populations that are in regular contact with others, genetic changes, even advantageous ones, are more likely to be swamped and to disappear strictly by chance. Since the source population for a new species is small and geographically peripheral, paleontologists will rarely observe it, and it is only ater speciation, when the species expands to overlap and sometimes replace its parent, that it crosses the threshold of paleontological visibility. his would explain why the fossil record rarely reveals a series of graduated transitional forms or “missing links” between parent species and their descendants. In the punctuationalist model, the driving force behind speciation remains natural selection, but its intensity varies over time, and it is episodically especially intense. Episodic climatic change is probably the most fundamental selective force, and in combination with mountain building, sea level changes, and other major geologic events, it could fragment a widespread species into isolated populations and thus hasten their tendency to diverge genetically. Pulsed climatic change is well-documented in the continuous sedimentary record of the deep-sea loor, as outlined in the next chapter, but the relation between pulsed change and species turnover in mammals is diicult to investigate. his is because there are few regions where large fossil samples are known to succeed each other closely over a prolonged interval. Successive rodent faunas dated between

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24 and 2.5 million years ago (Ma) in central Spain provide a notable exception, and mathematical analysis has shown that episodes of rodent species appearance and disappearance correspond closely to pulsed climate changes documented in the deep-sea record. Closer to the subject matter of this book, bursts of speciation in African antelopes and primates (including humans) may have occurred in response to episodes of profound climatic and environmental change about 5 Ma and again about 2.5 Ma. Fossil species are easier to recognize if cladogenesis is the rule and anagenesis the exception, but it is important to emphasize that it is the nature of the record rather than logic or experiment that most strongly supports punctuated equilibrium. In efect, punctuated equilibrium is a thoughtful generalization from data more than a theory, and there is no claim, even by strong advocates, that punctuated equilibrium should be assumed to characterize the evolution of people or any other particular species in the absence of empirical (fossil) support. SOURCES: punctuated equilibrium described (Tattersal 2002); forces that fragment species into isolated populations and hasten their genetic divergence (Vrba1995b); correspondence between rodent species turnover and pulsed climatic change in central Spain (van Dam et al. 2006); bursts of speciation in response to environmental change in Africa about 5 Ma and 2.5 Ma (Vrba 1993)

Phylogeny and Classification he noun phylogeny and its adjective phylogenetic are oten used as synonyms for evolution and evolutionary. However, strictly speaking a phylogeny is a tree diagram illustrating the evolutionary history and

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FIGURE 1.1. The process of speciation as visualized by gradualists (left) and by advocates of punctuated equilibrium (right) (redrawn after vrba [1980a], 62). The bell-shaped curves mirror the distribution of one or more morphological traits at different times. distinct species are indicated by the blank and shaded paths on which the curves rest. The gradualist model postulates a steady, cumulative shift in modal morphology along branches. When the shift is extensive enough, a new species is born. The punctuationalist model postulates essentially random noncumulative morphological change through time. a new species emerges only when innovations become ixed in a small peripheral population. The population then diverges rapidly before settling into a conservative mode where change once again tends to be random and noncumulative.

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FIGURE 1.2. a currently popular phylogeny of the hominins, illustrating proposed ancestor-descendant relationships between known fossil and living species (redrawn after Grine [1993a], ig. 13). The question mark in the phylogeny relects continuing debate about the origins of the Homo lineage. as discussed in chapters 3–6, some authorities believe the Homo lineage also comprises multiple branches.

relationships of a species or species group. It is thus to a species what a genealogy or family tree is to an individual. Figure 1.2 illustrates a currently popular phylogeny of the Hominini, the zoological tribe that includes all people, living and extinct. Like many phylogenies, it includes branches that do not extend to the present because species oten become extinct without issue. In fact, inal extinction (or termination) eventually afects all species and it should be conceptually separated from the situation, also relected in igure 1.2, where “extinction” results when one species evolves into another. Phylogeny is intimately tied to classiication, the assignment of species to higher categories based on their presumed evolutionary relationships. A phylogeny is essentially a diagrammatic representation of a classiication scheme, and insofar as phylogenetic reconstruction cannot occur without classiication, classiication is more fundamental. he categories used in biological classiication were devised in the seventeenth and eighteenth centuries and achieved essentially their present form in the tenth edition of Carolus Linnaeus’s Systema Naturae, published in 1758. he most basic unit of classiication is the species, which, as already discussed, is deined today as a population of organisms that look more or less alike and that can interbreed to produce fertile ofspring. Based on how recently species are thought to share a common ancestor (i.e.,

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their degree of evolutionary relationship), they are then classiied into progressively higher (more inclusive) categories, minimally including (from bottom to top) genus, family, order, class, phylum, and kingdom. hus, a genus comprises species that share a more recent common ancestor with each other than they do with species in other genera (the plural of genus); a family comprises genera that share a more recent common ancestor with each other than they do with genera in other families; and so forth, up to the level of the kingdom in which all the included species share a very distant common ancestor. Since Linnaeus’s time, many new levels have been inserted among the seven principal ones, mainly in an efort to better organize the great proliferation of known species. Depending on the species being classiied, twenty or more levels may be recognized today. Table 1.1 illustrates twenty-two that can be used to classify Homo sapiens according to the Linnaean system. Whatever the number of levels, the principle remains the same. A category contains a group of species whose overall degree of relationship is indicated by where the category occurs in the hierarchy. hus, although H. sapiens is the only surviving human species, chapters 4 and 5 note that it is commonly said to have at least two extremely close extinct relatives (H. habilis and H. erectus), which are therefore included in the same genus. Other extinct species that are more distantly related and that are human only in a broader sense are usually placed in the separate genus Australopithecus, and they are united with species of Homo only at the tribal level, in the Hominini. he Hominini in turn can be united with the chimpanzees (two species) and the gorilla in a common family, the Hominidae; the Hominidae with the other apes (several families living and extinct) in a common superfamily, the Hominoidea; the Hominoidea with the Old World monkeys (constituting a single superfamily, the Cercopithecoidea) in the infraorder Catarrhini; the Catarrhini with the New World monkeys (Platyrrhini) in a common suborder, the Anthropoidea (higher primates); the Anthropoidea with the tarsiers (suborder Tarsiiformes) in the semiorder Haplorhini; and inally the Haplorhini with the Strepsirhini (comprising the suborders Lemuriformes and Lorisiformes) in the order Primates. Figure 1.3 illustrates this hierarchy graphically, beginning with the order on the let and ending with the families on the right. Chapter 3 points out that details of this particular hierarchy are subject to debate and revision, but the broad outline is irm. From what has been said, it follows that each category above the species comprises a group of related species (except in those rare instances where a species has no close relatives, living or extinct, in which case a higher category may include only one species). For convenience, each species in the Linnaean hierarchy and each group of related species at any level is called a taxon (plural taxa). hus the species Homo sapiens is

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A classiication of living people involving twenty-two potential levels in the Linnaean hierarchy. A dash follows a level for which no taxon is in common use. Asterisks designate the seven obligatory and most basic levels in the Linnaean system. Technically, the species for living people is Homo sapiens, but sapiens is listed alone for simplicity’s stake.

TABLE 1.1.

*KINGDOM: Animalia *PHYLUM: Chordata SUBPHYLUM: Vertebrata SUPERCLASS: Tetrapoda *CLASS: Mammalia SUBCLASS: heria INFRACLASS: Eutheria COHORT: Unguiculata SUPERORDER: Euarchonta *ORDER: Primates SEMIORDER: Haplorhini SUBORDER: Anthropoidea INFRAORDER: Catarrhini SUPERFAMILY: Hominoidea *FAMILY: Hominidae SUBFAMILY: Homininae TRIBE: Hominini SUBTRIBE:———*GENUS: Homo SUBGENUS: (Homo) *SPECIES: sapiens SUBSPECIES: sapiens

a taxon, as are the genus Homo, the tribe Hominini, the family Hominidae, the superfamily Hominoidea, the infraorder Catarrhini, the suborder Anthropoidea, the semiorder Haplorhini, the order Primates, and even the kingdom Animalia. Species and genera are commonly referred to as lower taxa, while categories above the genus are known as higher taxa. Since it is taxa that are classiied, the term taxonomy is oten used as a synonym for classiication, though technically, it refers to the system of rules for constructing a classiication. Classiication can be problematic because the evolutionary relationships on which it depends are not directly observable but must be inferred from the degree of similarity among taxa. In general, the greater the similarity, the closer the assumed relationship. his explains how Linnaeus could devise the modern classiication system a century before Darwin popularized evolution and also how nonevolutionists or even antievolutionists can employ the system today. More important, the unavoidable reliance on degree of resemblance can be misleading, since factors other than common descent can produce similarities among taxa. Among other inluences, the most important is undoubtedly similar adaptation to shared environmental conditions. Resemblances due to similar adaptations are called analogies (or convergences) in distinction from homologies or similarities due to common descent. A highly

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ORDER

SEMIORDER

SUBORDER

INFRAORDER

SUPERFAMILY

FAMILY

Daubentonioidea Daubentoniidae Lemuriformes

(Lemuriformes)

Cheirogaleidae Lemuroidea

Strepsirhini

Indriidae Lemuridae

Lorisiformes

(Lorisiformes)

Lorisoidea

Lorisidae

Tarsiiformes

(Tarsiiformes)

Tarsioidea

Tarsiidae

Primates

Cebidae

Haplorhini Platyrrhini

Ceboidea

Aotidae Pitheciidae

Anthropoidea

Cercopithecoidea Cercopithecidae Catarrhini

Hylobatidae Hominoidea

conspicuous and ot-cited analogy is the streamlined, inned body form shared by sharks and dolphins, which, despite this morphological similarity, are not closely related. A less famous but equally clear analogy is the independent development in apes and some New World monkeys of an upper limb (arm) structure that facilitates the ability to hang below tree branches. In contrast, the numerous detailed similarities in head and body form between people and chimpanzees exemplify homologies inherited from a relatively recent common ancestor. In many instances (with sharks and dolphins, for example), careful scrutiny of multiple traits allows homologies to be separated from analogies. In other cases, the distinction can be diicult, particularly in fossil taxa, where the number of appraisable characters is limited. In addition, even when homologies have been identiied, they are not in themselves suicient to establish the degree of evolutionary relationship among taxa. A further distinction must be made between homologies that are widely shared and ones that have a narrower distribution within a taxonomic group. In general, widely shared characters have only limited taxonomic value, even if they are prominent and numerous. Characters with a limited distribution are more likely to reveal the basic evolutionary links among taxa in the group. he realization that diferent characters must be assigned diferent weights in evolutionary studies is central to the

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Hominidae

FIGURE 1.3. a hierarchical classiication of the living Primates down to the family level. Chapter 3 provides supporting detail and discusses alternatives.

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increasingly popular perspective on classiication known as cladistics. Following Hennig (1966), cladists usually distinguish between two fundamental kinds of homologous characters:

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1.

2.

Primitive (or generalized) characters (plesiomorphies or symplesiomorphies, in cladistic terminology), which arose early in the evolutionary history of a taxonomic group. hese will be extremely widespread and will therefore not help in dividing the group into lower-level taxa, that is, in determining their genealogical relationships. A well-known primitive feature in mammals, for example, is the occurrence of ive digits on the end of each limb. his feature has been retained in Primates, but its ubiquity renders it useless for subdividing the order. It cannot even be used to distinguish Primates from non-Primates, many of whom also retain ive digits as a result of distant, shared descent. Derived (or advanced) characters (apomorphies, to cladists) that arose relatively late in members of a group and will difer among them. In contrast to primitive characters, derived characters are useful for assessing genealogical links among taxa. For example, structural modiication of the rear limb to permit habitual bipedal (two-legged) locomotion is a key derived feature of the hominins (people, broadly understood) that ultimately underlies their separation from other closely related taxa within the larger hominoid group (apes and people). Derived features can be further subdivided into two basic types: shared derived characters (synapomorphies), which demonstrate a special evolutionary tie among taxa that have them, and unique derived characters (autapomorphies) or novelties, which distinguish a taxon from all others. Unique characters are not useful for inferring evolutionary connections, though they may eliminate one taxon from the ancestry of another that lacks them.

Applying cladistics to taxa produces a cladogram or tree diagram, which organizes the taxa according to the number of derived features they share. he larger the number of shared derived features, the more likely it is that the taxa will reside on branches that connect to each other before connecting to other branches farther down the tree. Taxa that reside on neighboring branches are commonly known as “sister taxa” to imply that they are each other’s closest evolutionary relative. In form, cladograms resemble traditional phylogenies; but unlike phylogenies, which place taxa at diferent points or branching nodes within a tree, cladograms arrange them at the ends of terminal branches. Also unlike phylogenies, cladograms do not consider time relationships, and they generally do not include a timescale. he essential diference is that cladograms illustrate perceived degrees of derived similarity among taxa, not ancestordescendant relationships. Figure 1.4 illustrates a cladogram for some living

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an m hu

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ril l

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tan gu or an

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m su s rh e

sp i

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on

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evoluTion, Clas s if iCaTion, and n om enClaTur e

higher primates based on the presumed degree of derived similarity in the structure of the beta-type globin gene. Chimpanzees and people in this cladogram are an example of sister taxa. Cladograms can portray the relationships among higher-level taxa, as well as among individual species, or among higher-level and lower-level taxa at the same time. Cladograms that portray multiple-level relationships simultaneously have branches that support additional branches (or smaller scale cladograms). Technically, all the branches in a cladogram are known as clades, and in a complex multilevel cladogram, lower-level clades are said to nest within higher-level ones. Cladistics requires that each clade be monophyletic, meaning that its members all share a common ancestor and that the clade includes all the descendants of that ancestor. If some descendants of the common ancestor actually reside in other clades, the clade is said to be paraphyletic, and if the clade includes members of diverse ancestry that should reside in diferent clades, it is called polyphyletic. Cladistic concepts can be extended to cultural or archaeological units, including, for example, the Aurignacian Culture, discussed in chapters 6 and 7. he Aurignacian is usually regarded as the earliest cultural manifestation of fully modern humans in central and western Europe, and most authorities believe it sprang from a single source, perhaps in southeastern Europe. However, some believe that it arose independently (convergently) in diferent places. In the irst instance, it would be monophyletic, in the second polyphyletic. Cladists usually regard all derived features as equally important, and this can lead to radical revisions in phylogenies and thus in classiication. For example, the chimpanzees and the gorilla have traditionally been joined with the orangutan in the family Pongidae and people have been assigned to their own family, the Hominidae. However, as discussed in chapter 3, in derived biomolecular characters, the chimpanzees and

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15 FIGURE 1.4. a cladogram illustrating the evolutionary relationships among humans (Homo), chimpanzees (Pan), gorillas (Gorilla), orangutans (Pongo), gibbons (Hylobates), rhesus monkeys (Macaca), and spider monkeys (Ateles), based on inferred derived similarities in the structure of the beta-globin gene (data in Goodman et al. [1990]). other biomolecular and morphological analyses usually produce similar cladograms, but some, particularly morphological studies, suggest that the chimpanzee and gorilla are more closely related to each other than either is to people. a cladogram that accepted this would place the gorilla and chimpanzee on branches that connected to each other before they connected to the human branch. With respect to the cladogram above, this could be effected by eliminating the solid line leading to the chimpanzee and replacing the dotted line between the gorilla and chimpanzee with a solid line.

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gorilla resemble people far more closely than they do the orangutan, and the implication is that the chimpanzees, the gorilla, and people share a much more recent ancestor with each other. Strictly speaking, this means that the chimpanzees and gorilla should be removed from the Pongidae and placed with people in the Hominidae. People could then be grouped in a tribe, the Hominini, within the Hominidae. However, it can be argued that people should remain alone in the Hominidae because they have departed farther than the chimpanzees and gorilla from the last shared ancestor. In essence, this is to say that the unique derived features that deine people—habitual bipedalism and especially the large and complex human brain—have produced a novel adaptation or grade of life. If this argument is accepted, it provides a plausible rationale for maintaining the status quo and for avoiding the confusion that inevitably follows terminological change. However, from a strictly logical standpoint the cladistic reclassiication is less arbitrary and easier to defend, and this book accepts the revision that separates people only at the tribal level. Cladistic procedure does not preclude the identiication of grades (similar levels of structural organization or of adaptation); it only excludes them from consideration in determining the level at which related species are classiied together. his is because grades need not bear on the degree of evolutionary relationship that classiication is supposed to relect. Moreover, in focusing exclusively on degree of similarity or difference, cladograms are less abstract than phylogenies, which must be more conjectural or hypothetical. If the principles behind cladistics are straightforward, however, the practice is oten problematic. Cladistic analysis may stumble if shared derived features that are actually analogies (or parallelisms) are mistaken for homologies. In addition, it is not always easy to determine whether a character is primitive or derived within a group. In fact it can be both, since character reversals are an occasional feature of the evolutionary process. Skull bone thickness, for example, changed from relatively thin in the earliest hominins (the australopiths and Homo habilis) to relatively thick later on (in H. erectus and early H. sapiens) back to thin (in later H. sapiens, including living people). In a situation like this, functional understanding of a character or character state would obviously be useful for determining its cladistic utility. A functional understanding is also crucial for determining whether two (or more) shared derived characters are truly independent. If not, they constitute only one character, and their implications for an especially close taxonomic relationship are diminished. Finally, for optimal results, cladistic methodology is best applied to discrete characters, that is, characters that are either present or absent. It produces more equivocal results when the characters difer among taxa mainly in their degree of expression or their frequency; yet such char-

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acters are the rule when the taxa are very closely related, as they are, for example, within the genus Homo. Some of the problems, together with the substantial fruits of cladistic analysis, are illustrated in subsequent chapters, though the emphasis there is on broad evolutionary trends, independent of classiication or phylogeny.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: cladistic fundamentals (Eldredge and Cracrat 1980; Skelton et al. 2002; Wiley 1981)

Nomenclature In theory, the naming of zoological taxa could be arbitrary, but long experience has shown that a rule-based system reduces potential ambiguity and improves communication. hus, all biologists adhere to a single naming scheme that essentially follows the practice of Linnaeus. In this scheme, taxa are always given Latin names or names whose form has been latinized. he Latin name of a taxon may be intended as a substantive characterization (Homo sapiens, for example, means “wise man”), but, in general, a name should be regarded simply as a label, not as a deinition. he name of a species always consists of two words—the genus (generic) name followed by the species (speciic) designation in the narrow sense. Grammatically, the genus name is a Latin noun and the species designation is a Latin modiier (either an adjective or a noun in the genitive singular) or another Latin noun in apposition. he name of a genus is always capitalized and italicized (or underlined in typescript), whether it stands alone or has a species modiier. he second part of the species name (the species designation narrowly understood) is italicized but not capitalized. If a genus has already been cited, a closely following citation of a species in the same genus may abbreviate the genus name, usually to its irst letter and a period: for example, H. erectus or H. sapiens closely following the mention of Homo habilis. In those relatively rare instances where a subgenus name is used, it is capitalized, italicized, and placed in parentheses between the genus and species terms, for example, Australopithecus (Paranthropus) robustus. If a subspecies name is used, it is italicized but not capitalized and appended to the full species name (e.g., Homo sapiens sapiens). Subspecies are breeding populations within a species that are morphologically and geographically distinct, but strictly speaking, they have no place in the Linnaean hierarchy except where there is reason to believe that they are actually species in the making (that is, that they are diverging progressively under the inluence of divergent natural selective pressures). No populations of living H. sapiens constitute such incipient species, but a case can be made for some extinct populations, such as the Neanderthals, which some specialists thus place in the subspecies H. sapiens neanderthalensis.

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Categories above the genus level are capitalized but not italicized. Higher taxa, up to the level of the superfamily, always derive their names from one of the genera they include. By convention, they also have endings that designate their level: (in vertebrates) -ini for tribe, -inae for subfamily, -idae for family, and -oidea for superfamily. Above the superfamily level, the only rule is that the name must be a Latin (or latinized) noun. he Latin names are usually anglicized by dropping the Latin ending (for example, hominins in place of Hominini). In their anglicized form, they are generally not capitalized and they can be used as either adjectives or nouns. Technically, the full name of a taxon at any level includes the name of its inventor(s) and the date when their invention was published. Species examples that are important in this book include Australopithecus africanus Dart, 1925, Paranthropus robustus Broom, 1938, Homo ergaster Groves and Mazak, 1975, and Homo sapiens Linnaeus, 1758. When species are transferred from one genus to another, the author and publication date of the original designation are enclosed in parentheses, followed by the author and publication date of the suggested transfer. Examples cited later in this book include Ardipithecus ramidus (White et al. 1994) White et al., 1995, for a species that was originally designated Australopithecus ramidus White et al., 1994, and Homo erectus (Dubois 1892) Mayr, 1944, for a species that was originally called Pithecanthropus erectus Dubois, 1892. In practice, specialists usually ignore the requirement to append author name(s) and date(s), particularly when, as in this book, taxonomy itself is not a primary focus. Ideally, all specialists would refer to the same taxon by the same name, but problems arise when one taxon has been given two or more names and there is disagreement about which has priority (that is, which was suggested or legitimized irst). Even if one name clearly has priority, many specialists may retain an alternative that has a long history of use. However, the most important nomenclatural disagreements do not low from names alone but from basic diferences of opinion about the evolutionary relationships among taxa. hus, as discussed in chapter 6, many specialists believe that the Neanderthals and modern humans should be assigned to the same species, Homo sapiens, while others, including the author of this book, place the Neanderthals in a separate species, Homo neanderthalensis. he diference matters, since the single-species alternative implies that the Neanderthals could have participated in modern human origins, whereas the separate-species alternative rules this out. As this book illustrates, paleontologists are particularly likely to engage in nomenclatural disputes that low from taxonomic (classiicatory) disagreements.

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The GeOLOGIC TIme Frame

2

Among all the variables behind human evolution, none is more crucial than time, and a discussion of the geologic time frame must precede any consideration of fossils and artifacts. In the best of all possible worlds, the ages of all ancient sites and fossils would be known in calendar years. In reality, the ages of many sites and fossils can be estimated only broadly, and much on-going work in paleoanthropology aims to provide more precise dates. he most important and widely useful techniques—numerical (or “absolute”) methods that can provide dates in years—were developed only in the late 1940s and 1950s, nearly a century ater the irst scientiic inquiry into human origins. Before numerical methods appeared, dating was limited to establishing the relative order in which sites formed, based mainly on the visible or inferred stratigraphic position of one site with respect to others. Even today many sites can be dated only in relative terms (rather than numerically in years), either by their stratigraphic location in a sedimentary sequence or, more oten, by the stratigraphic implications of their fossils or artifacts. Additionally, even at those sites where numerical dating is possible, stratigraphic context furnishes a vital cross-check on the results. Finally, there is a sense in which numerical dates are only unusually precise indicators of stratigraphic order (relative age). In sum, the concept of stratigraphy remains fundamental to research on human evolution.

Stratigraphic Units and the Geologic Timescale For dating purposes, any geological objects that can be arranged in a sequence from younger to older may constitute a stratigraphy. he most common objects are successive layers of rock whose distinctive qualities can be used to deine rock-stratigraphic (lithostratigraphic) units. Fossils extracted from rock layers provide the basis for biostratigraphic units,

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which subdivide between faunal and loral stratigraphic units or among particular kinds of faunal or loral units. hus, as illustrated below, biostratigraphies can be grounded in fossils of rodents, pigs, horses, or elephants or on some combination of the fossils of these taxa and others. Even human (or prehuman) fossils and artifacts can serve, though entities based on artifacts or other behavioral debris are probably best placed in a separate class of culture-stratigraphic units. Stratigraphic units are always based on materials found in rock layers, but diferent kinds of units need not correspond one-to-one. For example, a single biostratigraphic unit, perhaps deined by the presence of particular species of pig, may span several rock units. Diferent units that are assumed to have formed or existed at the same time are said to be correlated. he ultimate goal is to correlate particular rock units, biostratigraphic units, and so forth with time-stratigraphic or chronostratigraphic units deined purely in terms of time rather than the prsence of particular materials. Whereas units deined by rock type (lithology), fauna, lora, or other properties tend to be geographically localized, chronostratigraphic units have no spatial bounds, and diferent rock or biostratigraphic units from many diferent areas can correspond to a single chronostratigraphic unit. Chronostratigraphic units in turn correspond to periods of real geologic time, which are at once the most abstract and the most continuous components of the system. hey are not themselves stratigraphic units but are simply named timespans to which geologists ascribe particular chronostratigraphic units. In both theory and practice, the various kinds of stratigraphic units can usually be recognized at diferent scales, for which sets of hierarchical terms are available. hus, with respect to rock-stratigraphic units, the smallest deinable (“mappable”) unit is generally a bed, spatially and lithologically related beds are grouped into members, and related members are grouped into formations. he australopith (australopithecine) site descriptions in chapter 4 illustrate this principle. he hierarchy of words for biostratigraphic units is less formalized, but many specialists use terms such as faunal complex, stage, zone or biozone, sometimes preixed to indicate hierarchical rank or status. he smallest chronostratigraphic unit in common usage is the stage. A related group of successive stages constitute a series, related series constitute a system, and related systems constitute an erathem. he corresponding time terms (from least to most inclusive) are age, epoch, period, and era. Figure 2.1 presents a list of named eras and a partial list of included periods and epochs, spanning the entire history of the earth, from roughly 4.6 billion years ago to the present. he scheme is rooted mainly in biostratigraphy, while the boundary dates have been estimated mainly by the numeric dating methods discussed below. he table is less detailed for eras before the Cenozoic because this book addresses the evolution of

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Era

Period

Epoch Holocene

CENOZOIC

Quaternary

Pleistocene Pliocene Miocene

0

0.006 0.011 0.01 0.012 0.05 0.20 1.6–1.4 1.6 2.6 4.1 5.2 7-5 20–17 23.3

First cities First farmers First people in the Americas First fully modern humans Oldest firm evidence for fire First hominins in Eurasia Oldest stone artifacts & Homo Oldest known Australopithecus First hominins (humans sensu lato) Oldest known monkeys & apes

34

35–30

Oldest known higher primates

50

Oldest known primates of modern aspect

Oligocene

Tertiary

Eocene

MESOZOIC

Paleocene Cretaceous Jurassic Triassic Permian

PALEOZOIC

Ma Some Proposed Firsts

Carboniferous

56.5

65.5 65.5 120 145.5 160 199.5 220 251

First primates First placental mammals

299

300

First reptiles

370

First amphibians

475

First vertebrates (fish)

550

First chordates

359.2

Devonian

First birds First mammals & dinosaurs

416

Silurian 443.7

Ordovician 488.3

Cambrian 542

PRECAMBRIAN (= PROTEROZOIC and before)

800 1400 3500 4000 4600 4600 15,000

First multicellular life (sponges, algae) First nucleated cells (eukaryotes) First unicellular life (prokaryotes) First complex organic molecules Origin of solar system & Earth Origin of the Universe

the primates, which began only in the early Cenozic or the latest part of the Mesozoic era. he Miocene, Pliocene, and Pleistocene are the most critical in studies of Primate evolution, and they are further subdivided into Early, Middle, and Late stages (e.g., the Middle Miocene). Into the 1970s, treatments of human evolution accentuated the Quaternary (or last) period of the geologic timescale, partly because it seemed to coincide with the appearance and evolution of people and partly because it was considered climatically or faunistically unique. However, it is now clear that the base of the Quaternary, formally deined by local changes in marine fossils in Italy, dates to at most 2.1 Ma (million years ago) and perhaps to only 1.7–1.6 Ma (tentatively accepted here), whereas, as shown below, human evolution began more than 4.5 Ma. In addition, there is no sharp faunal or climatic break between the Quaternary, formally deined, and the immediately preceding Pliocene. A much sharper break, involving a signiicant drop in global temperature and increased

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21 FIGURE 2.1. The geologic timescale (modiied after Gradstein et al. [2004]) and schick and Toth [1993], 26). many specialists believe that the Tertiary and Quaternary Periods should be abandoned in favor of the Paleogene, which would incorporate the traditional Paleocene, eocene, and oligocene, and the neogene, which would incorporate the miocene, Pliocene, Pleistocene, and holocene. Those who favor retaining the Quaternary argue that it was climatically unique (“the ice age,” actually a series of ice ages separated by milder intervals) and that it was the timespan during which humans evolved (it is therefore sometimes known alternatively as the “anthropogene”). however, as conventionally deined, it postdates both earlier Cenozoic intervals of global cooling that resulted in expanded glaciers, particularly after 23 ma, and the initial divergence of humans from apes, 7–6 ma. The long-standing “type” section for the Quaternary is in northern italy, where its base is deined by substantial turnover in marine fossils, now dated to about 1.6 ma. This date is accepted in the table, although an increasing number of specialists prefer a basal date of 2.6 ma, coinciding with a sharp global downturn in temperature.

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continental aridity, occurs within the Pliocene, 2.6–2.5 Ma. he climatic shit can be detected globally, and many authorities argue it should be used to deine the Quaternary, which would then become the last 2.6 my. Other specialists would prefer to abandon the Quaternary totally and to divide the Cenozoic Era between only two periods, the Paleogene between 65.5 and 23.3 Ma and the Neogene between 23.3 Ma and the present. Given continuing disagreement on the deinition of the Quaternary, it will not be treated separately here, except to note that those who place its base at 1.7–1.6 Ma commonly subdivide it into three parts: (1) the early Quaternary, between 1.7 Ma and the beginning of the Brunhes Normal Polarity Chron, roughly 780 ka (thousand years ago); (2) the middle Quaternary between roughly 780 ka and the beginning of the Last Interglacial about 130 ka; and (3) the late Quaternary spanning the past 130 ky. he Brunhes Normal Chron and the Last Interglacial are discussed below.

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SOURCES: hierarchy of stratigraphic units (Hedberg 1976); dating the base of the Quaternary in Italian type section to 2.1 Ma (Kukla 1987) or 1.7–1.6 Ma (Tauxe et al. 1983); climatic break within the Pliocene (Denton 1999); controversial deinition of the Quaternary (Kerr 2008)

Relative Dating Methods he techniques for estimating the geologic antiquity of sites, fossils, artifacts, and other objects in geologic time divide between numerical (or absolute) methods that determine ages in years and relative methods that say only whether one item is older than another. Relative methods were developed irst and are more widely applicable. In addition, all numerical methods are obviously also relative ones, and the credibility of numerical methods depends in part on their consistency with other, standard relative methods. In short, relative dating methods remain basic to all historical geologic studies, including research on human evolution. Among relative methods narrowly deined, the most fundamental is the principle of stratigraphic superposition. his underlies the concept of stratigraphic units introduced in the previous section and in essence states that, all other things being equal, objects found in higher rock layers postdate objects found in deeper layers. he qualiication “all other things being equal” is necessary because burrowing animals, invading roots, and the like can displace objects into lower or higher layers, while crustal movements, landslides, and other geomorphic events can occasionally reverse a stratigraphic sequence, placing older layers on top of younger ones. Where such disturbances occur, however, their efects are oten minor, detectable, or both, and the principle of superposition has been fruitfully applied at countless archeological and fossil sites. Its main limitation is that, in the most literal sense, it cannot be used to date ob-

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jects in layers that do not physically overlap, that is, objects that occur at physically separate sites. In situations where physical overlap does not occur, stratigraphic dating depends on perceived similarities or diferences in the properties of two (or more) layers. he properties can be lithological, physical, chemical, or fossil, taken separately or in combination. In a sense, numerical or absolute dating methods simply illustrate the stratigraphic principle applied to the physicochemical composition of rock layers. Numerical methods are special not only because they indicate stratigraphic ages in years but also because the methods themselves usually do not require a special knowledge of local geological history and can be applied in the same way anywhere in the world. In contrast, relative dating methods require detailed local information, which restricts their application to limited geographic areas. he size of the area can vary from the immediate neighborhood of a site to a large region or even a continent. As illustrated below, relative methods that employ physicochemical analysis tend to be most tightly restricted, while ones that rely on fossils tend to have the broadest geographic application.

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Relative Dating by Chemical Content: The Fluorine Method

Among relative dating methods that depend on chemical analysis, the most inluential in human evolutionary studies has been the luorine method. It is based on the observation that buried bones adsorb luorine from groundwater. Bones that were buried in the same site at the same time should contain the same amount of luorine, and gross diferences in luorine content thus suggest noncontemporaneity. he luorine method was instrumental in unraveling the notorious Piltdown hoax, named for a site in Sussex, southern England, where a seemingly ancient skull and mandible were found in 1911–1912. he skull was thick-walled, but otherwise remarkably like that of a modern human while the jaw was very apelike in basic structure, including a bony (or simian) shelf behind the mandibular symphysis (chin region). he combination became increasingly incongruous as new fossil inds from elsewhere failed to replicate it, and in 1953 it was shown to be a forgery. A major indication was that the skull and mandible contained diferent concentrations of luorine and that both contained much less luorine than most of the associated animal bones that were the main evidence for great antiquity. It is now known that the skull came from a relatively recent human and the mandible came from an orangutan. In the Piltdown case, and in others where the luorine method has been used to check the contemporaneity of bones from the same site, the value of this method is plain. But it cannot be used to determine the relative ages of bones from diferent sites, because there is substantial

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geographic variation both in the lux of groundwater and in the amount of luorine it contains. he same limitation afects virtually all other relative dating methods based on the accretion or deletion of chemicals (for example, nitrogen or uranium) in buried objects.

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SOURCES: general description of lourine dating (Oakley 1969); Piltdown discovery (Chippindale 1990; Gardiner 2003; Spencer 1990; Tobias 1992, 1993), skull demonstrated as a forgery (Weiner 1955; Weiner et al. 1953), and its original constituents (Lowenstein et al. 1982)

Biostratigraphic Dating

In most instances, sites that do not physically overlap in some way and that are not amenable to absolute dating can be dated relative to one another only by their contents. here is an obvious danger of circularity here, if dating is based on objects whose relative age has not been independently determined. For example, it would be unwise to conclude that one site is older than another merely because it contains simpler artifacts, because irrefutable examples have shown that artifact change is not always progressive. Artifacts can be used for dating only when the cultural stratigraphy of a region has been established on independent empirical grounds. Likewise, fossils can be used only when the biostratigraphy of a region has been worked out in advance, but fossils have at least two clear advantages over artifacts. First, of course, they can be used to date sites where artifacts do not occur or that formed before artifacts were made. Second, at least with regard to the earlier stages of human evolution, they oten deine stratigraphic units that cover larger areas and shorter timespans than do units deined by artifacts. For biostratigraphic purposes, the most useful fossils come from taxa (species, genera, etc.) that spread quickly and widely, from taxa that appear to have died out over large areas at about the same time, or from taxa that were evolving rapidly, so that their stage of evolutionary development is itself a clue to their relative age. he aim is to deine biostratigraphic units that are as ine (brief) as possible and that transgress time as little as possible (that is, that correspond to the same time interval in every region). Some time transgression is inevitable, since no species can appear or disappear everywhere simultaneously and since evolutionary changes cannot occur at exactly the same time in all populations of a widespread species. But both theory and empirical research suggest that some taxa provide a basis for deining units whose time transgressiveness is negligible, at least compared with the antiquity of the human fossils or artifacts they can be used to date. In this context, in Eurasia the most productive biostratigraphy developed so far involves the microtine rodents, but large mammals are also useful for relative dating. In Africa, elephants, pigs, and horses have proved most helpful. SOURCES: biostratigraphic value of rapidly evolving taxa (Lister 1992)

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Microtine Biostratigraphy in Europe. he microtine rodents, including

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the voles, the lemmings, and their relatives, are a branch of the cricetids (hamsters) characterized by hypsodont (high-crowned) molars whose occlusal surfaces comprise a series of alternating, triangularly prismatic cusps (ig. 2.2). he heartland of microtine evolution has been in northern Asia, and over the past 5 million years, at times when climatic conditions were suitable, various microtine species have spread both eastward to North America and westward to Europe. hey are currently the most successful rodent group in the Northern Hemisphere, and their past success is relected in numerous fossil occurrences, particularly where bones were accumulated partly or wholly by owls or other birds of prey. he biostratigraphy of the microtines has been studied most thoroughly in Europe, from Russia and Ukraine on the east to France and Britain on the west. In each place, microtine biostratigraphic units can be deined by the irst appearance of particular taxa, by evolutionary changes within established lineages, or by both. Important irst appearances include the immigration (from northern Asia) of the Norway lemming, Lemmus, in the mid-Pliocene, perhaps 3.5 Ma, and of the collared lemming, Dicrostonyx, at or shortly ater the beginning of the Pleistocene, 1.7–1.6 Ma. (his date and others presented below depend mainly on correlations between microtine evolutionary events and the global paleomagnetic stratigraphy outlined later in this chapter.) Based on their historic distribution and climatic preferences, Lemmus and Dicrostonyx probably both migrated to Europe during periods of exceptionally cold climate. heir episodic expansion counts among the clearest biological evidence for repeated climatic deterioration (glaciation) later on, as discussed in the inal section of this chapter. In nearly all microtine lineages, evolutionary change involved the development of increasingly high-crowned molars. In some this was accompanied by a tendency for cementum (calciied tissue that forms on the surface of tooth roots) to pack between the triangular prisms of the molar crowns. Separately and together, these trends allowed more eicient feeding on grasses, which are an important but highly abrasive component of most microtine diets. Increases in crown height and cementum packing are particularly useful for characterizing successive stages in the evolution of the modern water vole, Arvicola, from its immediate ancestor, Mimomys. Based on changes in the irst lower molar (M1), igure 2.2 illustrates the major stages and shows two especially striking transitions, one about 3.25 Ma when cementum irst appeared between prisms and a second roughly 600–500 ka when the molars became ever-growing and ceased to develop roots, even in elderly individuals. As the igure suggests, the evolution of rootless molars is used to deine the appearance of Arvicola. Early populations of Arvicola still included some individuals with rooted molars. In later populations, ater roots were fully lost, successive

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ChaP Ter TW o

FIGURE 2.2. stages in the evolution of the voles Mimomys, Arvicola, and Microtus, based on morphological changes in the lower irst molar (modiied after heinrich [1987], 402). The horizontal arrows indicate approximate times of irst appearance for species in the Mimomys-Arvicola lineage. broken lines indicate two major transitions in this lineage, the irst approximately 3.2–3.3 ma, when cementum irst appeared between molar prisms, and the second nearer 600 ka, when ever-growing, rootless molars evolved. The rarity of archaeological or human fossil sites with (rooted) Mimomys fossils suggests that people were rare or absent across most of europe before 600 ka. among the rare sites with Mimomys, the most important is atapuerca Gd and atapuerca se, spain, dated to roughly 800 ka and 1.1 ma, respectively (bischoff et al. [2007]; additional discussion in chap. 6; Carbonell et al. [2008]; falguères [2003]; falguères et al. [1999]).

Ma

3

0

4

Ma

3

0

4

3

3

2

2

2

1

1

1

Arvicola terrestris Arvicola cantiana

1

5 4

2

1

Microtus arvalis

1

Mimomys savini

2

3 2 1

4 3 2 1

Allophaiomys pliocaenicus nutiensis Allophaiomys pliocaenicus deucalion

2 Mimomys pliocaenicus

5 cm

Mimomys

3

Mimomys polonicus

transition to rootless molars appearance of cementum between prisms cementum

dentine

Mimomys hajnackensis

4

1 mm enamel

Mimomys occitanus

1 mm

5

2

4 3 2

1

1

3

Mimomys davakosi

Mimomys savini Arvicola terrestris (rooted) (ever-growing) left lower M1s

evolutionary stages are recognized primarily by a tendency for the enamel to become thinner on the rear (convex) surfaces and thicker on the fore (concave) surfaces of the molar prisms. Figure 2.2 also illustrates stages in the evolution of the common (or ield) vole, Microtus. he sequence is founded mainly on a tendency for the irst molars to increase in length and on an associated increase in the number of triangular prisms or angles. Microtus perhaps evolved in Asia from a species of Mimomys and then migrated to Europe roughly 1.7–1.6 Ma. Like Arvicola, it was initially distinguished from Mimomys by the development of ever-growing, rootless molars. Together, Mimomys, Arvicola, and Microtus can be used to construct biostratigraphies that are directly relevant to human evolution and prehistory. Figure 2.3 shows

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The Geolo GiC Tim e f r am e

Regional Climatic Stratigraphy Holocene

Time Units

Central European Biostratigraphy (Arvicolidae)

Central Europe Holocene

Arvicola terrestris Zone

Weichsel Glaciation

Eem Interglacial Pleistocene

Quaternary

Important Fossil Sites, with Values for an Index of Molar Enamel Thickness in Mimomys and Arvicola

ArvicolaMicrotus Stage (Toringian) Arvicola cantiana Zone

Saale Complex

Holstein Complex

Southeastern Europe (Pannonian Basin)

Arvicola Euerwanger Buhl H: 83.03 Krockstein/Rübeld: 89.08 Kremathenhöhle: 89.23 Dzerová Skala: 92.04 Roter Berg/Saalfeld: 97.25 Burgtonna 2: 98.44 Untertürkheim: 100.81 Adlerberg/Nördlingen: 100.83 Taubach: 105.15 Schönfeld: 106.02

Pilisszánto: 84.48 Peskö: 89.31 Istállóskö: 89.54 Subalyuk: (96.43) Tata: 99.22

Hórvölgy: 101.91 Weimar-Ehringsdorf (lower travertine): 112.30

Solymár: 108.32

Dobrkovice 2: (123.21) Bilzingsleben: (132:52) Mosbach: (133.54) Hundsheim: 135.15

Budapest-Várbarlang 1: 123.08 Tarkö 4: (129.00) Vértesszöllös: (136.14)

Budapest-Várbarlang 2: (116.83)

Mimomys Elster & Cromer Complex

MicrotusMimomys Stage (Biharian)

Mimomys savini Zone

Přezletice: : 132.98 Voigtstedt: 139.09 Koneprusy C 718: 141.69

one that has wide application in central and southeastern Europe over the entire timespan (middle Pleistocene to Holocene) that people have occupied the area. SOURCES: North American heartland of microtine evolution (Repenning 1980, 1987); microtine biostratigraphy in Russia and Ukraine (van Kolfschoten and Markova 2005); north-central Europe (Fejfar 1976a; Heinrich 1982, 1987; Jánossy 1975; Koenigswald 1973; van Kolfschoten 1994), France (Chaline 1976; Chaline and Laurin 1986), and Britain (Stuart 1982; Stuart and Lister 2001); Mimomys-Arvicola transition (van Kolfschoten 1993); trends in the evolution of Arvicola (Heinrich 1982, 1987; Stuart 1982)

The Biostratigraphic Implications of Large Eurasian Mammals. Large

Eurasian mammals are less helpful for biostratigraphy than small ones because their remains are less abundant and they do not permit such ine stratigraphic subdivision. he microtines, for example, underlie three well-deined biostratigraphic stages within the past 1.7–1.6 my: the Villányan, Biharian, and Toringian. Figure 2.3 lists only the Biharian and Toringian because no unequivocal artifacts or human remains have been found with Villányan species. Over the same long period, large mammals deine only two stages: the late Villafranchian and the succeeding Galerian. he boundary between the two is oten placed at 0.9–1 Ma, where it would coincide broadly with the boundary between the early and late Biharian microtine stages. he distinction of the Galerian from

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FIGURE 2.3. middle and late Pleistocene biostratigraphy of central and southeastern europe, based on the voles Mimomys, Arvicola, and Microtus (modiied after heinrich [1987], 395). The far right column lists key sites with vole fossils, followed by average values for a biostratigraphically signiicant index of irst lower molar enamel thickness (thickness on the concave margin of the molar prisms/thickness on the convex margin × 100). sites shown in boldface type are those with important human fossils or artifacts.

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

ChaP Ter TW o

shed mature antlers of red deer

FIGURE 2.4. shed mature antlers of red deer (Cervus elaphus) from (left) the mosbach mid-Quaternary fossil site, Germany, and (right) the ilford late Quaternary site, england (redrawn after lister [1992], 335). antlers older than 500 ka typically terminate in a simple two-pronged fork, whereas younger specimens exhibit a multipronged “crown.” The difference can be used to date eurasian fossil and archaeological sites relative to each other when other evidence is lacking.

fork

“crown”

trez tine beam bez tine

brow tine

Mosbach (mid-Quaternary) 0

50 cm

Ilford (late Quaternary)

the late Villafranchian is fuzzy, however, and Villafranchian faunas may have given way to Galerian ones only gradually between 1 and 0.5 Ma. he failure of the Eurasian large mammal fauna to deine multiple stages ater 1.7 Ma does not bean that the fauna remained static. On the contrary, the alternation of glacial and interglacial episodes discussed below, coupled with the tendency for later glacial periods to be more intense, induced dramatic changes, including the progressive appearance ater 500 ka of distinctly cold-adapted or cold-resistant species like the arctic hare (Lepus timidus), arctic fox (Alopex lagopus), woolly rhinoceros (Coelodonta antiquitatis), woolly mammoth (Mammuthus primigenius), and reindeer (Rangifer tarandus). he recurrent appearance and disappearance of these in midlatitude Europe in fact counts as some of the strongest evidence for glacial/interglacial alternation ater 300 ka. hey may not deine a biostratigraphic stage, but individually they can be used with other taxa for broad temporal placement. he giant deer (Megaloceros giganteus), the red deer (Cervus elaphus), and the mammoth lineage (Mammuthus) provide particularly good cases in point. he giant deer (or “Irish elk”) ranged from Ireland to central Siberia, beginning roughly 400 ka. It appeared sporadically in diferent regions, relecting local environmental change, but it is unknown anywhere in Europe ater about 9.3 ka and anywhere in Siberia ater about 7 ka. his

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occlusal view

4

x

early M . primig

M. tron gonthe rii

ridiona lis late M. me

bplanif rons

x

x

x

x

late M. su

3

. subpla

2

nifrons

1

early M

millions of years ago

0

enius

lateral view

s

cm

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

late M. primige niu

25

15 20

10

early M . meridio nalis

5

1

29

m

x

x 8

10

12

14

16

18

20

22

24

number of plates FIGURE 2.5. The average number of plates on third molars in successive species of Mammuthus, from its irst appearance in africa before 4 ma until its extinction on the eurasian mainland, roughly 10 ka (redrawn after lister and bahn [1994], 150). a dwarf form survived until about 4 ka on Wrangel island in the siberian arctic (stuart et al. 2004). The earliest species, M. subplanifrons, was exclusively african, but later species, M. meridionalis, M. trogontherii, and M. primigenius, were eurasian. M. primigenius is known popularly as the woolly mammoth from its hairy coat, which late Paleolithic people depicted in their art and which is sometimes partially preserved in permanently frozen ground. The mammoths can be distinguished from other elephants by their domed braincases and their inwardly curved tusks, shown on a reconstructed woolly mammoth in the igure (redrawn after stuart [1982], 44). The molars of mammoths and other elephants comprise a series of subparallel enamel plates that are held together by cementum. each plate has an enamel shell surrounding dentine. occlusal and lateral views (top left) illustrate the basic structure on an upper third molar of Mammuthus primigenius from last Glacial deposits in britain (redrawn after stuart [1982], 47). The alternation of enamel and dentine produced a rough occlusal surface that helped to grind vegetal foods. in Mammuthus (and Elephas, as discussed in the text), natural selection for a more abrasive diet favored an increase in the number of plates through time. The increases occurred in geographically localized populations and spread to other populations through migration or gene low (lister et al. 2005b). Third molar plate numbers in last Glacial M. primigenius sometimes exceeded 25, as illustrated on the occlusal view in the igure.

is despite the existence of later sites where it might be expected to occur if it had survived, and its presence in an otherwise undated context thus implies a time before 10 ka or 8 ka, depending on the place. he red deer (a close relative of the American elk) is useful especially for a change in antler form about 500 ka, in which a simple double-pointed terminal fork evolved into a multipronged crown (ig. 2.4). Crowned antler tips indicate an age of less than 500 ka for the Swanscombe site in England

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Mammuthus

E. ekorensis

to Eurasia

PLEISTOCENE

to Eurasia

PLIOCENE

E. recki brumpti

L. atlantica angammensis L. exoptata

3

4

E. recki shungurensis E. recki atavus E. recki ileretensis E. recki recki

L. atlantica

L. africana L. adaurora kararae

1

2

Elephas

M. cf. meridionalis

Loxodonta

0

M. africanavus

Ma

M. subplanifrons

FIGURE 2.6. Time ranges of the elephantinae in africa (modiied after Cooke [1984], ig. 2). The successive stages of Elephas recki are particularly useful for establishing the chronological relations among otherwise undatable early and middle Pleistocene sites.

E. iolensis

ChaP Ter TW o

L. adaurora

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

30

5 Primelephas stock

with its famous fossil human skull, while simple two-point (acoronate) antlers imply a time before 500 ka for the well-known Ambrona and Torralba hand ax (Acheulean) sites in Spain. Finally, the mammoth lineage originated in Africa about 4.0 Ma and appeared in Eurasia between 3.5 and 3.0 Ma. Shortly thereater, mammoths became extinct in Africa and new species or subspecies arose primarily in eastern Asia, from which they subsequently spread to Europe. Like the fossil African elephants considered in the next section, successive species or subspecies of mammoths tended to evolve ever more complex molars, in which the enamel that surrounded individual molar plates became thinner, the plates became more tightly packed, and the number of plates per molar increased

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Ma 10

8

6

4

2

0

Cainochoerus africanus Nyanzochoerus sp. Nyanzochoerus devauxi Nyanzochoerus tulotos Nyanzochoerus waylandi Nyanzochoerus cookei Nyanzochoerus kanamensis Nyanzochoerus pattersoni Nyanzochoerus syrticus Nyanzochoerus australis Nyanzochoerus jaegeri Notochoerus euilus Notochoerus clarki Notochoerus capensis Notochoerus scotti Notochoerus harrisi Kolpochoerus deheinzelini Kolpochoerus afarensis Kolpochoerus heseloni Kolpochoerus cookei Kolpochoerus limnetes Kolpochoerus phacochoeroides Kolpochoerus paiceae Kolpochoerus majus Kolpochoerus morrocanus Kolpochoerus oldowayensis Metridiochoerus andrewsi Metridiochoerus modestus Metridiochoerus hopwoodi Metridiochoerus campactus Hylochoerus antiquus Hylochoerus meinertzhageni Phacochoerus antiquus Phacochoerus aethiopicus Potamochoerus porcus Sus scrofa

31 FIGURE 2.7. Time ranges of the fossil suids of africa after 10 ma (redrawn after Pickford [2006], igs. 5 and 6). Within each suid genus, some species were ancestral to others, and contemporaneous species within a genus often lived in different parts of africa. Nyanzochoerus and its probable descendant Notochoerus dominated the interval between about 10 and 3.5 ma, while Kolpochoerus and Metridiochoerus dominated between 3.5 ma and perhaps 800 ka. The extant genera Hylochoerus (giant forest hog), Phacochoerus (warthog), and Potamochoerus (bushpig) dominated thereafter. humans may have introduced Sus (eurasian wild boar) to northern africa within the last 10 ky. suids before 5 ma had generally low-crowned teeth and they were probably mainly browsers. many later suids had highcrowned teeth, and like those of contemporaneous bovids, they were probably more adapted to grazing. The shift from an emphasis on browsing to grazing relects the expansion of grassy vegetation in africa after 5 ma.

through time. hus, mean enamel thickness and the mean number of plates can oten date mammoth-rich sites relative to each other (ig. 2.5). SOURCES: iner stratigraphic subdivision allowed by small vs. large mammals (van Kolfschoten 1994); Villanfranchian/Galerian boundary at 1–0.9 Ma (Azzaroli et al. 1988) or less distinct between 1 and 0.5 Ma (Kahlke 2007; van Kolfschoten 1994, 1998); biostratigraphic implications of Megaloceros giganteus, Cervus elaphus, and Mammuthus sp. (Lister 1992); time range and extinction of Megaloceros (Gonzalez et al. 2000; Lister et al. 2005a; Stuart 1991; Stuart et al. 2004); transition from acoronate to coronate antler form in Cervus elaphus ka (Di Stefano and Petronio 2003; Lister 1986); changes in mammoth molars through time and space (Lister 1993; Lister and Bahn 1994; Lister et al. 2005b)

The Biostratigraphy of Elephants, Pigs, and Horses in Africa. he microtines can play no role in African biostratigraphy, for they occur only in Eurasia and North America. Many African groups can be used, however, and from a paleoanthropological perspective, the most noteworthy are the elephants, pigs, and horses. During the timespan of human evolution,

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32

* 2

EKORA

LANGEBAANWEG LOTHAGAM

LUKEINO

7

MPESIDA

Later 3

*

4

5

* 6

7

Later

6

*

H. turkanense

ATERIR MURSI KANAPOI

H. sitifense

Equus datum ?ST. ARNAUD

? AMAMA

8

M L E

1

8

NAKALI ?

Hipparion africanum

9

NGORORA 11 BEGLIA FM

9

10

11

Middle

10

BOU HANIFIA

PLEISTOCENE

E. zebra

?

Middle

*

PLIOCENE

*

0

Earlier

E. oldowayensis &

I

H. libycum

OLDUVAI

II

?

MIOCENE

5

CORNELIA OLORGESAILIE KANJERA

H. sp.

4

B A

III

H. afarense

3

IV

H. cf. africanum

2

L K J H G F E D C

LAETOLI

SHUNGURA FM (OMO)

1

KOOBI FORA FM

?

Ma

E. asinus

0

E. labeli

other deposits

capensis E. quagga E. burchelli

main deposits with dating control

Ma

E. grevyi

FIGURE 2.8. Time ranges of the later miocene to holocene horses (equidae) of africa, with some important sites where horse fossils occur (modiied after Cooke [1984], ig. 1). from a biostratigraphic point of view, the two most important events were the arrival and dispersal of three-toed horses (Hipparion) sometime between 12 and 10.5 ma and of one-toed horses (Equus) shortly before 2 ma.

CHEMERON

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

ChaP Ter TW o

? * 12

12

Hipparion datum

all three groups were characterized by important irst and last appearances and also by lineages for which directional evolutionary trends have been established. Fossil elephants, pigs, and horses provide the main basis not only for dating many important African sites relative to each other but also for cross-checking the validity of absolute dates when they are available. Figure 2.6 summarizes the biostratigraphic aspects of the elephants. Until perhaps 400 ka, the most common genus in Africa was Elephas, which appeared shortly before 4 Ma. An early species, Elephas ekorensis, was ancestral to the especially common Elephas recki, which evolved

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somewhat before 3 Ma and then underwent progressive change through time. his change is particularly obvious in the molars, which became increasingly high crowned while the enamel surrounding individual molar plates became thinner and more tightly folded. Based on molar change, several stages of Elephas recki have been deined, and where appropriate fossils occur, these can be used to bracket sites within an average range of about 500 ky. Pigs and horses provide a broadly comparable picture (igs. 2.7 and 2.8). Authorities disagree slightly on details with respect to the pigs, but both irst appearances and directional changes in molar dimensions and morphology can be used for biostratigraphy. he horses are most useful with regard to irst appearances, especially since most major events in horse evolution took place outside Africa and immigrant taxa then dispersed rapidly through much of the continent. Both the spread of the three-toed horse, Hipparion, and that of the modern one-toed form, Equus, are key time markers for biostratigraphic dating. hree-toed horses broadly referable to Hipparion evolved in North America about 15.5 Ma, spread through Eurasia 12–11 Ma, and dispersed from north to south through Africa around 10.5 Ma. One-toed horses evolved in North America about 3.7 Ma, dispersed through Eurasia around 2.5 Ma, and reached Africa shortly before 2 Ma. For 1.2–1.3 my they overlapped with threetoed species, whose inal known appearance was 800–900 ka. Together with pig taxa, the one-toed horse was especially crucial in demonstrating a large numerical (radiopotassium) dating discrepancy between the Koobi Fora and the Lower Omo sites, located only 100 km apart in eastern Africa. At irst, it seemed that a fauna including the one-toed horse had appeared roughly 2.6 ka Koobi Fora, but only about 2–1.8 Ma in the Lower Omo. As discussed in chapter 4, this led to further checking of the numeric dates, and it was ultimately shown that the original Koobi Fora radiopotassium estimates were in fact 700–600 ky too old.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: biostratigraphic implications of African elephants, pigs, and horses (Cooke 1984); change through time in Elephas recki molars (Beden 1979; Coppens et al. 1978; Maglio 1973); chronologic ranges of African late Cenozoic suids (Cooke and Wilkinson 1978; Harris and White 1979; Pickford 2006); origin and spread of three-toed horses (Cooke 1984; MacFadden 1992) and one-toed horses (MacFadden 1992; Palmqvist and Arribas 2001); resolution of the dating discrepancy between Koobi Fora and the Lower Omo (Hay 1980)

Numerical Dating Methods Although biostratigraphy and other relative dating methods remain basic to human evolutionary studies, the impact of numerical (“absolute”) dating methods cannot be overstated. his is especially true of isotopic (radiometric) methods, which have revolutionized paleoanthropology since their initial development in the 1940s and 1950s. Until 1960,

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ChaP Ter TW o

the best guesstimate for the timespan of human evolution was perhaps the past 1 million years or so. his notion changed radically in the early 1960s, when the irst radiopotassium dates from Olduvai Gorge, Tanzania, showed that people in the broad sense (hominins) already existed at least 1.75–1.85 Ma. More recently, as detailed in chapter 4, the radiopotassium method has been applied at many other east African sites, where it has combined with other dating methods and biostratigraphy to show that human evolution began substantially earlier, certainly before 4.4 Ma. he radiocarbon method has had a comparable impact at the recent end of the timescale, where, for example, it has helped to demonstrate that fully modern people did not appear everywhere at the same time. he implications of this are considered in chapters 6 and 7. he main purpose of this section is to summarize the principles behind radiopotassium dating, the radiocarbon technique, and other isotopic methods that have become so vital to human evolutionary studies. It also touches briely on nonisotopic numerical methods, of which the most important is paleomagnetism (paleomagnetic stratigraphy). Inevitably, it also points up some important limitations of numerical dating, above all the nagging diiculty of obtaining reliable dates between the minimum limit of the conventional radiopotassium method at perhaps 500–300 ka and the maximum limit of conventional radiocarbon dating at 40–30 ka. SOURCES: irst Olduvai radiopotassium dates (Evernden and Curtis 1965; Leakey et al. 1961); overview of isotopic dating methods (Ludwig and Renn 2000)

Some General Features of Isotopic Dating All isotopic dating methods rely on the decay of naturally occurring radioactive elements or isotopes (varieties of elements). In the decay proTABLE 2.1. Basic parameters of some isotopes used in numerical dating.

Isotope

Material Dated

Potential Range (Years)

Carbon-14

wood, charcoal, shell

100,000 theoretically; presently 5730 60,000 under ideal circumstances; 40,000 more commonly

Protactinium-231

deep sea sediment, shell, coral, travertine

200,000

32,700

horium-230

deep sea sediment, shell, coral, travertine

500,000

75,700

Uranium-234

coral

1 million

245,000

Chlorine-36

groundwater

500,000

310,000

Beryllium-10

deep sea sediment, polar ice

8 million

1,600,000

Potassium-40

volcanic ash, lava

(no practical limit)

1.25 billion

Uranium-238

igneous and metamorphic rocks

(no practical limit)

4.47 billion

Rubidium-87

igneous and metamorphic rocks

(no practical limit)

48.8 billion

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14

C 40 Ar/40K & 40Ar/39Ar Fission-track

+ 1-5% + 1-5% + 1-5%

Uranium-series 230

Th/234U Pa/234U 234 U/238U

+ 1-5% + 1-5%

231

+ 5-15%

Luminescence ESR AAR 102

+ 10-20%

103

104 105 Years B.P.

106

107

35 FIGURE 2.9. Time ranges covered by the numerical dating methods discussed in this chapter (adapted from schwarcz [1992], ig. 1). note that the horizontal scale (years before the present) is logarithmic. in practice, dates near the limits of each method may be dificult to obtain and their reliability may be especially questionable. This is perhaps particularly true of especially “old” dates from the 14C method and especially “young” dates from the 40K/40ar method.

cess, radioactive atoms are transformed from one isotope into another (not necessarily of the same element). For example, the justly famous radioactive isotope of carbon, carbon-14 (14C), decays to nitrogen-14 (14N). For each isotope, the rate of decay is a constant that is unafected by ordinary environmental variables such as temperature and humidity or by the chemical compound in which the isotope occurs. Each isotope’s decay rate is ordinarily expressed as its half-life, which is the average amount of time necessary for half the radioactive atoms in a sample to decay. Table 2.1 lists the half-lives of several radioactive isotopes that are important in isotopic dating. As an example, the table presents the half-life of 14C as 5,730 years, which means that roughly half the 14C atoms in a sample will disintegrate within 5,730 years, half of the remainder at the end of 11,460 (5,730 × 2) years, half of what is then let at the end of 17,190 (5,,730 × 3) years, and so forth. Technically, some 14C will always remain in a sample, though with conventional technology it is diicult to measure the amount ater a lapse of 25–30 ky. To obtain an isotopic date, several conditions must be met. First, of course, the object to be dated must contain a radioactive isotope with a known half-life. Second, at the time of formation, the object should in general have contained only the radioactive “parent” and none of the “daughter” into which it decays. In mathematical terms, at time of formation, the daughter/parent ratio should have been 0:1 (zero). Finally, it must be possible to measure the amount of parent and daughter in the object today, that is, to establish the modern daughter/parent ratio. If these conditions are met, the half-life of the parent (the known rate at which it decays to the daughter) may be used to calculate the time that

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has elapsed since the daughter/parent ratio was zero, that is, since the object formed. In practice, the condition that there be no initial daughter (that the daughter/parent ratio be zero) is frequently violated, but fortunately it is oten possible to separate original daughter from “radiogenic” daughter. From a paleoanthropological perspective, the most important isotopic dating techniques are the radiopotassium (K/Ar) methods, the ission-track method, the uranium-series method(s), and the radiocarbon method. his section also outlines the luminescence method(s) (TL and OSL) and electron spring resonance (ESR), since like isotopic methods narrowly deined, they also depend on radioactive decay. Figure 2.9 shows the approximate time ranges that each dating method covers and the percentage of error or uncertainty associated with the dates it produces.

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Radiopotassium (Potassium/Argon) Dating

his method relies on radiopotassium (40K), which makes up about 0.01% of naturally occurring potassium (K) in rocks. he 40K decays to argon40 (40Ar) and calcium-40 (40Ca) in a known ratio, and in theory, either the 40Ca/40K or the 40Ar/40K ratio could be used for dating. However, in practice the 40Ca/40K ratio is not useful because in most rocks radiogenic 40 Ca cannot be distinguished from original 40Ca, which is abundant in nature. In contrast, the 40Ar/40K ratio is extremely useful because rocks heated to a very high temperature (generally above 6,000°C) tend to lose any original argon (an inert gas) they contain. When they cool, radiogenic argon begins to accumulate again. Since the rate of accumulation is known (it is the rate at which 40K decays into 40Ar), the 40Ar/40K ratio in a rock is a direct function of the time elapsed since the rock cooled. Radiopotassium dating is more complicated in practice than in principle. It works best on rocks—or, more precisely, on minerals within rocks—that are rich in potassium. Some minerals tend to lose their argon under physical or chemical stress, independent of heating, and secondary heating may cause some but not all radiogenic argon to escape. A date for a rock that has been secondarily heated may thus confound two (or more) heating events. Because radiopotassium dating generally requires high temperatures to set the clock (daughter/parent ratio) to zero, its use is restricted mainly to volcanic extrusives, such as lavas and ash falls. Meteorites that were heated during passage through the atmosphere may also be dated, but they are much rarer than volcanic rocks. he extraordinarily long half-life of 40K (1.25 billion years) means that the radiopotassium method has no practical maximum limit (it can be used to estimate the age of the earth) but that in most cases it cannot be used to date rocks younger than a few hundred thousand years old. his is because they contain too little 40Ar

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for accurate measurement and because the statistical error associated with the age estimate may therefore be as large as the estimate itself. Since the late 1980s, the conventional radiopotassium (40K/40Ar) method has been largely replaced by a variant that oten allows more accurate and precise dates and that can be applied to otherwise undatable contaminated or mixed samples. In this variant, known as the 40Ar/39Ar method, the target sample is irradiated to convert its natural 39K content to 39Ar. he amount of resulting 39Ar is directly proportional to the content of 40K, and it can be measured more precisely than 40K content. he biggest advantage of the 40Ar/39Ar variant, however, is that it can be used to estimate the 40K/40Ar ratio in extremely small samples, including single grains of volcanic origin. It thus allows age estimates where grains from more than one volcanic event may be mixed in a single sample. A prominent example is the GATC tuf (volcanic ash layer) that immediately underlies the oldest known hominid fossils at the Aramis locality in the Middle Awash Valley of Ethiopia. Twenty-six individually dated feldspar crystals from a sample of this tuf divide between nine that formed an average of 23.6 Ma and seventeen that formed about 4.4 Ma. he 23.6my-old grains represent contaminants (xenocrysts) from a much older volcanic event, and only the 4.4-my-old grains truly bear on the age of the hominid fossils. In this instance, conventional K/Ar dating on the mixed sample gives a seemingly precise but obviously erroneous (mixed) age of about 15.5 Ma. he technology that underlies the 40Ar/39Ar method is complex, and it oten relies on lasers for the intense heat that is required to release argon from an irradiated sample. Like the conventional K/Ar method, the 40Ar/39Ar technique usually cannot produce dates that overlap the radiocarbon-time range within the past 50 ky because very young samples usually contain too little radiogenic 40Ar. In unusual circumstances, however, when a large or particularly potassium-rich volcanic crystal is available, 40Ar/39Ar can produce remarkably young dates. he most striking example is a date of about 1925 b.p. (years before the present) on large sanidine grains from the eruption of Mount Vesuvius that buried Pompeii. his closely approximates the known historic date of 1918 b.p. (a.d. 79). he radiopotassium method (broadly understood to include 40Ar/ 39 Ar) has been extensively applied in eastern Africa, where vulcanism has been nearly continuous since the early Miocene. Dates on volcanic extrusives, such as ash layers, that lie stratigraphically above or below a fossil or archeological site can be used to estimate the site’s age, as, for example, in the Middle Awash or at Olduvai Gorge, where a series of dates indicate that the oldest deposits, with their artifacts and bones, accumulated about 1.8 Ma. Another paleoanthropologically important application of the radiopotassium method is the dating of paleomagnetic reversals recorded in volcanic extrusives (see below). Since paleomagnetism

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is also recorded in nonvolcanic sediments, it can oten be used in turn to provide rough dates at fossil or archeological sites where direct radiopotassium dating is impossible.

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SOURCES: application of radiopotassium dating (Dalrymple and Lanphere 1969; Gentner and Lippolt 1969; Hall and York 1984); 40Ar/39Ar dating (Deino et al. 1998; Ludwig and Renne 2000; McDougall and Harrison 1988); 40Ar/39Ar dating of the GATC tuf (WoldeGabriel et al. 1994); 40Ar/39Ar dating of ash that buried Pompeii (Renne et al. 1997)

Fission-Track Dating

his method employs the “tracks” formed by spontaneous ission of uranium-238 (238U) in naturally occurring glasses and minerals. he tracks are annealed (erased) when a substance is heated to a high temperature, and they reform when it cools. Since the rate of track formation is directly proportional to the half-life of 238U, the age of the last cooling event can be determined from the ratio between the density of tracks and the amount of 238U a sample contains. he tracks are usually enlarged by acid etching in the laboratory and then counted under high magniication. A potential complication is that, depending on the substance, some annealing (track loss) can occur at low temperatures. his is particularly likely in natural glasses, which therefore tend to produce ages that are “too young.” Like the radiopotassium technique, ission-track dating is applicable mainly to volcanic extrusives. It requires minerals that are moderately rich in 238U (if there is too much, the tracks are too closely packed for counting). It has been used much less oten than radiopotassium, largely because it is more tedious. From a paleoanthropological perspective, its main importance has been to check radiopotassium dates at the important east African sites of Olduvai Gorge, Koobi Fora, Middle Awash, and Hadar. SOURCES: fundamentals of ission-track dating (Fleischer et al. 1969; Green 1979; Naeser and Naeser 1984; Wagner 1996); application at Olduvai Gorge (Fleischer et al. 1965), Koobi Fora (Gleadow 1980; Hurford et al. 1976), the Middle Awash (Hall et al. 1984), and Hadar (Walter and Aronson 1982)

Uranium-Series Dating

Uranium-series (U-series) dating is actually a set of methods based on the parent isotopes 238U, 235U, and thorium-232 (232h). All three decay, through a series of intermediate radioisotopes, to stable isotopes of lead. Some of the intermediate products decay too rapidly to be useful for dating geologic events, while those with longer half-lives tend to decay at rates about equal to the rates at which they are produced. his means they can be used for dating only when they are transferred from the system in which they were produced to another, where their removal or introduction sets the clock to zero.

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In general, U-series dating depends on the high solubility of uranium in water, whereas its decay products, including 230h (from 238U via 234U) and protactinium-231 (231Pa) (from 235U) usually precipitate out as they form. Uranium in seawater or lake water thus tends to remain in solution, while 230h and 231Pa become incorporated in sediments that accumulate on the loor below. Assuming that the 230h and 231Pa content was always the same as it is at the sediment surface today, the quantity of 230h or 231Pa in a buried layer can be used to calculate its age. his method is obviously applicable only to sea- or lake-bottom sediments, where it can, however, date climatic events that may bear on human evolution. here are several sources of potential error, including the possiility that terrestrial erosion introduced uranium to ancient lake or sea deposits and this then produced some of the 230h and 231Pa. In addition, the rate at which 230h and 231Pa settle out need not remain constant over time. In theory, the principles behind U-series dating of sea- or lake-loor deposits should also permit U-series dating of aquatic shells, which are usually built of carbonates that have been dissolved in the surrounding water. With the carbonates, there is also some dissolved uranium but virtually no (insoluble) 230h or 231Pa. he 230h /234U and 231Pa /235U activity ratios in a shell should therefore relect the time elapsed since the shell formed. U-series analysis of shells has dated ancient (raised) beaches in the Mediterranean that are stratigraphically related to archeological or fossil sites, but in general shell dates are problematic because shells tend to adsorb uranium from their burial environment. Bones commonly do too, since fresh bone contains only 0.1 ppm (parts per million) of uranium, while fossil bones typically contain 10–1,000 ppm. Diferent bones in the same layer can adsorb diferent amounts of uranium, resulting in discrepant dates, and diferent parts of large bones can even produce diferent dates. he bottom line is that U-series ages on bone should be regarded critically, and they are presented in succeeding chapters mainly when no alternative dating is available. Among carbonates of organic origin, those in corals provide the most reliable U-series ages, partly because coral carbonates tend to be relatively rich in uranium to begin with and partly because uranium intake ater death is rarer and relatively easy to detect. Fossil corals from various parts of the world have provided concordant dates of 125 ± 10 ka for a high sea level generally correlated with the Last Interglacial (discussed in the section on Cenozoic climates below). In addition, as discussed in the next section, accurate, high-precision U-series dates on coral show that radiocarbon dates oten underestimate true calendar ages, and the U-series ages may then be used to correct (or “calibrate”) the radiocarbon timescale to 23.5 ka and beyond.

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Finally, the 230h /234U and 231Pa /235U activity ratios can be used to date inorganic carbonates—limestones, cave lowstones (or speleothems, i.e., stalagmites and stalactites), and travertines—that precipitate out of solution in cave, spring, or lake deposits. Recently precipitated carbonates will contain coprecipitated uranium (from 0.1 to 10 ppm), but negligible amounts of thorium and protactinium, since they are relatively insoluble. hen 234U decay causes 230h and 231Pa to accumulate in the carbonate, and the 230h /234U and 231Pa /235U activity ratios can be used to estimate the time of carbonate precipitation. Maximum age estimation is constrained by the time it takes each ratio to reach 1:1 (equilibrium). his is about 500 ky for h/U and 200 ky for Pa/U. Both ratios, and especially h/U, can thus provide ages between the practical lower limit of radiocarbon and the common upper limit of radiopotassium. he relative abundances of 234U and its daughter products can be measured by counting atoms as they decay, mainly by alpha spectrometry, or before they decay, primarily by thermal ionization mass spectrometry or TIMS. he diference is analogous to the one between conventional and accelerator radiocarbon dating, discussed below, and it has the same important consequence: TIMS measurement is much more precise, which means it can provide older dates and also dates on smaller samples. Prior to TIMS measurement, the ranges of h/U and Pa/U dating were roughly the 350 ky and 130 ky, respectively, as opposed to 500 ky and 200 ky, as speciied above. U-series dating has been applied to inorganic carbonates in some key archaeological and human fossil sites, especially in Europe, but the results (reported in chaps. 5 and 6) are oten controversial. his is partly because the age estimates are not always internally consistent and partly because they oten seem to contradict dates implied by biostratigraphy or other methods. Sources of error include the possibility that the dated carbonate was contaminated by “detrital” uranium (in particles introduced by wind or lowing water) and the possibility that leaching removed uranium or its daughter products from the carbonate ater precipitation. he presence of 232h in a sample implies contamination, since 232 h does not result from 234U decay, and the amount of 232h can sometimes be used to estimate the proportion of 230h that also represents contaminant. he “corrected” 230h/234U ratio can then be used to estimate a true age. U-series dating is obviously most successful when it can be applied to sealed, pristine, uncontaminated carbonate deposits, and it is most compelling when samples from a single site provide replicable dates that are in proper stratigraphic order. From a paleoanthropological perspective, the most important application of U-series dating has probably been at Atapuerca SH (Sima de los Huesos), near Burgos, northern Spain. U-series analysis suggests that an SH lowstone (stalagmite) overlying human fossils formed at least 500 ka.

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he fossils represent proto-Neanderthals, and the implication is that the Neanderthal and modern human lines had already diverged by 500 ka. he 17 m thick sequence of travertines in the Abric (Rockshelter) Romaní near Barcelona, northeastern Spain, presents a less spectacular but especially persuasive application of U-series dating. During wet periods, cascades of carbonate-charged waters sprayed the loor of the shelter, and moss growth led the carbonate to precipitate out as travertine. During drier periods, Middle Paleolithic (or Mousterian) and early Upper Paleolithic (early Aurignacian) people occupied the shelter, and travertine layers thus sandwich successive Middle and early Upper Paleolithic horizons. U-series analysis has produced more than twenty stratigraphically concordant dates that begin at 70 ka near the bottom of the travertine sequence and end at 40 ka near the top. he dates near the top place the earliest Upper Paleolithic at about 43 ka, while accelerator (AMS) radiocarbon dates on charcoal from Romaní and other Spanish sites place the earliest Upper Paleolithic between 39 and 37 ka. However, for reasons discussed in the next section, radiocarbon dates in the 35,000–40,000 year range probably underestimate true calendar ages by 3,000 years or more, and even if the radiocarbon dates relected real calendar time, they need be only minima (40–35 ka or older). hus in a manner that may always elude radiocarbon, the Romaní U-series dates show, in a manner that may always elude radiocarbon, that the Upper Paleolithic replaced the Middle Paleolithic in northeastern Spain about 43,000 calendar years ago.

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SOURCES: principles behind U-series dating (Grün 2006, 8–14; Ku 1976; Schwarcz 1980, 1992, 1993; Schwarcz and Gascoyne 1984); U-series dating of raised Mediterranean beaches (Stearns and hurber 1965); diference between fresh and fossil bone in uranium content (Cook et al. 1982); inconsistent Useries ages for diferent parts of the same bone (Bischof et al. 1997a); U-series dates on last interglacial corals (Harmon et al. 1981; Stearns 1984); U-series calibration of radiocarbon dates (Bard 2001; Bard et al. 1990, 1993, 1998); U-series dating of inorganic carbonates (Ludwig and Renne 2000); U-series results for various European archaeological sites (Cook et al. 1982), Atapuerca SH (Bischof et al. 2003, 2007), and Abric Romaní (Bischof et al. 1988, 1994)

Radiocarbon (Carbon-14) Dating

he radiocarbon or carbon-14 (14C) method is the most celebrated of all isotopic techniques, thanks to its widespread application in archeology. It is based on three key physical indings: (1) that cosmic ray bombardment transforms nitrogen-14 (14N) into 14C in the upper atmosphere; (2) that for every atom of 14C in the atmosphere there are about one trillion atoms of 12C; and (3) that 14C is radioactive, while 12C is not. It further relies on four principal assumptions: (1) that the atmospheric ratio between 14C and 12C has remained constant through time; (2) that atmospheric 14C and 12C are equally likely to be oxidized into carbon dioxide (CO2); (3) that CO2 mixes rapidly in the atmosphere, so that the 14C/12C ratio is about the same everywhere; and (4) that most organisms do not

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discriminate between 14C and 12C when they build their tissues. Since most plants obtain all their carbon from atmospheric CO2, and since most animals obtain their carbon directly or indirectly from plants, the 14 C/12C ratio in plants and animals closely approximates the ratio in the atmosphere. Ater an organism dies and carbon assimilation ceases, however, the amount of 14C in its tissues decreases by half approximately every 5,730 years (the half-life of 14C). he 14C decays back to 14N, but age estimation depends on the 14C/12C ratio in a sample of dead tissue. his is compared to the 14C/12C ratio in the atmosphere, and the diference is a function of the time since death. his time is calculated in years and stated by convention in years before 1950 a.d. Two key assumptions behind 14C dating are clearly troublesome. First, the 14C/12C ratio in the atmosphere has varied through time. he combustion of vast amounts of coal, oil, and gas following the Industrial Revolution in the nineteenth century has released large quantities of “fossil” carbon into the atmosphere, reducing the 14C/12C ratio, while nuclear explosions since 1945 have had the opposite efect. Fortunately, these complications can be circumvented by measuring the 14C/12C ratio in calendrically dated objects that antedate the Industrial Revolution, and they do not materially afect dates obtained since 1955, when specialists adopted a standard ratio. However, 14C ages have also been repeatedly checked against the known calendar (solar) ages of Egyptian dynastic or other early historic objects and against ages from methods like tree-ring and U-series dating that provide ages in true calendar (solar) years. he results show that the atmospheric 14C/12C ratio has luctuated through deep time. Most ancient luctuations probably relect past variation in the strength of the earth’s magnetic ield and of the sun’s electromagnetic ield, both of which delect cosmic rays, and in the extent to which 14CO2 has been sequestered in the world ocean. Paired 14C and tree-ring dates allow the radiocarbon chronology to be corrected or “calibrated” over the past 12,400 calendar years, while paired 14C and high-precision U-series (230h/ 234U) dates on corals and 14C dates on planktonic foraminifera from annual sedimentary laminations (varves) in the Cariaco Basin northwest of Venezuela, extend the calibration to about 26,000 calendar years ago. he age of each lamination can be independently estimated from its position in the complete sequence. In contrast to uncalibrated ages, which are commonly presented in plain radiocarbon years, calibrated ones usually include the abbreviation “cal” as, for example, in 10,000 cal yrs b.p. (before the present, or more precisely before 1950 a.d.). In general, over the 26 ky interval for which calibration is reasonably well-established, 14C underestimates true calendar ages, and the degree of underestimation grows with time (ig. 2.10). he greatest discrepancy occurs before 20 ka, when 14 C ages are 3,000–3,500 years younger than calendar ages.

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FIGURE 2.10. a proposed curve for calibrating 14C ages to calendar (solar) years from 0 to 50 ka (modiied after fairbanks et al. [2005], ig. 3). Calibration before 26 ka is not yet irmly established. This is unfortunate for paleoanthropology because modern humans expanded from africa to eurasia between 50 and 35 ka, and in the absence of reliable calibration, the duration and the speed of the expansion remain debatable.

40

perfect agreement 30

proposed departure from perfect agreement (calibration curve)

20

calibration reasonably secure

10

43

calibration uncertain

0 0

10

20 30 Calendar Age (years Before Present x 1000)

40

50

Calibration of dates older than 26 ka remains uncertain because different studies provide discrepant results. he discrepant observations come from 14C dates on organic matter in annual varves at the bottom of Lake Suigetsu, Japan; paired 14C and U-series ages on carbonates (aragonites) laid down in Lake Lisan (the Last Glacial predecessor of the Dead Sea); paired 14C and U-series ages on corals and speleothems (cave carbonates) in various locations; and the correlation of the oxygen-isotope climatic signal in the Greenland Ice Sheet Project 2 (GISP 2) and European Greenland Ice Core Project (GRIP) ice cores with the signal in deep-sea cores. In the last instance, calendar ages estimated mainly from annual laminations in the ice cores are paired with 14C ages on foraminifera in the deep-sea cores. he diferent approaches agree that radiocarbon underestimates true calendar ages between 22 and 50 ka (meaning that the atmosphere was signiicantly richer in 14C than it was historically), but they disagree on the extent. he Lake Suigetsu varves, for example, imply an average underestimate of 2–3 ky, while observations on a Bahamian speleothem suggest the average is closer to 8 ky. he correlation of climatic events between the deep sea and the Greenland ice sheet suggests an average of roughly 5 ky, intermediate between the two extremes. he famous Chauvet Cave wall paintings and the modern human dispersal across Europe separately illustrate the importance of accurate calibration. he Chauvet paintings have an uncalibrated radiocarbon age of about 31 ka, but their calendar age would be 33, 36, or 38 ka depending on which dataset is chosen for calibration. Modern human dispersal in

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Europe provides a more fundamental example. Uncalibrated ages suggest that it occurred westward from the Balkans to western France over about 7 ky, between roughly 43 and 36 ka. Calibration based on speculative averaging of discrepant datasets narrows the dispersal interval to only 5 ky, between roughly 46 and 41 ka, and calibration based on deep-sea/ ice sheet correlations could narrow it still more. During their spread, modern humans progressively displaced Neanderthals locally, and specialists debate the extent to which the two groups exchanged genes or culture. Dispersal over 5–4 as opposed to 7 ky implies that modern humans expanded more rapidly—at a rate near 0.4 km versus 0.3 km/year, and it would thus suggest more limited opportunities for gene or culture exchange on the advancing front. In the absence of reliable calibration, however, the actual expansion rate remains unknown, and chapters 6 and 7, which address the modern human diaspora, present uncalibrated radiocarbon ages only. he second questionable assumption behind radiocarbon dating is that a sample obtained all its carbon from the atmosphere. Some living creatures (particularly some mollusks and water plants) are known to assimilate carbon from old rocks. hey will thus be enriched in 12C relative to the atmosphere, and if they are dated, their 14C ages will exceed their true calendar ages. A more serious and widespread problem arises when more recent carbon contaminates a sample in the ground. Humic acids from rotting vegetation, for example, can percolate downward into buried archeological objects, enriching them in 14C. he radiocarbon method will then underestimate their true (calendar) age, and the degree of underestimation grows with true age. he problem is particularly acute for samples whose actual age exceeds 4–5 14C half-lives (25–30 ky) because such samples will retain little of their original 14C, and even a minute addition of recent 14C will make them seem much younger than they really are. A 1% addition of modern carbon to a sample whose true 14C age is 67 ka, for example, will produce an apparent age of 37 ka. A 1% addition to a sample whose 14C age is 33 ka will produce an apparent age of 30 ka, and there are circumstances in which even this relatively small diference can be meaningful. he age of the last Neanderthals, discussed in chapter 6, provides an example. Most 14C dates from Neanderthal sites antedate 30 or even 35 ka, but some suggest that Neanderthals persisted until 30 ka or even later, perhaps especially in refuges on the extreme southeast and southwest. However, all the ages could be substantially greater than 30 ka if the younger ones were afected by only a tiny percentage of modern carbon contamination, and in general, this probably explains why in any given region ages for Neanderthal sites of 30 ka or less overlap with ages of 30 ka or more for the oldest Cro-Magnon sites. he general principle is that due to the ever-present possibility of minute contamination,

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14

C dates beyond 4–5 half-lives should be regarded as only minimum ages, until they have been replicated at multiple sites where they occur in the expected order between dates from younger and older layers. A corollary is that where 14C ages for the same ancient layer or cultural unit are discrepant, the oldest age is probably closest to the true age. Sample preparation procedures exist to detect and eliminate contaminants, but they are not foolproof, and the preferred procedure is to select materials that are easier to cleanse or less likely to be contaminated to begin with. Charcoal (charred wood) is probably best in this regard. he inorganic components of shell and bone are generally not useful because they tend to exchange carbon with their burial environment. he organic (protein) component of bone or shell is suitable but rarely survives in its original form. More commonly, only some amino acid constituents of protein remain, and these might come from the burial environment rather than from the bone or shell itself. he problem is particularly acute for ancient bones that retain little datable organic matter. he 14C dates on such material are oten younger than dates on associated materials like charcoal, implying that the organic extract was contaminated by more recent, intrusive carbon. Among the aminoacid constituents of bone protein, one—hydroxyproline—is especially sought for 14C dating because it is restricted to bone and does not occur in plants, soil microorganisms, or other likely contaminants. Unfortunately, it is highly soluble and is therefore oten leached from bones ater burial. Sophisticated, rigorous pretreatment of the protein remnant in bone to remove likely contaminants oten results in signiicantly older and probably more reliable dates, but in the main, when the dates exceed 25–30 ka, they should probably still be regarded as minimum age estimates only. Radiocarbon (and other isotopic) ages are properly stated with an estimate of statistical and analytical uncertainly, for example, 14,000 ± 120 years ago. he plus-or-minus igure is computed so that the chances are about 67% (one standard deviation) that the actual radiocarbon age of the sample has been bracketed. In the case of a date given as 14,000 ± 120 years ago, the bracketing interval would be 13,880–14,120 years ago. he error range will be larger for a sample that contains relatively little carbon. If the stated error range is doubled, the probability that the actual age of the sample has been bracketed increases to 96%. If the error range is tripled, there is no practical chance that the age of the sample lies outside the bracketing interval. he time range covered by the 14C method is constrained by the short half-life of 14C. In general, samples older than 30–40 ka (> 5 half-lives) contain too little 14C for accurate measurement by conventional technology. he conventional method estimates the 14C content of a sample by counting radioactive emissions (decaying 14C atoms), and it requires a

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statistically meaningful number of emissions during the time the sample is in the counter. For practical reasons, this time is usually limited to days, and in samples older than 30–40 ka, the number of counted emissions usually falls below the required threshold. he conventional methodology is now rapidly giving way to dating by accelerator mass spectrometry (AMS), in which linear accelerators functioning as ultrasensitive mass spectrometers count 14C atoms before they decay. Once a sample has been prepared, the AMS method is much speedier than the conventional technology, and it can also be applied to much smaller samples. In theory, it could provide reliable ages up to 100 ka, but in practice it is limited mainly to 40–50 ka. he reason is that sample preparation for AMS dating unavoidably introduces a tiny amount of modern carbon contaminant so that samples rarely appear older than 50 ka even when they clearly are. A sample preparation protocol known as acid-base wet oxidation with stepped combustion (ABOX-SC) allows reliable, inite AMS ages on charcoal up to 55 ka, but the rarity of charcoal in ancient sites and the ever-present possibility that ancient bone samples have been contaminated by a minute amount of undetectable, more recent carbon in the ground still means that paleoanthropologists must rely mainly on methods other than radiocarbon to estimate ages beyond 30–40 ka.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: basics of 14C dating (Gribbin 1979; Grün 2006, 4–8; Libby 1955; Taylor 1987, 1995–96; Terasmae 1984); luctuation of the atmospheric 14C/12C ratio through deep time (Seuss 1986; Taylor et al. 1996) and probable causes (Beck et al. 2001; Mazaud et al. 1991; Stuiver et al. 1991); radiocarbon calibration for the past 26 ky (Blackwell et al. 2006; Reimer et al. 2004); calibration before 26 ka (Bard et al. 2006; Fairbanks et al. 2005; Hughen et al. 2004; van der Plicht et al. 2004); the impact of calibration on the age of Chauvet Cave paintings (Bard et al. 2004) and on the timing of modern human expansion across Europe (Mellars 2006b); problems in dating bones that retain little organic matter (Staford et al. 1990; Taylor 1992); value of hydroxyproline (Gillespie et al. 1984); pretreatment to remove organic contaminant from degraded bone protein (Bronk Ramsey et al. 2004; Jacobi et al. 2006); AMS dating (Berger 1979; Taylor 1987, 1995–96); ABOX-SC (Bird et al. 1999)

The Luminescence Methods and Electron Spin Resonance

he luminescence dating methods and electron spin resonance (ESR) depend on the observation that irradiation by naturally occurring radioactive isotopes (mainly uranium, thorium, and radiopotassium) causes electrons to accumulate in defects within crystalline substances. he aggregate number of trapped electrons can be measured, and the rate at which they accumulated can be estimated from the level of background radioactivity to which a substance was exposed. he number of trapped electrons divided by their annual accumulation rate (the annual radiation dose) then provides the last time the crystal traps were empty. he luminescence methods are applied primarily to objects where heat or light emptied (or zeroed) the traps before burial, while ESR is applied mainly to dental enamel, where formation (precipitation during life)

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produced initially empty traps. he time range covered by the luminescence methods and ESR depends on the time it takes for traps to ill (or saturate) so that no more electrons can be added. his varies from substance to substance and from site to site, but in general the methods are applicable to materials that are between a few thousand and a few hundred thousand years old (ig. 2.9). he luminescence methods comprise thermoluminescence (TL), which has been applied mostly to objects like potsherds and occasional lint artifacts in which the crystalline traps were emptied by intense heat before burial, and optically stimulated luminescence (OSL), which has been applied mainly to quartz sand grains in which the traps were emptied by exposure to sunlight. TL and OSL difer mainly in the way they release and measure trapped electrons. TL relies on intense heat and OSL on intense light. In either case, the object glows as the electrons escape, and the intensity of the glow (luminescence) is directly proportional to the number of trapped electrons. Traps typically consist of two kinds— shallow ones from which the electrons are easily released and deeper ones from which they escape less readily. Either set of traps can provide a date, but the deep traps are less likely to have been fully emptied before burial, and they are thus more likely to provide a date that is “too old.” OSL is superior to TL for measuring the glow from shallow traps, and it is thus preferable whenever there is doubt about full zeroing. Such doubt is especially likely to afect sand grains as opposed to potsherds or burned lints. With regard to sand, TL is also more problematic because it generally requires many thousands of grains for a single date, whereas OSL can be applied to only a few dozen or even to single grains. his advantage makes OSL the method of choice, when there is some chance that older and younger grains have been geologically mixed in a single sample. TL and OSL rely equally on a sound estimate of the annual radiation dose, which depends partly on radioactivity within the target object and partly on radioactivity in the burial environment. Radioactivity within the target (the internal dose rate) is readily estimated, but radioactivity in the burial environment (the external dose rate) is more diicult, since it may have varied over time. Variation may occur because groundwater subtracted (leached) or added uranium through time, because variation in soil moisture content variably bufered buried objects from irradiation, or because the texture of the surrounding deposit changed. Finergrained deposits (for example, unconsolidated sand) inhibit irradiation less than lumpier ones (for example, undecomposed sandstone blocks). In short, the luminescence methods depend on site-speciic details that do not afect the radiocarbon, radiopotassium, or U-series disequilibrium methods, and it is for this reason that the uncertainty estimates associated with luminescence dates tend to be relatively large (ig. 2.9).

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In those situations where the annual radiation dose is likely to have luctuated greatly through time, the uncertainty estimate may not be reliably calculated or it may be comparable in magnitude to the reported date. In this circumstance, acceptance of the date becomes a matter of personal preference. Still, paleoanthropologists routinely rely on luminescence dating, for it oten provides the only means to obtain age estimates in the critical gap between the practical lower limit of radiocarbon at about 50 ka and the practical upper limit of radiopotassium at about 300 ka. he luminescence methods can also, of course, be applied where suitable materials for the radiocarbon, radiopotassium, or U-series methods are absent. At some sites, radiocarbon and luminescence dating have provided similar ages, and this has buoyed conidence in the luminescence methodology, particularly in TL when applied to burned lints. Unquestionably, the most important paleoanthropological application of TL so far has been in Israel, where dates on burned lints imply that modern or near-modern people were present between 120 and 90 ka, when Neanderthals were the sole occupants of Europe. he Israeli TL dates and the associated fossils provide crucial support for the Out-of-Africa theory of modern human origins, as presented in chapters 6–8. Electron spin resonance in its conventional application to dental enamel is arguably more problematic than the luminescence methods. It requires the same estimate of background radioactivity as TL and OSL, but unlike them it also must confront the problem that the target objects— teeth—tend to adsorb large quantities of uranium ater burial. Success in ESR dating thus depends on modeling the history of uranium uptake. Ordinarily two discrete models are considered, one in which the uranium is supposed to have accumulated shortly ater burial (the early uptake or EU model) and a second in which it is supposed to have accumulated more continuously (the linear uptake or LU model). he models produce particularly discrepant dates for teeth that are rich in uranium, and in the extreme case, a date based on the assumption of linear uptake can be twice as old as one based on early uptake. he true age is usually assumed to lie in between. ESR has provided dates at many of the same sites to which TL has been applied, but as discussed in chapter 6, the results are not always consistent. Most notably, ESR suggests a much shorter timespan than TL for the Middle Paleolithic in Israel. he problem may be largely that the history of uranium uptake is oten more complicated than the theoretical models allow, and it may oten include uranium subtraction (leaching). ESR dates are reported routinely below, and some are surely accurate. heir overall reliability remains to be established, however, and they should be carefully assessed site by site.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: dating ranges of the luminescence methods (Schwarcz and Rink 2001); application of TL (Aitken et al. 1986; Aitken and Valladas 1993; Richter 2007; Wintle 1980; Wintle and Huntley 1982) and OSL (Aitken 1994; Duller 2004; Feathers 1996; Huntley et al. 1985); application of OSL to single

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grains (Jacobs and Roberts 2007; Roberts et al. 1997); causes of variation through time in the external dose rate (Schwarcz and Rink 2001); site-speciic problems in the application of luminescence dating (Feathers 2002); TL and radiocarbon agreement at European late Paleolithic sites (Aitken et al. 1986; Valladas and Valladas 1987); TL dating of Israeli early-modern human fossils (Mercier et al. 1993; Valladas et al. 1988); requirements of ESR dating (Grün 1993, 2006, 14–24; Hennig and Grün 1983; Schwarcz and Grün 1993); discrepant TL and ESR dates at Israeli Middle Paleolithic sites (Mercier et al. 1995a, 1995b); diiculties in modeling uranium uptake for ESR dating (Zhao et al. 2001)

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

Nonradiometric Numerical Dating Methods he most important nonradiometric numerical dating methods are treering dating (dendrochronology) and varve dating. hey are highly accurate but can be applied only in certain restricted geographic regions, and they cover only the past 8,000–10,000 years. Only varve dating will be summarized here, but tree-ring dating follows broadly similar principles. An additional numerical dating method—amino acid racemization—also deserves mention because its application to ostrich eggshell has provided intriguing dates at some African sites. SOURCES: tree-ring dating in the southwestern United States (Towner 2002)

Varve Analysis

A varve is a distinctive band of sediment, oten made up of two subbands, that is laid down each year on the loor of a lake or other relatively calm body of water. In general the loor must also be anoxic (poorly oxygenated), so that the sediments are not disturbed by burrowing organisms. For dating purposes, the most useful varves are ones that form in quiet glacier-fed lakes. Each year, ater the spring thaw, material washed into such a lake consists of both coarse and ine particles. he coarse ones settle out irst, forming the lower subband of a varve. he iner particles settle out later, forming the upper subband. he overall characteristics of a varve, particularly its thickness, are determined by annually variable events, especially the intensity of the thaw. In an area with several glacier-fed lakes, each year will produce its own distinctive varve, alike from lake to lake. In some regions, paarticularly Fennoscandinavia, there are varved sediments that formed under lakes whose times of existence overlapped. he bottommost varves in one lake thus correspond to the uppermost varves in another. In Fennoscandinavia it has thus been possible, when starting with varves of known age, to establish a reliable varve sequence covering the past 10–12 ky. he varves can be used to date any materials they contain and to trace the retreat of the Last Glaciation ice sheet in its waning phases. Varves of known age have also been used to calibrate the 14C method, though they are less useful for this purpose than are tree rings. SOURCES: varve dating in Fennoscandavia (Tauber 1970)

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Amino Acid Racemization

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

he amino acids that make up proteins can exist in two mirror-image molecular forms: L (let-handed) and D (right-handed). In general, only L-amino acids occur in the tissues of living animals, but they are converted ater death to D-amino acids until an equilibrium 50/50 mix of L- and D-forms is reached. he ratio of L-to-D conversion that depends on ambient temperature, moisture, and pH (acidity), and it difers for each amino acid. he reaction that produces the conversion is known as racemization (or epimerization). Fossil shells and bones oten still contain some amino acids, and if the postmortem temperature and moisture history of a specimen are known, the D /L ratio for a particular amino acid is a measure of the time since death. Like U-series, luminescence, and ESR dating, amino acid racemization (AAR) is appealing because it can provide dates in the period between 40 and 100 ka or more that is not well-covered by either radiopotassium dating or radiocarbon dating (ig. 2.9). he lower limit is constrained by postdepositional temperature, but the method can extend to 200 ka in the tropics and to 1 Ma in cooler regions. he application of AAR to bones was briely popular in the 1970s, but it fell from favor, due in part to the erroneously ancient dates it provided for several prehistoric human skeletons from California. AAR suggested that some of these skeletons dated from between 70 and 40 ka, but archeological context, together with U-series determinations and 14C dates, sometimes on bones from the same skeletons, indicated that the skeletons dated from no more than 11 ka. In this instance (and others), the problem was probably a grossly inaccurate estimate of the racemization rate, which may proceed more rapidly in old bones that have lost most of their original amino acids. In other instances AAR, like 14C dating, can provide mistaken ages when foreign amino acids contaminate a buried bone, and the application of AAR to bone has been largely abandoned. AAR has been more fruitfully applied to mollusk shells, and ostrich eggshell (OES) also appears suitable. Like mollusk shell, OES retains its native organic material better than bone, it is less subject to postdepostional leaching and contamination, and its organic content commonly provides radiocarbon dates that agree closely with dates on associated charcoal. It is particularly abundant in African sites, but it also occurs across southern and central Asia to Mongolia and northern China, wherever ostriches ranged in the past. AAR dating of OES still requires an estimate of average temperature ater burial, but this can be obtained from the amino acid ratio in OES fragments that have been radiocarbondated to between 30 and 20 ka. Such fragments will have experienced a rough average of glacial and interglacial temperature extremes. he most important site to which AAR on OES has been applied is perhaps Border Cave, South Africa. Internally consistent AAR determi-

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nations bracket the Middle Stone Age sequence here between 145 and 70 ka. Other methods provide similar ages for comparable Middle Stone Age artifacts elsewhere. he Border Cave AAR dates may pertain to some well-known early-modern or near-modern human fossils, though the stratigraphic origin of some of these is uncertain or contested (chapter 6). OES is also associated with the even more famous modern or near-modern human remains at Qafzeh Cave, Israel. TL on associated burned lints suggests the Qafzeh human fossils date from about 92 ka, and AAR could provide a useful cross-check, since it relies on totally diferent assumptions. Alternatively, if the 92-ky age is accepted, AAR could be used to estimate the average temperature since burial. his could then be used for AAR dating at other Israeli Stone Age sites.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: overview (Grün 2006, 24–27); racemization rate (Elster et al. 1991); ratio of amino acid Dforms/L-forms to measure time since death of an organism (Hare 1980; Hare et al. 1993); AAR applied to bones (Bada 1985); discrepancies between AAR on bone and U-series (Bischof and Rosenbauer 1981) and radiocarbon dates (Taylor 1992); inaccurate estimation of the racemization rate in old bones (Bada 1985); application of AAR to mollusk shells (Wehmiller 1982) and ostrich eggshell (Johnson and Miller 1997); advantages of AAR on ostrich eggshell compared to AAR on bone (Brooks et al. 1990); AAR on ostrich eggshell from Border Cave (Miller et al. 1993)

Paleomagnetism (Paleomagnetic Stratigraphy) Paleomagnetic (or geomagnetic) dates are generally much less precise than those of other numerical dating methods, and paleomagnetism is perhaps best described as a cross between numerical and relative dating. In essence it provides the basis for a special kind of stratigraphy, with broadly the same kind of dating implications as biostratigraphy. he foundation for paleomagnetism are past luctuations in the intensity and direction of the earth’s magnetic ield. From a paleoanthropological perspective, the most important changes are in polarity, from times when a compass would point north to times when it would point south and vice versa. Reversals in currents within the earth’s luid core almost certainly cause these shits, though the currents are poorly understood and the timing of shits appears irregular. Shits do, however, clearly occur on two scales: long periods characterized by essentially the same polarity are punctuated by much shorter intervals of opposite polarity. he long periods, lasting hundreds of thousands or even millions of years, are called polarity chrons (formerly epochs). he shorter intervals, lasting no more than a few tens of thousands of years, are known as polarity subchrons (formerly events). Ancient polarity can be detected most readily in volcanic rocks and in ine-grained sediments that settled into place relatively slowly, for example, on the ocean loor. he alignment of ferromagnetic particles in a volcanic rock relects the direction of the ield when it cooled, while the alignment of particles in ine-grained sediments relects the direction

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at settling time. Subsequent compaction of the sediments prevents realignment. Polarity changes are also recorded in magnetic lineations or stripes that are ixed in oceanic crust as it spreads laterally from midocean ridges. Magnetic proiling of the oceanic crust has extended the paleomagnetic stratigraphy to 80 Ma and before. he sequence of polarity changes is especially well-known for the past 5 my. It comprises four polarity chrons and ten or more polarity subchrons (ig. 2.11). he chrons and subchrons have been dated from paired radiopotassium and paleomagnetic readings in volcanic rocks and from paired climatic and paleomagnetic determinations in deepsea sediments. he deep-sea record reveals a long sequence of cold/ warm oscillations whose ages can be calculated from the astronomical variables that probably forced them (the Milankovic cycles discussed briely below). he radiopotassium and astronomical datings agree closely where they can be cross-checked, for example, on the placement of the boundary between the Matuyama and Brunhes Chrons at 780–790 ka. he boundary dates between chrons remain subject to further empirical and statistical revision, but they are reasonably wellixed. For human evolution, the most important date is for the Brunhes/ Matuyama Boundary, for which an estimate of 780 ka is used here. he dates for subchrons are less secure because of their short duration, which may be less than the statistical uncertainty associated with radiopotassium dates. Even the existence of some subchrons remains controversial. Accurate measurement of remanent magnetism in ancient rocks or sediments requires not only proper instrumentation but also care in ield removal. Complications can be introduced by secondary heating of a volcanic rock and by postdepositional chemical processes that alter the behavior of ferromagnetic particles. Unlike either the irst appearance of a biological taxon (a species, genus, etc.), which may be time transgressive and regionally restricted, or global climatic change, whose impact may be harder to detect in some places than in others, a change in polarity afects the entire globe simultaneously, and given appropriate deposits, it should be detectable everywhere. his makes the geomagnetic stratigraphy attractive for deining boundaries between time periods, including, for example, the boundary between the Pliocene and the Pleistocene, which some specialists argued could be ixed at the top of the Olduvai Subchron, about 1.77 Ma, and the boundary between the early Quaternary and the middle Quaternary, now commonly equated with the boundary between the Matuyama Reversed Chron and the Brunhes Normal Chron at 780 ka. he geomagnetic stratigraphy is also very useful for bracketing sites in time that cannot be dated more directly. hree prominent examples

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0.1 0.3

Brunhes Normal Chron

0.5 0.7 0.9 1.1

Jaramillo Normal Subchron Cobb Mountain Normal Subchron

1.3 1.5

Matuyama Reversed Chron 1.7 1.9 2.1 2.3

Olduvai Normal Subchron Réunion I Normal Subchron Réunion II Normal Subchron

2.5 2.7 2.9

Gauss Normal Chron

3.1

Kaena Reversed Subchron

3.3

Mammoth Reversed Subchron

53 FIGURE 2.11. Global paleomagnetic stratigraphy for the past 5 my (based on Cande and Kent [1992]; mcdougall et al. [1992]; shackleton et al. [1995]; singer et al. [1999]). by convention, normal polarity (compass needle pointing north) is indicated in black, reversed polarity in white. Chrons are long intervals of normal or reversed polarity. subchrons are shorter intervals of opposite polarity within chrons. brief subchrons are dificult to detect and date, and the diagram speciically excludes geomagnetic excursions or events that may not qualify, either because their dating is problematic or because they may not represent full polarity reversals.

3.5 3.7 3.9 4.1 4.3 4.5

Gilbert Reversed Chron Cochiti Normal Subchron Nunivak Normal Subchron

4.7 4.9 5.1

Sidufjall Normal Subchron Thvera Normal Subchron

are the famous Peking man site of Zhoukoudian, northern China, and the Gran Dolina (GD) and Sima del Elefante (SE) sites, Atapuerca, Spain. Neither Zhoukoudian nor the Atapuerca sites have provided material that is ideally suited for isotopic age determination. At Zhoukoudian, paleomagnetic determinations from ine-grained parts of the ill have identiied the Brunhes/Matuyama boundary at a level below that containing the earliest Peking man fossils and artifacts. In combination with uranium-series dates on lowstones within the Zhoukoudian sequence, the paleomagnetic observations suggest that the deposits with human fossils and artifacts span the period from about 670 to 400 ka. At the Atapuerca sites, paleomagnetic analysis has identiied the Brunhes/Matuyama boundary in sediments above layers with the oldest

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artifacts and human remains. Based largely on this observation and on associated animal remains, the artifacts and fossils at GD and SE have been dated to about 800 ka and 1.1 Ma, respectively. Elsewhere in Europe, no site unequivocally demonstrates human occupation before 600–500 ka, but the Atapuerca sites imply that people were present at least sporadically before then. he Zhoukoudian and Atapuerca examples illustrate the most common application of paleomagnetism in archeology—that is, to determine whether a site formed before or ater 780 ka. Artifacts found in reversely magnetized sediments can generally be assigned to the Matuyama Reversed Chron between 2.6 Ma and 780 ka, since the oldest known artifacts are not older than 2.6 Ma. While it is theoretically possible that artifacts in reversed sediments could derive from reversed subchrons within the Brunhes Normal Chron, spanning the last 780 ka, these subchrons were probably too brief to be detected in archeological deposits. Artifacts in normally magnetized sediments are more ambiguously dated. hey could date to the Brunhes Chron or to one of the normal subchrons within the Matuyama Chron. Faunal remains (biostratigraphy) and paleomagnetic determinations from throughout a stratigraphic sequence (to detect shits between normal and reversed polarity that may represent subchrons) can help to narrow the alternatives.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: causes of paleomagnetic reversals (Cox 1969, 1972; Harland et al. 1990; Kappelman 1993); magnetic proiling of oceanic crust (Cande and Kent 1992); dating of chrons and subchrons in volcanic rocks (McDougall et al. 1992) and deep-sea sediments (Shackleton et al. 1995); agreement between radiopotassium and astronomical datings for the Brunhes/Matuyama boundary (Baksi et al. 1992; Bassinot et al. 1994; Sarna-Wojcicki et al. 2000); application of paleomagnetism to Zhoukoudian (Liu 1985; Zhou et al. 2000), to Atapuerca GD (Parés and Pérez-González 1995), and to Atapuerca SE (Carbonell et al. 2008)

Cenozoic Climates In addition to furnishing the chronological framework for human evolution, geologic studies can also illuminate the natural selective forces that drove it. Among these, none is potentially more important than climate or, more precisely, climatic change. For example, as discussed in the next chapter, the development of drier, more seasonal climates between 10 and 5 Ma (in the late Miocene) could at least partly account for the broadly contemporaneous emergence of the human tribe (Hominini or hominins) in that interval. Similarly, the factors that initiated Northern Hemisphere glaciation between 3 and 2.5 Ma (in the mid-Pliocene) could underlie the emergence of the genus Homo and the appearance of stone-tool technology. his section aims to summarize Cenozoic climatic change as it may bear on human evolution, with particular reference to the later Cenozoic Ice Age, which, in one of its milder phases, still grips the planet. he evidence documenting Cenozoic climatic change

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is incomplete and sometimes ambiguous or even contradictory, but the broad trends are clear. Only the details remain subject to revision. In the early Cenozoic, global climates were relatively warm, wet, and nonseasonal, and there was remarkably little diference between the equator and the poles. Global warmth meant that glaciers and sea ice barely existed, and relatively high, nonseasonal precipitation restricted the extent of deserts and grasslands. Forests were the dominant vegetation type and reached even into what are now ice-covered portions of the Arctic and Antarctic. World temperatures rose slightly during the Paleocene to a peak in the early Eocene, before beginning a long sporadic decline 51–50 Ma. he descent included at least ive especially abrupt downturns—in the early Oligocene roughly 34 Ma, the terminal Oligocene/early Miocene about 23 Ma, the mid-Miocene about 14.5–14 Ma, the late Miocene between 6.5 and 5 Ma, and the mid-Pliocene between 3.1 and 2.8 Ma (ig. 2.12). Cooling events were generally associated with a decline in atmospheric CO2, usually considered to be the most important of the greenhouse gases, but specialists debate whether and when CO2 variation was mainly cause or consequence. here is wide agreement that continental drit and associated tectonics played a causal role, as they redirected the oceanic currents, which transfer heat between latitudes, and promoted mountain building, which altered atmospheric circulation. Sealoor spreading and mountain building also inluenced chemical weathering rates and thus the concentration of atmospheric CO2. Key tectonic events that, ater varying lags, had an impact on climate include the northward drit of India into Asia and the subsequent uplit of the Himalayas and the Tibetan Plateau, beginning in the late Paleocene, about 55 Ma; the development of the Tasman passage between Australia and Antarctica and the Drake passage between South America and Antarctica, which together allowed the development of the Antarctic Circumpolar Current and the thermal isolation of Antarctica, beginning in the early Oligocene about 34 Ma; an acceleration in the uplit of the Tibetan Plateau in the early Miocene about 21 Ma; massive uplit of land adjacent to the east African Rit Valleys between 8 and 2 Ma; and the narrowing of the Panama seaway between South and North America, culminating in its full closure in the early Pliocene, roughly 5 Ma. Glaciers were rare or absent during the Paleocene and most of the Eocene, but small high-altitude glaciers probably formed in eastern Antarctica during the late Eocene, beginning about 36 Ma, and the sharp temperature plunge that occurred near the Eocene/Oligocene boundary, about 34 Ma, coincided with the formation of an expansive East Antarctic Ice Sheet that reached sea level. his ice sheet shrunk in the later Oligocene and earlier Miocene, but the mid-Miocene temperature drop, 14.5–14 Ma, revived it, and it has retained roughly its present dimensions ever since.

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56 FIGURE 2.12. Global temperature change over the past 70 my inferred from the 18o/16o ratios in bottom-dwelling, deep-sea foraminifera (modiied after Zachos et al. [2001], ig. 2; and billups [2005], ig. 1). The ratios are plotted as per mil deviations from a standard (δ18o), and the curve is based on adjusted observations from all the major ocean basins. larger values (to the left) imply cooler conditions with a greater likelihood for continental ice sheets. The vertical bars (right center) mark the duration of northern hemisphere and antarctic Glaciation, respectively. darker segments mark periods of especially intense glaciation. The temperature scale is for high-latitude surface water in an ice-free ocean and is realistic only for the period before about 34 ma when large-scale glaciation began in antarctica. The curve shows that cooling began about 50 ma and became particularly marked after 14–15 ma. for the entire period, the coolest temperatures, accompanied by wide luctuations between warmer and cooler intervals, occurred in the last 2 my or so.

4

∂18O (o/oo)

2

0

PlioPleistocene 10

> 50% present ice volume

millions of years ago

20

< 50% present ice volume

Miocene

Oligocene

30

Antarctic glaciation

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6

Northern Hemisphere glaciation

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40

50

60

70

Eocene

Paleocene

Cretaceous

glacial ice

0

generally ice free

4

8

12o Celsius

he cold episode at the end of the Miocene, between 6.5 and 5 Ma, was particularly acute, and it probably added a permanent West Antarctic Ice Sheet. During this cold period, the Mediterranean periodically dried up, leaving behind vast salt deposits that collectively deine the “Messinian salinity crisis.” Repeated glacier growth could have been responsible for this if it lowered sea level below the sill that separates the Atlantic and the Mediterranean, but tectonic (upward) displacement of the sill may have played the main role. In the period immediately following 5 Ma, global temperatures warmed, and the Mediterranean reilled. But in the mid-Pliocene, between about 3.2 and 2.8 Ma, temperatures plunged once more, and the northern continents were glaciated for the irst time. A sharp increase in debris ice-rated from polar glaciers to the surrounding seas about 2.8 Ma marks further temperature decline, and it coincided with the emergence of a highly conspicuous cycle in which

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glacial peaks returned on average every 41 ky. About 1 Ma, the average duration of glaciations expanded to about 100 ky, and their intensity increased sharply. Peak ice volume ater 1 Ma generally exceeded earlier volume by 50% or more. he mechanisms that drove late Cenozoic cold/warm alternation have been iercely debated. Vulcanism may have played an occasional role, and the sharp drop in global temperatures that occurred roughly 71 ka could have been inluenced by the eruption of Mount Toba, Indonesia, which occurred about the same time. he eruption released 850 cubic kilometers of ash into the atmosphere, and the sulfur aerosol it added to the stratosphere would have relected a major fraction of solar radiation back into space. he much smaller eruption of Mount Tambora, also Indonesia, in 1815, released only 20 cubic kilometers of ash. Yet, it led to a year without a summer, in which New England experienced snow in July. he Toba supereruption might have produced six similar years in succession, and plant and animal populations were surely afected. If the climate change actually extinguished many human populations, it could account for the remarkable lack of genetic diversity in people today, discussed in chapter 7. However, many other species came through without a similar loss of diversity, and artifactual continuity below and above Toba ash in southern India indicates that regional human populations survived. hus, despite the enormity of the eruption, its efects seem to have been short-lived. It may have contributed to a glacial onset, but it didn’t cause it. To explain glacial/interglacial (major cold/warm) alternations, most specialists today assign a fundamental role to astronomical variables or, perhaps more precisely, to their interaction with global oceanic and atmospheric circulation. he “astronomical theory” is commonly credited to the Serbian astronomer Milutin Milankovic, who reined it between 1915 and 1940. Milankovic suggested that glaciations began at times when Northern Hemisphere summers became relatively cool, and he noted that insolation (solar heat) reaching high latitudes in summer oscillates regularly, in correspondence with rhythmic change in three factors: the wobble of the earth’s spin axis (the precession of the equinoxes), which afects the time of year when the earth is nearest to the sun; the obliquity (tilt) of the spin axis, which alters the angle at which sunlight hits the surface; and the shape (eccentricity) of the earth’s orbit, which varies from nearly elliptical to nearly round. Insolation lows (times of reduced solar input) in response to the three key factors occur in cycles of 19/23, 41, and 100 ky, respectively. he durations of late Cenozoic glacial periods appear to match astronomical expectations, although the inluence of each cycle (or astronomical variable) has varied through time. Only the 19/23 ky (or precessional) cycle is apparent before 2.8 Ma, while the 41 ky (or obliquity) cycle dominated from roughly 2.8 Ma, when large

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ice sheets irst appeared in the Northern Hemisphere and highly conspicuous cold/warm oscillations began, until about 1 Ma. Ater 1–0.9 Ma, the 100 ky (or eccentricity) cycle became much more important, but the signal of the 41 ky cycle remained strong, especially in higher latitudes. he shit to dominance of the 100 ky cycle not only prolonged glaciations (to an average of 100 ky), it also initiated a sharp increase in their amplitude (intensity). Both the deep-sea records and the ice cores discussed below indicate that within the basic 100 ky cycle, there was a further shit about 450 ka to somewhat more intense glacial maxima and shorter but warmer periods between glacial intervals. he late Cenozoic cold periods, with their greatly enlarged ice sheets, are commonly called glacials or glaciations, and the intervening warmer periods, with their reduced ice sheets, are commonly called interglacials or interglaciations. he Holocene of igure 2.1 is simply the most recent interglacial and might better be known as the Present Interglacial. From a climatic perspective, it is not truly separate from the preceding Pleistocene, and some specialists would prefer to extend the Pleistocene to the present, equating it with the Quaternary. During the past 1–0.9 my, the cooling that characterized each glaciation occurred slowly and irregularly over 70–90 ky, whereas the terminal warming occurred much more abruptly, usually in less than 10 ky. Glaciations were generally interrupted by somewhat milder intervals or “interstadials” that alternated with periods of greater cold or “stadials.” Compared with interglacials, however, both stadials and interstadials were characterized by greatly expanded glaciers. he existence of glaciers far beyond their present limits was documented irst in Norwegian and Swiss mountain ranges in the early 1800s. Evidence for greatly expanded continental ice sheets was developed somewhat later in the nineteenth century, initially by the great Swiss zoologist Louis Agassiz. As land-based evidence for ancient glaciations accumulated in both North America and Europe, it seemed to indicate four major late Cenozoic glaciations, separated by three main interglacials (excluding the last or present one). he four main glaciations were named (from older to younger) Günz, Mindel, Riss, and Würm in a classic study by Penck and Bruckner of a small Alpine region south of Munich, Germany. Subsequently, the same glacial periods were recognized at a broad range of sites across Eurasia, and sites, fossils, artifacts, and such were commonly dated by reference to the presumed timespans of the glaciations or of the interglacials that separated them. Beginning in the mid1950s, however, studies of sediments from the deep-sea loor have shown that there were far more than four major glaciations. here were, in fact, eight glaciations during the past 780 ky alone and perhaps ity during the past 2.5 my, although those before 1–0.9 Ma were shorter and less intense. he four-glaciation Alpine scheme survives in some textbooks, but

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it should now be totally abandoned. It serves only to illustrate the risk of extending stratigraphic correlations across great distances without corroborating biostratigraphic markers or numeric dates. he reason climatic inferences drawn from deep-sea sediments are more reliable than those based on terrestrial deposits is that deposition on the deep-sea loor is more continuous, while erosion (which destroys the record) is rare. Major interruptions or discontinuities are more common in land-based sequences, especially in areas that were glaciated, since each succeeding glaciation tended to erase the evidence of its predecessors. In fact, the only land-based records that approach the deep-sea loor for continuity over long intervals come from ice drilled out of permanent glaciers in Greenland and Antarctica and from areas beyond the reach of the glaciers. Such areas include central Europe and central China, with their deep covers of wind-blown dust (loess), and the lake bottoms of Lake Bogotá in the Colombian Andes, Owens Lake in California, Grand Pile in northwestern France, Lake Phillipi in northern Greece, Lake Biwa in Japan, and Lake Baikal in south-central Siberia. For the periods these unusually long land-based records cover, they corroborate and expand the basic history of late Cenozoic climatic change now standardized on observations from the deep-sea loor. Physiochemical analysis of air bubbles trapped in the Greenland ice sheet, for example, has revealed that the North Atlantic region experienced more than twenty-four abrupt temperature swings between 100 and 15 ka, including a closely packed series of twelve-to-iteen between about 60 and 25 ka. hese swings had an impact on global ice volume, and they must have had dramatic effects on plants and animals, including humans. Each swing comprised a relatively warm peak, called a Greenland Interstadial (or DansgaardOeschger event ater the discoverers), when the temperature in Greenland rose by 8°–160°C. he intervening cold excursions are known as Greenland Stadials. In some cases, the shit from stadial to interstadial appears to have occurred within a few decades. he return to stadial conditions was generally less abrupt, but the stadials grew colder and more frequent with time, and some archaeologists have suggested that their cascade contributed to the extinction of the Neanderthals between 45 and 30 ka. he long deep-sea record for late Cenozoic climatic change has been established in sediment cores removed by sophisticated equipment that barely disturbs the sediment structure. Climatic change is inferred mainly from the species of foraminifera and other sea-dwelling microorganisms represented in diferent parts of a core and also from the changing chemical content of their shells, also known as tests. Variation in the content of oxygen-16 (16O) and oxygen-18 (18O) is particularly informative. hese are naturally occurring, stable (nonradioactive) varieties (isotopes) of oxygen, which foraminifera and other marine organisms extract from seawater and build into their tests. he 18O is heavier, and

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water molecules with 18O evaporate less readily than those with 16O. he molecules return to earth as rain or snow, but during glacial intervals, a signiicant proportion become permanently locked in the expanded ice sheets and are not returned to the oceans. he result is that seawater becomes richer in molecules with heavier 18O and poorer in molecules with lighter 16O. he 18O/16O ratio in microorganismal tests partly depends on the temperature and salinity of the water in which the microorganisms lived, but it mainly relects the original ratio in the water, and changes in the oxygen composition of tests from deep-sea cores therefore track the episodic growth and decline of the ice sheets. he climatic stages detected from oxygen-isotope analysis are designated by arabic numerals, with odd numbers for the interglacial stages and even numbers for the glacial ones (ig. 2.13). For the past 2.5–2.6 my when glacial/interglacial alternation was especially marked, specialists recognize a hundred stages, each distinguished by its stage number preceded by either OIS, for oxygen-isotope stage, or by MIS, for marine isotope stage. hus the Holocene or Present Interglacial is known as OIS 1. here is the complication, however, that a long, generally milder period within the Last Glaciation has been designated OIS 3 and that the Last Glaciation thus includes OIS 4 through OIS 2. OIS 5 corresponds to the Last Interglacial, though only its earliest substage, known as 5e (or 5.5), was comparable to OIS 1 in terms of ice-sheet reduction and global warmth (ig. 2.13). he later substages, 5d–5a, relect varying but generally cooler climate, and some authorities prefer to assign them to the early part of the Last Glaciation. Almost certainly, OIS 5d–5a correspond to some terrestrial deposits that have long been assigned to the early part of the Last Glaciation, for example, to the Würm I in France. he oxygen-isotope stages have been dated by a variety of methods, including 14C used near the top of deep-sea cores and a combination of paleomagnetism and extrapolated sedimentation rates used farther down. One important conclusion is that from roughly 900 ka to 430 ka, when the 100 ky cycle irst dominated glacial/interglacial alternation, interglacial periods tended to be relatively long but cool compared to OIS 5e (the Last Interglacial narrowly understood) or OIS 1 (the Holocene). Ater 430 ka, the interglacials became warmer, but shorter, and intervals as warm as OIS 1 rarely lasted more than 6,000 years. he Antarctic ice-core records corroborate this. In terms of both length and relative warmth, OIS 1 has only one obvious analog, OIS 11, between about 425 and 395 ka, but even considering OIS 11, the deep-sea and ice-core records show since 800–900 ka, people lived more than 95% of the time under cool to cold glacial conditions (ig. 2.13). he most obvious aspect of glaciations was the mushrooming of ice on the continents. At their maximum, the ice sheets incorporated 50 million cubic kilometers of water, and they covered nearly a third of

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(∂18O -o/oo) ka

2

1

0

-1

-2

-3

Isotope Stages 12 3 4

5a 100

5c

ka 11 24 57 71

5 5e

127 6 186

200

7

Brunhes Normal Chron

242 8 300

9 10

301 334 364

11

400

427 12 474 13

500

61 FIGURE 2.13. The δ oxygen-18 (δ18o) record for the past 900,000 years, based on deep-sea sediment cores md900963 in the tropical indian ocean and 677 in the equatorial Paciic ocean (modiied after bassinot et al. [1994], 103, 106). odd-numbered isotope stages indicate relatively warm periods, while even-numbered stages indicate colder ones. The boundary dates have been estimated primarily on the assumption that mathematically predictable variation in the relation between the earth and the sun forced the cold/warm oscillations.

528 14 568 15

600

621 16 659 17

700

712 18 19 20

800

warm

21

cold

22

760 787 806 865

900

the earth’s surface, mostly in the northern hemisphere (ig. 2.14). In partial compensation, however, sea level dropped by 130–160 m, and fresh land was exposed on the continental shelves. he emergent shelves not only became available for occupation but, in some cases, they connected previously isolated landmasses, allowing people and other animals to inhabit new areas. For example, it was the lowering of sea level during a glaciation between 2 and 1 Ma that allowed early humans (hominins) to occupy Java, and the broad land bridge that connected Siberia and Alaska during the Last Glaciation permitted or at least promoted the initial human colonization of the Americas. he impact of glacial climate, particularly ater 1 Ma, was worldwide. From a human point of view, one of the most important efects was a large-scale redistribution of plants and animals. Tundra and steppe (semiarid grassland) replaced forest in midlatitude Eurasia, and arctic animals such as reindeer penetrated far south of their interglacial range.

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ChaP Ter TW o

Camp Century Ice Core ■

NGRIP, GISP2 & ■ GRIP Ice Cores ■■ Deep-Sea Core ■ V28-14 Grand Pile

Clear Lake ■

■ Deep-Sea Core V23-82

■ ■

■

■

Deep-Sea Core MD952042 ■

L. Bonneville

■

Central European loess L. Phillipi Tadzhik loess ■

Owens Lake ■ Hawaii ODP Core 677

ODP Core 849 ■

■

■

■ Chinese loess

Deep-Sea Core V16-205 ■ ■ Barbados

■ Bogotá

Pleistocene ice sheets

■ L. Baikal

■ Deep-Sea Core MD900963 L. Malawi ■

■ L. Biwa Deep-Sea cores V28-238 and V28-239 Huon Peninsula ■ ■

main areas exposed by lower sea level major localities that document Pleistocene climatic change

■■

Lynchʼs Crater

L. George ■ ■ ODP Core 1119 0o

Ice-core localties in Antarctica

Dome Fuji -90

o

Vostok Dome C

90o

180o

FIGURE 2.14. Top: The maximum extent of Quaternary glaciation and associated sea-level change (modiied after roberts [1984], ig. 2.1). The named localities are places that have provided particularly long and informative records of Quaternary climatic change. The selection of deep-sea cores is arbitrary since there are now scores that largely corroborate one another. Bottom: antarctica with the main ice-core localities (modiied after Watanabe et al. [2003], ig. 1). The localities providing the most informative cores are named. one of the dome C cores is especially remarkable for the time it spans, chronicling eight consecutive glacial cycles over the past 740 ky (ePiCa Community members 2004). in all important respects, the dome C record conirms the comparable sections of the even longer records from the deep seabed, but it adds important detail on past atmospheric composition from air bubbles trapped in the ice.

Mean annual temperatures declined everywhere, by as much as 16°C in higher latitudes and perhaps as much as 3°C near the equator. In most places, precipitation also declined, probably mainly because less moisture evaporated from colder oceans. he efects of reduced precipitation are particularly obvious at lower latitudes (between roughly 30° S and 30° N), where large areas that are now rainforest became grassland or savanna, while grassland and savanna regions turned to desert. he reduced vegetation cover fostered wind erosion (delation) at low latitudes, and some of the wind-borne dust accumulated in deep-sea cores adjacent to the continents. Analysis of the cores shows that dust volume increased in step with ice volume ater 2.8 Ma, and like ice volume, it

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oscillated rhythmically in a 41 ky cycle between 2.8 and 1 Ma and in a 100 ky cycle aterward. Antarctic ice cores conirm that glacial intervals were extremely dusty because they were windier and more arid. he ice cores add a wrinkle because they contain trapped air bubbles, which show that periods of lowered temperature and greater dustiness were also times when the atmosphere was depleted in the greenhouse gases, carbon dioxide and methane. Greenhouse gas concentrations were much higher during warm interglacial periods, implying that they ampliied the effects of orbital forcing on glacial/interglacial alternations. In the part of the Holocene that preceded the Industrial Revolution, greenhouse gas levels were similar to those in other interglaciations over the past 420 ky, but modern levels are the highest on record because they are due in large part to human activity. he close correlation between glacial climate and signiicantly greater aridity falsiies an ill-founded earlier idea that wetter conditions (pluvials) characterized lower latitudes while glaciations afected higher ones. In fact, pluvial conditions were mainly an aspect of interglacials, though lakes did grow during glaciations in some midlatitude areas because of altered atmospheric circulation, lowered evaporation, or both. he recurrent and persistent aridity of the tropics during glaciations probably afected human populations as much as or more than increased cold did in higher latitudes, and the natural selection associated with stepwise reductions in humidity (and in the overall availability of surface water) may explain some of the important evolutionary events discussed in later chapters. hese events include the origin of the hominins between 7 and 5 Ma, the divergence of the genus Homo and of the robust australopiths roughly 2.6–2.5 Ma, the emergence of irst truly human species, Homo ergaster, 1.8–1.7 Ma, the extinction of the robust australopiths 1–0.9 Ma, and perhaps even the origin and dispersal of fully modern people about 50 ka.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: global climate change and human evolution (Feibel 1997); long-term trends in Cenozoic climate (Billups 2005; Denton 1999; Miller et al. 1987; Miller and Fairbanks 1985; Zachos et al. 2001); punctuated decline in Cenozoic temperatures (Kennett 1995); relation of Cenozoic climate change to CO2 variation (DeConto and Pollard 2003; Zachos et al. 2001) and continental drit (Laporte and Zihlman 1983); sealoor spreading, mountain building and the concentration of atmospheric CO2 (Denton 1999); tectonic events that promoted global climate change (Zachos et al. 2001) and aridiication in eastern Africa (Sepulcre et al. 2006); explanations for the Messinian salinity crisis (Denton 1999; Hodell et al. 1994); increase in ice-rated debris about 2.8 Ma (Jansen and Sjoeholm 1991) and the emergence of 41 ky glacial cycle (Tiedemann et al. 1994); increase in glacial duration and intensity about 1 Ma (Shackleton 1995); Mount Toba supereruption—impact on human populations signiicant (Ambrose 1998b, 2003; Rampino and Ambrose 2000) or limited (Gathorne-Hardy and HarcourtSmith 2003; Oppenheimer 2002); artifactual continuity in southern India before and ater the Toba eruption (Petraglia et al. 2007); astronomical explanation for glacial/interglacial alternation (Berger 1992; Paillard 1998); relation between times of reduced solar input and glacial cycles (Hays et al. 1976; Imbrie et al. 1984, 1993; Imbrie and Imbrie 1979; Shackleton et al. 1990; Zachos et al. 2001); shit in glacial intensity 450 ka (McManus 2004); deinition of the Pleistocene and early demonstration of expanded glaciers (Lowe and Walker 1997); initial deinition of the Alpine 4-glacial scheme (Penck

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and Bruckner 1909) and deep-sea evidence refuting it (Roebroeks and van Kolfschoten 1994; Shackleton 1995); ice-core records from Greenland (Dansgaard et al. 1993; Johnsen et al. 1972, 1992) and Antarctica (EPICA Community Members 2004, 2006; Jouzel et al. 1987, 1989, 1993, 2007; Petit et al. 1999; Watanabe et al. 2003); loess records in central Europe and central China (Kukla 1975, 1987; Kukla et al. 1988, 1990); climate records from Lake Bogotá (Hooghiemstra 1995; van der Hammen 1974), Owens Lake (Bischof et al. 1997b), Grand Pile (Beaulieu and Reille 1992; Guiot et al. 1989; Seret et al. 1992; Woillard 1978; Woillard and Mook 1982), Lake Philippi (van der Hammen et al. 1971; Wijmstra and van der Hammen 1974), Lake Malawi (Scholz et al. 2007), Lake Biwa (Fuji 1988; Kashiwaya et al. 1988; Meyers et al. 1993), and Lake Baikal (Williams et al. 1997); Greenland stadials and interstadials (Andersen and Members 2004), their probable efect on plants and animals (van Andel 2002), their abruptness (Cufey 2004), stadial intensiication with time (Arnold et al. 2002), and possible efect on the Neanderthals (Finlayson 2004; Tzedakis et al. 2007); deep-sea oxygen-isotope record for glacial/interglacial alternation (Shackleton 1967, 1975, 1987); numbering of oxygen-isotopic stages (Bassinot et al. 1994; Emiliani 1955, 1969; Shackleton 1975; Shackleton and Opdyke 1973); terrestrial deposits preferably assigned to the Last Interglacial vs. the early part of the Last Glaciation (Butzer 1986); duration and warmth of interglacials ater 800–900 ka (White 2004); biotic impact of global climate (Street 1980); dust and glacial cycles—in deep-sea cores (deMenocal 1995, 2004; deMenocal and Bloemendal 1995) and Antarctic ice cores (EPICA Community Members 2004; McManus 2004); greenhouse gas concentrations in the past 420 ky (Petit et al. 1999); glaciations and global humidity (Butzer 1978a); later Cenozoic moisture history and human evolution in Africa (Trauth et al. 2005)

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The PrImaTe BaCKGrOunD

3

he zoological order Primates includes about 230 living species classiied informally as lemurs, lorises, tarsiers, monkeys, apes, and humans. Many additional primate species are known only in fossil form, and many more probably await discovery. Knowledge of living and fossil nonhuman primates is crucial to an understanding of human evolution, and this chapter aims to provide the necessary background. It emphasizes, irst, the natural history of those living primates—particularly the great apes—that are most closely related to people and, second, the basic course of primate evolution. In general, it does not attempt to trace speciic ancestor-descendant relationships because, ironically, the growth of the fossil record has increased the number of plausible alternatives. In addition, the record now oten provides more potential ancestors than descendant species require. his is particularly true when the focus is on the ancestry of the living great apes and people. However, the core obstacle is that evolutionary relationships must be reconstructed from similarities among species, and specialists are more aware than ever that similarities can be ambiguous, even misleading, guides. It will be recalled from the opening chapter that the key similarities are ones that indicate a shared common ancestor, but to begin with, these can be diicult to distinguish from similarities that result from adaptation to shared circumstances (analogy) or from parallel evolution (convergence or homoplasy) in distantly related lines. Equally important, most authorities now recognize that similarities due to common descent must be divided between “primitive” ones that developed early in the history of a species or species group and “advanced” or “derived” ones that developed much later. Only shared derived features indicate a closely shared origin or a possibly close ancestor-descendant relationship between time-successive species. Yet, as discussed in this and later 65

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chapters, shared derived characters are oten hard to separate from shared primitive ones or even from shared characters that two species evolved independently as they adapted to similar environments. In these circumstances, a compelling ancestor-descendant relationship is far harder to establish than to disprove. In general, for disproof we need show only that the supposed descendant lacks derived or advanced features that occurred in its putative ancestor. he remainder of this chapter consists of three parts: a brief introduction to the skeleton, to deine basic anatomical terms that are relevant here and in later chapters; a longer discussion of the living primates emphasizing those aspects that are essential for understanding primate (including human) evolution; and inally, a brief summary of the nonhuman primate fossil record organized roughly by time, from the initial appearance of the primates 80–65 Ma (million years ago) to the emergence of the human line between 7 and 5 Ma.

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The Skeleton For descriptive purposes, the skeleton is conventionally divided into two main segments: the cranium (skull) and the postcranium (trunk and limbs). he cranium itself is oten divided into three principal sections: the braincase (the vault or neurocranium); the face (including the upper jaw or maxilla), which is attached to the braincase by bone; and the lower jaw or mandible, which is attached to the braincase and face only by sot tissue. he braincase and face are formed of discrete bones that grow separately and that fuse together only in adults, if at all. Figure 3.1 illustrates a cranium of the 2.5 million-year-old human relative, Australopithecus africanus, and names the main cranial elements. Teeth igure prominently in fossil studies because they are the most durable skeletal parts and they tend to dominate fossil samples. heir form also usually relects the dietary adaptation of a species, and since much evolution has involved dietary specialization and divergence, teeth play a key role in reconstructing evolutionary trends and relationships. Like other mammals, primates have four basic types of permanent teeth—known, from fore to rear, as incisors, canines, premolars, and molars (abbreviated as I, C, P, and M). he juvenile or deciduous dentition contains only incisors, canines, and premolars (abbreviated as dI, dC, and dP). he irst mammals had three permanent incisors, one canine, four premolars, and three molars on each side of each jaw. he sum is known as the dental formula, abbreviated as 3-1-4-3. By convention, from fore to rear, the incisors are designated I1, I2, and I3, the premolars P1, P2, P3, and P4, and the molars M1, M2, and M3. Superscripts identify upper

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The Pr im aTe b aCKGr ound

bregma

parietal bone

sagittal suture squamosal suture

coronal suture

lambdoidal suture

frontal bone lambda supraorbital region

67 FIGURE 3.1. reconstructed skull of Australopithecus africanus (sts 5, or “mrs. Ples”), showing the principal anatomical parts or regions (redrawn after le Gros Clark [1964], 130).

occipital plane of the occipital bone opisthocranion

maxilla temporal bone

premaxilla

nuchal plane of the occipital bone

zygomatic arch

ascending ramus of the mandible

mental symphysis

horizontal ramus of the mandible

Australopithecus africanus ("Mrs. Ples") teeth (e.g., I1) and subscripts identify lower ones (e.g., I1). In the course of evolution, most living mammals have reduced the number of teeth in one or more categories, but the remaining teeth retain the numbers of their primitive mammalian antecedents or homologues. Living primates, for example, have at least one less incisor and one less premolar than their remote ancestors, while people and their closest living relatives have two fewer premolars. he human dental formula is thus 21-2-3. he lost incisor is I3, while the lost premolars are P1 and P2. he remaining incisors on each side of an adult human jaw are thus labeled I1 and I2 and the premolars are called P3 and P4. Figure 3.2 illustrates the adult dentitions of some representative primates, with the abbreviations for individual teeth. he description of teeth oten requires an indication of their separate components or orientation, and for this purpose four terms are in common use: buccal, to indicate the portion of a tooth nearer the cheek (it is replaced by labial for teeth that abut the lips); lingual, for the portion nearer the tongue; mesial, for the portion nearer the front of the mouth (or nearer the midline depending on the tooth); and distal, for the portion nearer the rear of the mouth (or farther from the midline). At fossil sites, postcranial bones are usually less common than teeth because they tend to be soter and more fragile. hey are oten at least

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ChaP Ter Thr ee

tree shrew

lemur

I1 I2

I1

I2

I3 C P2 P3 P4 M1 M2

I2 C P2 P3

P4 M1 M2

I1

I1 I2 I3

C

M3

I2 C

C

P2 P3 P4 M1

M3

tarsier

M2

M2

I2

I1

M3

1 I2 I

C C

C P2 P3 P4 M1 M2 M3

P3 P4 M1 M2 M3

C P2 P3 P4 M1 M2

M2

M3

M3

chimpanzee

I1 I2

I2 C

C

P3 P4 M1 M2

P3 P4

M3

I1

M1 M2 M3

I1

I1 I2 C P3 P4 M1 M2 M3

equally informative, however, revealing behavioral aspects, such as locomotor habits, that are much harder to detect from teeth or skulls. Since much evolution has involved specialization and divergence in locomotor modes and other postcranial behaviors, postcranial bones supplement and complement teeth and skulls for the reconstruction of broad evolutionary patterns and relationships. All mammals have fundamentally similar postcranial skeletons, inherited from their common ancestor, and primates are distinguished mainly by a tendency to retain speciic elements that many other mammals have lost during their evolution. Figure 3.3 illustrates the skeleton of the 3.3 million-year-old human relative, Australopithecus afarensis, and names the main skeletal parts. Like teeth, individual postcranial bones can be described with respect to their orientation in a complete skeleton. he common terms are: proximal, for the portion of a bone closer to the skull; distal, for the portion farther from the skull; medial, for the portion closer to the midline of the body; lateral, for the portion further from the midline; anterior or ventral, for the portion closer to the front of the body; and posterior or dorsal, for the portion closer to the back. hese terms can be combined, as, for example, in “anterior distal humerus,” meaning the elbow end of the humerus as viewed from the front. Where context re-

FIGURE 3.2. right upper and lower dentitions of various primates (in each case, the upper dentition is to the left and the lower is to the right) (redrawn after schultz [1969], 102). The primitive mammalian dentition is thought to have comprised, on each side of each jaw, three incisors, one canine, four premolars, and three molars. in the course of evolution, all living primates have lost the irst premolar (P1), while all catarrhine primates have also lost the second (P2). both catarrhines and platyrrhines have lost the third incisor (i3), and the platyrrhines have either lost the third molar (m3) or retain it in reduced form, as in the capuchin monkey whose dentition is illustrated here.

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macaque monkey (cercopithecid)

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69 FIGURE 3.3. The reconstructed skeleton of Australopithecus afarensis, showing the names of the main skeletal elements.

clavicle scapula sternum

thoracic vertebrae humerus ribs ulna

pelvis lumbar vertebrae carpals

radius

sacrum

metacarpals manual phalanges

femur

patella fibula tibia calcaneum, talus & other tarsals metatarsals pedal phalanges

quires, other anatomical parts and terms are introduced and illustrated below.

Definition of the Primates he eighteenth-century inventor of modern biological classiication, Carolus Linnaeus, irst deined the Primates, and he chose the name to indicate their natural primacy, since they included people. Linnaeus’s original deinition has long since been discarded, but no universally accepted replacement has emerged. he problem is that, compared with the Cetacea (whales and dolphins), Rodentia (rodents), Carnivora (carnivores), Artiodactyla (even-toed ungulates), Perissodactyla (odd-toed ungulates), Proboscidea (elephants), and most of the other eighteen living

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orders of mammals, the Primates are diicult to characterize based on uniquely derived, shared traits. In distinction from other mammals, however, the living primates generally possess the following features:

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A body structure preserving the clavicle (collarbone), pentadactyly (ive digits on the end of each limb), and other primitive characters that have commonly been lost or modiied in other mammalian orders. Grasping extremities (hands and feet) with highly mobile digits, including a divergent hallux (big toe) and usually also an opposable pollex (thumb). Flattened nails replacing primitive mammalian sharp, bilaterally compressed claws on the hallux and usually also on other digits, associated with highly sensitive tactile pads on the tips of the digits opposite the nails. Besides enhancing the sense of touch, the pads bear friction ridges (dermatoglyphs) that facilitate grasping. Although nails themselves are not preserved in the fossil record, their presence is relected in characteristic (dorsoventral) lattening of the underlying terminal phalanges. Orbits (eye sockets) that tend to converge, that is, to be closely spaced and to face in the same direction, producing substantial overlap between the ields of vision and thus a high degree of stereoscopic, three-dimensional vision (depth perception). Also, in primates, unlike many other mammals, the orbits are completely surrounded by a bony ring (the post- or circumorbital bar), supplemented in higher primates by a bony wall (the postorbital plate or septum) separating the orbits from the skull behind. he high degree of stereoscopic vision relected in the orbits is associated with a unique neural apparatus for processing visual signals and with enlargement of the visual centers in the occipital and temporal lobes of the brain. A shortened muzzle or snout compared with that in most other mammals, generally associated with a more limited olfactory sense (sense of smell) and with a tendency for the olfactory bulbs in the brain to be relatively small. A fully bony auditory bulla or middle ear in which the ventral loor is composed either of an extension of the petrosal bone that encloses the inner ear or of a combination of the petrosal bone and the ectotympanic bone or tympanic ring (the bone across which the eardrum is stretched [ig. 3.4]). In other mammals, the bulla is generally loored by an independent entotympanic bone, which has been lost in primates. Reduction in the number of incisors and premolars compared with those in the earliest mammals and many living ones, combined with a relatively simple and primitive cusp pattern on the molars.

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cochlea and semicircular canals

tube of outer ear (auditory meatus)

petrosal bone tympanic ring middle ear cavity

8.

auditory bulla

A unique sulcal (issure) pattern on the surface of the cerebral cortex of the brain. Also, relative to body size, primates tend to have larger brains than other mammals.

SOURCES: diiculty of deining the Primates vs. other orders (Simpson 1955; Wible and Covert 1987); features that characterize the Primates (Andrews 1988; Cartmill 1975; Cartmill 1992; Le Gros Clark 1960; Martin 1986; Martin 1990)

Classification of the Primates Chapter 1 introduced the principles of biological classiication and noted that Carolus Linnaeus devised the modern scheme in the mid-eighteenth century. It is hierarchical, meaning that it comprises a series of levels and that units at any given level are combined to produce the units in the level just above. he species is the most basic unit, and it is commonly deined today as a group of organisms that look more or less alike and that can interbreed to produce fertile ofspring. Groups of related species are combined into genera (singular genus), genera into tribes, tribes into families, families into superfamilies, superfamilies into infraorders, infraorders into suborders, suborders into semiorders, semiorders into orders, and so forth. A species or group of related species at any level in the hierarchy is known as a taxon (plural taxa). Individual species and genera—for example, the species Homo sapiens and the genus Homo—are known as lower taxa. Groups of related species above the genus level— for example, the tribe Hominini, the family Hominidae, the superfamily Hominoidea, the infraorder Catarrhini, the suborder Anthropoidea, and the semiorder Haplorhini—are known as higher taxa. he rules for assigning a group of species to a level within the hierarchy are known as taxonomy, and the practitioners are taxonomists, although their interests commonly extend beyond taxonomy to ecology, physiology, and so forth. In modern biological classiication, the assignment of species to genera and higher taxa is meant to relect their evolutionary relationships,

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71 FIGURE 3.4. diagrammatic section through the right ear region of a therian mammal, seen from the front (redrawn after Cartmill [1975], ig. 3). in most nonprimates the auditory bulla either is cartilaginous or is formed by a separate entotympanic bone. in primates, it is formed by an extension of the petrosal bone. The primate character of the bulla has been used to assign the Paleocene group known as plesiadapiforms to the Primates, but it is conceivable that the loor of the plesiadapiform bulla was formed by an entotympanic that fused seamlessly with the petrosal. This occurs in some nonprimates today. The true character of the plesiadapiform bulla could be determined only from fetuses, which are lacking in the fossil record.

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such that species placed in the same higher-level taxon share a closer (more recent) common ancestor than species placed in diferent taxa at the same level. hus, when taxonomists include modern people, Homo sapiens, and the great apes, Pan troglodytes, Pan paniscus, Gorilla gorilla, and Pongo pygmaeus in the family Hominidae and they place the whitehanded gibbon, Hylobates lar, in the family Hylobatidae, the implication is that modern humans and the great apes share a more recent common ancestor with each other than either does with the gibbon. Similarly, when the Hominidae and the Hylobatidae are grouped in the superfamily Hominoidea, the assumption is that people, the great apes, and gibbons (broadly understood) are more closely related to each another than they are, for example, to the Old World (African and Eurasian) monkeys, which are usually placed in their own superfamily, the Cercopithecoidea. At a higher level yet, when the Hominoidea and the Cercopithecoidea are included in the infraorder Catarrhini, the implication is that they are more closely related to each other than they are to the New World (American) monkeys in the infraorder Platyrrhini. he system proceeds logically upward, so that all species included in the order Primates are presumed to share a more recent common ancestor with each other than they do with species in the eighteen remaining orders of living placental mammals. Until the 1960s, zoological classiication in general and primate classiication in particular depended mainly on anatomical comparisons among living species. he basic assumption was that the number of shared anatomical characters between two species relected their degree of evolutionary relationship. he more characters they shared, the more likely it was that they would be assigned to the same lower-level taxon. he practice continues, but since the 1960s, biomolecular, now mainly genetic, observations have played an every increasing role in taxonomic decisions. Simultaneously, new fossil discoveries and fresh analyses have shown that many fossil species cannot be readily accommodated in the same higher taxa as living species. As a result of recent genetic and fossil research, the long-standing traditional classiication of the Primates has been discarded. No new consensus has been reached, but for heuristic purposes most authorities would probably accept the breakdown into semiorders, suborders, infraorders, superfamilies, and families, presented in table 3.1. Unlike the traditional classiication, this one depends heavily on genetic observations, though it also involves anatomical characters, and it uses these to classify some fossil forms for which genetic data will probably never be available. It abandons the previous division of the Primates between two suborders—the Anthropoidea (“higher primates” or simians) and Prosimii (“lower primates” or prosimians) in favor of a higher-level division between two semiorders—the Haplorhini (simple- or dry-nosed

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TABLE 3.1. A provisional classiication of the Primates to the level of the suborder and to the level of the family within the suborder Anthropoidea (Modiied ater (Goodman et al. 1998, p. 594)). ◊‘s indicate extinct taxa.

Order Primates Plesion:◊Omomyiformes (Eocene tarsier-like species) Semiorder: Haplorhini Suborder: Anthropoidea (“higher primates”) Infraorder: Catarrhini (Old World higher primates) Plesion:◊Propliopithecoidea (late Eocene and early Oligocene catarrhines) Superfamily: Cercopithecoidea (Old World monkeys) Plesion:◊Victoriapithecidae (early and middle Miocene cercopithecoids) Family: Cercopithecidae Superfamily: Hominoidea (apes and people) Plesion:◊Pronconsulidae (early Miocene hominoids) Family: Hylobatidae (gibbons and siamangs) Family: Hominidae (great apes and people) Infraorder: Platyrrhini (New World higher primates) Superfamily: Ceboidea (New World monkeys) Plesion:◊Homunculidae (early Miocene platyrrhines) Family: Cebidae (capuchin monkeys, squirrel monkeys, marmosets tamarins) Plesion:◊Tremacebidae (late Oligocene platyrrhines) Family: Pitheciidae (sakis, titis, and uakaris) Family: Atelidae (spider, howler, and woolly monkeys) Suborder: Tarsiiformes (tarsiers) Plesion:◊Adapiformes (Eocene lemur-like species) Semiorder: Strepsirrhini Suborder: Lemuriformes (lemurs) Suborder: Lorisiformes (lorises and galagos)

primates) and the Strepsirhini (moist-nosed primates). In the previous classiication, the Prosimii included the Tarsiiformes (tarsiers), although their assignment to the Prosimii was always questionable because they share derived characters with the Anthropoidea. he speciic characters are outlined below. Genetics now conirms that the Tarsiiformes are more closely related to the Anthropoidea than to other supposed Prosimii, and to accommodate this inding, table 3.1 lists them as a suborder, alongside the Anthropoidea, within the Haplorhini. he suborder Prosimii disappears, and its two remaining former members—the Lemuriformes and Lorisiformes (or Loriformes)—are elevated to suborders within the semiorder Strepsirhini. he classiication in table 3.1 has the desirable characteristic that to the extent current knowledge allows, the taxa at each level in the hierarchy are all monophyletic, that is, each one comprises all the known descendants of a single common ancestor and the shared ancestor would also be included, assuming it were known. To accommodate fossil species that cannot be readily assigned to extant taxa and that may or may not include the ancestors of any extant form, table 3.1 employs the concept of the plesion—a group of species whose hypothesized rank and relationships are indicated by its indentation and position in the list. hus, the indentation and position of the

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plesion Omomyiformes, a group of Eocene tarsier-like species, is meant to suggest that they could represent a Primate semiorder and that this semiorder is more closely related to the extant Haplorhini than to the Strepsirhini. he position and indentation of the plesion Adapiformes, a group of Eocene lemur-like species, implies that they could also be a semiorder, though this semiorder would be most closely related to the living Strepsirhini. With regard to other plesions that are especially important to this book, the indentation and position of the Propliopithecoidea, a late Eocene/early Oligocene group, is meant to suggest they are an extinct catarrhine superfamily, and the indentation and position of the Proconsulidae—an early Miocene group—suggests they represent an extinct family within the hominoids. he classiication in table 3.1 resolves the issue of the tarsiers, but as presented, it leaves untouched a problem that is more central to this book—the division of the Hominoidea into lower taxa. Traditionally, specialists have recognized three hominoid families—the Hominidae, for people; the Pongidae, for the “great apes”; and the Hylobatidae, for the “lesser apes” (gibbons and siamangs). However, genetic analyses demonstrate conclusively that the living African great apes—the gorilla (Gorilla) and the chimpanzees (Pan)—are actually more closely related to people than either is to the living Asian great ape, the orangutan (Pongo). In technical terms, this means that the family Pongidae is polyphyletic— it includes members of two distinct evolutionary branches or clades, or if the emphasis is on its exclusion of people, it is paraphlyetic—it includes some but not all members of a single, deeper evolutionary clade. In either case, it may be a useful grade, meaning a group of species that are broadly similar in their level of ecological and behavioral organization, but it is not a valid taxonomic category. A minimal remedy that satisies basic taxonomic practice is to eliminate the Pongidae and to lump the great apes with people in the Hominidae. he principal objection to such a scheme is that it may impede communication, since generations of specialists and laypeople have become accustomed to using Hominidae (and its anglicized form, hominid) exclusively for people, living and extinct. he previous editions of this book accepted this rationale and retained the traditional usage. However, genetic and fossil evidence against it has continued to mount, and in deference to the evolutionary principles that are supposed to dictate classiication, this book adopts the scheme in table 3.2. It accepts that people share a particularly recent common ancestor with the chimpanzees, it reduces people from a family to a tribe, and it requires that they now be referred to formally as Hominini and informally as hominins. It represents a compromise between the long-standing traditional scheme and a more radical revision that would follow if all primate taxa at a

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TABLE 3.2. Subdivision of the Hominoidea, with special reference to the Hominidae (ater (Wood & Richmond 2000, p. 21)).

Superfamily: Hominoidea (“hominoids” = people and apes) Family: Hylobatidae (“hylobatids” = gibbons and siamangs) Genus Hylobates (gibbons and siamangs) Family: Hominidae (“hominids” = people and great apes) Subfamily: Ponginae (“pongines”) Genus: Pongo (orangutan) Subfamily: Gorillinae (“gorillines”) Genus: Gorilla (gorilla) Subfamily: Homininae (“hominines”) Tribe: Panini (“panines”) Genus: Pan (chimpanzees) Tribe: Hominini (“hominins” = people broadly understood) Subtribe: Australopithecina (“australopiths,” all extinct) Genus: Ardipithecus Genus: Australopithecus Genus: Paranthropus Genus: Kenyanthropus Subtribe Hominina (“hominans,” including living humans) Genus: Homo

given rank had to represent branches that were about equally old. In this instance, chimpanzees and living people might both have to be placed in the genus Homo, and Homo would also have to include all chimpanzee and human ancestors since their lines diverged. he result would not only confuse long-time observers, it would vastly complicate descriptions of human evolution like the one advanced in this book. SOURCES: classiication of the Primates (Goodman et al. 1998); biomolecular observations that link chimpanzees and the gorilla more closely to people than to the orangutan (Bailey et al. 1991; Goodman et al. 1990; Miyamoto and Goodman 1990; Pilbeam and Young 2004); subdivision of the Hominoidea (Wood and Richmond 2000, 21)

The Living Primates Primate (including human) evolution can be understood only by reference to the living primates, and any overview of fossil forms must start with a survey of living ones. he following brief survey is designed only to provide essential background information. It begins with those taxa that are most closely related to people and proceeds downward through table 3.1 to progressively more distant relatives, which it treats in progressively less detail. It is based mainly on information in the general references listed below. It should be read in conjunction with igure 3.5, which illustrates the geographic distribution of various living nonhuman primates. he igure shows that (excepting people) surviving primates are restricted almost entirely to the tropics and subtropics. his is particularly true of the most primitive forms (the Tarsiiformes and the

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100o

80o

60o

40o

20oW

0o

40o

20oE

60o

80o

100o

120o

140o

Cercopithecoidea 40o

40o

Cercopithecoidea Pan && Cercopithecoidea

20oN

Tarsioidea 20oN

Lorisoidea

0o

Ceboidea 20oS

, Pan & Gorilla, Cercopithecoidea

40o

0o

Hylobates & & Cercopithecoidea

Lorisoidea

20oS

Pongo, Hylobates && Cercopithecoidea Lemuroidea

40o

Cercopithecoidea Approximate distributions of the living nonhuman primates 120o

100o

80o

60o

40o

20oW

0o

20oE

40o

60o

80o

100o

120o

140o

Strepsirhini), and the sum suggests that primates originated under mild climatic conditions.

FIGURE 3.5. Geographic distribution of the living nonhuman primates (modiied after schultz [1969], 39, 41; and napier and napier [1967], ig. 4).

SOURCES: overviews of the living primates (Fleagle 1999; Martin 1990, chap. 1; Napier and Napier 1967; Pilbeam 1988, chap. 3; Richard 1985; Schultz 1969; Tuttle 1986)

Haplorhini: The Anthropoidea Basic Taxonomy and Phylogeny. In the classiication scheme employed

here, the living Anthropoidea or higher primates belong to three superfamilies—the Hominoidea, comprising people and apes; the Cercopithecoidea, comprising the Afro-Eurasian or Old World monkeys; and the Ceboidea, comprising the South and Central American or New World monkeys. he Old World and New World monkeys resemble each other in body and head form, but the shared characters are primitive, inherited from a distant common ancestor, and the Old World monkeys share other, more subtle features with apes and people, inherited from a more recent common ancestor. he derived characters that Old World Monkeys, apes, and people share include a common (2-1-2-3) dental formula, comprising two incisors, one canine, two premolars, and three molars on each side of each jaw; an external auditory meatus (bony tube) projecting laterally from the ectotympanic bone that supports the eardrum within the middle ear (ig. 3.4); and complete closure of the bony wall behind the orbit. In New World monkeys the dental formula involves the same number of incisors and canines, but there are three premolars on each side of each jaw (ig. 3.2), the third molar tends to be reduced in size, and in some species (the marmosets and tamarins discussed below), it has been completely lost. New World monkeys also lack the external

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auditory meatus, and their postorbital closure is less complete. Together with genetic observations, the sum implies that the Old World monkeys are actually more closely related to apes and people than they are to their New World counterparts. In addition to the cited bony diferences, the Old World and New World anthropoids also difer in the orientation of the nostrils (ig. 3.6). In general, in the New World monkeys these tend to be widely spaced and to face sideways, whereas in the Old World monkeys and the Hominoidea they tend to be closer together and to face more forward or downward. his has led to the use of the terms platyrrhine (lat-nosed) and catarrhine (downward-nosed) to distinguish the two groups. More formally, the Old World higher primates, living and extinct, are commonly placed in the infraorder Catarrhini, while the New World forms are assigned to the infraorder Platyrrhini. Within the catarrhines, the monkeys tend to be more primitive than apes and people. hus, they retain a primitive mammalian body plan involving a long, narrow, deep trunk (ig. 3.7); a vertebral column that is situated atop the rib cage; a shoulder structure that restricts arm movement mainly to a plane that parallels the body; and an external tail. In contrast, apes and people share a more derived body plan involving a short, lat, broad trunk; a vertebral column that is set within the rib cage; a dorsally placed scapula and laterally facing shoulder joint that together permit free rotation of the arms around the shoulder; and the absence of an external tail. he monkey body plan supports primitive quadrupedalism like that observed in many other mammals, and the monkeys are quadrupedal even in the trees, where they walk along the tops of branches. heir tails then function as balance organs to maintain the center of gravity directly over a branch. Apes and people may have inherited their derived trunk form and shoulder structure from a common ancestor that had given up monkeylike quadrupedalism in favor of underbranch suspension and arm-over-arm tree climbing with the body in an orthograde (upright) position. Some apes commonly use their arms to climb or swing through the trees today, while other apes and people more rarely do because they have shited mostly or entirely to life on the ground. Catarrhine monkeys also difer from the apes (and people) in their characteristic bilophodont lower molar form, which involves two pairs of relatively high cusps, each pair linked buccolingually by a crest or loph (ig. 3.8). In apes, lower molar form usually involves three rounded cusps on the lingual side and two slightly larger, shorter ones on the buccal side, separated by a Y-shaped groove or issure. his is oten known as the dryopithecine Y-5 pattern, because it was irst recognized in the European fossil ape Dryopithecus, discussed below. he diference between monkeys and apes relects a modal diference in diet that was probably particularly important when the two groups irst diverged: apes tend to

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New World Monkey Cebus

Old World Monkey Macaca FIGURE 3.6. representative new World and old World monkeys, illustrating the sideways orientation of the nostrils typical in the new World monkeys and the downward or forward orientation typical in the old World monkeys (redrawn after schultz [1969], 22).

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dorsal (back)

FIGURE 3.7. Cephalic (top down) views of the rib cage and right shoulder girdle of an adult macaque monkey and of an adult human, showing the deeper, narrower chest of the monkey and the different arrangement of the scapula and clavicle, which limits the monkey’s ability to rotate its arm around the shoulder (redrawn after schultz [1969], 81).

vertebral column scapula (shoulder blade) set atop the rib cage on the side of the rib cage narrow, deep rib cage

adult macaque monkey clavicle (collarbone) oriented downward

vertebral column set within the rib cage broad, shallow scapula (shoulder blade) rib cage on the back of the rib cage

adult human clavicle (collarbone) oriented backward

ventral (front)

FIGURE 3.8. lower molars of the early oligocene catarrhine Propliopithecus (Aegyptopithecus) and of the extant leaf-eating monkey Colobus (adapted from butler [1986], igs. 2 and 8). The monkey molar exhibits the bilophodonty typical of all cercopithecoids, with two pairs of cusps linked by shearing crests (lophs). The Propliopithecus molar shows a pattern of ive distinct cusps separated by a Y-shaped issure system that is broadly characteristic of all miocene to recent hominoids. The Y-5 pattern is believed to be primitive in catarrhines, and the bilophodont condition probably evolved from it.

buccal hypoconid protoconid hypoconulid distal

mesial

metaconid

entoconid lingual

The Y-5 pattern on a left lower molar of Propliopithecus buccal protoconid

hypoconid

loph distal

mesial

entoconid

metaconid lingual

Bilophodont pattern on a left lower molar of Colobus

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depend more on fruits that require crushing between relatively blunt molars, while monkeys oten rely more on leaves that require shearing between molars with sharp crests. he fossil record, as outlined below, implies that ape molar form relects the primitive catarrhine condition and that monkey molar form is derived from it. hus, in this particular feature, the monkeys are actually less primitive than the apes. As already noted, genes show that the name ape can be misleading, since some apes are actually more closely related to people than to other apes. he gibbons and the siamang, sometimes lumped together as the lesser apes, are distant not only from people but also from the two species of chimpanzee, the gorilla, and the orangutan, oten known as the great apes. A more surprising result is that, despite appearances, the chimpanzees and the gorilla actually share a much more recent ancestor with people than they do with the orangutan. he anatomical features that unite the great apes and distinguish them from people must thus be either primitive retentions inherited from a distant common ancestor or specialized characters that the various ape species acquired independently as they adapted to similar circumstances. Among ape characters that surely represent shared, primitive retentions (lost uniquely in people), the most notable are a relatively small brain, lower limbs that are not constructed for eicient bipedalism, and a dentition in which the anterior teeth (incisors and canines) are large relative to the posterior ones (premolars and molars). he canines are especially large and, unlike human canines, they tend to wear along the mesial and distal (fore and rear) surfaces rather than at the tips. Associated with this, the lower third premolar (P3) just behind the lower canine is sectorial (elongated mesiodistally and essentially unicuspid). When the mouth is shut, the mesial surface of the upper canine shears between the distal surface of the lower canine and the mesial edge of the sectorial premolar (ig. 3.9). he lower canine its in a gap (diastema) between the upper canine and the upper lateral incisor (I2). In the chimpanzees, the gorilla, and the orangutan, the canines are not only large overall but in further distinction from people, males tend to have much larger canines than females. he dental diferences that consistently diferentiate the great apes from people are particularly useful for separating their fossils, since jaws and teeth dominate the fossil record. Some anatomical features and occasional genetic analyses suggest that the chimpanzees and the gorilla share a more recent common ancestor with each other than either does with people, but the majority of genetic analyses link people and chimpanzees most closely. he reason for the apparent inconsistency is that the branching events that initiated the gorilla, chimpanzee, and human lineages were closely spaced, all probably between roughly 8 and 6 Ma, and the genetic variation in their last shared ancestor, from which the gorilla line diverged irst, survived

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FIGURE 3.9. occlusal relation between the upper canine, lower canine, and lower third premolar (P3) in a nonhuman catarrhine primate (left) and a human being (right) (early wear above, later wear below) (redrawn after le Gros Clark [1964], 182). in catarrhine monkeys and apes, the upper canine is part of a shearing complex involving the lower canine and the elongated unicuspid (sectorial or “cutting”) lower third premolar. When the jaws close, the canines interlock and wear occurs mainly on their fore and rear surfaces. in people, the canines do not interlock and wear occurs primarily at the tips. in addition, in people the lower third premolar is bicuspid and more rounded in outline.

ape and catarrhine monkey condition (illustrated from a monkey)

modern human condition upper canine

upper canine lower canine lower canine

sectorial P3

P3

VERY EARLY WEAR

LATER WEAR

until the subsequent divergence of the chimpanzee and human lines. As a result, the genomes of the gorilla, chimpanzees, and people overlap signiicantly, and diferent genes or diferent individual gorillas, chimpanzees, and humans will always suggest diferent, contradictory branching sequences (diferent phylogenetic relationships between species pairs), sometimes linking the gorilla most closely to chimpanzees or even to people, even if the large majority of genetic comparisons show that the closest relationship is between chimpanzees and people. he principal objection to an especially close evolutionary relationship between the chimpanzees and people is probably that the chimpanzees and the gorilla share derived anatomical features that people lack. hese may include some aspects of dental enamel development and structure and especially a shared, highly specialized mode of terrestrial locomotion. When the chimpanzees and the gorilla stand or move on all fours, they rest their forelimbs on the dorsal surfaces (backs) of their middle phalanges, and their gait is thus oten known as “knucklewalking,” as discussed again below. It difers conspicuously not only from human bipedalism but also from the terrestrial posture and locomotion of orangutans, who tend to rest their forelimbs on the sides of their balledup ists. he wrist morphology of living humans and of the fossil hominins, Australopithecus anamensis and A. afarensis, may imply a knucklewalking ancestor, but the evidence is debatable, and if it is discounted, chimpanzees and gorillas either inherited knuckle-walking from an ancestor they did not share with people or they developed it independently. Some aspects of wrist morphology and the closer relationship of chimpanzees to people suggest that independent development is more likely. he remainder of this section sketches the living anthropoids, with emphasis on those that are closest to people. SOURCES: biomolecular/genetic indications of an especially close relationship between chimpanzees and gorillas (Deinard et al. 1998; Djian and Green 1989; Marks 1993, 1995) vs. one between chimpanzees and people (Bishop and Friday 1986; Caccone and Powell 1989; Goodman et al. 1989, 1990; Hol-

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mes et al. 1989; Horai et al. 1992; Pilbean and Young 2004; Ruvolo 1997; Ruvolo et al. 1991, 1994; Satta et al. 2000; Sibley and Ahlquist 1984, 1987; Sibley et al. 1990); implications of closely spaced branching events and genetic variability in the last shared ancestor of the chimpanzees, the gorilla, and people for assessing their phylogenetic relationships (Pääbo 2003; Rogers 1993); enamel features shared between chimpanzees and gorillas (Andrews and Martin 1987a; Dean and Delson 1992); deinition of knuckle walking (Tuttle 1969); wrist anatomy in living humans that may imply a knuckle-walking ancestor (Corruccini and McHenry 2001); distal radius morphology in Australopithecus anamensis and A. afarensis that may imply a knuckle-walking ancestor (Richmond et al. 2001; Richmond and Strait 2000) and arguments against (Dainton 2001; Lovejoy et al. 2001); parallel evolution of knuckle walking in chimpanzees and gorillas (Dainton and Macho 1999)

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The Chimpanzees. he chimpanzees comprise two species, known pop-

ularly as the common chimpanzee (Pan troglodytes) and the pygmy chimpanzee or bonobo (P. paniscus). Historically, common chimpanzees inhabited forest and woodland across equatorial Africa, from Gambia on the west to the shores of Lake Victoria and Lake Tanganyika on the east. Bonobos were restricted to the forests of the Zaire, south of the Congo River. Common chimpanzees vary somewhat in size from place to place, but on average, free-ranging adult males appear to weigh about 54 kg and females about 40 kg. Despite the “pygmy” appellation, bonobos are not signiicantly smaller, and they difer from common chimpanzees mostly in their relatively longer trunks, longer legs, and shorter arms. hey are also less dimorphic, particularly in major skeletal dimensions. Both species of chimpanzees feed mainly on ripe fruits, which they climb trees to obtain. hey usually ascend with the upper body upright and depend mainly on their long arms for lit and stability. Young individuals oten brachiate (arm swing) from branch to branch, but adults are too heavy to do this routinely. hey travel between trees mainly on the ground, where they commonly assume a four-legged posture, with the feet lat and the hands curled, so that the weight of the forequarters rests on the knuckles (ig. 3.10). heir habitual locomotion is thus quadrupedal (four-footed), though it could equally be called quadrumanous (four-handed), since chimpanzee feet are very handlike from a human perspective. Quadrumanism is perhaps a particularly apt description for chimpanzee climbing, in which both the hands and feet are used to grasp branches. Free-ranging common chimpanzees form loosely knit communities ranging from fewer than twenty to more than one hundred individuals. Males usually remain in their natal groups for life, whereas adult females tend to disperse to other groups. Each female forages through a small territory either by herself or with her dependent young, while males form bands that patrol the territories of several females, fending of neighboring males. Female and mixed-sex aggregations occur at particularly fruitful trees, but individuals scatter when the food is exhausted. Bonobos have not been as thoroughly studied, but their social behavior seems

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FIGURE 3.10. Quadrupedal postures in (clockwise) an old World monkey, a human, a gorilla, and a chimpanzee (redrawn after schultz [1969], 55). note that the monkey and human are standing with their palms lat, whereas the chimpanzee and gorilla are standing with their knuckles curled. note also that the chimpanzee and gorilla have much longer arms relative to their legs.

adult pig-tailed macaque

human child

adult chimpanzee

juvenile gorilla

broadly similar. he most conspicuous diferences are that they forage more oten in small mixed-sex groups and their communities tend to be more stable or cohesive. Compared with common chimpanzees, bonobos are also marked by stronger bonds between females, weaker ones among males, markedly fewer aggressive interactions, and a greater inclination to share food. he diferences might occur in part because, unlike common chimpanzees, bonobos nowhere overlap with gorillas. When the fruits they favor become seasonally scarce, they thus can more readily fall back on the leaves and shoots that gorillas otherwise preempt. Bonobos’ reduced susceptibility to food stress may partly explain why they are less aggressive toward each other, and reduced aggression among males could explain why they are less dimorphic. In their high level of male aggression, common chimpanzees are nearly unique in the animal kingdom, and the species they resemble most closely is probably Homo sapiens. hus, common chimpanzee males in neighboring groups tend to be mutually hostile, and if a male patrol from one group encounters an isolated male from another, their reaction is to attack and kill. he result over time can be the elimination of all

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the males in a second group and the appropriation of their territory and resources, including their females. Conceivably, common chimpanzee and human males inherited their shared tendency to intergroup violence from their last shared ancestor 6–7 Ma, although this would require an evolutionary loss in bonobos, ater they split from common chimpanzees about 2.5 Ma. A more serious objection is that the pattern of male violence difers signiicantly between common chimpanzees and people in those historic human societies that are most relevant for comparison. hese are hunter-gatherer communities that are sometimes referred to as “simple” for their egalitarian social structure, small average group and total population size, luid group composition, and nomadic lifestyle. Until 50 ka, all people probably lived in such societies, and as recently as 10–12 ka, most probably did. Historically, homicide rates in such societies sometimes approached those in modern industrial communities, but unlike “panicide” rates, they were due mainly to sexual jealousy between males within a group, not to conlicts between males of diferent groups. he diference arises primarily because unlike neighboring chimpanzee communities, neighboring hunter-gatherer groups usually share access to key resources. More complex human societies generally do not, and it is their competition that gives rise to a high level of intergroup male violence. he sum suggests that male violence in common chimpanzees and living humans has diferent roots. However, if it is assumed that the last shared ancestor was more like the common chimpanzee in its socioecology and behavior, then intergroup male violence in hominins may have gone from frequent in the earliest species to infrequent when simple hunter-gatherer societies irst emerged to frequent again when human societies became larger, more complex, and more competitive. Anatomical and archaeological observations summarized in chapter 5 tentatively place the emergence of simple hunter-gatherer societies around 1.7 Ma in the evolution of Homo ergaster and the Acheulean Cultural Tradition. Greater social complexity appeared much later, mainly ater 12–10 ka with the advent of agriculture. hus, human male intergroup violence probably came to resemble the common chimpanzee pattern only late in human evolution and it resulted from social change more than from shared genes. Beginning with Jane Goodall in Gombe Stream National Park, western Tanzania, observers have now repeatedly recorded two other derived behaviors—hunting and tool use—that common chimpanzees share with living humans and that are more likely to have been inherited from the last shared ancestor. Common chimpanzees hunt monkeys and other small mammals with whom they share their range, and some chimpanzee communities consume several hundred kilograms of meat per year.

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In addition, like human hunter-gatherers, they do not hunt only to eat. Males are the principal hunters, and they trade meat for sexual access. hey may also share meat with other males to cement political alliances. Tool use, as detailed below, sometimes involves the modiication of natural objects, which makes it even more humanlike. he speciics of common chimpanzees hunting and tool use vary from place to place, and in this sense again, they recall humans. For example, in the Taï National Park, Ivory Coast, chimpanzees commonly initiate hunts before the prey are in range, they oten stalk in groups, and they regularly share the spoils. In contrast, at Gombe, they hunt much more opportunistically, they more rarely cooperate in stalking, and they share more reluctantly. Similarly, in western Africa, many chimpanzee groups routinely use stone or wooden hammers and anvils to crack nuts, at least some modify twigs to pluck marrow from the bones of hunted prey, some use their incisors to sharpen the tips of branches that they jab into tree hollows where bushbabies or other small mammals may be resting, and most if not all probe insect nests with stems, twigs, or pieces of vine (the insects that cling to the invading probes are extracted and eaten). In contrast, no east-central African chimpanzees use tools to crack nuts or sticks to extract marrow, none have been reported to sharpen twigs with their incisors, and they vary in how much they use modiied vegetal probes to obtain insects. Since nuts, marrow bones, and insect nests occur nearly everywhere, some observers argue that regional diferences in hunting, tool use, and other behaviors at least partly relect chimpanzee cultures that anticipate human ones. Unlike humans, however, chimpanzees transmit their cultures exclusively through direct observation or imitation, which is to say that only behavior is passed between generations, not ideas or meanings. In this regard chimpanzees are like lions or songbirds whose ofspring also learn speciic behaviors by observing their elders. Perhaps even more important, chimpanzee tools are so simple that it is easy to imagine their reinvention if observation and imitation failed to promote their transmission, and variation in the frequency and intensity of tool use among chimpanzee groups suggests it could be lost with only minimal impact on species survival. In contrast, human tools and other socially transmitted behaviors are oten far more complex, and even in the simplest human societies, they relect a compounding and accumulation of innovations over countless generations. If they were somehow lost, they would thus be far harder to regenerate from scratch, and their loss would usually be catastrophic for the group. With these facts in mind, it is clear that chimpanzees lack both Culture and cultures in the fundamental human sense. heir behavioral resemblance to people is striking, but the diference is even more remarkable, and a major aim of this book is to outline its development.

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SOURCES: common chimpanzee social organization (Wrangham 1987); diferences in social organization between common and pygmy chimpanzees (Susman 1987; White 1996a, 1996b) and the possibility the diference relects lack of overlap between pygmy chimpanzees and gorillas (Wrangham et al. 1996); male violence in common chimpanzees (Manson and Wrangham 1991; Wrangham and Peterson 1996) and in simple human forager societies (Knaut 1991); pioneering research on common chimpanzees (Goodall 1986); volume of meat-eating by chimpanzees (Stanford 1995); hunting for purposes other than food by human hunter-gatherers (Hawkes et al. 1991, 1997b) and by chimpanzees (Stanford 1995, 1996); regional variation in chimpanzee hunting or tool use (Boesch and Boesch 1989; Boesch-Achermann and Boesch 1994; Hernandez-Aguilar et al. 2007; McGrew 1992; van Schaik et al. 1999); west African chimpanzee tool use and manufacture (Mercader et al. 2007; Pruetz and Bertolani 2007); regional variation in chimpanzee behavior as an indication of chimpanzee cultures (Lycett et al. 2007; McGrew 1992, 2007; Small 1993)

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The Gorilla. Gorillas overlap chimpanzees in distribution, but gorillas

require much denser forest, and they are far less widespread and abundant. hey concentrate in two distinct enclaves about 1,000 km apart—a larger one, comprising perhaps 70,000 individuals in the lowland forests around the Gulf of Guinea in west-central Africa and a smaller one, comprising perhaps 700 individuals, in the mountainous areas of eastern Democratic Republic of the Congo, western Uganda, and western Rwanda in east-central Africa. he separate populations are known, respectively, as the lowland and mountain gorillas, and they are usually regarded as subspecies of a single species, Gorilla gorilla. Some authorities further divide the lowland gorilla between a more western and a more eastern subspecies. Gorillas are the largest living apes, but the sexes difer strikingly in size. Free-ranging adult males reach 180 kg or more, while females average closer to 90 kg. Young gorillas are adept brachiators, but the adults are so large that they rarely enter trees, except occasionally to sleep. Like chimpanzees and bonobos, gorillas are quadrupedal on the ground, and they move from place to place with the feet lat and the hands resting on the knuckles. hey eat fruit where it’s available, but they oten subsist mostly on leaves and shoots. hey difer in this regard from both chimpanzees and orangutans, both of which rely far more heavily on fruit. heir characteristic social groups number between nine and twenty-two individuals including a single dominant mature male (oten designated a “silverback” from the mat of gray white hairs found on the back), some subadult males, and several unrelated females with immature young. “Surplus” adult males live alone or in small bachelor groups. Males compete iercely for females, but gorilla communities are more stable and coherent than their chimpanzee counterparts, perhaps because gorilla food staples are more abundant and evenly distributed. Studies of freeranging gorillas indicate that unlike chimpanzees, gorillas rarely if ever eat meat or use tools. SOURCES: gorilla taxonomy (Groves 1986, 2001); gorilla socioecology (Harcourt and Stewart 2007a, 2007b); rarity of meat-eating or tool use by gorillas (Fossey 1983; Schaller 1963)

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The Orangutan. In contrast to the chimpanzees and the gorilla, which are

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exclusively African, the orangutan is exclusively Asian. here is only one living species, Pongo pygmaeus, which was narrowly restricted in historic times to the southeast Asian islands of Borneo and Sumatra. Fossils, however, show that during the Pleistocene the same species occurred in Java, Cambodia, hailand, Vietnam, Laos, and southern China, over an area that was roughly comparable in extent to the ranges of the African great apes. Orangutans are like chimpanzees in size but like gorillas in the degree of sexual dimorphism. Free-ranging adult males probably average about 65 kg, while females weigh only about half as much. In sharp contrast to both gorillas and chimpanzees, orangutans are almost exclusively arboreal and come to the ground relatively rarely. Young orangutans actively brachiate, but adults climb through trees very cautiously, so as to avoid potentially fatal falls. hey are mostly quadrupedal on the ground, but unlike chimpanzees or gorillas, they tend to walk on the sides of their ists rather than on their knuckles. hey feed mainly on fruits, supplemented to some extent by insects. Many of their preferred fruits are hard coated, which may explain why they have thicker dental enamel than either chimpanzees or gorillas. hey seem to be essentially solitary (nongregarious), and the only coherent social unit is a female and her dependent young. Adult males apparently defend small territories containing a handful of adult females with whom they mate. Free-ranging orangutans rarely if ever eat meat, but like chimpanzees, they sometimes use twigs or other natural objects as tools to acquire food. Also like chimpanzees, orangutans vary from place to place in the frequency of tool use, probably because like chimpanzees and people, orangutans are better mimics than inventors. Since reinvention is rare, tool behaviors are most likely to persist where high population density and social tolerance provide good opportunities for copying, and they can disappear strictly by chance where density is low and sociability is minimal. SOURCES: orangutans—Pleistocene distribution (Louys et al. 2007); socioecological overview (van Schaik 2004); tool use in free-ranging populations (van Schaik et al. 1996) and reasons tool use varies from place to place (Fox et al. 2004; van Schaik et al. 1999, 2003; van Schaik and Knott 2001)

The Lesser Apes (Hylobatids). As the vernacular name lesser apes im-

plies, the hylobatids are relatively small. Depending on the species, adults weigh between 4 and 13 kg, and males and females are generally similar in overall size and appearance. In their near total lack of sexual dimorphism, the lesser apes contrast strongly with the great apes, from which they are also readily distinguished by the extraordinary length of their arms relative to their trunks and by hardened pads of skin (ischial callosities) that overlie broadened, roughened margins (ischial tuberosities) on the ischial bones of the pelvis. he pads permit prolonged sitting on

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the haunches and are also found in the Old World monkeys. hey may therefore represent a primitive catarrhine feature that was lost in the great apes only ater the line leading to the lesser apes had diverged. Historically, the lesser apes were distributed from southern China through the forests of southeastern Asia, including the ofshore islands of the Indonesian Archipelago where they overlapped the orangutan. As many as eight species are recognized, divided between two subgenera of the genus Hylobates. he single species placed in the subgenus H. (Symphalangus) comprises the largest of the lesser apes, oten known as the siamangs. he several species placed in the subgenus H. (Hylobates) include a variety of smaller apes commonly known as the gibbons. Like the term lesser apes, however, the term gibbons is oten used to embrace both the gibbons and siamangs. Gibbons and siamangs rarely leave the trees, where they use their powerful, elongated arms to brachiate from branch to branch and tree to tree. he gibbons proper eat mainly ripe fruits, supplemented with insects and birds’ eggs, whereas the siamangs concentrate more on fresh leaves and shoots, supplemented with fruits and insects. Unlike other apes, both gibbons and siamangs have a social organization involving male-female pairs that bond for life. Each pair and its immature ofspring inhabit a territory that they defend against neighboring pairs. Adult males do not regularly compete for females, which perhaps explains the limited amount of sexual dimorphism.

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The Old World Monkeys. he cercopithecoid or Old World monkeys are

far more widespread and diverse than the apes, though this is a relatively recent development, and the fossil record shows that apes were once much more diverse than monkeys. Historically, cercopithecoid monkeys were found more or less throughout Africa and southern Asia, except in extreme deserts. During parts of the Pleistocene they also occurred in Europe, and an isolated population persists today on the Rock of Gibraltar. he taxonomy of the cercopithecoids is disputed, but there are perhaps seventy-ive living species in seven genera usually divided between two subfamilies—the Cercopithecinae and the Colobinae—within a single family, the Cercopithecidae. he cercopithecines are the common monkeys of sub-Saharan Africa (Papio, heropithecus, Cercocebus, Erythrocebus, and Cercopithecus), with only a single genus (Macaca) in northern Africa and Eurasia. In contrast, the colobines are more abundant in Asia (Nasalis, Presbytis, Pygathrix, and Rhinopithecus), though one genus (Colobus) is restricted to Africa, where it is locally common. Cercopithecines and colobines contrast sharply in what they eat: whereas cercopithecines tend to focus on fruits (both ripe ones and ones not ripe enough for ape consumption), colobines emphasize leaves. To

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Papio mandrillus

FIGURE 3.11. skulls of male and female adult mandrills (Papio mandrillus), illustrating the extraordinary degree of sexual dimorphism, in both overall size and canine development, that characterizes many cercopithecoid monkeys (redrawn after schultz [1969], 202).

male

0

5 cm

female

subsist on leaves, which are relatively high in nonnutritive bulk (iber), colobines have evolved elaborate, sacculated stomachs. hey also have relatively long, sharp crests on their molars, which are thus well-suited for shearing leaves—unlike the shorter, blunter crests on cercopithecine molars, which are better suited for crushing fruits. All cercopithecoids are primarily quadrupedal and arboreal, though some appear equally at home on the ground and a few cercopithecines are largely or wholly terrestrial. Knuckle walking is unknown; most arboreal species tend to rest and move on their palms and soles, while largely terrestrial species commonly stand and walk on the tips of their digits. Some colobines routinely hang from branches by their arms, but they are anatomically incapable of the arm-swinging locomotion (brachiation) observed in gibbons and immature great apes. Terrestrial species tend to be larger than arboreal ones, and body size varies from the highly arboreal, 1–2 kg talapoin monkey (Cercopithecus talapoin) to the largely or wholly terrestrial baboons (Papio and heropithecus), in which adult males reach 40 kg. Most species are organized in mixed sex troops, numerically dominated by females. Adult males compete vigorously for females, which perhaps explains why they usually have much larger bodies and canines (ig. 3.11). The New World Monkeys. he ceboid or New World monkeys may be di-

vided among three families: the Atelidae (Ateles, Brachyateles, Alouatta, and Lagothrix), the Pitheciidae (Pithecia, Chiropotes, Callicebus, and Ca-

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cajao), and the Cebidae (Cebus, Saimiri, Aotus, Callimico, Callithrix, Cebuella, Leontopithecus, and Saguinus). Species of all three families tend to resemble the Old World monkeys in overall form, diet, and sociality, but there are no large terrestrial forms (comparable to the baboons) or dedicated leaf eaters (comparable to the colobines). Instead, they are all more or less strictly arboreal, and most feed mainly on fruits, variably supplemented by leaves and insects. hey are all primarily quadrupedal, but like the colobines, the atelids have developed shoulder and arm specializations that allow them to hang from branches. hey also have a prehensile tail that functions as a ith limb in underbranch suspension. Like most colobines, they occupy forests where their suspensory abilities allow them to feed near the ends of slender branches that strictly quadrupedal forms could not reach. he cebids divide between two subfamilies, the Cebinae, comprising the capuchin, squirrel, and owl monkeys (the irst three genera listed in the previous paragraph) and the Callitrichinae, comprising the marmosets and tamarins (the last ive genera). Despite their close phylogenetic relationship, the cebids are remarkably diverse in behavior and ecology. Capuchin monkeys have semiprehensile tails that broadly recall those of the Atelidae, and owl monkeys are the only higher primates that are more or less exclusively nocturnal. On average, the marmosets and tamarins tend to be much smaller than other cebids, and in size and some morphological attributes they recall lower primates (strepsirhines). Perhaps most notable, like the lower primates but unlike other higher primates, they have a mix of nails and claws. he claws represent an evolutionary reversal since they evolved from nails, but they are bilaterally compressed in the manner of true claws, and they occur on every digit but the big toe. hey function to secure the body on branches that small hands and feet could not grasp. In addition, like most lower primates, some marmoset and tamarin species feed mainly on insects or gum, as opposed to fruits, nuts, or leaves, on which the other New World monkeys focus. True lower primates are absent in the New World, and to a degree the marmosets and tamarins ill their niche. In key features of head and body, however, the marmosets and tamarins closely resemble the other cebids, and like nearly all other higher primates, they are also diurnal and highly social. In Africa and Asia, where lower primates overlap higher primates in the same way that marmosets and tamarins overlap other ceboids, the lower primates are nocturnal and solitary.

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SOURCES: Ceboid systematics and phylogeny (Opazo et al. 2006; Rosenberger 2002)

Haplorhini: The Tarsiiformes

he Tarsioidea or tarsiers are tiny (100–200 g) animals represented by a single genus (Tarsius) with three or four species. hey are restricted to

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Sumatra, the southern Philippines, and the Celebes Islands of southeastern Asia where they inhabit bush and forest. hey are almost totally nocturnal in their activity pattern, and they have enormous eyes for night vision. hey feed primarily on insects, supplemented with small vertebrates and perhaps with vegetal material. hey take their name from their highly elongated tarsus (ankle), which they use to propel themselves in long leaps between near-vertical branches to which they cling in an upright position. his distinctive mode of locomotion has been called vertical clinging and leaping, and it is aided by a long nonprehensile tail that can be shited in midair to ensure an upright landing. Like the lorises and some lemurs described below, tarsiers appear to be mainly solitary, although semipermanent male-female pairs have sometimes been reported. As already indicated, the taxonomic and phylogenetic status of the tarsiers has been a matter of intense disagreement because they resemble the strepsihrines in features like their laterally placed incisors, their relatively sharp nonmolarized premolars, and their lack of sexual dimorphism. However, unlike strepsirhines and in common with the anthropoids, they lack both a naked rhinarium (a spot of hairless, glandular moist skin around the nose) and an accompanying clet in the upper lip, their orbits are partly separated by bone (a partial postorbital plate) from the jaw musculature just behind, and their right and let frontal bones are completely fused. hey also resemble the anthropoids in key characters of reproductive biology, including the presence of an advanced hemochorial type of placenta. he features they share with anthropoids are all derived, and together with their genes, they justify placement with the anthropoids in the Haplorhini.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: disagreement over the taxonomic position of tarsiers (Aiello 1986; Schwartz and Tattersall 1987); derived features that tarsiers share with higher primates (Fleagle 1994, 18; Martin 1990, 663– 670); assignment of the tarsiers to the Haplorhini based on anatomy (Beard et al. 1991; Hofstetter 1988; Martin 1990; Miller et al. 2005a; Szalay et al. 1987; Szalay and Delson 1979) and molecules (Baba et al. 1980; de Jong and Goodman 1988; Koop et al. 1989; Miyamoto and Goodman 1990; Pollock and Mullin 1987; Sarich and Cronin 1980; Schmitz et al. 2005)

Strepsirhini Basic Taxonomy. he strepsirhines or lower primates divide between

two suborders—the Lorisiformes with a single superfamily, the Lorisoidea, and Lemuriformes with two, the Lemuroidea and the Daubentonioidea. In times past, the Tupaiiformes with the superfamily Tupaioidea might also have been included in the strepsirhines, but they now occupy their own order, and they are considered below because they may constitute the best available living model for the last shared ancestor of the Primates.

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The Lorises and Lemurs. he Lorisoidea or lorises live in Africa and

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southern Asia, while the Lemuroidea or lemurs and the Daubentonioidea or aye-ayes are conined entirely to Madagascar and the nearby Comoro Islands. In keeping with their limited geographic distribution, they are oten known as the Malagasy lemurs. Among the shared anatomical features that imply a single (monophyletic) origin for the lorises and lemurs, the most notable are the alignment and forward protrusion of the lower canines and incisors (ig. 3.2) to form a “dental comb” used in grooming, and the presence of a claw on the second toe, also used for grooming. In contrast to the haplorhines, both the lorises and the lemurs lack a postorbital plate, and they tend to have faces like those of nonprimate mammals, with a relatively long snout, a rhinarium, a clet upper lip bound down to the gum, relatively large and free-moving ears, and an immobile facial expression. hey further difer from the anthropoidea in having more laterally placed incisors; relatively sharp, nonmolarized premolars; and in most species a mandible in which the two halves remain unfused at the symphysis, or midline (ig. 3.12). hey are also much less sexually dimorphic than most anthropoids. In external features, they resemble anthropoids most obviously in their hands and feet. hese are functionally adapted for grasping, with mobile digits tipped by lattened nails except, as already mentioned, for the second toe, which bears a grooming claw. he lorises are commonly divided between two major types, treated here as subfamilies—the Lorisinae or lorises in the narrow sense and the Galaginae or galagos. he lorises are both south Asian (two species in two genera) and equatorial African (two species in two genera). he galagos or “bushbabies” (four-to-six species in one genus) live exclusively in Africa, where they have a broader distribution than the lorises, extending far to the south in savanna and bush. In sharp distinction from the exclusively diurnal catarrhine monkeys and apes that share their range, the lorises and galagos are all nocturnal. hey are also relatively unsociable, and individuals usually forage alone. All lorises are almost entirely arboreal, but the lorises in the narrow sense tend to be slow-moving climbers, while the galagos are more energetic runners and jumpers. Like the tarsiers and some lemurs, the galagos have powerfully built hindlimbs that aid in vertical clinging and leaping, but galagos vary in how much they cling and leap, and some species are predominantly quadrupedal. Both lorises and galagos include species that feed primarily on insects and others that focus more on gum or fruits. he lemurs are a much more diverse group of animals; this relects the minicontinental size of Madagascar, its environmental diversity, and its separation from Africa beginning roughly 170 Ma. It is currently 430 km east of Mozambique, and this distance has been constant since

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ANTHROPOID (Cebus monkey)

FIGURE 3.12. Comparison between the skulls of a malagasy lemur (Lemur) and a new World monkey (Cebus) to illustrate some of the features that commonly distinguish prosimians from anthropoids (redrawn after rosenberger [1986], ig. 4).

frontal bones fused

fully enclosed orbit shortened face

advanced grinding stroke of the chewing cyle

frontal bones fused postorbital closure

cancellous petrosal bone

fused mandibular symphysis

incisors enlarged and located frontally, premolars blunted

PROSIMIAN (Lemur) frontal bones unfused

no postorbital closure long face

simpler grinding stroke of the chewing cyle

frontal bones unfused no postorbital closure

noncancellous petrosal bone

unfused mandibular symphysis

incisors smaller and more laterally emplaced, premolars more pointed

130–120 Ma, long before lemurlike animals or any Primates had probably appeared on the African mainland. he molecular diversity of the living lemurs suggest that their last shared ancestor reached Madagascar between 80 and 50 Ma, most probably around 60 Ma, and the only viable, if seemingly improbable explanation is that it arrived on a loating rat of vegetation. Molecular evidence actually allows for two separate rating events, one for each of the lemur superfamilies. he stochastic (or “sweepstakes”) nature of rating events precluded the arrival of most other modern mammals, and the ancestral lemurs thus evolved in isolation from monkeys, apes, and many other potential competitors. his isolation allowed them to radiate into niches that lower primates could not occupy elsewhere.

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Specialists debate the number of extant lemur species, but at present there are minimally thirty-six species in sixteen genera, and at least fourteen additional species and seven genera probably became extinct only within the past 1,500 years, ater humans arrived in Madagascar. Most lemurs are arboreal, but some are more terrestrial, and to judge from their morphology and large size, some recently extinct forms may have been totally terrestrial. In feeding, some species focus on insects or gum while others emphasize fruit or leaves. hey also vary greatly in locomotor pattern, preference for nighttime versus daytime activity, social behavior, and other features; and though many species recall lorises in their overall level of organization, others are more reminiscent of monkeys. In keeping with contrasts between monkeys and lorises in Africa and Asia, the more monkeylike lemurs tend to be larger bodied, more diurnal, more dependent on leaves or fruit (vs. insects or gum), and more social (vs. solitary) than their less monkeylike relatives. he monkeys and monkeylike lemurs are oten cited as an example of parallel evolution, whereby creatures with similar genetic backgrounds exposed to similar environmental conditions have evolved in similar or parallel fashion.

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SOURCES: taxonomy of strepsirhines in general (Masters et al. 2005; Rumpler 2004; Yoder 1997), lorises (Masters et al. 2006), and Malagasy lemurs in particular (Del Pero et al. 2006; Tattersall 2007); initial lemur colonization of Madagascar (Stankiewicz et al. 2006); extinction of lemurs within the past 1,500 years (Burney and MacPhee 1988; Dewar 1984); resemblances between the monkeylike lemurs and true monkeys (Martin 1990)

Tupaiiformes

he only extant superfamily of the Tupaiiformes is the Tupaioidea or tree shrews, which are widely distributed throughout southern and southeastern Asia, including the ofshore islands. In total, there are perhaps ive genera, all resembling true shrews (order Insectivora) or tropical squirrels (order Rodentia) in overall appearance. Most are highly active, largely terrestrial, diurnal inhabitants of forest undergrowth, but one genus is arboreal and mainly nocturnal. hey all feed primarily on insects, supplemented with vegetal material and small vertebrates. Like the undoubted primates already discussed, tree shrews have a complete post- or circumorbital bar, and they possess a few minor features of the teeth, eye, brain, and limbs that suggest some relationship to lemurs. However, they lack the bony ear structure (petrosal bulla discussed above) that all other primates share, the orbits are not notably convergent in most species, the digits all have claws, and the hands and feet are not especially well-adapted for grasping. hese and other nonprimate features exclude tree shrews from the Primates, and they are now commonly placed in their own order, the Scandentia, which is then grouped with the Primates and the Dermoptera (colugos or “lying

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lemurs”) in the superorder Euarchonta. he Dermoptera may actually be more closely related to the Primates, but the Scandentia are still important to primate evolution, particularly in cladistic analyses of primate relationships where they serve as a useful “outgroup,” meaning a taxon to which all Primates are equally unrelated because it branched away from the Primates before they diversiied internally.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: morphological similarities between tree shrews and lemurs (Kay et al. 1992; Le Gros Clark 1960); assignment of the tree shrews to the Scandentia (Butler 1972); tree shrew systematics and phylogeny (Olson et al. 2005; Sargis 2004); relationships of Scandentia, Dermoptera, and Primates (Janečka et al. 2007)

Primate Evolution and the Molecular Clock he genes and proteins of extant primates not only inform on their evolutionary relationships, they can also be used to estimate the time(s) when extant taxa last shared a common ancestor. he method is oten called the molecular (or biomolecular) clock since it relies on the assumption that DNA or protein diferences accumulate between taxa at a more or less constant (linear) rate. Roughly speaking, the rate is the degree of genetic or protein diference between two taxa divided by the length of time the fossil record indicates they have been separate. Once the rate has been established, it can be applied in reverse to estimate the divergence times between taxa for whom only the degree of molecular divergence is known. he molecular clock was irst applied to human evolution in the 1960s and 1970s, and it suggested that the human and chimpanzee lines probably separated around 4–5 Ma and no more than 6–8 Ma. his contradicted fossils from the Siwalik Hills on the India/Pakistan border, which implied that the human line had separated by 12–14 Ma. he specimens were jaws and teeth assigned to the genus Ramapithecus, and they led specialists to doubt the validity of the molecular clock. However, in the 1980s, fresh fossils suggested that Ramapithecus actually documented the divergence of the orangutan lineage from a joint African ape and human line. African fossils discussed in the next chapter now support the 8–5 Ma molecular estimate for African ape and human divergence, and few specialists now ignore the molecular clock. he use of the clock has ballooned, particularly since the mid-1990s, with the development of technologies that produce genetic sequences ever more quickly and cheaply. However, it can never provide precise divergence dates, if only because the determination of its ticking rate depends on fossil estimates of taxonomic divergence, and few of these are irm. Most fossil estimates for initial appearance of a taxon are minima, and in many cases, the actual time could be signiicantly older. his is particularly true for estimates in deep time, focused on the earliest oc-

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Primates Anthropoidea Hominoidea

Ma 0

HUMAN

CHIMP

GORILLA

Cercopithecoidea

ORANG

BABOON MACAQUE

VERVET

Ceboidea

SQUIRREL TITI MONKEY MARMOSET MONKEY

Strepsirrhini

MOUSE LEMUR

LEMUR

GALAGO PLEIST.1 PLIO.2

10

8.6 (7.7-9.2)

*6.6 (6.0-8.0) 9.9 (8.9-11.8)

MIOCENE

* 6.6 (6.0-7.0)

17.1 (15.0-20.5)

18.3 (16.3-20.8) 20

30

OLIGOCENE

20.8 (18.2-24.9)

30.5 (26.9-36.4)

40

40.9 (35.3-51.0)

EOCENE

42.9 (37.3-52.4)

CENE

57.1 (49.4-71.4) 60

PALEO-

50

70

77.5 (67.1.2-97.7) 80

CRETACEOUS

K-T BOUNDARY

FIGURE 3.13. The divergence dates between representative great apes, old World monkeys, new World monkeys, and prosimians (strepsirrhini), estimated from reined statistical analysis of an extensive region of the genome homologous in each taxon to human chromosome 7 (from steiper and Young [2006], 189). divergence rates were estimated from published fossil divergence times between chimpanzees and people, between rhesus macaques and anubis baboons, between the rodent genera Mus and Rattus, and between the carnivore genera Felis and Canis. The divergence dates in this igure resemble those derived from other molecular data, and they are also broadly consistent with the fossil record.

currence of higher-level taxa. In addition, the ticking rate is now known to difer between taxa and between diferent parts of the genome. Within the Primates, for example, repeated studies have shown that the rate tends to be signiicantly slower in hominoids than in cercopithecoids. In recognition of the known problems, the clock methodology itself has evolved to employ multiple fossil divergence estimates and mathematical models that allow for disparate evolutionary rates, both within the genome and

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between taxa. he result is greater congruence between divergence dates based on diferent genes or on diferent portions of the genome and, with this, increased conidence that the dates approximate reality. Figure 3.13 presents a state-of-the-art efort to date the divergence between major primate taxa. It was based on a substantial section of the genome, it took account of known rate variation among diferent primate lineages, and it employed four fossil divergence estimates: (1) the split between the chimpanzee and hominin lineages, assuming that Sahelanthropus, dated to between 7 and 6 Ma and discussed in the next chapter, is a basal hominin; (2) the split between rhesus macaques (Macaca mulatta) and anubis baboons (Papio anubis), ixed by fossils between 8 and 6 Ma; (3) the divergence between the lines leading to “mice” (Mus) and “rats” (Rattus), estimated by fossils to have occurred between 14 and 8.8 Ma; and (4) the split between the lines leading to “cats” (Felis) and “dogs” (Canis), bracketed by fossils between 65 and 45 Ma. he use of ranges for fossil divergence times means that the molecular estimates must also be ranges, but they are relatively narrow, and with their associated midpoints, they probably approximate the actual times of taxonomic divergence as well as fossil inds, summarized in the next section.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: initial application of the molecular clock to human evolution (Sarich 1971, 1983; Sarich and Wilson 1967); Ramapithecus irst as evidence for the divergence of the human and great ape lines (Pilbeam 1972; Simons 1972; Simons and Pilbeam 1965) and later for the divergence of the orangutan and African great ape/human lines (Pilbeam 1986) 12–14 Ma; molecular clock methodology and estimated divergence dates for primates (Pilbeam and Young 2004; Steiper and Young 2006)

Primate Evolution prior to 6–7 Ma he two previous editions of this book attempted detailed summaries of primate evolution before the emergence of hominins in the late Miocene, between 8 and 5 Ma. his followed on a long-standing tradition in which paleoanthropologists sought to trace the lines leading to extant primate taxa, including humans, as far back in time as possible. he goal remains valid, but as the fossil record has burgeoned, it has grown ever more elusive. he problem is mainly that the expanded record shows that the derived traits that deine extant primate taxa appeared piecemeal rather than in neat packages, and the individual derived traits that fossil taxa share with each other or with extant taxa are diicult to separate between ones that probably relect close common ancestry and ones that relect convergent or parallel evolution. Many must relect convergent evolution, since any given taxon at any given time, including the historic present, can have only one ancestor at any previous time, yet based on shared, derived features, the fossil record oten appears to provide multiple possible ancestors for later taxa.

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With the interpretative diiculties in mind, the following summary is much less detailed than its predecessors, and it focuses on the emergence of ever more advanced primate taxa, culminating in the emergence of the hominins. he fossil record is now complete enough to indicate when progressively more advanced taxa had appeared, although in each case, the oldest known occurrence of a taxon or a lineage may substantially postdate its earliest representative. his is not only because the fossil record is chronologically and geographically uneven or because individual fossils tend to be fragments, mainly teeth and jaws, but also because the derived features that unambiguously deine a lineage may be impossible to detect in its earliest representatives. he emphasis on the appearance of ever more advanced primates is not meant to imply that their evolution was inevitable, for, even if each stage required a predecessor, nothing demanded its successor. On the contrary, natural selection and the other forces that drive evolution could have produced an entirely diferent result at any time, and there might then never have been hominins—or any other species—self-consciously seeking pattern in the fossil record.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: challenges to reconstructing primate evolution from the fossil record (Miller et al. 2005a)

Primate Origins he zoological class Mammalia, to which the Primates belong, evolved from mammal-like (therapsid) reptiles during the Triassic period of the Mesozoic era, roughly 220 Ma. he early mammals soon diversiied into several stocks, including one leading to the heria, a branch or subclass of the mammals that includes the marsupials (Metatheria), placentals (Eutheria), and some other, extinct infraclasses. he marsupial and placental lineages diverged during the early Cretaceous period, between 130 and 100 Ma. he earliest placentals were diverse but all were variously similar to shrews, moles, tenrecs, or other living creatures that have oten been lumped into the order Insectivora. No Cretaceous fossils can be unambiguously assigned to any extant placental order, and the fossil record and the molecular clock together suggest that the extant orders, including the Primates, originated ater 65.5 Ma, in the Paleocene or early Eocene. Still, it is surely from some Cretaceous insectivore-like creature that the Primates sprang, and some background on the Cretaceous world is thus relevant to primate origins. Cretaceous geography was very strange by modern standards (ig. 3.14). In the very early Cretaceous, South America was still joined to Africa, but the two continents began to drit apart roughly 125 Ma, and by the end of the Cretaceous, 65.5 Ma, when the Primates perhaps emerged, the continents were separated by a narrow but expanding South Atlantic.

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Late Triassic (ca. 200 Ma)

North America

Eurasia North America Equator Tethys Sea

South America

Africa

Madagascar India

Eurasia

Tethys Sea

Africa

Gondwana

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Early Cretaceous (ca. 135 Ma) Laurasia

FIGURE 3.14. Changing positions of the continents from the late Triassic, roughly 200 ma, to the middle eocene, roughly 44 ma (modeled after Zihlman [1982]). in the late Triassic, the modern continents were essentially joined in a single supercontinent known to geophysicists as Pangaea. subsequent fragmentation (drift) divided Pangaea into a northern hemisphere landmass known as laurasia and a southern hemisphere mass known as Gondwana. Yet further fragmentation divided laurasia and Gondwana into separate parts, foreshadowing the modern continents. The conjunction of north america and eurasia in the Paleocene and early eocene, between roughly 65.5 and 45 ma, accounts for the close similarity of their early primate faunas, as discussed in the text.

Pangaea

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India

South America

Australia Antarctica

Madagascar

Mid-Paleocene (ca. 58 Ma) Paleocene sites with plesiadapiform or possible plesiadapiform fossils

Antarctica

Australia

Mid-Eocene (ca. 44 Ma) Eocene fossil primate sites

South America and North America were separated throughout the Cretaceous, but North America was connected to Europe via Greenland. Also, global climate was remarkably mild more or less throughout, and there was relatively little temperature diference between the equator and the poles. In both the Southern and Northern Hemispheres, temperate forests thrived at high latitudes. he Cretaceous is oten known as the “age of dinosaurs” because they were its most conspicuous, if not its most numerous, vertebrates. But it also witnessed important evolutionary developments and diversiication in other kinds of reptiles, in birds, in early mammals, and not least in plants. Before the Cretaceous, the principal plants were gymnosperms (nonlowering plants such as conifers, palms, and cycads). Angiosperms (lowering plants, including trees, grasses, herbs, etc.) arose in the early Cretaceous and subsequently radiated to become the dominant plant forms in the succeeding Paleocene. heir success created niches for creatures that could feed on the nectar, nuts, berries, or fruits that lowering plants produce. One result was an explosion in insects, particularly ones that promote plant pollination. Birds and mammals diversiied to exploit the increase in both edible plant parts and insects. Virtually all Cretaceous mammals had sharp cutting ridges on their molar teeth that were well-suited for slicing through insect tissue. he earliest primates probably continued to eat insects, but their molars de-

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Ma

FIGURE 3.15. Two interpretations of early primate phylogeny (adapted from Cartmill [1975], ig. 8). in both, the Paleocene Plesidapiformes, taken here as archaic Primates, diverged from their insectivore (or insectivore-like) ancestors by a greater emphasis on arboreality and herbivory. in both, the irst primates of modern aspect—dating from the eocene and labeled here omomyiformes and adapiformes—evolved following a shift back toward insectivory, based now on binocular vision and grasping extremities. finally in both, increasing emphasis on herbivory was a crucial element in the evolution of the anthropoidea. The main difference between the two interpretations is that in (A), the omomyiformes and adapiformes share a single Paleocene ancestor between 65.5 and 55.5 ma, whereas in (B) they have separate ones in which binocular vision and grasping extremities evolved independently. on present evidence, A is more likely. both interpretations suggest that the omomyiformes and not the adapiformes include the ancestors of the anthropoidea, but this issue remains debatable. The circled x’s mark the loss of the irst premolar (P1) in the Plesiadapiformes. This is one of the specializations that makes them unlikely ancestors for either the omomyiformes or the adapiformes.

EOCENE

34

OLIGOCENE

Ma increasing herbivory

increasing herbivory

Anthropoidea

Anthropoidea

34

Omomyiformes Adapiformes

Omomyiformes

Adapiformes

56.5 PALEOCENE

56.5

LATE CRETACEOUS

65.5

visual predation; 2 binocularity; grasping feet

Plesiadapiformes

Plesiadapiformes

ƒ

loss of P1

ƒ

visual predation; 2 binocularity; grasping feet

arboreality, 1 herbivory

ƒ loss of P1

65.5

arboreality, 1 herbivory

INSECTIVORA INSECTIVORA

A

B 120

120

veloped lower and less pointed cusps, blunter ridges, and other distinctive features that suggest they also fed on fruits, seeds, and other vegetal items. In order to specialize on insects or fruit, early primates were probably at least partly arboreal, and it has long been assumed that a primeval adaptation to life in the trees accounts for such characteristic primate features as grasping extremities (hands and feet), relatively sophisticated vision (relected particularly in orbital convergence), a diminished sense of smell, and a relatively large brain. Grasping extremities assist in movement over branches, and vision is more useful than smell for locating food or for moving safely and rapidly in a three-dimensional, arboreal habitat. Much of the brain in all species is dedicated to muscular control and coordination, and arboreal life places a special premium on the ability to coordinate various muscle groups with each other and with the eyes. However, squirrels prove that active arboreal life does not demand primate grasping extremities with nails, specialized vision involving convergence of the orbits, and expanded brains, and in their extremities, vision, and brains, the Cretaceous ancestors of the primates were probably more like squirrels than like later primates. It is thus necessary to ind a more speciic adaptive explanation for primate specializations. Perhaps the most plausible alternatives are that they promoted visual predation on insects among slender branches in forest undergrowth, that they facilitated access to fruits and lowers growing at the ends of slender

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angiosperm branches, or that they promoted both at the same time. he last possibility is not simply a compromise, since fruits and insects oten occur together, and the tropical American opossum, Caluromys derbianus, uses lower primate-like vision and extremities to feed simultaneously on insects and fruits in terminal branches. Figure 3.15 incorporates the hypothesis stressing enhanced visual predation on insects, and it illustrates two similar but competing views of early primate evolution linked to major dietary shits.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: initial divergence of placental and marsupial mammals (Ji et al. 2002; Novacek 1992); dating the emergence of extant placental orders (Smith and Peterson 2002); early radiation of the angiosperms (Sussman 1991, table I); probable nature of early primate teeth (Rose and Fleagle 1981; Szalay 1972); arboreal adaptation as an explanation for primate specializations (Le Gros Clark 1960) and alternative explanations—visual predation on insects in forest undergrowth (Cartmill 1974, 1975, 1992), access to fruits at the ends of slender branches (Sussman 1991), or access to both insects and fruits on terminal branches (Rasmussen 1986, 2002b)

Plesiadapiformes: The Earliest Primates? he dinosaurs disappeared at the end of the Cretaceous Period, perhaps extinguished by an asteroid impact that initiated a chain of environmental crises. If such an impact occurred, the prime catastrophe was a period of weeks or months when the atmosphere was illed with ine debris that blocked out the sun. In a cladistic sense, the dinosaurs did not vanish, since fossils indicate that birds and dinosaurs were sister taxa, more closely related to each other than dinosaurs were, for example, to crocodiles. However, in the conventional sense, no dinosaurs survived the end of the Cretaceous, and their passing opened ecological opportunities for their bird relatives and for mammals. Both burgeoned in the succeeding Paleocene Period, between 65.5 and 55.5 Ma. he mammals included the Plesiadapiformes, which are usually regarded either as Primates or as close to the line that produced the Primates Most specialists recognize at least sixteen genera of Plesiadapiformes divided among six-to-twelve families. he most proliic plesiadapiform sites are in western North America and in western Europe, which were connected in the Paleocene by a broad, forested land bridge, stretching over what is now the subpolar ocean and the islands in between (ig. 3.14). he climate across the land bridge was subtropical, and the plesiadapiform faunas at both ends were similar. By modern primate standards, the plesiadapiforms were all relatively small, varying from shrew size (100 g or less) to house cat size (roughly 5 kg). hey may all have been omnivorous to a degree, but diferences in body size, dental form, or both suggest that some concentrated more on insects, others more on seeds or fruits. heir skulls, known especially for the type genus Plesiadapis, were primitive, with large snouts and laterally placed (nonconvergent) orbits that indicate a well-developed sense

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Paleocene plesiadapiform (Plesiadapis)

Eocene adapiform (Notharctus) circumorbital (= postorbital) bar

orbit lacking a circumorbital (= postorbital) bar

large, procumbent incisors

reduced number of premolars

small, more vertically emplaced incisors

circumorbital (= postorbital) bar

cm

small, more vertically full complement of premolars emplaced incisors

full complement of premolars

distally short and broad calcaneum laterally compressed, strongly recurved, claw-bearing terminal phalange

Eocene omomyiform (Tetonius)

distally elongated calcaneum

distally hyperelongated calcaneum

broad, flat nail-bearing terminal phalange

FIGURE 3.16. Top: skulls of the Paleocene plesiadapiform Plesiadapis, the eocene lemur-like adapiform Notharctus, and the eocene tarsier-like omomyiform Tetonius (redrawn to the same scale after rose and fleagle [1981]). Middle: reconstructions of Plesiadapis, Notharctus, and Tetonius (redrawn to different scales after rose [1995], 162). Bottom: Terminal (third) phalanges and calcanea (heel bones) of the same taxa (redrawn to different scales after rose [1995], 162). note the large snout, laterally placed, nonconvergent orbits open to the side, large procumbent incisors, and reduced number of premolars in Plesiadapis. Terminal phalanges show that Plesiadapis had claws, whereas Notharctus had nails. Notharctus and especially Tetonius also had elongated calcanea, indicating that they were well-adapted for leaping from branch to branch. in its enhanced ability to leap, Tetonius probably resembled the living tarsier or the african galagos.

of smell and limited overlap between the ields of vision (ig. 3.16). he orbits were open to the sides, as in primitive mammals generally, rather than surrounded by a post- or circumorbital bar, as in later primates. he postcranium appears to have been more variable. In Plesiadapis, the elbows and ankles were relatively mobile and probably facilitated climbing, but bilaterally compressed, pointed terminal phalanges indicate that the digits all retained sharp claws (vs. nails) (ig. 3.16), and there is no evidence for typical primate grasping hands and feet. In contrast, in another well-known genus, Carpolestes, the hallux or big toe carried a nail in place of a claw, and the hallux could be opposed to the other toes for primate-like grasping. Carpolestes had a typical plesiadapiform skull, without primate visual specializations, and if its hallux implies that it was a primate, it might also mean that primate grasping served initially to facilitate feeding on fruits and lowers in terminal branches and perhaps only later to aid visual predation on insects.

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he plesidapiforms are linked to later primates by details of premolar and molar morphology, by the derived aspects of the Carpolestes foot, and by the possibility that Carpolestes and perhaps other genera possessed the characteristic primate petrosal bulla (or middle ear). Yet in overall form, the plesiadapiforms probably resembled modern tropical squirrels, and their dentitions were more specialized than those of their immediate primate successors, discussed in the next section. hus, while their successors tended to have small generalized incisors and to retain the full primitive complement of incisors and canines, the plesiadapiforms had large procumbent central incisors, perhaps used to grasp food, and they had variously reduced the number of lateral incisors, anterior premolars, or both. Barring a highly unlikely evolutionary reversal, this precludes them from the ancestry of later primates, and some or all should arguably be removed to their own order, the Plesiadapiformes or Proprimates, that shared a close Cretaceous ancestor with the Primates. Alternatively, some or all might be allocated to the order Scandentia with the tree shrews or to the order Dermoptera with the colugos. If they are retained in the Primates, they should probably be placed in a separate semiorder and they could be known in the vernacular as archaic primates to distinguish them from the euprimates or primates of modern aspect which are known only ater 55 Ma. Dental specializations not only exclude the known plesiadapiforms from the direct ancestry of the euprimates, they also mean that North America or the combined North American/European landmass is unlikely to have been the euprimate birthplace. South America can probably also be excluded, since its relatively well-known late Cretaceous to Eocene fossil record contains no early primates or likely primate ancestors. Asia has provided at least one plesiadapiform, and it remains a possibility. However, Africa is probably the best candidate, since it hosted so many later major events in primate evolution. An African origin for the euprimates is diicult to investigate because African Paleocene fossil sites formed mainly on the continental margin, and they tend to be poor in terrestrial mammals. Ten isolated teeth from Adrar Mgorn 1, southern Morocco, dated to roughly 60 Ma, suggest the presence of primates, but they are insuicient to determine whether the species involved was a local plesiadapiform or perhaps a more advanced euprimate-like species.

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SOURCES: plesiadapiform diversity (Bloch et al. 2007; Bown and Rose 1991; Conroy 1990; Covert 1986; Fleagle 1999; Rose and Fleagle 1981; Simons 1972; Szalay 1972; Szalay and Delson 1979); similarities between American and European plesiadapiform communities (Gingerich 1973); questionable plesiadapiform status of the Microsyopidae and Apatemyidae (Gingerich 1986b); Paromomyid plesiadapiforms as possible Dermoptera (Beard 1990; Kay 1990; Kay et al. 1992) or not (Bloch and Silcox 2001; Krause 1991); plesidapiform auditory bulla (Bloch and Silcox 2006; Kay et al. 1990, 1992; MacPhee et al. 1983; Szalay and Delson 1979); plesiadapiform diets (Covert 1986); Plesiadapis ankles and elbows (Szalay et al. 1975, 1987) and terminal phalanges (Gingerich 1986b); Carpolestes—foot (Bloch and Boyer 2002; Sargis 2002), skull (Bloch and Silcox 2006), and the possibility that Carpolestes and later pri-

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mates evolved foot features in parallel (Kirk et al. 2003); plesiadapiform specializations lacking in later undoubted primates (Cartmill 1974, 1975; Fox 1993; Rose and Fleagle 1981; Simons 1972); dental resemblances between plesiadapiforms and later primates (Rose 1995); deinition of archaic primates (Fleagle 1999; Szalay and Delson 1979), euprimates (Hofstetter 1977), and Proprimates (Gingerich 1989, 1990b); Asian plesiadapiforms (Smith et al. 2004b); likelihood of euprimate origins in Africa (Gingerich 1986a, 1986b; Hofstetter 1974); African Paleocene fossil sites (Cooke 1978b; Savage and Russell 1983); Adrar Mgorn possible plesiadapiform (Gingerich 1990a; Hooker et al. 1999; Sigé et al. 1990)

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The First Primates of Modern Aspect Most plesidapiforms became extinct before the Paleocene/Eocene transition, 56.5 Ma, and those that survived disappeared soon ater, perhaps in part because of unsuccessful competition with evolving primates of modern aspect. It will be recalled from the last section that the earliest unquestioned primates are known in the vernacular as euprimates. heir oldest widely accepted fossils are jaws and teeth from early Eocene deposits, dated to about 55 Ma, but somewhat later specimens that clearly represent the same taxa include skulls and limb bones (ig. 3.16). hese exhibit a suite of derived primate characters, including a petrosal bulla, a post- or circumorbital bar, orbital convergence, large brains relative to body size, an expanded visual cortex and reduced olfactory bulbs (detected in endocranial casts), and grasping hands and feet with nails on most, if not all, the digits. hey unequivocally document Primates, and their appearance coincides closely with the appearance of the oldest widely accepted representatives of other extant mammalian orders. he early Eocene was geographically and climatically similar to the Paleocene. Western North America and western Europe were still connected by a forested land bridge, and subtropical forests extended as far north as the modern English Channel. he land bridge allowed early Eocene euprimates to disperse widely, and the initial euprimate faunas of North American and Europe were taxonomically similar. Continental drit disrupted the land connection about 50 Ma (ig. 3.14), and the North American and European euprimate communities then became more distinct as they continued to diversify. Specialists disagree on the precise taxonomy of the euprimates, but the especially abundant Euramerican fossils come from more than sixty genera that are commonly divided between two groups—ones that were more lemur- or loris-like and ones that were more tarsier-like. Less abundant fossils demonstrate that one or both groups existed at the same time in southern and eastern Asia and northern Africa. heir presence on the Indian subcontinent is particularly striking, since it was a large island in the Eocene, separated from mainland Asia and Africa by substantial stretches of ocean (ig. 3.14). he implication is that like the Malagasy lemurs discussed above and the New World monkeys discussed below,

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the Indian euprimates must descend from forms that colonized isolated land masses by chance, on rats of loating vegetation. In times past, the lemur-/loris-like euprimates were commonly assigned to the family Adapidae and lumped with the living lemurs in the infraorder Lemuriformes. he more tarsier-like forms were placed in the family Omomyidae and joined with the tarsiers in the Tarsiiformes. Today, most authorities agree that the two euprimate groups were too diverse to be classiied only as families and that the similarities between them and living taxa were overstressed. For present purposes, following the scheme in table 3.1, they are separated as the Adapiformes and the Omomyiformes and listed as if they were Primate semiorders, more closely related to the Strepsirhini and the Haplorhini, respectively. he adapiforms tended to be larger than the omomyiforms, and in overall body size some approached the larger extant Malagasy lemurs. Based on cheek tooth and jaw morphology, most species in both groups probably fed mainly on fruits, but some of the smaller species emphasized insects, and some of the larger adapiforms focused on leaves. In both groups, postcranial bones imply that some species were hindlimbdominated, highly active arboreal quadrupeds like most living lemurs, others were relatively slow arboreal quadrupeds like living lorises, and some had elongated tarsal bones that imply a tarsier- or galago-like ability to leap. Relatively small orbit size indicates that most adapiforms were diurnal, while much larger orbits suggest that most omomyiforms were nocturnal like the living tarsiers or galagos. he adapiforms shared a unique, derived ear structure with living lemurs and lorises, while the omomyiforms shared derived ear features with the tarsiers. Some adapiforms also resembled lemurs and lorises in derived features of the wrist and ankle. Overall, neither the adapiforms nor the omomyiforms exhibited unique specializations that would preclude them from the ancestry of the living lemurs/lorises and tarsiers, respectively, and they difered from living forms most conspicuously in the retention of primitive features. In the adapiforms, these included four premolars on each side of each jaw (vs. three in living lemurs and lorises) and generalized incisors and canines (vs. the protruding, elongated ones that form a specialized dental comb in lemurs and lorises). In the omomyiforms, relative to the tarsiers, the primitive features included a rhinarium (indicated by a large gap between the upper incisors) and the absence of a bar or plate separating the orbit from the skull behind. It thus seems reasonable to suppose that a so far unknown adapiform, accidentally transported to Madagascar on a rat of vegetation 55– 60 Ma, was the direct ancestor of the Malagasy lemurs. Africa is usually assumed to have been the source, but India (or more precisely, the Indian continental plate) is also possible, since it was closer to Madagascar then than it is now (ig. 3.14). Conceivably, the same adapiform or a closely

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related one was also ancestral to the modern African and Asian lorises. A parallel case can be made for tracing the ancestry of the tarsiers to the omomyiforms, although only an early species would qualify, since a partial cranium and isolated teeth from the Shanghuang issure-illings, China, show that the Tarsiiformes themselves (genus Tarsius) were present by roughly 45 Ma. On strictly anatomical grounds, the issue of which group was closer to the ancestry of the anthropoids is more thorny. Omomyiform teeth and skulls are similar to anthropoid ones in some respects, but adapiforms shared features with early anthropoids that neither shared with the omomyiforms. hese include a relatively long snout with nearly parallel tooth rows (vs. a short snout and more divergent tooth rows in omomyiforms); symphyseal fusion of the two halves of the mandible in several adapiforms and all anthropoids but in no omomyiforms; short, vertically placed, spatulate incisors in adapiforms and anthropoids (vs. more procumbent, more sharply pointed ones in omomyiforms); large, interlocking, sexually dimorphic canines in some adapiforms and in anthropoids (vs. small, nondimorphic, premolarlike canines in omomyiforms); and the lack of an external auditory meatus (tubular ectotympanic) in adapiforms and early anthropoids (vs. its presence in omomyiforms). he seeming contradiction relects a common problem in reconstructing ancestral-descendant relationships referred to above—the dificulty of distinguishing between derived similarities that result from close common descent and similarities that evolved in parallel (convergently) as relatively distant lineages adapted to similar circumstances. On fossils alone, a compelling distinction might never be made in the adapiform/omomyiform/anthropoid case, but if the tarsiers derive from an omomyiform and it is accepted on molecular grounds that the tarsiers and anthropoids are sister taxa, more closely related to each other than either is to lemurs or lorises, then the similarities between adapiforms and early anthropoids are likely to be parallelisms, and omomyiforms were probably closer to anthropoid ancestry.

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SOURCES: plesiadapiform extinction (Maas et al. 1988); extant mammalian orders that appear about the same time as the Primates (Beard 2002b); primates of modern aspect deined (Simons 1972); euprimate endocranial casts (Radinsky 1975); similarities among early Eocene euprimates (Gingerich 1984b, 1984c, 1986a; Rose 1995; Rose and Bown 1991); taxonomy of Euramerican euprimates (Conroy 1990; Covert 1986, 2002; Fleagle 1999; Rose and Fleagle 1981; Simons 1972; Szalay and Delson 1979); Asian euprimates (Conroy 1990; Covert 1986; Fleagle 1999; Marivaux et al. 2006; Rose and Fleagle 1981; Simons 1972; Szalay and Delson 1979); African euprimates (Simons and Rasmussen 1994a); diet and ecology of the adapiforms and omomyiforms (Covert 1986, 2002; Soligo and Martin 2006; Strait 1997, 2001b); diversity of adapiforms (Gebo 2002) and omomyiforms (Gunnell and Rose 2002); locomotion in adapiforms (Covert 1986; Rasmussen 1986; Rose 1995; Rose and Walker 1985) and omomyiforms (Covert 1986; Gingerich 1984a); derived similarities between adapiforms and lemurs in ear structure (Martin 1990) and in the ankle and wrist (Beard et al. 1988; Martin 1988, 1990); derived similarities between omomyiforms and tarsiers in ear structure (Beard et al. 1991; Martin 1991b); Chinese, mid-Eocene Tarsius (Beard et al. 1994; Rossie et al. 2006); probable adapiform ancestry of the lemurs and lorises (Gingerich and Schoeninger 1977; Hofstetter 1974); ancestry of the anthropoids—adapi-

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form (Gingerich 1980; Rasmussen and Simons 1988) vs. omomyiform (Delson and Rosenberger 1980; Hofstetter 1974; McKenna 1980; Rosenberger and Szalay 1980; Szalay et al. 1987; Szalay and Delson 1979); primitive features in omomyiforms (Aiello 1986; Martin 1991a); derived features shared between adapiforms and early anthropoids (Gingerich 1980; Gingerich and Schoeninger 1977; Rasmussen 1986; Simons and Rasmussen 1989)

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The Oldest Known Anthropoids Nearly all adapiforms and omomyiforms disappeared before the Eocene/ Oligocene transition, roughly 34 Ma, perhaps in part because terminal Eocene global cooling eliminated much of their habitat and in part because the emerging Tarsiiformes and Anthropoidea appropriated their niches. As mentioned above, the Tarsiiformes had appeared in Asia by the mid-Eocene, 45 Ma, and if their sister-group relationship to Anthropoidea is accepted, the Anthropoidea must also have been present by this time. he oldest known anthropoids may be represented by isolated teeth from Algeria, dated to 50–46 Ma, by jaws from China, dated to 45 Ma, by jaws from Myanmar (Burma), dated to 44–40 Ma, by jaws and teeth from hailand, dated to perhaps 40 Ma, and by jaws and teeth from Egypt dated to roughly 37 Ma. he argument in each instance depends on derived dental characters that the fossil taxa share with later undoubted anthropoids, but the precise characters difer from taxon to taxon, and the possibility exists that they relect functional or adaptive convergence rather than closely shared descent. If not, the anthropoids must have radiated extensively in the early Eocene, before 45 Ma, and the root of the radiation might have been in Asia. A decision on this will require additional fossils, especially skulls with features that appear to be uniquely derived in anthropoids. he single most widely accepted feature is postorbital closure—the presence of a bony plate or septum between the orbit and the braincase behind. his is uniformly lacking in adapiforms, omomyiforms, and strepsirhines, and it may have been a structural response to the static stresses that resulted when early anthropoids repeatedly used their front teeth to open hard-coated fruits or large nuts. Such feeding might also explain why anthropoids early on developed mandibles in which the two halves were fused at the symphysis (“chin”) and in which the symphysis was oten buttressed internally by bony bars or tori (plural of torus). he oldest known skulls that exhibit postorbital closure come from the Fayum Depression of northern Egypt (location in ig. 3.17). Correlation of the Fayum paleomagnetic sequence to the global magnetostratigraphy suggests that the skulls date from 35–34 Ma. he Fayum Depression is desertic today, but sedimentology, vertebrate fossils, and siliciied tree trunks show that it was well-watered and heavily vegetated when early anthropoids were present. Similar conditions extended across northeastern Africa, and the Ashawq Formation in Oman on the Arabian Penin-

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Shanghuang ● ● Ashawq Fm Yangu & Rencun Bir el Ater ● ● Chambi (Thaytiniti & Taqah) ● ● Pondaung Glib Zegdou ● Jebel Qatani Fm ● (Fayum) Wai Lek ● Malembe ●

T ethy s Sea

107 FIGURE 3.17. Top: eocene and oligocene localities with possible or probable fossils of early higher primates (anthropoidea) (redrawn after simons and rasmussen [1994b], 130). by far the most important locality is the fayum, egypt. Bottom: africa during the late eocene and early oligocene, before the development of the red sea that separates africa and the arabian Peninsula today (modeled after Zihlman [1982]).

Ara Pen bian insu la

late Eocene/early Oligocene

sula has provided a smaller sample of fossils that include some of the same early anthropoids found in the Fayum. At the time when the anthropoids were emerging, the Red Sea did not yet exist, and the Arabian Peninsula formed the northeastern corner of Africa. he Fayum and Ashawq fossils not only conirm that anthropoids had evolved by 35–34 Ma, they also document the simultaneous decline of other, more primitive primates. he Fayum fossil primate assemblage numbers more than 1,000 specimens, but the overwhelming majority come from early anthropoids. A handful comes from a possible tarsiiform and from the oldest known lorisiforms (strepsirrhines). Anthropoids also dominate the smaller Ashawq sample. he phylogenetic relationships of the Fayum early anthropoids are not fully resolved, and at least one genus (Arsinoea) is not easily grouped with any others. However, for present purposes, the remaining genera can be divided among three superfamilies: the Proteopithecoidea (Proteopithecus, Serapia), the Parapithecoidea (Parapithecus, Apidium, Qatrania, and Biretia), and the Propliopithecoidea (Oligopithecus, Catopithecus, and Propliopithecus [including Aegyptopithecus]). As discussed near the end of this section, the proteopithecoids may be near the ancestry of the ceboid monkeys (platyrrhines). In times past, some specialists

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I1 I2 C

0

I1 I1I2 C P2 P3 P4 M1 M2

I2 C P3 P4 M1

P3 P4 M1

M2

M2

M3

M3

M3

Apidium phiomense

Propliopithecus chirobates

5 cm

FIGURE 3.18. mandibles of the parapithecoid, Apidium phiomense, and of the propliopithecoids, Propliopithecus chirobates and P. (Aegyptopithecus) zeuxis (partially restored) (drawn by Kathryn Cruz-uribe from photos in szalay and delson [1979], ig. 156; and fleagle and Kay [1983], igs. 4, 9, 10). note the presence of three premolars on each side of the jaw in A. phiomense, versus two in P. chirobates and P. zeuxis. The mandibles and teeth of Propliopithecus are strikingly apelike in overall form and differ from each other only in size and some basic proportions, particularly in the relatively larger size of m1 in P. chirobates. based on mandibular and dental morphology, Propliopithecus could include the ancestors of all later catarrhines.

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placed the parapithecoids near the line leading to the cercopithecoid monkeys, but most today place them in a lineage of basal anthropoids that diverged from the line leading to catarrhines and platyrrhines before they diverged from each other. In contrast, for reasons outlined below, most authorities consider the propliopithecoids to have been primitive catarrhines from which both the cercopithecoids and the hominoids could have evolved. To relect this possibility, table 3.1 lists them as if they were an extinct superfamily within the Catarrhini. Fossils of the early propliopithecoid, Catopithecus, dated to 35–34 Ma, include some of the oldest known primate skulls to exhibit postorbital closure, but it is fossils of the later propliopithecoid, Propliopithecus (including Aegyptopithecus) that shed the least controversial light on catarrhine origins. Depending on what paleomagnetic correlation is accepted, they could date from 33 Ma or they could be 2–3 my younger. All Fayum anthropoids were small by modern anthropoid standards, but Propliopithecus included the largest species, whose estimated male body mass (5.9 kg) approximated that of living gibbons. In its skull, Propliopithecus exhibited a remarkable mix of primitive and derived features that could never have been anticipated in advance. It was derived in the direction of later catarrhines, for example, in its reduced 2-1-2-3 dental formula (ig. 3.18); in the degree to which its canines and lower anterior premolars (P3’s) difered in size between the sexes; and above all in the Y-5 occlusal pattern of its lower molars (ig. 3.8). It will be recalled that the Y-5 pattern characterizes all later undoubted apes, including living ones, and that it contrasts with the bilophodont pattern of the cercopithecoid monkeys. Molar form implies that Propliopithecus fed mainly on fruits, like most later apes, including

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5 cm

white-handed gibbon (Hylobates lar)

banded-leaf monkey (Presbytis melalophus)

Propliopithecus (Aegyptopithecus) zeuxis

living ones, and on molar form alone, it might be regarded as an ancestral ape. However, the oldest known cercopithecoid monkeys, dated to 19–17 Ma, had molars that possessed a ith cusp and incomplete lophs and even cercopithecoids dated to as late as 14 Ma had dentitions that were intermediate between those of apes and those of extant cercopithecoids. he cercopithecoid dental morphology is thus plausibly derived from the more hominoid-like form observed in Propliopithecus, and Propliopithecus is now commonly regarded as a generalized catarrhine that antedated the divergence of the hominoid and cercopithecoid lines. Among other features in which Propliopithecus was derived in the anthropoid direction, the most notable are fusion of the two halves of the mandible at the symphysis, which was buttressed internally by both inferior and superior transverse tori; fusion of the right and let frontal bones; olfactory bulbs that were signiicantly smaller and a visual cortex that was signiicantly larger than in most strepsirhines; and as already indicated, cuplike orbits that, unlike those of strepsirhines or euprimates, were closed of from the skull behind by a postorbital plate or septum (ig. 3.19). his was somewhat less complete than in extant catarrhines, resembling the more primitive condition that platyrrhines retain. he

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FIGURE 3.19. Top: facial and lateral views of the skull of Propliopithecus (Aegyptopithecus) zeuxis (drawn by K. Cruz-uribe from photos in simons [1967]). The mandible has been partially reconstructed from pieces that were not directly associated with the skull. Bottom: lateral view of the same skull compared with skulls of extant catarrhines of roughly similar size (redrawn after bown et al. [1982], ig. 2). note that Propliopithecus zeuxis had a signiicantly longer snout, more laterally placed orbits, and a braincase that was absolutely and relatively smaller than that in the other catarrhines.

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FIGURE 3.20. anterior and distal views of the distal left humerus in a black-andwhite colobus monkey, a red howling monkey, the primitive fayum catarrhine Propliopithecus (Aegyptopithecus) zeuxis, and a patas monkey (redrawn after bown et al. [1982], ig. 3). note that like all extant catarrhines, the colobus and patas monkeys lack an entepicondylar foramen, which does occur in some platyrrhines and in P. zeuxis. note also that in basic distal humerus structure P. zeuxis was more like the colobus and howling monkeys, which are arboreal, than like the patas monkey, which is terrestrial.

black-and-white colobus monkey (Colobus gueraza) arboreal entepicondylar foramen

red howling monkey (Alouatta seniculus) arboreal entepicondylar foramen

Propliopithecus zeuxis

Patas monkey (Erythrocebus patas) terrestrial

orbits were relatively small, and they show that like all living catarrhines, Propliopithecus was diurnal. Among strikingly primitive characters, Propliopithecus had a snout that was somewhat variable in size but that was always longer and more protruding than in later anthropoids; a brain that was bigger for body size than in strepsirhines but still below the lower limit for other anthropoids; more extensive postorbital constriction (narrowing of the skull behind the orbits) than in any other known anthropoid, recalling the condition in some Eocene adapiforms; orbits that faced more laterally than in other known anthropoids; and a bony ear that lacked an external auditory meatus (the tubular extension of the ectotympanic bone that supports the eardrum within the middle ear) (anatomy illustrated in ig. 3.4). Propliopithecus was more uniformly primitive in its postcranium. he known postcranial bones exhibit no specializations for underbranch suspension or arm-over-arm climbing, but they imply instead that Propliopithecus was a slow-moving arboreal quadruped broadly similar to some living monkeys. It probably also had a monkeylike tail. he best known postcranial bone, the humerus, was particularly primitive in retaining an entepicondylar foramen at the distal end (ig. 3.20). his occurs in strepsirhines, in some platyrrhine monkeys, and in some later fossil apelike creatures, but it was lost in the evolution of the extant cercopithe-

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coids and hominoids. In sum, the postcranium shows no specializations that align Propliopithecus exclusively with later catarrhines, but neither does it exhibit any that would preclude it from their ancestry. Propliopithecus exhibited some primitive features that are retained in the platyrrhine monkeys, but it could not have been near their ancestry, if only because it had already developed the reduced dental formula of the catarrhines. It will be recalled that the platyrrhines are more primitive in the retention of a third premolar on each side of each jaw, resulting in a 2-1-3-3 dental formula. he platyrrhines appeared abruptly in South America sometime between 26 and 23 Ma, and by 10 Ma, they had radiated into forms that are remarkably similar to extant species. heir ancestry is murky, but molecular analyses indicate that they share a common anthropoid ancestor with the catarrhines as opposed to a separate euprimate or strepsirhine ancestor. Since some early platyrrhines exhibit striking dental similarities to Fayum early catarrhines, the shared ancestor probably lived in Africa, and it might have been one of the Fayum anthropoids that retained the more primitive platyrrhine dental formula. he proteopithecoid, Proteopithecus, noted above and dated to roughly 35–34 Ma, meets the dental criterion, and it may also anticipate the platyrrhines in its skull and postcranium. However, if a proteopithecoid was ancestral to the platyrrhines, there is still the question of how it reached South America, which was separated from Africa by hundreds of kilometers of open ocean. he best available answer is that like the ancestors of the Malagasy lemurs, it arrived on a rat of loating vegetation, disgorged from the mouth of an African river. Most such rats would have been lost at sea, but over time an occasional one would complete the crossing, and the rare success could also explain how the ancestors of the South American caviomorph rodents arrived from Africa at roughly the same time as the ancestral platyrrhines.

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SOURCES: latest adapiforms (Gingerich and Sahni 1984; Pan 1990); putative early anthropoid fossils from Algeria (Godinot and Mahboubi 1992, 1994), China (Beard et al. 1994; Beard and Wang 2004), Myanmar (Ciochon 1985; Ciochon et al. 1985; Jaeger et al. 1999; Maw et al. 1979; Takai et al. 2001), and hailand (Chaimanee et al. 1997; Ducrocq 1998, 1999; Ducrocq et al. 2006); possibility that some putative Burmese early anthropoids were advanced adapiformes (Gingerich 1980); possible Asian origin of the anthropoids (Beard 2002a; Jaeger and Marivaux 2005; Marivaux et al. 2005) and alternatives (Miller et al. 2005a); functional signiicance of postorbital closure (Rosenberger 1986); Ashawq early anthropoids (Rasmussen and Simons 1992; homas et al. 1991); oldest known strepsirrhines (Seifert et al. 2003, 2005b); dating of Fayum fossiliferous deposits (Fleagle et al. 1986a, 1986b; Gingerich 1993; Kappelman 1992, 1993; Kappelman et al. 1992; Rasmussen et al. 1992; Seifert 2006; Simons 1967, 1984, 1989; Van Couvering and Harris 1991); Fayum primates that possibly or probably date from the late Eocene, before 34 Ma (Rasmussen and Simons 1992; Seifert et al. 2000, 2005a; Simons 1990, 1993; Simons and Rasmussen 1989, 1994a, 1994b); Fayum primates that date from the early Oligocene ater 34 Ma (Fleagle 1986b; Fleagle et al. 1986b; Simons et al. 1986, 1987); Fayum primates—paleoenvironment (Bown et al. 1982; Bown and Kraus 1988; Olson and Rasmussen 1986), taxonomy (Beard 2002a; Fleagle et al. 1986b; Fleagle and Kay 1985; Gingerich 1977, 1980; Kay et al. 1981; Rasmussen 2002a; Rasmussen and Simons 1988; Simons et al. 1987; Simons and Rasmussen 1991), body size and ecology (Fleagle 1978; Fleagle and Kay 1985; Kay and Simons 1980), locomotion (Gebo 1989; Gebo and Simons

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1987), diurnal habitus (Fleagle and Kay 1985), relationship to later primates (Gebo and Simons 1987; Simons 1967, 1972, 1987), as generalized anthropoids (Andrews 1985a; Fleagle 1986b, 1994; Fleagle et al. 1986b; Fleagle and Kay 1983; Harrison 1987; Kay et al. 1981; Simons 1984; Simons and Rasmussen 1994b); parapithecoid dentition (Kay and Simons 1983; Simons 1986); parapithecoids as a unique anthropoid branch (Fleagle and Kay 1987; Harrison 1987); dental resemblances between Propliopithecus and early cercopithecoids (Beneit and McCrossin 2002; Delson 1979); dietary implications of bilophodonty in the earliest cercopithecoids (Andrews 1981c; Temerin and Cant 1983); Propliopithecus—unexpected mix of primitive and derived characters (Kay et al. 1981; Simons 1987), brain (Radinsky 1977; Simons et al. 2007), locomotion—(Bown et al. 1982; Fleagle 1983), and dental variation (Kay et al. 1981); Proteopithecus and platyrrhine origins (Miller and Simons 1997; Simons 1997; Simons et al. 1999; Simons and Seifert 1999); oldest known platyrrhine fossils (Fleagle and Tejedor 2002; Hofstetter 1969, 1974, 1980; MacFadden 1990; Takai et al. 2000; Takai and Anaya 1996; Wolf 1984) and their dating (Kappelman 1993; MacFadden 1985, 1990; MacFadden et al. 1985; McRae 1990; Naeser et al. 1987); subsequent diversiication of the platyrrhines (Bown and Larriestra 1990; Fleagle 1986a, 1990, 1999; Gingerich 1980; Hartwig and Meldrum 2002; Hofstetter 1980; Kay 1990; Kay et al. 1987; MacFadden 1990; Rosenberger et al. 1990, 1991; Schrago 2007); dental similarities between Fayum early anthropoids and platyrrhines (Fleagle 1986a); geographic origin of platyrrhine ancestors (Ciochon 1985; Ciochon and Chiarelli 1980; Gingerich and Schoeninger 1977; Hartwig 1994; Hofstetter 1974, 1980; Rosenberger 1986) and probable rating across the south Atlantic (Gingerich and Schoeninger 1977; Hofstetter 1974; McKenna 1980); the broadly contemporaneous rating of caviomorph rodents to South America (Fleagle 1986b; MacFadden 1990; Poux et al. 2006)

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Hominoid Origins he biomolecular clock suggests that hominoids diverged from cercopithecoids in the early to middle Oligocene, around 30 Ma (ig. 3.13). However, the oldest fossils to document the split come from early Miocene sites in eastern Africa, dated between 22 and 17 Ma. Both hominoids and cercopithecoids are represented, though cercopithecoids are much rarer. he hominoids are plausibly derived from Fayum early Oligocene catarrhines, but 9–10 my separate the two groups (ig. 3.21). So far, only three isolated teeth and two fragmentary jaws from Lothidok, northwestern Kenya, provide a potential intermediary. Radiopotassium dating places the Lothidok fossils between 27 and 24 Ma, and they have been assigned to the genus Kimoyapithecus. Dental similarities to later east African hominoids, particularly Afropithecus, dated to roughly 17 Ma, suggest that Kimoyapithecus may also have been a hominoid. Hominoids burgeoned in the Miocene, between roughly 23 and 6 Ma, and the reasons have much to do with Miocene geography. During the earliest Miocene, before 20 Ma, the continents reached basically their current positions, but the Arabian Peninsula still formed the northeastern corner of Africa, and Africa was separated from Eurasia by the protoMediterranean or Tethys Sea (ig. 3.22). Sometime between 20 and 17 Ma, the northward drit of the African (or Afro-Arabian) plate brought it into contact with Europe and southwestern Asia. Initially, the land connection was intermittent, but between 15 and 14 Ma it became permanent. As the Tethys Sea shrunk, its climatic inluence on the adjacent continents was reduced. Southern Eurasia became generally cooler and drier,

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The Pr im aTe b aCKGr ound

Ma

late

Pliocene

15

20

Miocene

10

middle

5

Pleistocene

early

0

Pan

Gorilla

Homo

Homo Paranthropus Australopithecus Ardipithecus Orrorin

Sahelanthropus

Pongo Hylobates

Ma 0

Homo Gigantopithecus

Graecopithecus Oreopithecus Ankarapithecus Dryopithecus

hominoid distributions sharply reduced in E. Asia Lufengpithecus

5

10

Sivapithecus

Pliopithecus & Anapithecus Pierolapithecus

Otavipithecus

Kenyapithecus Nyanzapithecus Equatorius & Nacholapithecus ? Simiolus Afropithecus & Turkanapithecus Morotopithecus Limnopithecus, Micropithecus & Rangwapithecus ? Dendropithecus Proconsul

Griphopithecus first hominoids in Europe & W. Asia?

15 Dionysopithecus first hominoids in E. Asia?

20

25

Kimoyapithecus

Oligocene

EASTERN ASIA ?

hominoids probably extinct in Europe & W. Asia

meager fossil record, but presumed origin Nakalipithecus & Samburupithecus of hominins, Chororapithecus Pan & Gorilla

25

30

EUROPE & WESTERN ASIA

AFRICA

limited fossil record, but major catarrhine radiation

30 Propliopithecus Catopithecus

35

35

FIGURE 3.21. Temporal distribution of fossil hominoid genera discussed in the text (based partly on information in begun [2002]; harrison [2002]; Kelley [2002]; Ward and duren [2002]). note the gaps in the african fossil record between roughly 31 and 22 ma and between roughly 12 and 5 ma. The irst gap was almost certainly a time of major catarrhine evolution when the hominoids differentiated from the cercopithecoids, and the second gap includes the time when the line leading to hominins diverged from the lines leading to the chimpanzee and the gorilla. hominoids are unknown in eurasia before about 17 ma, probably because they evolved in africa and were unable to reach eurasia until the contact between africa and eurasia was signiicantly broadened about 17–16 ma. in the late miocene, about 8 ma, hominoids became extinct in europe and their numbers and distribution were greatly reduced in asia, probably because climatic change eliminated suitable habitat in middle latitudes.

and the trend was accelerated by broadly simultaneous mountain building in the Mediterranean and Himalayan regions. Equally important, the newly developed land connection promoted an unprecedented degree of faunal interchange. Among the African species that migrated to Eurasia were hominoids. By 15 Ma, they had spread widely through Eurasian broad-leaved forests, from Spain on the west to China on the east. he acceleration of faunal exchange between Africa and Eurasia roughly 17 Ma separates two phases in hominoid evolution—an early one in which hominoids were exclusively African and were so primitive that their hominoid status can be questioned and a later one in which they were also widely distributed in Eurasia and some anticipated the living great apes in their skulls, bodies, or both. SOURCES: Miocene history of the Tethys Sea (Laporte and Zihlman 1983; Rögl 1999); dating of Lothidok (Boschetto et al. 1992); description of Kimoyapithecus (Leakey et al. 1995b)

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FIGURE 3.22. relative positions of africa and eurasia in the early and middle miocene (modiied after laporte and Zihlman [1983], igs. 3 and 4). The oldest eurasian higher primates probably evolved from african forms that dispersed to eurasia when the northward drift of the african plate created a land connection between 20 and 17 ma.

●

●● ● ●●

●

● ●●

● ●● ● ● ●● ●● ● ●

●

●

● ●

●

Tethys Sea ● ●●● ●●

● catarrhine

fossil site

Early Miocene (ca. 22 - 17 Ma)

●

●

hominoid fossil site

Middle Miocene (ca. 16 - 10 Ma)

The Oldest Known Hominoids

So far, early Miocene hominoid fossils come overwhelmingly from sites in shallow sedimentary basins near modern Lake Victoria in western Kenya and neighboring Uganda (ig. 3.23). he fossiliferous layers are interstratiied with alkaline volcanic ashes and lavas that helped to preserve the bones from acid dissolution and that have provided radiopotassium dates conirming an early Miocene age, between about 22 and 17 Ma. Associated plant and animal fossils indicate that the east African early Miocene hominoids occupied tropical forest and woodland. Forest prevailed and stretched along the Equator across the continent into areas that became woodland, bushland, or savanna ater 17–16 Ma. Hominoids dominate the primate samples, but prosimians similar to modern lorises and galagos also occur, and there are occasional cercopithecoid monkey jaws and teeth. he monkeys had primitive molars that were incompletely bilophodont. he best known hominoids had a primitive monkeylike postcranium, and the relative rarity of early Miocene cercopithecoids suggests that the more abundant hominoids occupied niches that monkeys took over later. At least early on, cercopithecoids may have been more at home in less forested environments, such as those that existed in the early Miocene of northern Africa, where cercopithecoids occurred without hominoids. he major cercopithecoid adaptive radiation probably occurred only much later, between 10 and 4 Ma, at a time when hominoid fortunes were in decline, owing at least in part to progressively cooler, drier global climate and to the accompanying growth of savannas at the expense of forests and woodlands. Despite a growing fossil sample and more thorough, more sophisticated analyses, the taxonomy of the Miocene hominoids has been revised many times, and highly qualiied specialists continue to hold widely divergent views. Some disagreement is inevitable, since taxonomic judgments must oten be based on small samples, dominated by partial jaws and isolated teeth. Skulls and postcranial bones are much rarer, and it is oten

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30o

35o

40o

FIGURE 3.23. approximate locations of the main african miocene fossil hominoid sites (partly after Kunimatsu et al. [2004], 367).

Ad Dabtiya ■ Chorora

115

■

Berg Aukas ■

SUDAN

5o

Lothidok (late Oligocene) Kalodirr ■ Napak ■ ■ Moroto

D. R. CONGO

■ Bukwa

ETHIOPIA

■ Buluk

■ Nachola ■ Nakali & Samburu ■

■ Ngorora & Kipsaramon (Tugen Hills) Songhor & Mteitei Maboko ■ ■■ Fort Ternan ■ Meswa, Legetet & Koru Mfwangano ■■ Rusinga

UGANDA 0o

❂ Nairobi

KENYA

L. Victoria TANZANIA

5o

■ early Miocene locality ■ mid or late Miocene

N

0

locality highlands over 1500 m 300 km

diicult to ascertain which postcranial bones belong with which teeth. In these circumstances, the number of characters available for comparison tends to be small, and it can be hard to separate interspeciic variability from intraspeciic variability, particularly intraspeciic variability due to sexual dimorphism. he problem is exacerbated by the possibility that some fossil hominoid species may have been more dimorphic than their living counterparts, which are thus misleading guides. Finally, and perhaps most intractable, there is the perennial diiculty of determining which evolutionary novelties that fossil taxa share relect close common descent (homology) as opposed to common adaptation (analogy) or evolutionary parallelism (homoplasy). Evolutionary parallelism or convergence is especially likely to occur among closely related species adapting to similar conditions, and it could explain, for example, why postcranial features that anticipate those of living hominoids recur in Miocene species whose crania or geographic locations imply divergent ancestries.

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With these challenges in mind, at least provisionally, most specialists would probably accept a division of early Miocene hominoid fossils among eight genera: Proconsul, Rangwapithecus, Afropithecus, Turkanapithecus, Morotopithecus, Limnopithecus, Micropithecus, Dendropithecus, and Simiolus (temporal positions in ig. 3.21). Proconsul was primitive enough to be on or near the line leading to most subsequent hominoids, including the living great apes and humans. Morotopithecus might have evolved from an early form of Proconsul, and it is the oldest known species to exhibit some postcranial specializations of the living great apes. Dendropithecus had lightly built limb bones like those of the living hylobatids (gibbons and siamangs), and it was similar to them in size. However, it possessed no derived hylobatid anatomical features, and together with Rangwapithecus, Limnopithecus, Micropithecus, Afropithecus, Turkanapithecus, and Simiolus, it is noteworthy mainly for revealing the extraordinary diversity of early Miocene hominoids. Afropithecus, Turkanapithecus, and Simiolus underscore the point, since they come mainly from sites in northern Kenya where Proconsul, Morotopithecus, Dendropithecus, and the other genera are unknown. An Afropithecus-like form is also represented in deposits dated to about 17 Ma at Ad Dabtiyah, Saudi Arabia, and the sum suggests that early Miocene hominoid diversity would be even greater if it were possible to sample everywhere in Africa where early Miocene hominoids probably lived. Proconsul and Morotopithecus deserve special consideration for their potential bearing on later hominoid evolution. Proconsul has produced the large majority of early Miocene hominoid fossils, and they have been irmly dated to between 20 and 17 Ma. Dentitions and isolated teeth dominate, but there are also some partial skulls and postcranial bones. Size alone implies that Proconsul comprised at least three diferent species. Average adult weight ranged from 11 kg in the smallest (siamang size) to 87 kg in the largest (orangutan or female gorilla size). he two largest species were far larger than any known Oliogocene catarrhine, and they highlight the evolutionary gulf that separates the early Miocene hominoids from even the most advanced Oligocene species. Proconsul heseloni, which was siamang size, and P. nyanzae, which was pygmy chimpanzee size, illustrate Proconsul anatomy most fully. In its skull (ig. 3.24, bottom), Proconsul was far advanced over Oligocene Propliopithecus, and it resembled the living apes in the full forward rotation of the orbits, the limited amount of postorbital constriction, the small size of the snout relative to the braincase, and the large size of the brain, both absolutely and relative to body size. Dentally, Proconsul resembled both Propliopithecus and the living apes, and in both cranial and dental features, it is a plausible Miocene link between them. It lacked the secondarily enlarged incisors (for peeling fruit) of the chimpanzee and orangutan and the especially high-cusped molars (for masticating

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117 FIGURE 3.24. Above: reconstruction of Proconsul heseloni (rusinga island specimen no. 2036 in the collections of the national museums of Kenya) (redrawn after Walker and Pickford [1983]). Below: reconstructed skull of female P. heseloni (redrawn after Walker et al. [1986b]). P. heseloni was formerly called P. africanus (Walker et al. 1993).

25 cm

0

5 cm

0

Proconsul heseloni (KNM-RU 2036) leaves) of the gorilla and was thus generalized enough to be ancestral to all three. Its primitive, low-cusped, thin-enameled molars suggest a basically frugivorous diet. In its postcranium (ig 3.24, top), Proconsul was more complex. Its arms and legs were subequal in length, as in living monkeys, and the wrist was adapted to bear the weight of the forequarters, probably on the palms rather than on the knuckles. he upper humerus shat was curved to accommodate a deep, narrow monkeylike chest, and the lumbar (waist) portion of the vertebral column was monkeylike in its length and lexibility. Limb proportions, torso shape, and other features suggest that Proconsul was mainly pronograde (with the spinal column parallel to the ground) and quadrupedal in the manner of living monkeys. However, it also had relatively mobile joints, particularly at the elbow, that may have allowed it to hang below branches, and it may have lacked a tail. Its nearest living analogs may be those New World monkeys that

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combine arboreal quadrupedalism with an enhanced capacity for hanging below branches. Like the skull and dentition, the postcranium exhibits no specializations that exclude Proconsul from the ancestry of the living apes. Morotopithecus is much less well-known than Proconsul and less irmly dated. he fossil sample, which comes exclusively from the Moroto site, Uganda, includes a palate and partial face, two fragmentary mandibles, parts of two femurs, a nearly complete middle lumbar vertebra, fragments of four other vertebrae, and a proximal scapula. Body weight, inferred mainly from the femora, was probably between 40 and 35 kg, similar to a female chimpanzee. Geologic age could be greater than 20.5 Ma, based on disputed 40Ar/39Ar dates, or 17.5 Ma, based on the associated mammalian species. Dentally, Morotopithecus broadly resembled other early Miocene hominoids, including Proconsul and Afropithecus, but postcranially, it may have been much more like a modern ape. he complete lumbar vertebra suggests a short, stif waist region, the scapula hints at an apelike ability to rotate the arm around the shoulder, and the femurs suggest signiicant hip and knee mobility, less than in living apes, but greater than in monkeys. he sum implies that Morotopithecus was equipped for apelike hand-over-hand climbing and underbranch suspension, and its anatomy and body size together might then place it near the ancestry of the living great apes. his would imply that the great apes split from the hylobatids as much as 20.5 Ma, in contrast to biomolecular estimates that variously place the split between 18 and 14 Ma. In bodily form and function Morotopithecus was advanced over not only Proconsul and Afropithecus but also over all their known immediate successors, including, for example, the African mid-Miocene genera Equatorius and Nacholapithecus, for both of which there are partial skeletons that imply Proconsul-like bodies. Subsequent to Morotopithecus, the oldest known hominoids with apelike postcrania are the European genera Pierolapithecus, Dryopithecus, and Oreopithecus, all of which postdate 14 Ma (temporal positions in ig. 3.21). he absence of other more apelike species before this time may mean that Morotopithecus independently evolved its apelike postcranial features and that it had nothing to do with great ape ancestry. However, if parallel or convergent evolution is ruled out and apelike postcranial specializations are the hallmark of the hominoids, then to the degree that Morotopithecus and Proconsul overlapped in time, Proconsul and other equally primitive hominoids might have to be removed to a divergent, extinct family, the Proconsulidae, that does not bear on the ancestry of any living species. hey could even be removed from the Hominoidea altogether and placed in a separate superfamily, the Proconsuloidea, that inherited some similarities to the hominoids from a shared Oligocene catarrhine ancestor and

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that may have evolved others in parallel. he issue is unsettled, but table 3.1 accepts Proconsulidae within the Hominoidea.

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SOURCES: East African early Miocene sediments (Bishop 1971; Pickford 1986b; Van Couvering and Van Couvering 1976), environments (Andrews 1981a, 1981b, 1992; Andrews and van Couvering 1975; Pickford 1983), prosimians (Walker 1978a), and cercopithecoids (Beneit and McCrossin 2002; Harrison 1987; Strasser and Delson 1987); north African early Miocene cercopithecoids (Delson 1979); late Miocene/early Pliocene cercopithecoid radiation (Jablonski 2002); Miocene hominoid taxonomy— Africa (Harrison 2002; Ward and Duren 2002), Europe and western Asia (Begun 2002), eastern and southern Asia (Kelley 2002), and general (Andrews 1985b, 1992; Begun 1991; Ciochon 1983; Finarelli and Clyde 2004; Harrison 1987; Kelley and Pilbeam 1986; Pilbeam 1996; Young and MacLatchy 2004); overviews of Proconsul and other east African Miocene hominoids (Walker and Shipman 2005), Afropithecus (Leakey et al. 1988a; Leakey and Leakey 1986a; Leakey and Walker 1997), Turkanapithecus (Leakey et al. 1988b; Leakey and Leakey 1986b), Morotopithecus (MacLatchy 2004), Limnopithecus, Micropithecus, and Dendropithecus (Andrews 1980; Fleagle 1984; Fleagle and Simons 1978), and Simiolus (Leakey and Leakey 1987); Ad Dabtiyah (Andrews 1992; Andrews and Martin 1987a, 1987b; Andrews et al. 1987); Proconsul—body size (Raferty et al. 1995), overall anatomy (Walker and Pickford 1983; Walker et al. 1986b, 1993; Ward et al. 1993), advanced features in the skull (Walker et al. 1983), dentition and diet (Fleagle and Kay 1985), wrist (McHenry and Corruccini 1983), rib cage and vertebral column (Ward 1993; Ward et al. 1993), locomotion (Andrews 1992; Fleagle 1983; Rose 1983, 1994; Walker and Pickford 1983; Ward 1993), tail absent (Ward et al. 1991) or present (Harrison 1998); Morotopithecus—geologic age (Pickford et al. 1999, 2003), dentition (Patel and Grossman 2006), lumbar vertebra (Sanders and Bodenbender 1994; Ward 1993), and femur and scapula (Gebo et al. 1997; MacLatchy et al. 2000); biomolecular vs. anatomical estimates for the divergence of great apes and hylobatids (Young and MacLatchy 2004)

The Oldest Known Hominoids of Modern Aspect

During the middle and later Miocene, beginning 16–15 Ma, global climatic change produced generally cooler conditions in middle and upper latitudes and drier ones in lower latitudes, at least seasonally. In eastern Africa, drier and probably more seasonal climate combined with largescale crustal movement (riting) to reduce the extent of tropical forest, which was largely replaced by more open tropical woodlands in the middle and later Miocene. As drier, more seasonal woodlands spread, the distribution of fruits probably became patchier and more seasonal, and other vegetal items became more attractive as dietary staples. hese staples included nuts, seeds, subterranean tubers, or other hard or gritencrusted foods that could help account for dental novelties in the east African hominoid Kenyapithecus and its possible Eurasian descendants, discussed below, and also leaves that could help explain the greater abundance of cercopithecoid monkey fossils in east African mid-Miocene sites. he molars were more fully bilophodont than in earlier Miocene cercopithecoids, but only one cercopithecoid genus has been identiied, and it appears to antedate the divergence of cercopithecines and colobines. he biomolecular clock places the divergence sometime between 17.9 and 14.4 Ma, and undoubted (derived) colobine fossils from eastern Africa and Namibia show that it had occurred by 10.5 Ma.

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ChaP Ter Thr ee SOURCES: middle and late Miocene vegetational change in eastern Africa (Andrews 1981b; Andrews and Kelley 2007; Laporte and Zihlman 1983; Pickford et al. 1983); increased abundance of cercopithecoid fossils in mid-Mocene sites (Harrison 1987; Pickford and Kunimatsu 2005); taxonomy of midMiocene cercopithecoids (Beneit and McCrossin 1997, 2002; Beneit and Pickford 1986; Delson 1979; Delson and Andrews 1975; McCrossin and Beneit 1994; Strasser and Delson 1987); cercopithecine/colobine divergence—biomolecular dating (Raaum et al. 2005) and relevant fossils (Andrews et al. 1996; Barry et al. 1982; Beneit and Pickford 1986; Conroy et al. 1996; Heinz et al. 1981)

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Africa In eastern Africa, hominoid fossils that date ater 17 Ma and especially ater 12 Ma are less abundant than earlier ones, but they show that taxonomic diversity remained high. he number of genera is debatable, but there were probably at least seven: Nachalopithecus, Equatorius, Kenyapithecus, Nyanzapithecus, Chororapithecus, Samburupithecus, and Nakalipithecus. Sparse fossils suggest that Proconsul or a descendant may have persisted in eastern Africa until 14 Ma or even later. Eurasian fossils, addressed briely below, demonstrate that by 16–15 Ma hominoids occurred far outside the range of any living species, and Otavipithecus, dated to 14–13 Ma at Berg Aukas, northern Namibia, shows that this was true even in Africa (ig. 3.25). Since many potentially relevant African regions have not or cannot be sampled, mid-Miocene hominoids were surely more diverse than the fossil record currently indicates. Nacholapithecus and Equatorius are represented by partial skeletons, which identify them as monkeylike quadrupeds that difered only in detail from Proconsul. Kenyapithecus (narrowly deined to exclude specimens formerly assigned to Nacholapithecus and Equatorius) so far lacks informative limb bones, but it shared a robust mandible, relatively short, stubby canines, enlarged upper premolars, thick enamel, and other derived dental features with species of the early hominin, Australopithecus. he next chapter shows that Australopithecus comprised multiple species and they were all hominins by deinition, since they were habitual bipeds. If it could be assumed that hominins are equally unique in their dentitions, then Kenyapithecus might be regarded as the oldest known form. However, it is dated to 14–12 Ma, and biomolecular estimates indicate that hominins diverged from chimpanzees only ater 7–8 Ma. More important, Kenyapithecus also shared its Australopithecus-like dentition with the Eurasian later Miocene hominoids Graecopithecus (Ouranopithecus), Sivapithecus, and Gigantopithecus, discussed below but not with the African terminal Miocene/early Pliocene hominoid Ardipithecus. As noted below, facial similarities suggest that Sivapithecus was on or near the line that led to the orangutan, while leg bones indicate that Ardipithecus was a bipedal hominin, perhaps directly ancestral to Australopithecus. he sum implies that Kenyapithecus and Australopithecus evolved their shared dental features independently, probably as they adapted to similar tropi-

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FIGURE 3.25. fossil hominoid localities dating between about 16–15 and 8–7 ma in relation to the historic distribution of the chimpanzees and the gorilla (redrawn after Conroy et al. [1992, 356]). note the concentration of fossil sites in equatorial eastern africa and their spread across midlatitude europe from spain to China.

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cal or subtropical woodlands where they came to rely on relatively hard or gritty vegetal foods, at least seasonally. Kenyapithecus remains a plausible ancestor for Sivapithecus and the other dentally similar late Miocene Eurasian genera, but the case is not compelling, and convergent evolution could once again explain the shared dental features. Among the remaining genera, Samburupithecus, Chororapithecus, and Nakalipithecus are notable because each was in the right place— eastern Africa—at the right time—between 11 and 9 Ma—to have been on or near the line that included the last shared common ancestor of the African great apes and hominins. Each also existed at a time when less wooded, more seasonal conditions were spreading at the expense of forests in equatorial Africa, and as discussed below, the resulting change in selection pressures may lie behind the separation of hominins from the African great apes narrowly understood. However, at present, each genus is too sparsely represented—by a partial maxilla (Samburupithecus), nine whole or partial teeth (Chororapithecus), and a partial mandible and eleven isolated teeth (Nakalipithecus)—for persuasive phylogenetic placement. Chororapithecus has been likened dentally to the gorilla, but

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the fossil support is slim, and only the emergence of hominins by 6–5 Ma demonstrates that the gorilla line existed before then. If Chororapithecus is placed aside, the gorilla is known only in living form, and the same is true of the chimpanzee, except for three-to-four teeth dated between 540 and 500 ka (thousands of years ago) in the Lake Baringo basin of southwestern Kenya and the possibility, mentioned in the next chapter, that the Ardipithecus sample includes fossils of an early chimpanzee. It may always be diicult to document early chimpanzee evolution, if as seems likely, most skeletal elements of early chimpanzees closely resembled those of early hominins.

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SOURCES: possible mid-Miocene persistence of Proconsul (Hill 1994; Hill et al. 1985); Nacholapithecus (Ishida et al. 2004; Kunimatsu et al. 2004; Nakatsukasa et al. 1998, 2003); Equatorius (Kelley et al. 2002; Sherwood et al. 2002b; Ward et al. 1999b); Kenyapithecus (in the narrow sense) (Pickford 1985, 1986a); Nyanzapithecus (Harrison 1986a, 1986b); Chororapithecus (Suwa et al. 2007b); Samburupithecus (Ishida and Pickford 1997; Pickford and Ishida 1998); Nakalipithecus (Bernor 2007; Kunimatsu et al. 2007); Otavipithecus (Conroy 1994; Conroy et al. 1992, 1993a, 1993b, 1996; Schwartz and Conroy 1996; Singleton 2000); Ardipithecus (Haile-Selassie 2001; Haile-Selassie et al. 2004b; White et al. 1994,1995; WoldeGabriel et al. 2001); Baringo chimpanzee teeth (McBrearty and Jablonski 2005)

Eurasia It was pointed out above that a land connection developed between Africa and Eurasia sometime between 20 and 17 Ma, that it became permanent about 15 Ma, and that it allowed hominoids to spread from Africa into broad-leaved woodlands that stretched across southern Eurasia from Spain on the west to China on the east. hey lourished in southern Eurasia until 8–7 Ma, when adverse climatic change eliminated them from Europe and reduced their Asian distribution to the extreme southeast, including southern China. he irst African hominoid to have crossed the land bridge from Africa may have been a small-bodied Dendropithecus-like or Micropithecuslike form that gave rise to Dionysopithecus, dated to 18–16 Ma at sites in hailand and southern China. Dionysopithecus could have been near the ancestry of the mid-Miocene, mainly European hominoids Pliopithecus and Anapithecus, while other African emigrants gave rise directly or indirectly to at least nine additional middle and late Miocene Eurasian genera: Griphopithecus, Pierolapithecus, Dryopithecus, Oreopithecus, Graecopithecus, Ankarapithecus, Sivapithecus, Gigantopithecus, and Lufengpithecus (temporal positions in ig. 3.21). he relationships of the various Eurasian hominoids to each other and to living forms are controversial. Like east-African early Miocene Dendropithecus, Pliopithecus resembled the hylobatids in its body size and lightly built skeleton (ig. 3.26). However, again like Dendropithecus, it was a monkeylike quadruped, and it exhibited no derived hylobatid features. In some respects, including the incomplete ossiication of its

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123 FIGURE 3.26. reconstructed skeleton of Pliopithecus vindobonensis from deposits dated to ca. 15 ma in slovakia (redrawn after Zapfe [1960], ig. 106). in its short broad face, slender postcranial bones, and other features, P. vindobonensis resembled the living gibbons and siamangs, but it lacked their extraordinarily long arms and other morphological specializations for brachiation. it also retained some remarkably primitive features, such as an incompletely ossiied external auditory meatus and an epicondylar foramen on the distal humerus. most authorities now regard it as one of a group of closely related small hominoids or hominoid-like anthropoids that inhabited western and central european forests between roughly 16 and 11 ma and that bear no relation to any later hominoids.

external auditory meatus and the retention of an entepicondylar foramen on the distal humerus, it was remarkably primitive. he peculiar mix of features suggests that Pliopithecus, the closely related Anapithecus, and their putative ancestor Dionysopithecus, should be assigned to their own mid-Miocene family, the Pliopithecidae, within the Hominoidea or, perhaps, within the Propliopithecoidea, separate from the Hominoidea. Placement within the Propliopithecoidea would imply descent from Propliopithecus or a related Oligocene catarrhine through an as yet uncertain early Miocene form. he removal of Dendropithecus, Pliopithecus, Dionysopithecus, and like genera from hylobatid ancestry leaves the hylobatids without a fossil record before the appearance of modern or near-modern Hylobates at sites that postdate 2 Ma across southeastern Asia. Griphopithecus, dated to roughly 16.5 Ma in Turkey, was roughly chimpanzee-size, and it may have been the irst large-bodied hominoid to reach Eurasia. It is known mainly from isolated teeth, but there are also a few partial jaws and some hand and foot bones. In its robust mandible, thick enamel, and reduced canines, it was similar to Kenyapithecus, and if it was actually geologically more ancient, as the available dates suggest,

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it might imply that Kenyapithecus evolved in southeastern Europe and spread back to eastern Africa. Dryopithecus was one of the irst fossil hominoids to be described (or the irst, if Pliopithecus is removed to the Propliopithecoidea). he genus was devised for French fossils in 1856, and it now comprises four-toive siamang-to-chimpanzee size species found at sites that date between about 12 and 8 Ma in western and central Europe. As noted previously, the Y-5 cusp and issure coniguration that characterizes hominoid (as opposed to cercopithecoid) molars is oten known more fully as the dryopithecine Y-5 pattern because it was irst recognized in Dryopithecus. Also as noted before, the same pattern characterized the Oligocene catarrhine Propliopithecus, and cercopithecoid bilophodonty is thought to have evolved from it. In its cranium and dentition, Dryopithecus broadly resembled earlier Miocene hominoids, including Proconsul. In its postcranium, however, it diverged sharply from Proconsul in the direction of the living great apes. Recall that Proconsul had a primitive, monkeylike torso and limb proportions, which combine to show that it was habitually pronograde and quadrupedal. In contrast, Dryopithecus had a derived, apelike body, with a shortened, relatively inlexible lumbar vertebral (waist) skeleton, a broad, lat thorax (chest), scapulae (shoulder blades) that were situated behind the thorax (rather than alongside it), and long powerful arms that could rotate around the shoulder joint. In living apes, these features promote orthograde (upright) posture (with the spinal column perpendicular to the ground) and an enhanced ability to hang below branches or to climb hand over hand. Pierolapithecus, which could be immediately ancestral to Dryopithecus and Oreopithecus, which could be its descendant, had broadly similar apelike bodies, and Oreopithecus certainly possessed the same apelike ability to climb and hang. Whether Pieralopithecus, Dryopithecus, and Oreopithecus evolved their apelike features independently in Europe or inherited them from an older shared African ancestor like Morotopithecus, they would surely qualify as great apes if they had survived to the present. Dental peculiarities indicate that Oreopithecus could not be on the line leading to any living great ape, but Pierolapithecus and especially Dryopithecus may have been closely related to the last shared ancestor of the African apes and hominins. Graecopithecus was a chimpanzee-size ape known by jaws, teeth, and a partial face from deposits dated between 11 and 9 Ma in Greece and Turkey. It was remarkably Australopithecus-like in its dental characters, including thick enamel, round, swollen molar cusps, the absence of a honing facet (from contact with the upper canine) on the mesial (anterior) face of the lower third premolar (P3), and substantial sexual dimorphism in the cheek tooth size (especially marked in the earliest

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Pongo pygmaeus

nasal cavity

Sivapithecus indicus (GSP 15000)

Pan troglodytes

Australopithecus). Its postcranium remains undescribed, but if teeth were all that mattered, it could be placed on the direct line to Australopithecus. In this role, it would, however, face the same objections noted previously for Kenyapithecus, and it is more likely that Graecopithecus and Australopithecus evolved their dental similarities in parallel, as they adapted to tropical or subtropical woodlands, where they fed, at least seasonally, on similar hard or abrasive vegetal items. Among the remaining Eurasian genera, Sivapithecus deserves special mention for its bearing on orangutan origins and calibration of the molecular clock, and Gigantopithecus for its apparent overlap with Homo. Sivapithecus comprised two-to-four siamang-to-orangutan-size species represented in deposits dated between roughly 12.7 and 8.5 Ma at sites in the Siwalik Hills on the India-Pakistan border. As noted above, it shared thick enamel, relatively short, stubby canines, and other derived dental features with Australopithecus, and into the late 1970s, one if its species, then assigned to the now defunct genus Ramapithecus, was widely considered to be an early hominin, perhaps on the line to Australopithecus. It was dismissed from this position when biomolecular analyses showed that it probably existed before the hominins had diverged from the African great apes and when fresh fossils demonstrated that its face was remarkably orangutan-like (ig. 3.27). It is now widely thought to indicate that the orangutan lineage had emerged by 12 Ma. In a turnabout, this fossil-based estimate is now oten used to calibrate the biomolecular clock. here is the problem, however, that the postcranial bones of Sivapithecus indicate it was a monkeylike quadruped, perhaps broadly like

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125 FIGURE 3.27. Partial skull of Sivapithecus indicus (GsP 15000) compared with skulls of an orangutan, Pongo pygmaeus, and of a chimpanzee, Pan troglodytes (drawn by K. Cruz-uribe from photographs and casts). in its high, narrow orbits, narrow interorbital region, smooth (nonstepped) transition from the loor of the nasal cavity to the subnasal plane, and large size of the central incisors compared with the lateral ones, the Sivapthecus skull closely resembles that of the orangutan and differs from those of the african great apes. This suggests that Sivapithecus was near the ancestry of the orangutan. GSP = Geological survey of Pakistan.

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Proconsul and most other early and middle Miocene hominoids. Particularly telling is the curvature of the upper humerus shat, which suggests that the humerus lay alongside a deep, narrow, monkeylike chest and that it was not routinely raised for hand-over-hand climbing or for hanging below branches. In its lack of derived (apelike) postcranial features, Sivapithecus poses an evolutionary conundrum. If Dryopithecus, which possessed a straight humerus shat and other derived apelike postcranial features, was near the line leading to the African great apes and Sivapithecus was on or near the line leading to the orangutan, then the extant African great apes and the orangutan must have developed their shared postcranial specializations in parallel. he alternative is that the facial similarities of Sivapithecus and the orangutan represent parallelisms or shared primitive features and that the orangutan evolved from another mid-Miocene hominoid. Ankarapithecus, Lufengpithecus, and a Lufengpithecus-like form, Khoratpithecus from hailand, are possible, if not compelling alternatives. Ankarapithecus was an orangutan-size ape that has been dated to 10–9 Ma in Turkey, and it anticipated the orangutan in some features, including its relatively closely spaced orbits and the large size of its central incisors relative to its lateral ones. It difered from both the orangutan and Sivapithecus in many other key characteristics, but the similarities and its age might allow it have been ancestral to both Sivapithecus and the orangutan. Lufengpithecus, known from partial skulls and numerous dentitions dated to roughly 8–7 Ma and perhaps later in southwestern China, illustrates a recurrent dilemma in reconstructing hominoid ancestral-descendant relationships: the conlicting signal of diferent skeletal characters. he teeth of Lufengpithecus resemble those of the orangutan far more closely than do those of Sivapithecus (or Ankarapithecus), but its face and skull resemble those of the orangutan far less. If only teeth were known and geographic proximity igured prominently in reconstructing evolutionary relationships, Lufengpithecus would surely be preferred to Sivapithecus as a likely orangutan ancestor. Gigantopithecus was perhaps the largest primate that ever lived, exceeding even the gorilla in size. Its oldest occurrence may be documented in the Siwalik Hills between 7.8 and 7.3 Ma, but it is best known from cave deposits dated between 1 and 0.5 Ma in southern China and adjacent Vietnam. he Chinese sample so far comprises three mandibles and more than 1,000 isolated teeth. here are no limb bones, but dental size alone implies that Gigantopithecus was entirely terrestrial. Like Sivapithecus, it shared thick enamel, stubby, low-crowned canines, and other dental specializations with Australopithecus, and it may have relied on similar vegetal foods. Its extinction has been speculatively attributed to unsuccessful competition with Homo erectus, with which it appears to have overlapped in both China and Vietnam.

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SOURCES: late Miocene extinction of hominoids in Europe (Agustí et al. 2003) and the Siwaliks (Patnaik et al. 2005); Dionysopithecus (Ducrocq et al. 1994; Harrison and Gu 1999); Pliopithecus and Anapithecus (Andrews et al. 1996; Begun 1989; Ginsburg 1986; Harrison 1987; Nagatoshi 1987); oldest Hylobates (Jablonski et al. 2000; Louys et al. 2007); Griphopithecus (Andrews and Kelley 2007; Begun et al. 2003; King et al. 1999; Martin and Andrews 2003), Dryopithecus—craniodental features (Begun 1992a, 1992b; Köhler et al. 2001), body form (Moyà-Solà and Köhler 1996), and possible close relationship to the African great apes (Begun 1994, 2002); Pieralopithecus (Begun and Ward 2005; Moyà-Solà et al. 2004); Oreopithecus—geologic antiquity (Azzaroli 1985; Azzaroli et al. 1986; Rook et al. 2000), postcranium, posture, and locomotion (Fleagle and Kay 1985; Harrison 1986a; Harrison and Rook 1997), and dentition (Alba et al. 2001; Delson and Andrews 1975; Rosenberger and Delson 1985; Szalay and Delson 1979); Graecopithecus (Ouranopithecus) (Andrews 1992; Bonis et al. 1981, 1986, 1998; Bonis and Koufos 1994; Bonis and Melentis 1984; Dean and Delson 1992; Güleç et al. 2007; Koufos and Bonis 2005, 2006); Ankarapithecus (Alpagut et al. 1996; Begun et al. 2003; Begun and Gülec 1998); Sivapithecus—antiquity and included species (Kelley 2005), dentition (Andrews and Tekkaya 1980; Kelley 1988; Kelley and Pilbeam 1986; Simons 1981), face (Andrews and Tekkaya 1980; Pilbeam 1982; Ward and Pilbeam 1983), postcranium (Morbeck 1983; Pilbeam et al. 1990; Raza et al. 1983; Rose 1983, 1984, 1986), and implications of a monkeylike body and an orangutan-like face (Andrews 1992; Beneit and McCrossin 1997; McCrossin and Beneit 1994; Pilbeam 1996; Pilbeam et al. 1990); inclusion of Ramapithecus in Sivapithecus (Greenield 1979; Pilbeam 1982); Gigantopithecus—questionable identiication in the Siwaliks (Patnaik et al. 2005), dental characteristics (Daegling and Grine 1994; Dean and Schrenk 2003; Olejniczak et al. 2008; Szalay and Delson 1979; von Koenigswald 1952; Wu and Poirier 1995) and possible overlap with Homo erectus in China and Vietnam (Ciochon et al. 1996; Louys et al. 2007; Wang et al. 2007); Lufengpithecus (Badgley et al. 1988; Kelley 1986; Kelley and Plavcan 1998; Schwartz 1990, 1992, 1997; Schwartz et al. 2003); Khoratpithecus (Chaimanee et al. 2003, 2004, 2006)

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Summary and Conclusion he primate fossil record consists mainly of fragments, particularly partial jaws and teeth, and the preceding account shows that these are unevenly spread through space and time. he spotty, oten sparse record helps explain why informed specialists commonly disagree on what fossils to assign to what species and on the relationships among fossil species and between fossil species and living ones. Some disagreement also stems from diferent advance expectations or theoretical perspectives, which themselves are partly determined by the state of the record. Much research on fossil primates has understandably been driven by a desire to trace the origins of living ones, but as the fossil record has improved, especially since the 1960s, tracing the descent of living primates has become more problematic, not less. his is because the fuller record shows that few known fossil primates possessed derived features or specializations that link them unequivocally and uniquely to any living form. Many fossil species, in fact, had their own unique specializations that efectively eliminate them from the ancestry of later species and that suggest they became extinct without issue. To the extent that fossil forms resemble living ones (or each other), it is oten diicult to determine if the shared features are actually primitive retentions inherited from a relatively distant common ancestor or, alternatively, are parallelisms that genetically similar taxa developed independently as they adapted to sim-

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es

a at re eg

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late Cretaceous

Plesiadapidae

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lemur/loris stock

ntidae Picrodo Carpolestidae

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Eocene "lemuroid" stock

s

pe

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io

n ce

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anth ropo

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M

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Europ e "lemuan Eocene roids " Eo N. A cen m e " e ri c lem an uro i ds "

Oligocene

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ne Eoce " ican arsioids mer t N. A ocene " -Olig Europea n Eo "tarsioid cene s"

"lo

riso

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Pliocene

e cen es Mioyrrhin t pla

5.2

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ids " "le Asiat mu ic roi ds"

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Lemurs

ae Paromomyid

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catarrhine stock

anthropoid stock

tarsier stock tarsioid/anthropoid stock

Plesiadapiform ("archaic primate") stock

euprimate stock

?

98

FIGURE 3.28. a provisional phylogeny of the Primates (modiied after martin [1990], 46). The times when major groups diverged are disputed, but the order of divergence is wellestablished.

ilar circumstances. Parallelism (convergence or homoplasy) is especially problematic and diicult to detect, yet it surely explains why diferent anatomical regions sometimes seem to imply contradictory evolutionary relationships. On top of this, the fossil record presents many more potential ancestors than the living Primates require, and this underscores the point that many fossil forms have no living descendants. But even if it is not possible to draw a full and compelling phylogeny linking successively younger fossil primates to living species, it is possible to outline it, along with the provisional times when major primate groups diverged from each other (ig. 3.28). Equally important, fresh discoveries and increasingly sophisticated analyses have revealed the main stages of primate evolution, which can be conceptualized as a series of adaptive radiations preceding the emergence of hominins: 1.

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An initial radiation beginning probably in the late Cretaceous, between 98 and 65.5 Ma, leading to the Paleocene Plesiadapiformes. hese were distinguished from their insectivore (or insectivore-like) ancestors primarily by the morphology of their cheek teeth and perhaps by their ear structure. hey barely difered from insectivores in their brains, special senses, or locomotion, and their main behavioral distinction was perhaps the signiicant addition of seeds, fruits, or other vegetal matter to a basically insectivorous diet.

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A second radiation beginning in the Paleocene, between 65.5 and 56.5 Ma, or possibly before, culminating in a wide variety of much more advanced lemur-like and tarsier-like forms. hese were distinguished from the Plesiadapiforms and from their own more primitive ancestors by the development of typically primate grasping hands and feet with nails instead of claws and by a reorganization of the brain and face to emphasize vision over olfaction. he selective advantage of these morphological advances was perhaps an enhanced ability to hunt insects among slender branches in forest undergrowth and to obtain fruits and lowers growing near branch tips. A third radiation beginning in the early or middle Eocene, before 45 Ma, producing a variety of primitive higher primates. Like their lemur-like or tarsier-like ancestors, they were mainly arboreal and quadrupedal, but they may have spent more time walking on top of branches and less time leaping between them. hey probably also focused less on insects and more on fruits and leaves, and their brains tended to be larger, with an even greater emphasis on vision over olfaction. A fourth radiation or set of radiations during the late Oligocene or early Miocene, between 30 and 20 Ma, in which higher primates diferentiated into forms anticipating extant apes and monkeys. To begin with, the main diference between apes and monkeys was dental, and this may mean that the earliest monkeys difered from their apelike contemporaries mainly in their greater tendency to eat leaves. he early apes and monkeys were probably very similar in their palm-lat, quadrupedal, nonsuspensory habitual postures and locomotion. Almost certainly both were basically arboreal, though the early monkeys may have been relatively more terrestrial. To begin with, the apes were much more abundant and diverse than the monkeys, probably because forested, fruit-rich settings dominated the tropical and subtropical region of Africa where monkeys and apes irst diverged. A ith radiation in the middle and later Miocene, ater 17–16 Ma, that produced apes whose bodies and limb mobility imply a modern apelike tendency for orthograde posture and hand-over-hand climbing. Some apes also developed novel dentitions combining thick enamel, enlarged molar surfaces, and low-crowned, noninterlocking canines. hese dental specializations may have been a response to an increased reliance on hard or grit-encrusted foods when fruits became seasonally less abundant. he dietary shit in turn was probably sparked by a middle-to-late Miocene trend toward cooler, drier, more seasonal continental climates that promoted the spread of woodlands and savannas at the expense of forests, particularly in tropical and subtropical latitudes. he novel dental complex is especially noteworthy

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because it anticipated the complex that characterized the early hominin Australopithecus, ater 4.1 Ma.

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It is highly likely that the hominin and chimpanzee lines leading to people and chimpanzees diverged in equatorial Africa between 8 and 5 Ma and that the last shared ancestor inhabited woodlands that increasingly replaced forests during the late Miocene, ater 10 Ma. To ind food or to move between food sources in these woodlands, the common ancestor was probably at least partly terrestrial, and the hominin lineage emerged when one population adopted bipedalism as its habitual mode of terrestrial locomotion. Arguably a particularly sharp reduction in tree cover near 6 Ma provided the selective stimulus for bipedalism. Unfortunately, the African fossil record between 10 and 6 Ma remains stubbornly sparse, and the morphology of the protohominins remains conjectural. If Australopithecus, described in the next chapter, is a reasonable guide, the protohominins had chimpanzee-like bodies and skulls that they inherited from a middle-to-late Miocene predecessor. hey may also have had thick enamel, enlarged molars, and low-crowned, noninterlocking canines, but this inference is questionable because it would mean that the chimpanzees and the gorilla must have evolved thin enamel, relatively small molars, and enlarged canines separately, as opposed to inheriting them from the last ancestor they shared with hominins. Even more important, as discussed in the next chapter, meager fossils dated between 6 and 5 Ma in eastern Africa suggest that the earliest hominins may have had chimpanzee-like enamel and canines. If these fossils include the ancestor of Australopithecus, Australopithecus must have evolved thick enamel and associated dental features on its own, as it adapted to broadly the same environmental circumstances that favored these characters in some of its Miocene predecessors. he expanding fossil record will one day allow an informed choice. SOURCES: interaction between advance expectations and the state of the fossil record (Cartmill 1982; Fleagle and Jungers 1982)

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The auSTraLOPIThS anD HOMO HABILIS

In 1939 Gregory and Hellman (1939) proposed that ancient primitive human fossils from South Africa be allocated to the subfamily Australopithecinae, in distinction from a second subfamily, the Homininae, containing later, more advanced humans. he Australopithecinae and the Homininae together comprised the family Hominidae separate from the family Pongidae, which included the great apes. he formal terms Australopithecinae and Homininae never found favor, and their use is now precluded by a taxonomic revision that lumps the great apes with people in the family Hominidae and that separates people and the African great apes only at the tribal level. In this revised scheme, people broadly understood are no longer hominids (anglicized from Hominidae) but rather hominins (anglicized from Hominini). Earlier and later people are separated only at the generic level: Ardipithecus, Australopithecus, Paranthropus, and Kenyanthropus for earlier more primitive forms, and Homo for later, more advanced ones. Since at least one of the earlier genera was probably more closely related to Homo than it was to the other genera, there can be no formal, higher-level systematic distinction between the earlier genera and Homo. Still, informally, for the sake of convenience, many specialists continue to apply the term australopithecine to the earlier hominin genera, which difer from Homo more or less equally in adaptive grade, if not in degree of evolutionary relationship. It is in this informal sense that the term australopithecine is used here, abbreviated to australopith to limit the possibility for taxonomic confusion. he australopiths are the oldest known undoubted hominins, with a fossil record that extends from before 4.4 million years ago (Ma) to at least 1.2 Ma. In the vernacular, they have sometimes been called “manapes” because they combined bipedal locomotion—the deining characteristic of the hominin tribe—with small, ape-sized brains and apelike skulls and faces. It is becoming increasingly clear that most if not all

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australopiths also resembled apes in other respects, including an apelike upper body form with long powerful arms, and that they probably mixed human-style bipedal locomotion on the ground with ape-style agility in the trees. Further, in those australopith species for which body size can be estimated, males were probably substantially larger than females, suggesting an apelike social organization in which the sexes did not cooperate economically and males competed vigorously for females in estrus. No australopith species appears to have let an archaeological record, and if they relied on technology, it was probably only to the limited extent observed among living chimpanzees. In short, morphology and behavior together suggest that the australopiths were bipedal apes more than primitive humans. Humans in the narrow sense—assigned to the genus Homo for their larger brains, their smaller faces, and other signiicant departures from the ape condition—appeared only about 2.5 Ma, and it is probably no coincidence that the oldest known stone tools date from about the same time. he evolutionary origins of the australopiths remain vague because of a sparse African fossil record between 8 and 4.5 Ma. he probable rate at which genetic diferences accrue imply that the human and chimpanzee lineages diverged during this interval, but pertinent fossils remain scarce. he total so far comprises eighteen fragmentary specimens (mainly isolated teeth) from the Middle Awash region of Ethiopia, assigned to Ardipithecus kadabba; a skull, four fragmentary mandibles, and four isolated teeth from the Djurab Desert, northern Chad, assigned to Sahelanthropus tchadensis; and twenty specimens (partial dentitions, isolated teeth, and postcranial fragments) from near Lake Baringo, westcentral Kenya, assigned to Orrorin tugenensis (references in table 4.7). At the moment, among the three species, O. tugenensis is the most plausible ancestor for the australopiths, since it is the only one for which postcranial bones imply bipedalism, but the case for bipedalism is arguable, and future inds from the other species could muddy it. he phylogenetic relations of unquestioned australopith species to each other and to the earliest widely recognized species of Homo, H. habilis, are also uncertain, and fresh discoveries in Africa have expanded the possibilities. More important, however, they have revealed the basic morphological and behavioral course of early hominin evolution. his chapter focuses on the known pattern.

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History of Discovery: South Africa he irst australopith fossil was discovered by Raymond Dart, a Britishtrained anatomist who in 1922 was appointed professor of anatomy at the University of the Witwatersrand in Johannesburg, South Africa. here are minor discrepancies between the discovery as recounted by Dart and

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Ahl al Oughlam Sahabi

Bahr el Ghazal (Koro-Toro) Toros-Menalla Hadar & Dikika Gona Middle Melka Kunturé Awash Konso Omo Fejej West Turkana Koobi Fora Lothagam & Kanapoi Allia Bay Nyabusosi Kanjera Chesowanja Chemeron Senga Peninj Olduvai Gorge Laetoli

Atlantic Ocean

Rift

Eastern

Rif Western

t

Equator

Malema Uraha Gondolin, Gladysvale, Drimolen, Coopers, Swartkrans, Kromdraai & Sterkfontein Makapansgat Taung

Indian Ocean

Langebaanweg

FIGURE 4.1. approximate locations of african Plio-Pleistocene hominin fossil sites and of the nonhominin fossil sites at ahl al oughlam (morocco), sahabi (libya), and langebaanweg (south africa). most east african sites lie near the eastern branch of the Great rift valley, where tectonic activity associated with rifting created numerous basins that could trap and preserve sediments and bones. While the sediments were accumulating, regional volcanoes often erupted alkaline ashes that promoted bone preservation and that can be dated by the potassum-argon method and its 40 ar/39ar variant. finally tectonic activity, often sparse vegetation, and episodic and frequently violent rainfall have encouraged erosion that exposes fossils that would otherwise lie deeply buried. The rift system does not extend to southern africa, where sedimentary basins tend to be rare and shallow. The southern african Plio-Pleistocene hominin sites are all in limestone caves, and they lack volcanic materials for dating. The known caves cluster mainly in an intensively explored area between Johannesburg and Pretoria, south africa. ahl al oughlam, sahabi, and langebaanweg have provided no hominin fossils, though they variously date between 5 and 2.5 ma, when hominins were present closer to the equator, and they are as rich or richer in mammalian fossils than any of the other sites on the map. They differ from the other sites primarily in their location in temperate zones near the northern and southern margins of africa, and they imply that the earliest hominins were conined to tropical or subtropical latitudes.

as reconstructed by his equally illustrious successor, Phillip V. Tobias. Dart’s account is summarized here. Dart planned to create a museum in his department, and he encouraged students to collect suitable fossils during their vacations. In 1924 a student brought him a fossil baboon skull from the Buxton lime quarry at Taung (then Taungs) about 320 km southwest of Johannesburg, on

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FIGURE 4.2. facial skeleton and endocast of Australopithecus africanus from Taung, viewed from the right side (drawn by Kathryn Cruz-uribe from photos and casts). The mainly deciduous dentition shows that the skull represents a juvenile whose age at death is now thought to have been three-to-four years. The small deciduous canines are among the features that identify the skull as hominin rather than ape. The location of the lunate sulcus may also imply hominin status, if it was far to the rear as in humans (dart 1925) rather than closer to the front as in apes (falk 1989). a position further to the rear would imply a human-like expansion of the parietal and temporal association cortex toward the rear of the brain. arguably, however, the position of the lunate sulcus cannot be unequivocally determined from the endocast (holloway 1984), and most authorities today do not use it to determine the phylogenetic afinities or neural organization of the Taung child.

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endocranial cast

approximate position of the lunate sulcus (Dart)

cm

deciduous canines

Taung permanent first molars (erupting)

the eastern edge of the Kalahari Desert in what is now the Northwest Province of South Africa (ig. 4.1). hrough a geologist colleague who visited the lime quarry shortly aterward, Dart obtained two crates of fossiliferous breccia—rock-hard blocks of sandy sediment and fossils cemented together by limy glue—from a small cave exposed in the quarry. he fossils in the crates mainly represented additional baboons, but one crate contained a natural endocast (mold of the inside of a skull) that Dart immediately realized came from a hominoid primate. Among the breccia blocks in the crates, Dart found one with a depression into which the endocast itted. Inside the block, he saw traces of bone that he hoped was the front of the skull associated with the endocast. he front was present (ig. 4.2), but it took Dart many weeks to expose it with hammer, chisels, and a sharpened knitting needle. he dentition showed that the skull came from a juvenile whose irst molars were just erupting, and various other features showed that in important respects the creature was intermediate between apes and people. For example, it was obviously apelike in the small size of its brain—or more precisely in its endocranial (internal skull) volume, which Dart estimated would have reached 525 cc at adulthood. Subsequent reestimates have reduced this to between 404 and 440 cc, but even Dart’s igure was much closer to the average for common chimpanzees (ca. 400 cc) and gorillas (ca. 500 cc) than to the average for living people (1,350–1,375 cc). At the same time, the deciduous canines were much smaller and less projecting than those in apes, and the foramen magnum (“large hole”), through which connec-

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external occipital protuberance

supraorbital ridge

nuchal crest

gorilla

nuchal musculature

orbital aperture infraorbital foramina

occipital condyle

axis of the position of the foramen magnum foramen magnum on the base of the skull

mental foramen

superior temporal line

135 FIGURE 4.3. Gorilla and modern human skulls, illustrating the contrasting position of the skull with respect to the spinal column (adapted from le Gros Clark [1967], 67). in people, the skull is balanced on top of the spinal column, and the foramen magnum consequently lies much farther forward on the base of the skull. The orientation of its axis is also more vertical. The more posterior position of the foramen magnum in the gorilla, together with its larger, more protruding face, also requires larger, more powerful nuchal (neck) muscles to stabilize the head.

modern human external occipital protuberance

position of the foramen magnum on the base of the skull

axis of the foramen magnum

tions passed from the spinal column to the brain, was more downwardly oriented on the base of the skull. his suggested that the skull was balanced on top of the spinal column, as in people, where this position is a natural concomitant of upright, bipedal locomotion (ig. 4.3). In short, the position of the foramen magnum implied that the Taung individual was bipedal like people rather than quadrupedal-like apes. Dart published a description of the skull in the February 7, 1925, issue of Nature and concluded that it came from a previously unknown species “intermediate between living anthropoids and man” (Dart 1925). He named the species Australopithecus africanus (“African southern ape”) and stressed both the skull’s intermediate or transitional morphology and its discovery far outside the geographical ranges of the living apes. Taung lies in a nonforested region where living apes could not survive, and from geological studies Dart argued that the regional environment was

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probably similar when the fossils accumulated. he ancient vegetation was thus inappropriate for an ape in the narrow sense. It was, however, clearly suitable for an ape that was developing human traits, especially bipedal locomotion. Rare bone fragments that may have come from a nyala antelope (Tragelaphus angasi) now suggest that the environment may have been more wooded than Dart believed, but baboon and rock hyrax fossils dominate heavily, and they continue to imply an environment that apes would generally avoid. In the following issue of Nature, Dart’s assessment was severely criticized by several anthropologists. A consensus soon developed that Australopithecus was simply a fossil ape, with no special relevance to human evolution. According to a prominent eyewitness to the debate, the skeptics were inluenced by nonscientiic considerations, including Dart’s lowery, rhetorical writing style, his reputation for coming to hasty conclusions, and his unbelievably good luck in inding such a spectacular fossil, just two years ater arriving in Africa. However, there were fundamental scientiic concerns as well. here was very little from which to estimate the antiquity of the Taung skull, but to some authorities, the associated baboon fossils implied it was too recent to be a human ancestor. In addition, the discovery of primitive human fossils (“Pithecanthropus”) in Java in 1891 had convinced many specialists that Asia, not Africa, was the cradle of humanity. Finally, the skull did not it either of the current theories of human evolution. he irst held that human ancestors should be equally primitive in all traits, whereas Dart’s ind showed a mix of advanced and primitive features. he second theory, supported by the famous Piltdown skull (later exposed as an elaborate fake), proposed that the singular human brain had evolved before other uniquely human traits. he Taung skull, as interpreted by Dart, suggested just the reverse—that the uniquely human mode of bipedal locomotion had evolved before the brain. Beyond these theoretical obstacles, some specialists disputed Dart’s claim that the lunate (or simian) sulcus, a issure or furrow that delineates the primary visual cortex, lay far back on the surface of the brain, as it would in humans, rather than far forward as it does in apes. Dart inferred the position from the endocast, and it remains arguable to this day whether the position can be unambiguously detected. If it were far back as Dart suggested, it would imply a humanlike expansion of the parietal and temporal associational areas on the cerebral cortex. Still more problematic was the very young age of the Taung individual at the time of death. his was initially estimated at ive-to-seven years, based on the maturation rate and dental eruption schedule of living people. It has now been reestimated at only three-to-four years, since growth increments in australopith enamel, together with the relative timing of crown and root development in various teeth, suggest that the australopiths matured

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at about the same rate as modern apes, perhaps 30%–50% faster than modern people. he crucial point is that very young apes resemble very young humans much more than adult apes resemble adult humans, and some of Dart’s critics suggested that if the Taung child had grown to adulthood, it would have been much more apelike in all features, including the orientation of the foramen magnum. Finally, there was the problem that Dart was inferring bipedalism from a portion of the anatomy not directly involved in locomotion. Without actual bones from the locomotor skeleton—especially from the pelvis or leg—it could be argued that Australopithecus may not have been bipedal. Unfortunately, Dart did not visit the Taung site to seek additional fossils, especially adult skulls or postcranial bones that could have won his case, and by the time qualiied specialists did go, the cave that had contained the juvenile skull had been quarried away. Dart continued to remove breccia from the skull, and in 1929 he succeeded in separating the upper and lower jaws to permit a more complete assessment of the dentition. He distributed casts of the dentition to many specialists, and W. K. Gregory, an eminent American authority on mammalian teeth, concluded that it came from (what we now call) a hominin rather than from an ape. Gregory’s support aside, however, additional specimens were obviously necessary to convince many other authorities, and Dart was not personally prepared to make the search. his task fell to Robert Broom, a Scottish-born physician and authority on fossil reptiles who had settled in South Africa. He visited Dart’s laboratory two weeks ater the formal announcement of the Taung ind in Nature and was an early convert to Dart’s opinion. But he had other work to complete, and it was not until the 1930s that he was able to initiate research on Australopithecus. In 1936, working from his base at the Transvaal Museum in Pretoria, about 90 km north of Johannesburg, Broom was directed to the fossiliferous breccia in a cave on the farm Sterkfontein, roughly 25 km northwest of Johannesburg and 9.6 km north-northwest of the town of Krugersdorp in what is now the Gauteng Province of South Africa (ig. 4.1). he site was being actively mined for lime, and in August 1936, on Broom’s third visit, the mining supervisor handed him the partial skull and endocast of an adult australopith. Between 1936 and 1939, Broom made numerous other visits and recovered additional skulls, jaws, teeth and some limb bones. In 1938, during one of his periodic trips to Sterkfontein, Broom heard of another promising cave at Kromdraai, approximately 1.6 km to the east-northeast. Again he was quickly rewarded with an adult skull, but it difered in important respects from the earlier Sterkfontein skull. He concluded that it represented a diferent kind of australopith, a view that is still widely accepted.

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Having found adult skulls, by 1939 Broom could show that adult australopiths were no more apelike than the Taung child. Equally important, at Sterkfontein he had recovered a distal femur and at Kromdraai an astragalus (talus) that showed the australopiths probably were bipedal, as Dart had argued. Partly for economic reasons and partly because of World War II, research in the Sterkfontein region largely ceased between 1939 and 1945. In collaboration with G. W. H. Schepers, Broom spent the war years preparing a monograph on the Sterkfontein and Kromdraai inds. his appeared in 1946 and was widely read and acclaimed. Above all, it helped to convert the eminent British anatomist and paleoanthropologist, Sir Arthur Keith, from a skeptic on the ancestral human status of the australopiths to a supporter, and many others followed Keith’s lead. Some remained critical only of Broom’s tendency to divide the known fossils among more taxa than seemed warranted: he assigned the Sterkfontein and Kromdraai inds to their own genera—Plesianthropus (near man) and Paranthropus (alongside man), respectively—distinct from Dart’s original Australopithecus. Many specialists later assigned all the fossils to the single genus Australopithecus, divisible between two species—A. africanus (Taung and Sterkfontein) and A. robustus (Kromdraai). At present, however, a growing number have resurrected the category Paranthropus as originally deined by Broom, and Paranthropus is used here. In the vernacular, A. africanus is oten called the “gracile (or slender) australopith” and Paranthropus (or Australopithecus) robustus is the “robust australopith,” although the main diference was that P. robustus had a more massive face and mandible, related to more powerful chewing muscles. In body mass and stature, it closely resembled A. africanus. In 1947–1948, assisted by John T. Robinson, Broom renewed work at Sterkfontein, with spectacular success. Among the crucial specimens they recovered was a partial australopith skeleton, including a pelvis that demonstrated beyond all doubt that the owner was bipedal. hey also found a nearly complete adult skull (Sts 5) that is probably the most famous South African australopith fossil ater the original Taung ind. It was regarded as a female Plesianthropus and became known popularly as Mrs. Ples. (Figure 3.1 illustrates a reconstruction.) In 1947 James Kitching, participating in an expedition organized by Dart, found a partial australopith skull at the Makapansgat Limeworks Cave near the town of Potgietersrus in what is presently the Limpopo Province of South Africa, approximately 300 km north of Johannesburg, and, in 1948, Broom and Robinson made the irst discoveries in a cave on the farm Swartkrans, about 1.2 km west of Sterkfontein. Research in the late 1940s at Sterkfontein, Makapansgat, and Swartkrans greatly enlarged the available australopith sample, producing many skulls or partial skulls, dentitions, and isolated teeth, plus a small but diagnostic

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sample of postcranial bones that pointed consistently to bipedalism. By 1950, the mounting evidence had convinced nearly all skeptics that the australopiths were primitive, early representatives of humanity. Debate continued regarding their relation to later humans, however, and the issue is still not settled. Sterkfontein, Kromdraai, Makapansgat, and Swartkrans retain fossiliferous breccias, and each is being investigated or has recently been investigated. In 1992, they were supplemented by the discovery of two australopith teeth from the long-known Gladysvale Cave site, about 13 km northeast of Sterkfontein; in 1994 and aterward by the recovery of spectacular robust australopith fossils at the previously unstudied Drimolen Cave, roughly 7 km north of Sterkfontein; in 1997 by the discovery of a robust australopith molar in brecciated sediments at Gondolin Cave, about 20 km northwest of Sterkfontein; and in 2001 by the excavation of a handful of robust australopith fossils at Coopers Cave, between Sterkfontein and Kromdraai (references below). Yet other sites undoubtedly exist, particularly in the Krugersdorp region, and Drimolen demonstrates that new sites can be as productive as established ones. Together the southern African sites have provided a large sample of australopith bones, comprising more than thirty-two skulls or partial skulls, roughly one hundred jaws or partial jaws, hundreds of isolated teeth, and more than thirty postcranial bones. Since 1959, however, southern Africa has been largely eclipsed by spectacular discoveries of australopith fossils in eastern Africa.

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SOURCES: discovery of the Taung australopith according to Dart (1925; Dart and Craig 1959) and Tobias (1984, 1985, 2000); endocranial capacity of the Taung skull (Falk 1987; Falk and Clarke 2007; Holloway 1970); Taung paleoenvironment (Berger and Clarke 1995, 1996); early nonscientiic reasons for rejecting Dart’s interpretation of the Taung child (Le Gros Clark 1967); failure of the Taung skull to it then-current theories of human evolution (Washburn 1985); debate on the position of the lunate sulcus on the Taung endocast (Falk 1983b, 1985a, 1989, 1991; Holloway 1984, 1991a); age of the Taung child estimated at three-to-four years (Conroy and Vannier 1987); enamel growth increments (Beynon and Wood 1987; Bromage 1987; Bromage and Dean 1985), timing of root and crown development (Smith 1986), and rapid maturation in australopiths (Dean 2006; Dean et al. 2001); 1946 monograph on the australopiths (Broom and Schepers 1946); conversion of Keith (1947) and then others to the hominin status of the australopiths (Tobias 2002); inventory of South African australopith fossils (Day 1986b; Grine 1993b; Howell 1978a; Susman 1993)

History of Discovery: Eastern Africa East African early hominin sites now occupy the limelight for two important reasons. First, in eastern Africa early hominin fossils oten occur in relatively friable ancient stream or lake-edge deposits rather than in rockhard cave ills. At some sites, such as Olduvai Gorge, the fossils can even be excavated using dental picks and brushes in the accepted archaeological manner. At the southern African sites, dental picks and brushes have been subordinate to dynamite, pneumatic drills, hammers, and chisels.

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Second, and at least equally important, many of the east African sites are stratigraphically related to volcanic extrusives (mainly ancient volcanic ash layers or tephra) that can be dated in years by the radiopotassium and ission-track methods. So far, the southern African sites generally lack reliable material for numeric dating, and they have been placed in time mainly by the mammal species they contain, compared to species from dated layers in eastern Africa. In 1935 and 1939, Laetoli (Garusi) near Olduvai Gorge, northern Tanzania, provided an australopith canine and a partial australopith maxilla, respectively, and in 1955, Olduvai Gorge itself produced two australopith teeth. he Laetoli fossils are now assigned to Australopithecus afarensis, and the Olduvai teeth to Paranthropus (or Australopithecus) boisei. However, they were not initially appreciated for what they were, and the irst discovery of an australopith in eastern Africa is usually dated to 1959. he specimen was a well-preserved partial skull that Mary and Louis Leakey excavated from near the base of Olduvai Gorge, and it was important not only because it extended the known geographic range of the australopiths but also because it occurred in deposits that were soon dated to about 1.75 Ma by a pioneering application of the radiopotassium method. he 1.75 my estimate was the irst indication of the true antiquity of the australopiths, and it nearly doubled the total timespan that most specialists had allowed for human evolution. Louis Leakey initially assigned the 1959 skull to Zinjanthropus boisei (“Boise’s East African Man”), but he and others quickly realized its close resemblance to the South African robust australopiths and reassigned it to Paranthropus (or Australopithecus) boisei. he third molars were erupting at time of death, implying it came from an adolescent, and its extraordinary robusticity suggested it was male. It is still known in the vernacular as “Zinj,” and following its discovery, the Leakeys obtained inancial support that allowed them to uncover numerous other early hominin fossils at Olduvai, including the irst remains of Homo habilis, the oldest known species of Homo. Equally important, the Leakeys’ success prompted fruitful searches for australopiths and earliest Homo at other sites in eastern Africa (ig. 4.1), beginning with Peninj (Lake Natron) (1964) and continuing with Chemeron (1965), the lower Omo Valley (1966), East Turkana (Koobi Fora) (1968), Chesowanja (Chemoigut) (1970), Hadar (1973), Laetoli (1974), the Middle Awash (1981), West Turkana (1984), Fejej (1989), Konso (originally Konso-Gardula) (1991), Kanapoi (1994), Gona (formerly Kada Gona) (1999) and Woranso-Mille (2002). Several of these sites, including Olduvai, continue to produce signiicant fossils, early artifacts, or both, and many new sites undoubtedly await discovery. So far, eastern Africa has provided only about half as many australopith specimens as southern Africa, but many of the east African fossils are

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remarkably complete and they originate from sediments whose age can oten be estimated numerically.

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SOURCES: irst discovery of australopith fossils at Laetoli (Leakey et al. 1967) and Olduvai (Leakey 1958); irst K/Ar dating of an australopith at Olduvai (Leakey et al. 1961); Zinjanthropus boisei (Leakey 1959) and its reassignment to P. boisei (Leakey et al. 1964; Robinson 1960; Tobias 1967a); discovery of H. habilis (Leakey et al. 1964)

Geology of the Southern African Australopith Sites he southern African australopith sites are all limestone caves in which sands, silts, natural stones, fossils, and sometimes artifacts accumulated in unsorted jumbles or breccias. Calcareous glue precipitated from groundwater commonly hardened the breccias to a cement-like consistency. he sites were discovered mainly during mining for bands of calcitic lowstone (travertine) that occur in and around the breccias. Since the breccias tend to be rock hard, the excavators usually could not employ standard archaeological techniques, which are applicable only to much soter deposits. For the most part, the cave ills cannot be securely dated by radiometric methods, and their antiquity has been estimated mainly by their faunal contents (compared with faunal assemblages from radiometrically dated sites in eastern Africa). In addition, at the most thoroughly studied caves, australopith fossils and others appear to have been introduced mainly by carnivores or raptors (at Taung), by carnivores or natural pitfall trapping (at Kromdraai), by hyenas (at Makapansgat), by large cats (at Sterkfontein and Swartkrans), and by unspeciied carnivores (at Drimolen). Strictly speaking then the caves can tell us little about australopith behavior. However, the abundance of australopiths, cercopithecoid monkeys, or both at three sites (Sterkfontein, Kromdraai, and especially Swartkrans) may mean that they were attractive for rocky ledges or other inaccessible sleeping places. Free-ranging baboons oten seek out such ledges to reduce the risk of nighttime predation, but even if predation was only occasionally successful, it would still produce a local bone assemblage in which the sleeping species was common. he taxonomy of the South African australopiths is not completely settled, but most authorities recognize just two species—Australopithecus africanus (at Taung, Makapansgat, Sterkfontein, and Gladysvale) and Paranthropus (or Australopithecus) robustus (at Swartkrans, Kromdraai, Drimolen, Gondolin, and Coopers). he antiquity of A. africanus is unsettled, but a range between 3 and 2.5 Ma seems likely. Some datings, discussed below, suggest it persisted until 2.2 Ma or even later. P. robustus was probably present between 1.8 and 1.2 Ma, but it may have appeared earlier, since no South African site unequivocally samples the period between 2.5 and 1.8 Ma. Its appearance may have coincided with an increase

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142 FIGURE 4.4. ages (in millions of years ago) of some important south african hominin fossil sites, based on their faunal contents (modiied after vrba [1982], 741). also shown are the known time ranges of some important mammalian taxa used in the dating, as well as the inferred time ranges for fossil hominin species. (Parapapio and Papio are baboons, Equus and Equus burchelli are zebras, and the remaining species are buffalo or antelopes.)

Hominin-associated assemblages

0

0.5

1.0

1.5

2.0

2.5

3.0 Ma

Makapansgat Mbs 3 & 4 Sterkfontein Mb 2 Sterkfontein Mb 4 Taung Kromdraai B East Mb 3 Swartkrans Mb 1 Drimolen Sterkfontein Mb 5 Swartkrans Mbs 2 & 3 Elandsfontein Kabwe (Broken Hill)

Hominin taxa

Cave of Hearths Australopithecus africanus Paranthropus robustus H. ergaster

H. sapiens

? H. habilis Parapapio

Papio Antidorcas

Other taxa

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

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Equus Equus burchelli Parmularius angusticornis Rabaticeras arambourgi Pelorovis antiquus Megalotragus priscus

Makapania Simatherium kohllarseni

in grassy vegetation that occurred across much of eastern and southern Africa around 2.5 Ma. he concluding section of this chapter considers what this vegetational shit might mean for early human evolution. he essential features of the following site-by-site presentation are summarized in igures 4.4–4.6 and in the summary section below on the geologic antiquity and geographic range of the earliest hominins. he

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2.6

2.4

2.2

Gauss Chron

2.0

1.8

1.6

Olduvai Subchron

Oldowan

1.4

1.2

Matuyama Chron

1.0

0.8

0.6 Ma Brunhes Chron

Jaramillo Subchron

Acheulean & Developed Oldowan

Pakefield Le Vallonet Atapuerca GD and SE Fuente Nueva 3 & Barranco Léon (Orce) Isernia Monte Poggiolo Agnani-Colle Marino Kärlich A Kärlich Bb Prezletice & Stránská Skála Korolevo VII & VIII

Erk-el-Ahmar

Europe

‘Ubeidiya

Western Asia

Evron Quarry Gesher Benot Ya’akov Latamne

Dmanisi

Central & South Asia

Kuldara Riwat Dina Bori Nihewan Donggutuo, Xiachangliang & Majuangou Maliang & Cenjiawan Lantian (Gongwangling & Chenjawo) Ban Mae Tha Mata Menge (Flores) Casablanca sequence

Eastern Asia

Northern Africa

Ain Hanech Mansourah Gona

Gona Hadar (Localities 666 & 894)

Middle Awash Daka Bodo Melka Kunturé Gomboré 1B Garba IV & XII (lower) Simbirro III, Garba II & XII Gadeb 2 & 8 Konso Fejej

Omo Shungura G and F

Lokalelei Mb

Koobi Fora (East Turkana) KBS Karari West Turkana (Nachukui Fm) Kalochoro Mb Kaitio Mb Natoo Mb Nyabusosi

Nariokotome Mb Kilombe

Chesowanja (Chemoigut) Kanjera South 1, 2 & 3

Kariandusi Olorgesailie

Peninj Olduvai Bed I

Eastern Africa

Olduvai Bed II

Olduvai Beds III/IV & Masek

Mwimbi (Chiwondo Beds)

Southern Africa

Sterkfontein Mb 5 Swartkrans Mbs 1-3

2.6

2.4

2.2

2.0

1.8

1.6

1.4

1.2

1.0

0.8

0.6 Ma

FIGURE 4.5. dating of the earliest artifact industries in africa and eurasia (in a format proposed by isaac [1986], ig. 13.2). dotted lines indicate possible or probable dates based mainly on geologic inference or faunal correlations. boldface marks the oldest secure sites on each continent. The most compelling evidence for human emergence from africa before 1 ma comes from dmanisi, Georgian republic (lordkipanidze et al. 2005, 2007). Paleomagnetic dating suggests that artifact makers may have appeared in the nihewan basin, northern China at almost the same time, 1.6–1.7 ma (Zhu et al. 2004).

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1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

FIGURE 4.6. Left: Time spans of the most commonly recognized hominin species between 4.4 and 1.0 ma (in a format suggested by Kimbel [1995]). Right: dating of some key behavioral and anatomical traits (adapted from harris [1983], ig. 1). broken lines imply uncertain or insecure records.

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5.0

4.8

4.6

4.4

4.2

4.0

3.8

3.6

3.4

3.2

3.0

2.8

2.6

2.4

2.2

2.0

1.8

1.6

1.4

1.2

Au. garhi

Hadar, Dikika, Gona (Kada Hadar Mb) Middle Awash (Belohdelie & Maka), Laetoli, ?Omo-Usno Fm, ?Omo-Shungura B, ?Koobi Fora (Tulu Bor Mb), ?W. Turkana (lower Lomekwi Mb), & ?Lothagam (Kaiyumung Mb)

Au. afarensis

Au. anamensis

Ar. ramidus

Gona (As Duma), Middle Awash (Aramis), ?Lothagam (Apak Mb) & ?Chemeron (Tabarin)

H. habilis

H. rudolfensis

P. boisei P. aethiopicus

Gona (Dana Aoule North & Busidima) Middle Awash (Daka-Bouri) Konso, Omo-Shungura K, W. Turkana (Nariokotome), Koobi Fora (?U. Burgi, KBS & Okote Mbs), Olduvai II, Swartkrans Mb 1 & ?Kromdraai East Mb 3

Konso, Omo-Shungura G-L, W. Turkana (Kaitio Mb), Koobi Fora (U. Burgi, KBS Middle Awash Hadar, Koobi Fora (U. Burgi & Okote), Chesowanja, (Bouri-Hata) & Okote Mbs), Olduvai I & II Olduvai I & II, Peninj & Malema & ?Sterkfontein Mb 5 Omo-Shungura C & ?D-G & W. Turkana (Lokaleilei Mb) ?Omo-Shungura Mbs E-G Koobi Fora (U. Burgi Mb) Taung, Sterkfontein (Mbs 2 and 3), ?W. Turkana (Nachukui Fm), Makapansgat (Mbs 3 & 4) & ?Chemeron (JM85) & Uraha Gladysvale

Au. africanus W. Turkana (Kataboi & Lomekwi Mbs)

K. platyops

Middle Awash (Asa Issie & Aramis 14) ?Fejej, E. Turkana (Allia Bay) & Kanapoi

P. robustus Kromdraai B, Swartkrans (Mbs 1-3), Drimolen, Gondolin, Coopers D & ?Sterkfontein Mb 5

bipedalism brain expansion posterior dental reduction stone artifact manufacture “home base” behavior

5.0

4.8

4.6

4.4

4.2

4.0

3.8

3.6

3.4

3.2

3.0

2.8

2.6

2.4

2.2

2.0

1.8

1.6

1.4

1.2

1.0

Ma

key anatomical & behavioral traits

consumption of large mammals

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Ma

H. ergaster/ erectus

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site descriptions refer to artifact assemblages using the terms Oldowan and Acheulean, which are deined fully only in the following sections. Readers who do not need or want a detailed introduction to the sites may wish only to skim the individual sketches.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: bone accumulation agency at Taung (Berger and Clarke 1995, 1996), Kromdraai (Brain 1981; Vrba 1981; Vrba and Panagos 1982), Makapansgat (Maguire 1985; Maguire et al. 1980), Sterkfontein and Swartkrans (Brain 1981, 1993a), and Drimolen (Keyser et al. 2000)

Taung

he child’s skull that is the type specimen of Australopithecus africanus remains the only australopith fossil ever found at Taung. he skull came from a cave that was largely quarried away before geologists could study it, but subsequent studies show it was part of an extensive system formed within the oldest of four limestone aprons (tufas) mantling the Gaap Escarpment near Taung. Reddish sands, silts, and occasional bones entered the cave from the surface through issures. he materials were subsequently cemented together by calcite (crystallized calcium carbonate) precipitated from water that periodically permeated the deposits. A nearby cave that may have been part of the same system contains carnivoredamaged fossil baboon skulls, and these suggest that leopards or other large cats could have accumulated bones at Taung. he African crowned eagle (Stephanoaetus coronatus) or another large raptor may also have played a role. Nearly all the Taung animals, including the child, fall within the preysize range of an eagle, and some of the Taung bones, including the child’s skull, exhibit the kind of damage that eagles inlict with talons or beaks. he geologic age of the Taung skull is diicult to establish, in part because its geologic provenance is so poorly documented. An estimate of the time when geomorphic change irst opened the cave, combined with uranium-series dates of 942 ka on the limestone apron in which the cave occurred and of 764 ka on the next youngest apron, suggest an antiquity of 1 Ma or less. However, the geomorphic dating requires assumptions that are impossible to verify, and uranium is now known to have entered the aprons ater they formed. he U-series readings thus underestimate the true age, perhaps by a substantial interval. he Taung fauna contains relatively few species that have been dated elsewhere and its stratigraphic association with the skull is unclear, but it includes cercopithecoid monkey fossils that resemble east African ones dated to about 2.3 Ma. he Taung skull itself has igured in the dating controversy, since at an age of 1 Ma or less, it would be the latest known Australopithecus africanus, probably by more than 1 my. his raises the possibility that it actually derives from the robust australopith species Paranthropus robustus, which, on evidence from Swartkrans (presented below), may have survived to 1.2 Ma or later. However, details of the Taung dentition and most aspects

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of the endocast align it with Sterkfontein and/or Makapansgat fossils of A. africanus that surely antedate 2 Ma. he antiquity of the Taung skull will always be speculative, but a model of how caves formed within the limestone apron, combined with the known or probable stratigraphic positions of the child’s skull and key cercopithecoid fossils, suggests that the cercopithecoid fossils are younger. In this case, the child’s skull could have reached Taung between 2.8 and 2.2 Ma, the maximum timespan during which fossils of A. africanus probably accumulated at the Sterkfontein site, discussed immediately below.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Taung geologic context (Butzer 1974b, 1980; Partridge 1982b; Peabody 1954; Tobias 1985); carnivore-damaged baboon skulls from a nearby cave (Brain 1985b); possible eagle role in the Taung bone accumulation (Berger 2006; Berger and Clarke 1995, 1996; McGraw et al. 2006; Sanders et al. 2003); age estimate for Taung—by geomorphology (Partridge 1973, 1982b), U-series (Tobias et al. 1993; Vogel and Partridge 1984), and associated baboon fossils (Delson 1984, 1988; Williams et al. 2007); resemblance of Taung to Sterkfontein A. africanus based on the dentition (Grine 1985a) and endocast (Falk and Clarke 2007); possibility that the Taung baboon fossils postdate the child’s skull (McKee 1993a, 1993b)

Sterkfontein

At Sterkfontein, silts, sands, and other materials fell or were washed through shats that descended from the surface into a solution cavern within the local dolomitic limestone. Once inside, the foreign debris, together with fragments of the cavern roof and walls, were cemented to a rock-hard breccia by calcite, which also occurs as pure interbeds and lenses within the breccia. Erosion later removed most of the cavern roof, exposing the breccia at the surface. In places, subsequent decalciication has transformed the breccia into a more friable mix of sand, rocks, and bones. On average, the Sterkfontein cave ill (or Sterkfontein Formation) comprises about 20 m of deposit, divided among six successive members. hese are numbered 1–6 from oldest to youngest, but they are not neatly stacked. he contacts are oten complex, partly because deposition oten occurred at an angle on talus slopes, partly because sediments sometimes subsided into younger, lower-lying solution cavities, and partly because solution pockets and cracks, illed with later material, sometimes penetrate the members. he result is that younger sediments can lie immediately alongside or even under older ones. his problem particularly afects Members 4 and 5, which are sometimes separated primarily on their content. Dating has proceeded mainly by faunal comparisons with eastern Africa. So far, only Members 4–6 have provided adequate numbers of fossils, and many were unfortunately collected before the members were deined. heir stratigraphic provenance is thus uncertain, and some mixing between adjacent members is likely. Mixing from Member 5 could especially explain occasional elements of Equus in Member 4. his complica-

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tion of mixing aside, however, faunal dating suggests that Member 4 (“the type site”) probably accumulated sometime between 2.8 and 2.3 Ma; that Member 5 (“the extension site”) probably formed between 2 and 1.4 Ma, perhaps in discontinuous bursts; and that Member 6 accumulated much later, ater 200 ka (ig. 4.4). he faunal estimate for Member 4 is broadly corroborated by an average electron spring resonance (ESR) age of 2.1 ± 0.5 Ma on enamel from eight bovid teeth and by paleomagnetism. he fauna from Member 2 is sparse, but it includes both lion (Panthera leo) and leopard (P. pardus), which suggest an age of 3.5 Ma or less, assuming that Panthera appeared in southern and eastern Africa about the same time. he oldest east African record, at about 3.5 Ma, is at Laetoli, northern Tanzania. Paleomagnetism, constrained by the presumed 2.8–2.3 Ma age of the higher-lying Member 4, has been used to suggest that Member 2 formed between 3.58 and 3.22 Ma; the ratio between cosmogenic aluminum-26 (26Al) and beryllium-10 (10Be) in quartz grains has supplied an estimate closer to 4.2 Ma; and the ratio between uranium-238 (238U) and lead-206 (206Pb) in lowstone bands near the fossil hominin StW 573 (discussed below) has provided an age near 2.2 Ma. he 26Al/10Be age is probably the least reliable because it depends on the unproven assumption that the dated quartz grains moved abruptly from exposure on the surface to inclusion in the cave deposit. 238U/206Pb, or more simply U/Pb dating, is a classic radiometric method that was not addressed in chapter 2 because the 4.47-billion-year half-life of the parent isotope (238U) means that it is applied mainly to rocks that long antedate hominin presence. Its application to Sterkfontein Member 2 is so far the only published exception, and the resulting age may be the most reliable one for the site. However, it is at odds with the faunal estimate of 2.8–2.3 Ma for overlying Member 4, and it would mean that climatically forced woodland reduction near Sterkfontein occurred ater 2.2 Ma. Similar reduction elsewhere in Africa coincided with the onset of Northern Hemisphere glaciation between 3 and 2.5 Ma, addressed in chapter 2. he paleomagnetic age estimate could probably be adjusted to it any radiometric age, and the bottom line is that the actual age of Member 2 remains debatable. For the moment, a time near 3 Ma may it all the data best. Members 2, 4, and 5 have produced hominin fossils, while Member 5 has also provided stone artifacts. he hominin fossils from Member 2 include twelve foot and lower leg bones extracted from breccia blocks in 1979–1980. hese are now known to associate with a skull and skeleton (Stw 573), discovered in 1997 still encased in breccia. he skeleton is remarkably complete, and it will allow irm assessment of limb and general body proportions. he foot and leg bones include eight articulating elements of the let ankle and foot (designated “Little Foot”) that were initially said to imply an apelike degree of divergence between the big toe

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and the others. Systematic comparisons to the relevant joint in people and apes have failed to conirm a signiicant diference from people, and the foot was probably specialized for bipedalism. he skeleton is most likely to represent Australopithecus africanus or its immediate ancestor. Member 4 has provided about 650 hominin specimens that most authorities attribute exclusively to A. africanus, although the degree of craniodental variability raises the possibility that multiple species are represented. hus, besides specimens of A. africanus, Member 4 may also have provided numerous specimens of a larger-toothed species that could be assigned to Paranthropus and rarer ones of one or more species that anticipate or represent Homo. Member 5 has produced roughly twenty specimens from Homo habilis, H. ergaster/erectus, or both and three teeth that probably came from Paranthropus robustus. he Member 5 artifacts have been divided between an Oldowan assemblage (3,245 pieces with no bifaces), putatively dated between 2 and 1.7 Ma, and an Early Acheulean assemblage (635 pieces including bifaces) that postdates 1.7 Ma. Much later Middle Stone Age artifacts rest on top of Member 5 but have been found in place only in the adjacent Lincoln Cave, where the deposits probably originated partly from Sterkfontein Member 5. Antelope species indicate that the environs of Sterkfontein changed from relatively moist and wooded in Member 4 times to much grassier and drier aterward. Fossil wood fragments imply that large tropical or subtropical trees grew nearby when Members 2 and 4 accumulated. he especially abundant wood fossils from Member 4 indicate that Sterkfontein lay within a riverine gallery forest composed of large closely spaced trees that supported lianas and an understory of small trees and shrubs. he mammalian fauna implies that less dense vegetation existed further from the cave. he Jacovec Cavern, which adjoins the main Sterkfontein site on the east, contains stratigraphically separate but fundamentally similar brecciated deposits. Sediments that have been dated by 26Al/10Be to about 4 Ma have provided a partial australopith skull, ive isolated teeth, and ive postcranial fragments. However, the associated fauna is said to resemble the fauna from Sterkfontein Member 4, which would imply an age closer to 2.5 Ma.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: description of the Sterkfontein Formation (Clarke 2006; Partridge 1978); complexity of the Sterkfontein deposits (Clarke 1994b; Partridge and Watt 1991); separation of Members 4 and 5 mainly by content (Kuman and Clarke 2000); faunal dating of the Sterkfontein Formation members (Clarke 1994b, 2002b; Delson 1988; McKee 1993a; Partridge 1982b, 1986; Vrba 1982, 1985a, 1995b); dating of Member 4 by ESR (Schwarcz et al. 1994) and paleomagnetism (Jones et al. 1986); Panthera in Member 2 (Pickering et al. 2004a) and the antiquity of Panthera in eastern Africa (Barry 1987; Turner and Anton 1997); dating of Member 2 by paleomagnetism (Partridge et al. 1999), by 26Al/10Be (Partridge 2005; Partridge et al. 2003), and by 238U/206Pb (Walker et al. 2006); hominin fossils from Members 2, 4, and 5 (Clarke 1994b; Clarke and Tobias 1995; Tobias 1978); stone artifacts from Member 5 (Kuman 1994a, 1994b; Robinson and Mason 1957; Stiles and Partridge 1979); StW 573 (Clarke 1998, 1999, 2002a; Clarke et al. 2003; Partridge et al. 1999); the StW 573 foot—apelike (Clarke and Tobias 1995; Dugard

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1995) or more human (McHenry and Jones 2006); craniodental variability in Member 4 hominin fossils (Lockwood and Tobias 2002); possible Paranthropus ancestor (Clarke 1988, 1994b) and Homo (Kimbel 1995; Kimbel and Rak 1993; Moggi-Cecchi et al. 1998) among the Member 4 fossils; division of Member 5 artifacts between Oldowan and early Acheulean (Kuman 1994a, 1994b, 1996; Kuman and Clarke 2000); MSA artifacts in Lincoln Cave (Reynolds et al. 2003); antelope fossils implying change in environment between Member 4 and later (McKee 1991; Vrba 1974, 1975, 1980b, 1988); Member 4 fossil wood (Bamford 1999); Jacovec cavern—australopith fossils (Clarke 2006); 26Al/10Be dating (Partridge et al. 2003), and associated fauna (Berger et al. 2002; Kibii 2007)

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

Swartkrans

he geology of Swartkrans is better understood than that of any other southern African australopith site, thanks to the insight and persistence of C. K. Brain. Like nearby Sterkfontein, Swartkrans began as a subterranean dolomite cavern into which sediments fell or were washed through a shat descending from the surface. he accumulating ill or breccia was cemented by calcite and was exposed at the surface when erosion later removed most of the cavern roof (ig. 4.7). he stratigraphy is complex because later materials sometimes illed erosion or solution hollows within earlier breccia, and objects at the same depth may thus difer greatly in age. he Swartkrans cave ill (or Swartkrans Formation) comprises ive members, numbered 1–5 from bottom to top. Member 1 includes two distinct parts—the Lower Bank below and the Hanging Remnant above— that were separated when erosion removed the intervening deposit. Member 2 subsequently illed the void and its presence both below and above Member 1 underscores the stratigraphic complexity that Brain had to unravel. Based on faunal comparisons with eastern Africa, Member 1 probably accumulated sometime between 1.8 and 1.5 Ma (ig. 4.4). Composite ESR/U-series analysis of two Paranthropus dental fragments and two bovid teeth bracket the Hanging Remnant between roughly 2.1 and 1.6 Ma. Member 2 is more diicult to date by faunal comparisons because the faunal sample almost certainly contains many elements inadvertently mixed in from Member 3, which ills a gully incised through Member 2 into the Lower Bank of Member 1. Both Members 2 and 3 appear broadly similar in fauna and artifacts to Member 1, however, and they may have formed immediately aterward, between perhaps 1.5 and 1 Ma. ESR analysis of two bovid teeth from Member 2 provided ages between 200 and 100 ka, and the incongruity suggests the teeth may actually derive from Member 4. Member 4 contains Middle Stone Age artifacts, suggesting that it postdates 250 ka. Bones from Member 5 have been radiocarbon dated to about 11 ka. Members 1–3 have provided about 300 mostly craniodental fragments from more than a hundred individuals of a robust australopith. When the irst specimens were found in 1948, Robert Broom assigned them to Paranthropus crassidens, distinct from the previously known Paranthropus

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FIGURE 4.7. Three stages in the evolution of the swartkrans australopith cave (modiied after brain [1993c], 31–32). in the irst stage, outside sediment has just begun to funnel down a recently formed shaft. in the second, erosion has partly removed earlier sediment, and fresh material has illed the erosional gap. in the third stage, erosion has totally removed the roof, and the complex ill is exposed to the elements.

dolomitic limestone

Approximately 1.8 Ma: A shaft over the southeast wall is contributing the initial sediments of Member 1 (”Lower Bank”).

travertine Mb 1 - “Lower Bank”

water Mb 1 - “Hanging Remnant” Mb 2 Mb 1 - “Lower Bank”

Shortly after 1.5 Ma: Erosion has divided Member 1 into two parts (the “Lower Bank” and the “Hanging Remnant”, and Member 2 has filled the space in between.

dolomitic limestone water

The historical present (before mining and excavation). Erosion has removed the roof of the cave, exposing a complex sequence of breccias at the surface.

Mb 1 - “Hanging Remnant” stratified Mb 2 Mb 3 Mb 2 Mb 1 - “Lower Bank”

dolomitic limestone

Three Stages in the Evolution of Swartkrans Cave robustus at nearby Kromdraai. Some specialists continue to advocate for two species, but most now regard P. crassidens as a junior synonym of Paranthropus (or Australopithecus) robustus. he large majority of robust australopith fossils come from Member 1, and erosion of Member 1 could have introduced some into Members 2 and 3. he P. robustus fossils that are in place in Member 3 could represent the last robust australopiths, dating from near 1 Ma. Members 1 and 2 have also furnished sixteen craniodental pieces that most authorities now assign to Homo ergaster or H. erectus. he specimens include a mandible (SK 15) that was the type for the now-defunct species “Telanthropus capensis” and that provided the irst indication that Paranthropus overlapped with a more advanced species. he most complete specimen of Homo is a partial skull (SK 847/45) assembled from what were once thought to be parts of separate individuals. All three Swartkrans members have provided postcranial elements

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that are diicult to separate between Paranthropus and Homo, though in advance, it seems likely that most represent P. robustus. his is particularly true for the postcranial bones in Member 1, where 97% of the diagnostic craniodental fragments come from P. robustus. Members 1–3 have yielded more than 875 laked stone artifacts and seventy-seven bone fragments that appear polished or worn from use. he stone artifacts recovered in place do not include any unequivocal bifaces, and they have been assigned to the Developed Oldowan “Culture.” Member 4 contains Middle Stone Age artifacts, while Member 5 is artifactually sterile. Occasional burned bones irst appear in Member 3, perhaps signaling incipient human control over ire. he burned bones are not locally concentrated, however, and there is no evidence for hearths. he antelopes, monkeys, and other mammals represented in Members 1–3 imply relatively dry, grassy conditions like those indicated for the broadly contemporaneous deposits of Sterkfontein Member 5.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Swartkrans—geology (Brain 1976, 1981, 1982, 1985a, 1988, 1993b, 1993c; Brain et al. 1988; Butzer 1976a), dating by faunal comparisons (Vrba 1982) and by ESR/U-series (Curnoe et al. 2001); robust australopith inventory (Brain 1981; Grine 1993b; Watson 1993a); P. crassidens—original diagnosis (Broom 1949) and subsequent recognition (Grine 1993a; Howell 1978a); fossils of H. ergaster from Members (Mbs) 1 and 2 (Grine 2005); “Telanthropus capensis” (Broom and Robinson 1949); SK 847/45 (Clarke et al. 1970); assignment of most postcranial specimens to P. robustus (Susman 1993); stone artifacts (Clark 1993b) and utilized bone fragments (Brain and Shipman 1993) from Mbs 1–3; burned bones from Mb 3 (Brain 1993d; Brain and Sillen 1988; Sillen and Hoering 1993)

Kromdraai

Kromdraai comprises two adjacent fossil caves, Kromdraai B, the “apeman site,” which has provided both australopith and animal bones, and Kromdraai A, the “faunal site,” which has provided animal bones and artifacts but no hominin fossils. Kromdraai A and B may have been part of a single cavern system into which sediments fell or were washed through shats descending from the surface, as at Sterkfontein and Swartkrans. Subsequent cementation of the cavern ills by calcite and removal of the roof(s) by erosion exposed the hardened cave ills (breccias) at the surface. he geology of Kromdraai B is better known, thanks to a systematic study associated with excavations between 1977 and 1980. Kromdraai B comprises two spatially distinct areas, called East and West. To date, hominin fossils have been found only at Kromdraai B East, where they occur in Member 3 within a sequence of ive members comprising the Kromdraai B East cave ill (or Kromdraai B East Formation). he irst fossils included an adult skull and associated partial skeleton (Transvaal Museum [TM] 1517) that are the type (and arguably the geologically oldest) specimens of Paranthropus robustus. he sample now totals about twenty hominin specimens from perhaps seven individuals of P. robustus and one of early Homo. he specimen attributed to early

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Homo is a juvenile dentition, and if the specimen has been correctly assessed, it would add to evidence from Sterkfontein, Swartkrans, and Drimlen for overlap between P. robustus and early Homo. Kromdraai B East 3 has not yet provided artifacts, but these occur in one or both of the overlying members (4 and 5). he fauna is not especially useful in dating, but the morphology of the Paranthropus fossils suggests that Member 3 accumulated about 2 Ma (ig. 4.4). It could thus fall between Sterkfontein Member 4 and Swartkrans Member 1. An age of 2 Ma is consistent with the mainly reversed polarity of Kromdraai B sediments, which places them within the Matuyama Reversed Chron, between 2.48 and 0.78 Ma. he Member 3 animal bones come mainly from cercopithecoid monkeys, including leaf-eating (colobine) forms whose presence implies woodland nearby. ESR analysis suggests that a bovid tooth of unspeciied stratigraphic origin within Kromdraai B could be 666 ky old. he artifacts from Kromdraai A include seventy-nine laked stones and twenty-one unmodiied pebbles or rock fragments that only people could have introduced. he associated animal bones come mainly from ungulates that indicate relatively dry, grassy conditions. Since the composite sequences from Sterkfontein and Swartkrans suggest increasing aridity over the period when the Kromdraai deposits formed, the fauna may imply that the Kromdraai A artifacts postdate the Kromdraai B Paranthropus remains. he Kromdraai A assemblage includes large lakes (> 10 cm in maximum length) and other pieces that broadly recall early Acheulean artifacts from Sterkfontein Member 5, and it may date from the same period, between roughly 1.7 and 1 Ma.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Kromdraai B geology (Partridge 1982a, 1982b) and associated excavations (Vrba 1981; Vrba and Panagos 1982); type specimen of P. robustus (Broom 1938); possible presence of Homo in Kromdraai B East Member 3 (Braga and hackeray 2003); morphology of the Kromdraai robust australopiths (Grine 1982) and their dating (Vrba 1982); Kromdraai B—paleomagnetism (Jones et al. 1986) and ESR dating (Curnoe et al. 2002); Kromdraai A artifacts (Kuman et al. 1997)

Coopers

he Coopers (or Cooper’s) site complex includes three or four spatially distinct fossiliferous breccias, labeled A-B, C, and D. Cooper’s A-B has provided two possible australopith teeth (a molar and an incisor) of uncertain stratigraphic origin and geologic age. Excavations in decalciied breccia at Cooper’s D have produced three isolated teeth (two deciduous, one permanent), a toothless partial mandible, cranial fragments, and at least one postcranial bone from Paranthropus robustus. More than 9,000 associated mammalian remains represent species that suggest an age between 1.9 and 1.6 Ma, coeval with Swartkrans Member 1 and Kromdraai A. Bones of canids and suids are more abundant than at any other local australopith site, suggesting a unique mode of bone accumulation.

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SOURCES: general description and in situ hominin fossils (Berger et al. 2003); hominin teeth of uncertain provenience (Berger et al. 1995; Tobias 2000)

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

Drimolen

he brecciated cave ill at Drimolen broadly resembles the inill at Swartkrans, and its faunal content implies that it accumulated in parallel with Swartkrans Member 1, between roughly 1.8 and 1.5 Ma. he hominin remains also recall those from Swartkrans in two key respects: both Paranthropus robustus and an early species of Homo are represented, and P. robustus dominates. he bones assigned to P. robustus comprise a nearly complete skull with mandible (DNH 7), a second complete mandible, three highly fragmentary skulls, twelve partial mandibles and maxillae, thirty-three isolated teeth, an ulna associated with one of the partial dentitions, and a partial pelvis and associated sacrum. he association between pelvis and sacrum is unique for the australopiths, and the ensemble serves to conirm bipedalism in P. robustus. he specimens assigned to Homo include a partial mandible, ive isolated teeth, and a radius and ulna associated with the partial mandible. An additional eleven postcranial bones have not yet been separated between P. robustus and Homo. Bones of juveniles, including infants and even neonates, are common in both the P. robustus and Homo samples. No stone artifacts have been reported, but there are but twenty-three bone fragments with polished tips like those from Swartkrans Mbs 1–3. he Drimolen sample of P. robustus augments the sample from Swartkrans and Kromdraai and is directly important for two more speciic reasons. First, in morphological and metrical characteristics, the Drimolen teeth overlap those from both Swartkrans and Kromdraai, and they therefore suggest that P. robustus should be retained as a single species and not divided between P. robustus in the narrow sense (from Kromdraai) and P. crassidens (from Swartkrans). Second, the mandible associated with the complete skull is much smaller than the second complete mandible, and the skull lacks the bony (sagittal) crest along the midline of the vault that is conspicuous on some other P. robustus skulls. he sum suggests that the skull represents a female, and the size contrast between the mandibles implies a high degree of sexual dimorphism, comparable to that inferred for P. boisei in eastern Africa. SOURCES: hominin fossils geologic antiquity of the associated fauna (Keyser 1998, 2000; Keyser et al. 2000); Paranthropus partial pelvis and sacrum (Gommery et al. 2002)

Gladysvale

Gladysvale is the remnant of a cave system that broadly resembled the systems at nearby Sterkfontein, Swartkrans, and Kromdraai. Blocks of

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breccia from mining dumps have provided an extensive fauna, including two isolated teeth assigned to Australopithecus africanus. Excavation of intact breccia has produced a hand phalange assigned to Homo, many additional mammalian fossils, and a hand ax, which is so far the only artifact. Species representation, augmented by paleomagnetic readings and ESR determinations on antelope teeth, suggests that the deposits span an exceptionally long interval from before 1.3 Ma until ater 200 ka. If the A. africanus fossils were associated with the more archaic elements in the fauna, they probably date from between 3 and 2 Ma. he phalange of Homo and the hand ax may date from around 780 ka.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: (Berger 1993; Berger et al. 1993; Hall et al. 2006; Lacruz et al. 2003; Plug and Keyser 1994; Schmid 2002)

Gondolin

Gondolin was a large subterranean cavern that received sediments through a vertical, steeply sloping shat like those that existed at the longer-known and more productive australopith caves near Sterkfontein. Abundant fossils occur in breccia blocks dumped by miners and in breccia that remains in place. Examination of blocks in the dump known as Gondolin A (GDA) has produced two hominin molars. One is too fragmentary for taxonomic assignment, but the other is an exceptionally large, let lower second molar that probably came from a robust australopith. Breccia that was excavated in place has provided a rich mammalian fauna, including a form of the extinct warthog-like species, Metridiochoerus andrewsi, that suggests an age between 1.9 and 1.5 Ma. his is the same interval implied for the robust australopith Paranthropus robustus at Swartkrans, Kromdraai, Drimolen, and Coopers. he fauna relects the abundance of rocky slopes nearby, but it implies a grassy regional environment like the one inferred for Swartkrans Members 1–3. SOURCES: hominin molars (Menter et al. 1999); mammalian fauna from intact breccia (Adams and Conroy 2005); antiquity based on Metridiochoerus (Watson 1993b)

Makapansgat

Whereas the Krugersdorp australopith caves (Sterkfontein, Swartkrans, Kromdraai, Gladysvale, Drimolen, Gondolin, and Coopers) were probably subterranean receptacles linked to a gently undulating surface by strongly sloping or even near-vertical shats, the Makapansgat Limeworks cave was more likely a tunnel-like cavern with a nearly horizontal entrance from the lank of a steep-sided valley. Relative to gravity, water low probably played a greater depositional role at Makapansgat than at the Krugersdorp sites, and some of the older deposits probably accumulated under standing water. In addition, animals, especially hyenas, may

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have regularly penetrated deeper into Makapansgat than into the Krugersdorp caves. As in the Krugersdorp sites, however, calcium carbonate, precipitated from groundwater, hardened the Makapansgat sediments ater they were deposited. he Makapansgat cave ill (or Makapansgat Formation) has been divided among ive members, numbered 1–5 from bottom to top. It is possible, however, that this scheme seriously oversimpliies much greater stratigraphic complexity and that supposedly successive members consist in part of broadly contemporaneous depositional variants or facies. Member 3 (the “gray breccia”) has provided roughly twenty-ive hominin fossils, and Member 4 has produced at least four. he irst Member 3 australopith fossil was allocated to a new species, Australopithecus prometheus, named in the erroneous belief that black staining on some of the Makapansgat animal bones implied human ability to make ire. he black color is now known to derive from manganese in the surrounding limestone. Following John Robinson, most authorities now assign the entire Makapansgat hominin sample to Australopithecus africanus, although it arguably includes a few specimens of a larger-toothed species that may also occur in Sterkfontein Member 4. A proposal that Australopithecus modiied and used many of the accompanying animal bones has been discounted, and Makapansgat has produced no indisputable artifacts. Faunal comparisons with eastern Africa imply that Member 3 formed about 3 Ma, and Member 4 is probably only slightly younger (ig. 4.4). Makapansgat is the most suitable southern African australopith site for paleomagnetic dating because the lower members were laid down partly underwater, and the sediments were not heavily disturbed aterward. he available paleomagnetic readings support an age near 3 Ma for Member 3. he Makapansgat A. africanus sample is thus probably older than the one from Sterkfontein Member 4, and it could approach the age of the one from Sterkfontein Member 2. he species composition of the fauna from Makapansgat Member 3 suggests a mix of bush and grass, perhaps similar to the historic regional savanna woodland. A broadly similar, moderately wooded environment inferred for Sterkfontein Member 4 underscores the likelihood, based on apelike upper body proportions, that A. africanus preferred or required trees for feeding, refuge, or both.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Makapansgat Cave—original form (Maguire 1985) and sedimentary sequence (Partridge 1979, 1982b); possibility that supposedly successive Mbs are actually depositional variants (Maguire 1985); hominin fossil inventory (Reed 1997); diagnosis of A. prometheus (Dart 1948); assignment of all Makapansgat hominins to A. africanus (Robinson 1954b) or partly to a larger-toothed species (Clarke 1994a); proposal that A. africanus used animal bones as tools (Dart 1957; Dart and Craig 1959) and counterargument (Brain 1981); dating by faunal comparisons (Maguire 1985; Partridge 1982b; Vrba 1982); suitability of Makapansgat sediments for paleomagnetic analysis (Rayner et al. 1993) and paleomagnetic dating (Brock et al. 1977; McFadden 1980; McFadden and Brock 1984; Partridge 1986); paleoenvironmental implications of the Mb 3 fauna (Reed 1997; Sponheimer et al. 1999)

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Geology of the East African Australopith Sites In contrast to southern Africa, where the australopith fossils come exclusively from caves, in eastern Africa, they originate from ancient stream or lake deposits that have been exposed by recent gullying. he known sites are closely associated with the eastern branch of the Great Rit Valley (ig. 4.1), which began forming more than 20 Ma, when tension between massive continental plates forced a strip of land more than 2,000 km long and 40–80 km wide to fall with respect to its sides. Associated tectonic activity has repeatedly created and destroyed natural dams, alternately creating and draining lakes and redirecting streams. he same tectonic activity frequently faulted or tilted preexisting lake and river deposits, while crustal movement, oten sparse vegetation, and episodically violent rainfall have encouraged erosion, which has favored fossil exposure and discovery. Vulcanism, also linked to tectonic activity, has repeatedly contributed lavas and ashes (tufs) that can be used to date enclosing lake and river deposits. In many places lowing water displaced fossils from their original resting places even before burial, and most were discovered only ater they had eroded out. In some places, however, fossils and artifacts occur in near primary position, in ancient “living sites.” his is especially true at Olduvai Gorge, Koobi Fora (East Turkana), the Lokalalei site (West Turkana), and in the Gona region of the Awash Valley (Ethiopia), as discussed below. he western branch of the Rit Valley has been much less productive, in large part because the associated deposits are less richly fossiliferous and the fossils are less frequently exposed. As with the previous section on southern African sites, a reader who does not need or want detailed information on speciic east African sites may prefer to skim the individual sketches, attending mostly to the accompanying igures. he sites are summarized in rough order from south to north. he principal conclusions are summarized in igures 4.5 and 4.6 and in the text sections that follow this one.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: geologic history of eastern branch of the Great Rit Valley (de Heinzelin 1994)

Chiwondo Beds (Uraha, Malema, and Mwimbi)

he Chiwondo Beds cover an area of about 70 × 10 km on the western margin of the Rit Valley in northern Malawi. hey formed under the predecessor of historic Lake Malawi and in streams that fed it. One hundred forty-ive discrete fossiliferous localities have been identiied, but they have yielded only about 1,000 vertebrate specimens. hese include two hominin fossils—a mandible (UR 501) from Uraha in the south and a maxillary fragment (RC 911) from Malema in the north. Associated mammals place both specimens between 2.5 and 2.3 Ma. he mandible has been assigned to Homo rudolfensis, and it is one of only ive speci-

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mens older than 1.9 Ma attributed to Homo. Table 4.11 lists the others. he maxillary fragment has been tentatively assigned to Paranthropus boisei, but it is too incomplete for irm diagnosis, and it might represent the older P. aethiopicus. he Mwimbi locality has produced Oldowan artifacts that are thought to have accumulated between 2.4 and 1.6 Ma.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: geology and fossils (Sandrock et al. 1999; Schrenk et al. 1995); UR 501 mandible (Bromage et al. 1995; Ramirez Rozzi et al. 1997; Schrenk et al. 1993); RC 911 maxilla (Kullmer et al. 1999) and its uncertain assignment within Paranthropus (Wood and Strait 2004); Mwimbi artifacts (Kaufulu 1990; Kaufulu and Stern 1987)

Olduvai Gorge

Olduvai Gorge is a normally dry valley on the western margin of the Eastern Rit Valley in northern Tanzania. It was irst investigated in 1913 by Hans Reck who reported fossils that later caught the attention of Louis Leakey. In 1931 Leakey, Reck, and A. T. Hopwood returned for further work, and in 1935 Louis and Mary Leakey initiated repeated visits that recovered large samples of artifacts and animal bones and that culminated in the discovery of the famous “Zinjanthropus” australopith skull in 1959. Between 1960 and 1973, Mary Leakey excavated many early archaeological sites at Olduvai, some of which provided additional human fossils. he Leakeys’ research revolutionized paleoanthropology, not only because it provided key fossils and artifacts at Olduvai but also because it encouraged others to tap the great paleoanthropological potential of other east African sites. Olduvai itself is not exhausted and specialists following in the Leakeys’ footsteps have discovered additional human fossils, animal fossils, and artifacts. he Gorge comprises two usually dry branches, known as the Main Gorge and the Side Gorge (ig. 4.8). hey have a combined length of about 50 km and they expose up to 100 m of lacustrine, luvial, and eolian deposits within a shallow basin. Important sites oten occur where gullies cut into the sidewalls of the Gorge, and the sites are oten named for a combination of their discoverer and the Swahili word for gully (Korongo), as in “Frida Leakey Korongo,” abbreviated FLK. he deposits are divided among seven successive mappable stratigraphic units known from bottom to top as Bed I, Bed II, Bed III, Bed IV, the Masek Beds, the Ndutu Beds, and the Naisiusiu Beds (ig. 4.9). At various levels within the sequence, volcanic tufs aid in horizontal correlation and mapping, which has been complicated by extensive faulting. K/Ar dating, 40Ar/39Ar dating, paleomagnetism, and 14C dating ix the Olduvai sequence between 2.03 Ma and 15 ka. During the accumulation of Bed I, the Olduvai Basin contained a broad, shallow saline and alkaline lake that luctuated up to 3 m in depth and varied between 7 and 15 km in diameter. Most archaeological

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158 FIGURE 4.8. The location of olduvai Gorge on the serengeti Plain (redrawn after hay [1990], ig. 1).

N 0

10

20 km major fault

SERENGETI PLAIN Main Olduvai Gorge

Masek Lake

Side Gorge

Laetoli

al

FLK

alb

3o S

DK MK

Lake Ndutu

Ol b

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

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VOLCANIC HIGHLANDS Ngorongoro Crater

35o E

and paleontological sites occur along the southeastern lakeshore where freshwater streams lowed into the lake from nearby volcanic uplands. K/Ar and 40Ar/39Ar dates indicate that volcanic deposits at the base of Bed I formed about 2.03 Ma but that most of the sedimentary deposits, including all the fossiliferous sites, accumulated rapidly between about 1.86 and 1.75 Ma. Lava lows and associated sediments at the base of Bed I exhibit reverse magnetism, but the overlying deposits into the lower part of Bed II show normal magnetism. he normally magnetized deposits deine the Olduvai Normal Subchron, which interrupted the Matuyama Reversed Chron between about 1.95 and 1.77 Ma. he lake persisted during the deposition of Lower Bed II, roughly until the accumulation of a sequence of eolian tufs known as the Lemuta Member. A major disconformity above the Lemuta Member marks the beginning of widespread faulting and folding in the central part of the basin. his forced the lake to shrink, and it disappeared entirely shortly before the end of Bed II deposition. he disconformity also marks an abrupt faunal change at Olduvai, ater which some archaic large mammals represented in Bed I and Lower Bed II no longer occur and several new species make their irst appearance. Another disconformity separates Bed II from overlying Bed III and is believed to relect a major phase of Rit Valley faulting that has been radiopotassium dated elsewhere to 1.20–1.15 Ma. hus Bed II is believed to have formed between about 1.75 and 1.20 Ma. he top of the Olduvai Normal Subchron oc-

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100 erosion alternating Naisiusiu Beds with fluvial and eolian deposition Ndutu Beds 90 gorge incision

Masek Beds

alluvial plain

shrinking lake

80

Beds III & IV

major volcanic markers paleomagnetism & 40Ar/39Ar ages (Ma) and inferred ages (Ma) ? H. sapiens Norkilili Mb

Jaramillo Normal Subchron

Tuff IVB Tuff 4

70

Tuff 2

60

Lemuta Mb w/ Tiff IIa Tuff IF

Tuff IID

Bed II “faunal break”

50

40

lake Bed I

30

20

1.749 + 0.007

Tuff IE

1.75 + 0.020

Tuff ID

1.76 + 0.028

lava Tuff IA

0.99 1.07

1.77/1.79

Paranthropus boisei Homo habilis

Oldowan

Olduvai Normal Subchron

Tuff IC

Tuff IB

Middle & Later Stone Age

H. ergaster Acheulean & (or H. erectus) Developed Oldowan Matuyama Reversed Chron

thickness (meters)

1.798 + 0.004 1.859 + 0.007 1.865 + 0.020 1.976 + 0.015

10

1.95 0

Naabi Ignimbrite

2.029 + 0.005

curs just below the Lemuta Member, and the disconformity above it that separates Lower Bed II from Upper (or Middle and Upper) Bed II must thus date to about 1.7–1.6 Ma. he widespread faulting that produced the disconformity between Bed II and Bed III led to substantial erosion of Bed II. he Olduvai Basin was transformed into an alluvial plain, and Beds III and IV were primarily stream laid. hey are readily distinguished only on the eastern side of the basin. he Masek Beds are mainly luvial deposits similar to those of Beds III and IV and were the last sediments to accumulate before major incision of the gorge. An early paleomagnetic study suggested that Bed IV incorporated the boundary between the Matuyama Reversed Chron and the Brunhes Normal Chron at 780 ka. Fresh research suggests that Bed IV sediments are reversed more or less throughout, that the lower part of the Masek Beds is reversed, and that the upper part is again normal. It seems most likely that the upper part of the Masek Beds formed during the Jaramillo Normal Subchron between 1.07 and 0.99 Ma, in which case the entire underlying Olduvai sequence formed within the Matuyama Reversed Chron. he Ndutu Beds are luvial and eolian deposits that accumulated over a lengthy period of intermittent faulting, erosion, and partial illing of the gorge. he upper unit of the Ndutu Beds contains eolian tufs that

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FIGURE 4.9. schematic stratigraphy of olduvai Gorge, showing the principal beds, their depositional environment, their paleomagnetism, the main volcanic stratigraphic markers, their 40ar/39ar ages, and the local stratigraphic extent of hominin taxa and artifact industries (modiied after leakey and hay [1982], ig. 2; and Tamrat et al. [1995], ig. 6).

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are mineralogically similar to but much more weathered than tufs in the Naisiusiu Beds, whose age has been established by the radiocarbon method. If a constant weathering rate is assumed, the upper unit of the Ndutu Beds has a mean age of 75 ka, from which it has been suggested that the Lower Ndutu Beds accumulated between roughly 400 ka (the top of the Masek Beds) and 75 ka. Finally, the Naisiusiu Beds are mainly eolian tufs deposited ater the Upper Ndutu Beds had been severely eroded and ater the gorge had been incised to its present level. Radiocarbon dates ix the Naisiusiu Beds between about 22 ka and 15 ka. Past climates at Olduvai can be inferred from the proportionate representation of windblown sediments, the stable oxygen- and carbon-isotope composition of soil carbonates, pollen, and vertebrate remains. Overall, the vicinity of the gorge became drier from 2 Ma to the present, but the trend was interrupted by wet periods that may correspond to Northern Hemisphere interglacials. he upper part of Bed I (between Tufs IB and IF in ig. 4.9), which probably accumulated in less than 20 ky, records a dramatic shit from wetter to drier and back to wetter. he abruptness of the change recalls equally rapid shits that are recorded in late Pleistocene and Holocene deposits in eastern Africa. Fossils of a robust australopith (Paranthropus boisei) have been found in Bed I, Lower Bed II, and Upper Bed II, while fossils assigned to Homo habilis occur in Bed I and Lower Bed II. Fossils of H. ergaster (or H. erectus) have been found in Upper Bed II, Bed III, and Bed IV, while highly fragmentary fossils that may derive from early H. sapiens are known from the Masek and Lower Ndutu Beds. A skull of early H. sapiens from Lake Ndutu, at the headwaters of Olduvai Gorge, came from deposits that are probably coeval with the Upper (Norkilili) Member of the Masek Beds. Stone artifacts occur throughout the Olduvai sequence: Oldowan in Bed I and Lower Bed II; Developed Oldowan and Acheulean in Upper Bed II to Bed IV; Acheulean in the Masek Beds; Middle Stone Age near the top of the Lower Ndutu Beds and in the Upper Ndutu Beds; and Later Stone Age in the Naisiusiu Beds. he Lake Ndutu skull, from deposits that may correlate with the Upper Masek Beds, was associated with Acheulean artifacts.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: investigation of Olduvai Gorge by Reck (1914), M. D. Leakey (1971; 1975; Leakey and Roe 1994), and others aterward (Blumenschine and Masao 1991; Blumenschine et al. 2003; Johanson et al. 1987); Olduvai Gorge stratigraphy (Hay 1976, 1990; Leakey 1978, 1980; Leakey and Hay 1982) and dating (Evernden and Curtis 1965; Leakey et al. 1961; Tamrat et al. 1995; Walter et al. 1991); paleomagnetism of Bed IV and the Masek Beds (Tamrat et al. 1995); Olduvai paleoclimate inferred from windblown sediments (Hay 1976), stable oxygen and carbon isotopes in soil carbonates (Cerling and Hay 1986), pollen (Bonneille 1984), and vertebrate remains (Fernández-Jalvo et al. 1998; Kappelman 1984); possible correlation between Olduvai wet intervals and northern hemisphere interglacials ater 2 Ma (Kappelman 1986); rapid climatic shit in Upper Bed I (Walter et al. 1991); Ndutu skull (Leakey and Hay 1982)

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Laetoli

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

he Laetoli locality is approximately 45 km south of Olduvai Gorge in northern Tanzania (ig. 4.8). Technically, it was the irst site in eastern Africa to provide australopith fossils, comprising a lower canine found in 1935 and a small fragment of a right maxilla, with premolars and an isolated third molar, found in 1939. However, the canine was initially misidentiied as cercopithecoid, and the signiicance of the maxillary fragment and molar was appreciated only much later, following carefully controlled ieldwork at the site between 1974 and 1979. he controlled research produced twenty-three australopith fossils, including isolated teeth, jaws, and a highly fragmentary immature skeleton. Most of the fossils were found on the surface, but they clearly derive from the upper part of the Laetolil Beds (also known as the Garusi or Vogel River Series), which consist mainly of eolian and air-fall volcanic tufs covering an area of about 70 square km in the Laetoli region. Based on their indspots, the individual Laetoli australopith fossils can be broadly related to eight widespread marker tufs, numbered 1–8 from bottom to top. hree of the systematically recovered specimens came from below Tuf 5 and twenty from between Tufs 5 and 8. he age of the entire sample is ixed between 3.76 and 3.46 Ma by radiopotassium estimates from below Tuf 1 and from Tuf 8, respectively (ig. 4.10). All twenty-six known hominin fossils have been assigned to Australopithecus afarensis, for which a mandible (Laetoli Hominid 4) that probably slightly antedates 3.46 Ma is the holotype (type fossil). A. afarensis was probably responsible for some spectacular human footprints that formed about 3.6 Ma on a paleosurface within Tuf 7 (the “footprint tuf ”). Surfaces within the tuf also preserve abundant prints from a wide variety of animals. In addition to australopith fossils, the Laetoli locality has provided a massive, heavily abraded mandible (Laetoli Hominid 29), tentatively assigned to Homo erectus (or H. ergaster as used here) and an archaic Homo sapiens skull (LH 18). he skull came from the Upper Ngaloba Beds, which cap the local stratigraphic sequence, and the mandible probably came from the somewhat older Lower Ngaloba Beds. A girafe vertebra from the same level as the skull has been dated to 129 ± 4 ky ago by 230h and to 108 ± 30 ky ago by 231Pa. hese dates must be provisional, but they support a previous age estimate for the skull. his was based on the skull’s position just above a tuf that is mineralogically very similar to one near the top of the Lower Ndutu Beds at Olduvai. Based on both its stratigraphic position and the extrapolated sedimentation rates at Olduvai, the correlated Ndutu Tuf appears to have an age of 120 ± 30 ka. No stone artifacts have been found in the Laetolil Beds. However, Acheulean or Developed Oldowan bifaces and other artifacts occur in

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162

ChaP Ter f our

FIGURE 4.10. Left: schematic stratigraphic column for the laetoli area, showing the position and age of dated horizons. Right: dated tuff horizons within the upper unit of the laetolil beds (redrawn after drake and Curtis [1987], igs. 2.10, 2.11).

Schematic stratigraphic column for the Laetoli area showing the position and age of dated horizons Age (Ma + s.d.) Ngaloba Beds Olpiro Beds Naibadad Beds Ogol Beds Upper Unit Ndolanya Beds Lower Unit Ndolanya Beds Upper Unit Laetolil Beds

2.26 + 0.06 2.41 + 0.12

Dated tuff horizons within the upper unit of the Laetolil Beds Age (Ma + s.d.)

3.49 + 0.12 3.46 + 0.12 3.49 + 0.11 3.56 + 0.2

8 7

Phonolite air-fall tuff

6 3.76 + 0.03

Lower Unit Laetolil Beds

5 4 3 2

Claystone Tuff 20 m

Melitite-carbonatite air-fall tuff Horizon of lapilli & blocks Eolian tuff 20 m

Lava flow Erosional surface 4.32 + 0.06

1

10

0

3.76 + 0.03

the Olpiro Beds, which unconformably overlie the Laetolil Beds and which may have formed about the same time as Olduvai Bed II, ater 1.7 Ma. he Ngaloba Beds, unconformably overlying the Olpiro Beds, contain Middle Stone Age artifacts; some were directly associated with the Ngaloba skull (LH 18). SOURCES: initial canine (White 1981) and maxillary fragment and isolated molar (Leakey 1987a) of A. afarensis; Laetoli deposits (Harris 1985; Hay 1987; Leakey 1980; Leakey and Hay 1982; Leakey et al. 1976) and dating (Drake and Curtis 1987); holotype for A. afarensis (Johanson et al. 1978); footprint tuf (Hay and Leakey 1982; Leakey 1987b; Leakey and Hay 1979); LH 29 and LH 18 (Leakey 1987c); dating of LH 18 (Hay 1987); Acheulean or Developed Oldowan in the Olpiro Beds (Harris and Harris 1981)

Peninj

he Peninj site is west of Lake Natron, approximately 80 km northeast of Olduvai Gorge in northern Tanzania. he fossiliferous sediments at Peninj accumulated mostly in and around a lake within a broad, relatively shallow basin. he deposits have been divided into two major units—the Humbu Formation (older) and the Moinik Formation (younger). Paleomagnetic readings and radiopotassium determinations on a basalt near the base of the Moinik Formation and on a basalt within the Humbu Formation suggest that the Humbu Formation probably dates from be-

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tween 1.6 and 1 Ma. his age is consistent with the Humbu Formation fauna, which resembles that from Upper Bed II at Olduvai Gorge. he mandible of a robust australopith was found in Humbu Formation deposits that are probably just older than 1.6 Ma, and Acheulean artifacts occur in deposits that are probably only slightly younger. hey are exceeded in antiquity only by Acheulean artifacts from West Turkana and the Gona Region, Middle Awash.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: geology (Domínguez-Rodrigo et al. 2002; Isaac 1967), dating (Isaac and Curtis 1974), and fauna (Geraads 1987)

Tugen Hills, Baringo (Lukeino, Chemeron, and Chemoigut)

he Tugen Hills, west of Lake Baringo, central Kenya, contain more than 3,000 m of luviatile and lacustrine sediments spanning the period between roughly 16 Ma (mid-Miocene) and near present (Holocene). he sequence has been divided among twelve successive geologic formations, including seven that have provided hominoid (including hominin) fossils, artifacts, or both. Table 4.1 lists the sedimentary units and their fossil or artifactual contents, from older to younger. For the purposes of this chapter, the most important units are the Lukeino Formation (Fm) with fossils of Orrorin tugenensis, dated to roughly 6 Ma; Member 1 of the Chemeron Fm with a mandible fragment possibly from Ardipithecus ramidus, dated to about 4.42 Ma; Member 4 of the Chemeron Fm with a temporal fragment of early Homo, dated to 2.4 Ma; and the Chemoigut Fm with two skulls and some fragmentary teeth of Paranthropus boisei, dated between 2 and 1.5 Ma. As discussed below, the fossils of O. tugenensis include a nearly complete proximal femur with features that imply bipedalism, and if this assessment is correct, O. tugenensis is presently a plausible ancestor for all later hominins. If the taxonomic assignment and geologic antiquity of the Chemeron temporal fragment have been correctly assessed, it is one of only ive fossils (listed in table 4.11) that possibly or probably document Homo before 1.9–1.8 Ma. he fossils are not only rare but incomplete as well, and they are useful only for demonstrating the presence of Homo by 2.5–2.4 Ma. As discussed below, Paranthropus appears to have emerged simultaneously, and the implication is that a branching speciation event, with Australopithecus afarensis at its root, occurred between 2.8 and 2.5 Ma. It is the oldest branching event that nearly all authorities accept. he fossils of Paranthropus boisei from the Chemoigut Fm are broadly associated with Oldowan artifacts, but for reasons discussed below, early Homo was the probable artifact maker. At one Oldowan site, the artifacts were directly associated with baked clay fragments that may represent the oldest traces of humanly controlled or produced ire.

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Sedimentary units with hominoid (including hominin) fossils, artifacts, or both in the Tugen Hills (Baringo) region, central Kenya. Geologic ages have been established mainly from K/Ar and 40Ar/39Ar determinations, supplemented by paleomagnetism and mammalian associations. TABLE 4.1.

Geologic Unit

Fossils and Artifacts (Where Present)

Geologic Antiquity

References

Muruyur Fm (or Beds)

Various large hominoid fossils, including a partial skeleton and 38 isolated teeth from at least 4 other individuals of Equatorius africanus (Kipsaramon site complex)

16 - 13 Ma

(Behrensmeyer et al. 2002; Hill 1994; Hill et al. 1985; Kelley et al. 2002; Sherwood et al. 2002b; Ward et al. 1999b)

Ngorora Fm

2 or perhaps 3 teeth from at least two large hominoids, including possibly the youngest representative of Proconsul.

13 - 8.5 Ma

(Hill 1994; Hill et al. 1985; Hill et al. 2002)

Mpesida Beds

(No hominoid specimens)

7 - 6.2 Ma

(Kingston et al. 2002)

Lukeino Fm

44 mapped fossiliferous localities, including 4 which have provided 20 fossils of Orrorin tugenensis.

6.2 - 5.6 Ma

(Deino et al. 2002; Galik et al. 2004; Gommery & Senut 2006; Pickford & Senut 2001; Pickford et al. 2002; Sawada et al. 2002; Senut et al. 2001)

Chemeron Fm

Mb 1: A hominin proximal humerus (KNM-BC 1745) and a partial mandible (KNM-TH 13150) tentatively referred to Ardipithecus ramidus (Tabarin locality). Mb 4: a temporal fragment (KNM-BC 1) assigned to Homo sp. (JM85 locality).

5.6 - 1.6 Ma

(Deino & Hill 2002; Deino et al. 2002; Feibel et al. 1992; Hill 1989; Hill et al. 1985; Hill et al. 1992b; Pickford et al. 1983; Sherwood et al. 2002a; Ward & Hill 1987)

Chemoigut Fm

Two fragmentary skulls and some fragmentary teeth assigned to Paranthropus boisei; artifacts of the Chemoigut variant of the Oldowan industry; baked clay fragments possibly representing the oldest traces of humanly controlled ire.

2 - 1.5 Ma

(Bishop et al. 1978; Bishop et al. 1975; Carney et al. 1971; Gowlett et al. 1981; Hooker & Miller 1979)

Chesowanja Fm No hominin remains. Losokweta < 1.42 Ma variant of the Acheulean or Developed Oldowan Industrial Tradition.

(Gowlett et al. 1981; Harris & Gowlett 1980)

Kapthurin Fm

(Cornelissen et al. 1990; Day 1986b; Deino & McBrearty 2002; Leakey et al. 1969a; McBrearty 1999; McBrearty et al. 1996; McBrearty & Jablonski 2005; Solan & Day 1992; Tallon 1978; Tryon & McBrearty 2002; van Noten 1983; Wood & van Noten 1986)

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Two mandibles, a phalanx, a right metatarsal, and a right ulna assigned here to “early Homo sapiens” or H. heidelbergensis. Also four teeth assigned to Pan, probably P. troglodytes. Late Acheulean and Middle Stone Age artifacts.

164

700 - < 200 ka

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However, natural baking—for example, below a smoldering tree stump— cannot be ruled out.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES (also table 4.1): on Tugen Hills sequence (Bishop 1971, 1978; Bishop et al. 1971; Hill 1994, 2002; Hill et al. 1985); baked clay and ire in the Chemoigut Formation (Clark and Harris 1985)

Lothagam and Kanapoi

Lothagam and Kanapoi are approximately 75 km apart, in the drainage of the Kerio River southwest of Lake Turkana in northern Kenya (ig. 4.11). At both sites, the sequence comprises lacustrine sediments from a greatly expanded Lake Turkana and luvial sediments from when the lake had substantially shrunk. he 900 m thick sedimentary sequence at Lothagam has been subdivided (from bottom to top) among the Nabwal Arangan Beds, the Nawata Formation, the Nachukui Formation, and the Galana Boi Formation. he Nachukui Formation was irst described in the region northwest of Lake Turkana, where it has provided important hominin fossils and some of the oldest known archaeological sites, both discussed below. he Nawata and Nachukui Formations at Lothagam have produced abundant mammalian fossils, including herbivore teeth whose carbon-isotope composition documents the shit from C3 (less grassy) to C4 (grassier) vegetation that occurred in tropical Africa beginning about 8 Ma. Carbon isotopes in Lothagam soils record the same change, and Lothagam data thus support the idea that an expansion of grassy settings prompted hominin emergence. Lothagam has so far provided no early artifact occurrences and only seven possible or probable hominin fossils, listed from older to younger in table 4.2. he specimens are all too incomplete for unambiguous taxonomic assignment, but the partial mandible (KNM-LT 329) shares relatively thin molar enamel with Ardipithecus ramidus and a molar root pattern and other features with Australopithecus afarensis and especially Au. anamensis. If its suggested age near 5 Ma is correct, it could be oldest known specimen of Australopithecus. At Kanapoi, the sequence consists of a basal luvial unit overlain by lacustrine deposits covered in turn by luvial sediments. A deltaic facies (variant) of the lake deposits provided a hominin distal humerus in 1965 and a highly fragmented juvenile skull, eight partial mandibles, two partial maxillae, about thirty isolated teeth, a tibia, a proximal hand phalange, and a wrist bone (capitate) between 1994 and 1997. 40Ar/39Ar dating brackets all the hominin fossils between about 4.2 and 4.0 Ma. hey have been assigned to Australopithecus anamensis, which has also been identiied in 4.2–4.1-my-old deposits in the Middle Awash (Ethiopia) and in 3.9-my-old deposits at Allia Bay on the eastern shore of Lake Turkana in the Middle Awash, described below, associated mammalian fossils imply a relatively moist, wooded paleoenvironment, while at Kanapoi

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Usno Riv er

e

ar-K okk Ham m let

km

Il

ILERET

Lake Turkana

0

oi

k Ko

1000

ap ate G

Bak

Koobi Fora Fm. ▲ KUBI ALGI

KAT AB OI

S

▲ SIBOLOT ▲ JARIGOLE

HU RH ILL

to Narioko

Lothidok Range

0o

Er

Labur Range

me

Marua Rith Range ol

ak Kal

sin

Guragha Mts.

Turkana Basin

Ba

Omo River

10 N

ali

Lorienetom Range

o

CHEW BAHIR

ri Kara

WEST TURKANA Nachukui Fm.

en f Ad

o Gulf

Hadar Region

(Lake Stephanie)

LOKITAUNG

4o N

ea dS

Shungura Fm.

Re

LOKOMARINYANG

38o 40oE

Usno Fm.

KIBISH

5o N

37o

So m

36o

eR a ng

Mursi & Nakalabong Fm.

ei

FIGURE 4.11. The lake Turkana basin showing the geological formations (in boldface) that have provided fossils of australopiths, early Homo, or both (redrawn after brown [1994], 287). Collectively, localities within the Koobi fora formation are often referred to as east Turkana, those within the nachukui formation as West Turkana, and those within the usno and shungura formations as omo or lower omo. in total, they have provided more than 70,000 vertebrate fossils, including more than 450 hominin specimens.

g Sure

KALAKOL

▲ MOITI

NORTH HORR

Koobi Fora Fm. GUS

MARSABIT BARRIER VOLCANICS ▲ MT. NYIRU

IYU

SUG PLA TEA UTA U VAL LEY

Rive Kerio

KANAPOI

LOR

Turkwell River

r

EKORA

ie As

T ER ES ID

LOTHAGAM ▲

LOWASERA

B AL CH

Allia Bay

LODWAR

3o N

Mt. Kulal

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

2o N

0

20

40

60 km

Hominin fossils from Lothagam, northern Kenya. Dating is based on K/Ar, 40Ar/39Ar, mammalian associations, estimated sedimentation rates, or a combination of these.

TABLE 4.2.

Geologic Unit

Fossils

Estimated Geologic Age

References

Upper Nawata Fm

a let M3 and a right I1 (?Hominini indet.)

somewhat more than 5.23 Ma

(Leakey & Harris 2003)

Apak (basal) Mb of the Nachukui Fm

a right mandible fragment (KNM-LT 329) with heavily worn M1 and roots for the P4, M2, and M3 (?Ardipithecus ramidus).

ca. 5 Ma

(Kramer 1986; Leakey & Walker 2003; Patterson et al. 1970; White 1986b)

Kaiyunmung Mb of the Nachukui Fm

partial teeth tentatively identiied as a right dP3, a right M3, a let M2, and an indeterminate P (?Australopithecus afarensis)

ca. 3.5 Ma

(Leakey & Harris 2003)

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and Allia Bay, they suggest a mosaic of bush and grass, with strips of forest or woodland along water channels. Stable-isotope analyses of fossil soil carbonates at Kanapoi and of mammalian teeth at Allia Bay conirm this reconstruction, but they suggest that trees and shrubs may have been more common than they were in historic times.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: initial research at Lothagam and Kanapoi (Behrensmeyer 1976); Lothagam sequence (Feibel 2003); mammalian fossils from Lothagam (Leakey and Harris 2003) and their carbon-isotope implications (Cerling et al. 2003); resemblance of KNM-LT 329 to A. afarensis (Hill 1994; Hill and Ward 1988; Hill et al. 1992a) and A. anamensis (Leakey and Walker 2003); Kanapoi sediments (Leakey et al. 1995a), distal humerus (Patterson and Howells 1967), and remaining hominin fossils (Leakey et al. 1995a, 1998; Ward et al. 1997, 2001); stable-isotope analysis of carbonates at Kanapoi (Wynn 2000) and of mammalian teeth at Allia Bay (Schoeninger et al. 2003)

Koobi Fora (East Turkana, a.k.a. East Rudolf)

he localities collectively called Koobi Fora are on the east side of Lake Turkana (formerly Lake Rudolf) in northern Kenya (ig. 4.11). he fossiliferous sediments at Koobi Fora were laid down mainly by streams lowing toward Lake Turkana from uplands to the east. Lacustrine sedimentation in the Turkana Basin began shortly before 4 Ma and was probably initiated by the eruption of basalts that disrupted previous drainage patterns. Between 4 and 2.5–2 Ma, the basin was sometimes illed by a large lake, but it was more oten dominated by an extended Omo (or “Turkana”) River that lowed south and east to the Indian Ocean. A lake like the historic one was present more continuously ater 2 Ma. Vulcanism and tectonic movement determined the alternation between lake and river, but when a lake was present, climatic factors afected its volume. More than 130 layers of volcanic tuf (or tephra) have been identiied in the Lake Turkana Basin. hese provide material for radiopotassium, 40 Ar/39Ar, and ission track dating, and their geochemical signatures are oten unique. hey thus allow stratigraphic correlations within the Koobi Fora region and between Koobi Fora and other areas, including the lower Omo River Valley to the north, the West Turkana region to the west, the Kerio Valley to the southwest, the Hadar and Middle Awash regions of east-central Ethiopia, and even deep-sea cores from the Gulf of Aden. he Plio-Pleistocene sequence in the Koobi Fora region comprises over 560 m of luvial, lacustrine, and deltaic deposits that are grouped into the Koobi Fora Formation. his has been divided in turn among eight successive members, each named for a tuf at its base (igs. 4.12– 4.14). A combination of K/Ar and 40Ar/39Ar dates, paleomagnetic readings, and faunal correlations indicates that the deposits accumulated discontinuously between about 4.34 and 0.7 Ma. In retrospect, the Koobi Fora Formation provides a highly instructive example of how radiometric dates may be checked by faunal remains. Initially, dates of approximately 2.6 Ma for the KBS Tuf implied that various mammal taxa

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Chari 1.39 Okote 1.56 KBS 1.89 U. Burgi

K Okote

J

KBS Lorenyang

H

9

L. Burgi Tulu Bor 3.32 Lokochot 3.5 Moiti 3.89

8 6 4

7 5

G

2.2 2.68

Molluscan biozonation of W illiamson 10

L

Chari

Loxodonta africanus

C

Metridiochoerus andrewsi

B

Mesochoerus limnetes

A

Notochoerus capensis

Mb

TUFFS Silbo

Vertebrate biozonation of Maglio Metridiochoerus compactus

MEMBERS > 0.6

Metridiochoerus andrewsi

Koobi Fora Formation

Notochoerus scotti

Vertebrate biozonation of J. M. Harris Shungura Fm

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

ChaP Ter f our

3

Burgi

C 2

Hasuma B Lokochot

A 1

Moiti

Lonyumun < 4.3

FIGURE 4.12. schematic stratigraphy of the Koobi fora formation, with correlations to the shungura formation of the lower omo river valley and to biostratigraphies suggested by harris, maglio, and Williamson (modiied after brown and feibel [1986], ig. 9). Notochoerus and Metridiochoerus are genera of extinct pigs. Loxodonta africana is the living african elephant.

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(Equus, for example) had appeared at Koobi Fora at least 600 ky before they irst appeared in the lower Omo River Basin (Shungura Formation) at the north end of Lake Turkana. his seemed unlikely and prompted redating, which eventually showed that the original 2.6 my estimate was about 700 ky too old. he Koobi Fora Formation has provided fossils of australopiths or early Homo at Koobi Fora proper, where they are dated between roughly 2.1 and 1.3 Ma, and at Allia Bay, about 40 km to the south, where they are dated to about 3.9 Ma. Table 4.3 lists the individual hominin taxa represented, the geologic units from which they come, and their estimated geologic age. he table includes modern or near-modern Homo sapiens, based on a skull and a robust femur that may be as old as 300 ka. he specimens are considered in chapter 5 on the Neanderthals and their contemporaries. he oldest stone artifacts found at Koobi Fora occur just below and within the KBS Tuf, where they are dated to about 2–1.8 Ma. hey have been assigned to the KBS Industry, and they resemble broadly contemporaneous Oldowan artifacts from Bed I at Olduvai Gorge. Somewhat younger artifacts from the Okote Member, dated to roughly 1.4 Ma, have been assigned to the Karari Industry, which is broadly similar to the Developed Oldowan at Olduvai. An early Acheulean site has been excavated in deposits that probably correlate with the Chari Member, slightly postdating 1.4 Ma. However, the early Acheulean is poorly represented at

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Lower Omo: Shungura & Usno Formations L K J H G-G-14 G-G-13 F E D C B A

East Turkana: Koobi Fora Formation

West Turkana: Nachukui Formation

Chari

Nariokotome

Okote

Natoo

KBS U. Burgi

Kaitio Kalochoro

L. Burgi Lokalalei

Basal

Tulu Bor

Lomekwi

Lokochot Moiti Lonyumun

Lonyumun

10 20 30 40 50 60

Kataboi

10 20 30 40 50 60 70 80 90

10

Number of early hominin fossils

Koobi Fora, particularly compared with its abundance at Konso, roughly 200 km to the northeast. he diference may relect the rarity of suitable raw material for hand ax manufacture at Koobi Fora. SOURCES: origin of Koobi Fora fossiliferous sediments (Vondra and Bowen 1976); history of lacustrine sedimentation in the Turkana Basin (Brown 1994, 1995); climate impact on past Lake Turkana volume (Rogers et al. 1994); tephra correlations between Koobi Fora and other regions (Brown 1994; Brown et al. 1985b, 2006; Feibel et al. 1989); subdivision of the Koobi Fora Formation (Brown and Feibel 1986, 1991; Feibel et al. 1989); redating of the KBS Tuf (Brown et al. 1978; Hay 1980); hominin fossils from Koobi Fora (Walker 1981b; Walker and Leakey 1978) and Allia Bay (Coing et al. 1994; Leakey et al. 1995a, 1998; Ward et al. 2001); near-modern hominin fossils from Koobi Fora (Bräuer 2001); Koobi Fora artifacts (Harris and Isaac 1976; Isaac and Harris 1978; Isaac et al. 1997); Konso early Acheulean (Asfaw et al. 1992)

169 FIGURE 4.13. The approximate numbers of hominin fossils by member within the principal formations of the omo Group, lake Turkana basin (modiied after feibel et al. [1989], 615). The fossils are mainly fragmentary jaws, isolated teeth, or fragmentary limb bones, but some (especially in the Koobi fora and nachukui formations) are more complete, including a partial skeleton (from the nachukui formation). bars for members within the Koobi fora and nachukui formations are scaled to correspond to like-aged members within the shungura and usno formations. it is thus obvious that the majority of fossils from the lower omo are older than the majority from Koobi fora.

The Lower Omo River Basin

he localities collectively referred to as Lower Omo occur in the southern valley of the Omo River, north of where it enters Lake Turkana in southwestern Ethiopia (ig. 4.11). he Lower Omo preserves more than 1,000 m of Plio-Pleistocene sediments, deposited in a subsiding basin mainly by the meandering proto–Omo River. he sequence also records four or ive episodes of mainly lacustrine sedimentation under a greatly expanded ancient Lake Turkana. As at Koobi Fora, numerous interbedded tufs provide material for radiometric dating and also serve as markers for stratigraphic correlations, not only within the Lower Omo region but also between the Lower Omo and other localities, especially Koobi Fora and West Turkana (igs. 4.12, 4.14). As a result of extensive paleontological collecting, preceded by meticulous and thorough geologic mapping and dating, the Lower Omo has provided the standard PlioPleistocene biostratigraphic sequence with which other east African records are routinely compared.

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170

K

L. Nariokotome Chari L. Koobi Fora Natoo

J H

Members NARIOKOTOME

NATOO

500

G

KALOCHORO G F E4 E D C9

200

C4 C

F E

B A

Kalochoro Kokiselei Lokalalei Emekwi

D C

B

100

0

L. Koobi Chari Fora Okote Black Pumice Malbe KBS Lorenyang

“Orange” Malbe KBS

400

300

Kale Silbo

OKOTE

1.0

?

1.5

KBS 2.0 BURGI 2.5

LOKALALEI

LOMEKWI Waru Tulu Bor Lokochot Moiti Topernawi

Members CHARI

KAITIO

Ma 0.5

Ma 0.5

Tuffs

U. Burgi Burgi Ingumwai Ninikaa Hasuma Waru

Allia Toroto Tulu Bor Lokochot

TULU BOR

3.0

LOKOCHOT 3.5 MOITI

KATABOI

Moiti LONYUMUN 4.0

Acheulean

Kale

Oldowan

Tuffs

L

Homo habilis Homo ergaster

600

L3 L K J7 J4 J H4 H2 H

Paranthropus boisei

700

Tuffs Members

Koobi Fora Fm.

Paranthropus aethiopicus

m

Nachukui Fm.

Australopithecus afarensis

Shungura Fm.

Australopithecus anamensis

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1.0

1.5

2.0

2.5

3.0

3.5

4.0

LONYUMUN A BASAL

FIGURE 4.14. Correlation of the Plio-Pleistocene successions in the lower omo river valley (shungura formation), West Turkana region (nachukui formation), and Koobi fora region (Koobi fora formation) (modiied after brown [1994]; brown et al. [2006]; feibel et al. [1989]). The formations and their constituent members are scaled according to thickness, not according to age. isotopic dates on tuffs determine the ages indicated on the timescale to the right of the Koobi fora formation. Tuffs can be identiied in more than one region, based on their distinctive geochemical signatures. Continuing research has progressively reined both the ages and the correlations.

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Radiometric dates and complementary paleomagnetic readings show that the Plio-Pleistocene deposits of the Lower Omo span the period between roughly 4.1 and 0.8 Ma. he sequence includes four principal, spatially discrete geologic formations: the Mursi Formation, dated to about 4.1 Ma; the Nkalabong Formation dated to about 3.95 Ma; the Usno Formation bracketed between 4.1 and 2.97 Ma; and the Shungura Formation (the classic “Omo Beds”), with exposures spanning the period from approximately 3.6 Ma to perhaps 1 Ma. hese formations are now grouped with the Koobi Fora Formation and other correlative deposits in the Lake Turkana Basin into the Omo Group. Paleontologically, the Shungura Formation is by far the most productive, and it is the principal source of information on dated Plio-Pleistocene faunal change in eastern Africa. It comprises about 760 m of luvial, lacustrine, and deltaic sediments that have been divided among twelve members, labeled (from bottom to top) the Basal Member and Members A–L (excluding I, which was skipped). Except for the Basal Member, each member includes a tuf at its base with the same letter designation. At most localities, Shungura Formation sediments have been deformed by tilting and subsequent faulting, so that their bedding planes intersect the surface at a distinct angle. hey are overlain by much younger (late mid-Pleistocene to mid-Holocene) horizontal sediments assigned to the Kibish (or “Omo-Kibish”) Formation.

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TABLE 4.3. Hominin taxa (and artifact traditions) represented in the Koobi Fora Formation, northern Kenya, the

geologic units from which they originate, and their geologic ages, estimated mainly from K/Ar and 40Ar/39Ar determinations. Figure 4.14 presents the same information graphically for the entire Turkana Basin. Hominin Taxon (and Artifact Traditions, if any)

Geologic Unit

Estimated Age

Australopithecus anamensis

Lonyumun Mb, 5 m below the Moiti Tuf

3.9 Ma

?Australopithecus afarensis

Tulu Bor Mb

3.3–3.2 Ma

Paranthropus boisei (Oldowan)

Mainly in the KBS and Okote Mbs

from shortly before 2 Ma to ca. 1.4 Ma

Homo habilis (including H. rudolfensis) (Oldowan)

Upper Burgi Mb, just below the KBS Tuf and between the Lower Ileret and Chari Tufs

2 to ca 1.44 Ma

Homo ergaster (= African H. erectus) KBS Mb, just below and above the (Developed Oldowan and/or Okote Tuf Acheulean)

from somewhat before 1.7 Ma to perhaps 1.3 Ma

Homo sapiens (possibly Middle Stone Guomde Locality, Ileret; Upper Age or Later Stone Age) Chari Mb of the Koobi Fora Fm or possibly much younger Galana Boi Fm

sometime between 300 and 10 ka

Most early hominin fossils from the Lower Omo come from the Shungura Formation (236 specimens), but a small number have also been recovered in the Usno Formation (twenty-three specimens) (ig. 4.13). In contrast to Koobi Fora, where the hominin fossils are mostly younger than 2 Ma, the Lower Omo specimens are mainly from between 3 and 2 Ma. he reason for the limited overlap is not clear, but the records are thus more complementary than supplementary. he Lower Omo fossils also tend to be much more fragmentary because they accumulated mainly in higher-energy (luvial) environments. he vast majority (225) are only isolated teeth, and this hinders both sorting and diagnosis. However, at least ive taxa are recognized at present, from Australopithecus afarensis through Homo ergaster (table 4.4). he oldest well-dated artifacts in the Lower Omo succession come from Shungura Members F and G and probably date to 2.4–2.3 Ma. Potentially older artifacts, dating to 2.5–2.4 Ma, may occur in Member E or even Member C, but this has yet to be conirmed by excavation. In addition, the specimens attributed to Member E lack unequivocal attributes of human laking, and they may all be geofacts (stones fragmented by stream action or weathering). he unquestioned artifacts from Members F and G it within the Oldowan Industrial Complex, as originally deined at Olduvai Gorge, and they imply that the artifact makers had mastered the principles of stone knapping to the same remarkable degree as the people who made like-aged or somewhat older artifacts at Gona, Hadar, and West Turkana.

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Hominin taxa and their stratigraphic origin in the Lower Omo River Basin. Isolated teeth dominate the hominin sample from the Shungura Formation (Suwa et al. 1996). Figure 4.14 presents the same information graphically for the entire Turkana Basin. TABLE 4.4.

Taxon

Unit

Age

Australopithecus afarensis

Usno Formation and Shungura Formation Member B

ca. 3 Ma

Paranthropus aethiopicus

Shungura Formation Member B and perhaps Members D - G

2.6 - 2.3 Ma

Paranthropus boisei

Shungura Formation Members G - L

2.3 - ?1.2 Ma

Homo sp. (? H. rudolfensis)

Shungura Formation Members E - H

2.3 - ?1.3 Ma

Homo ergaster (= African H. erectus)

Shungura Formation Member K

ca. 1.4 Ma

Homo sapiens

Kibish Formation Member 1

between 196 and 104 ka

In addition to fossils of australopiths and early Homo, the Lower Omo has provided fossils of near-modern or modern Homo sapiens. hese come from Member I of the Kibish Formation, which comprises up to 100 m of horizontal, tectonically undisturbed, mainly deltaic deposits that accumulated disconformably on the Nkalabong and Mursi Formations of the Omo Group. Ar40/Ar39 dating of underlying and overlying volcanic tufs has bracketed the human fossils between 196 and 104 ka, and probable correspondences between depositional pulses within the Kibish Formation and dated organic-rich mud deposits (sapropels) on the Mediterranean sealoor suggests the fossils probably date to 196 ka. hey may include the oldest known representatives of H. sapiens narrowly understood, and they are discussed further in chapter 6. SOURCES: overivew of Omo sediments (Chavaillon 1982; Howell 1978b); Omo Tufs (Brown et al. 1985b; Feibel et al. 1989; Harris et al. 1988a); Omo Group (de Heinzelin 1983); faunal change within the Shungura Formation (Bobe and Behrensmeyer 2004; Bobe et al. 2002); Kibish Formation (Butzer 1976b); predominance of isolated teeth in the Omo hominin sample and taxonomic attribution (Suwa et al. 1996); artifacts from Members F and G (Chavaillon 1976; de la Torre 2004; Howell et al. 1987; Merrick and Merrick 1976); possibility that Member E artifacts are geofacts (de la Torre 2004); dating of hominin fossils in the Kibish Formation (McDougall et al. 2005)

West Turkana

West Turkana is the name informally applied to a ten-kilometer strip of land on the northwest shore of Lake Turkana, northern Kenya (ig. 4.11), which contains up to 715 m of Plio-Pleistocene luvial and lacustrine

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deposits. hese have been assigned to eight successive members of the Nachukui Formation. he members are delineated mainly by volcanic tufs that also occur in the Koobi Fora Formation east of Lake Turkana and in the Shungura Formation in the Lower Omo Valley north of the lake. In total, the Nachukui, Koobi Fora, and Shungura Formations share twenty-three tufs that allow precise correlations (ig. 4.14). Tufs directly dated at West Turkana and correlations to tufs dated elsewhere show that the Nachukui Formation accumulated between about 4.3 and 0.7 Ma. he mammalian fauna from the Nachukui Formation (more than a thousand mammalian fossils from at least ninety-four species) corroborates and supplements the basic biostratigraphy previously established in the Lower Omo River Valley and at Koobi Fora. he Nachukui sequence is particularly notable for fossil assemblages that accumulated between roughly 3 and 2 Ma, a period that elsewhere in Africa is well-controlled only in the Lower Omo. Table 4.5 provides a listing of the Nachukui Fm hominin fossils by taxon. he sample is smaller than those from the Lower Omo or Koobi Fora (ig. 4.14), but it includes some remarkably complete specimens:

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

• a distorted but otherwise well-preserved skull (KNM-WT [Kenya National Museum–West Turkana] 40000), dated to 3.5 Ma, that is the type specimen of Kenyanthropus platyops. If the skull has been correctly diagnosed, it shows Australopithecus afarensis was not the only hominin species between 4 and 3 Ma. Fossils tentatively assigned to Au. afarensis accompany other fossils assigned to K. platyops in deposits dated to 3.3 Ma. • the famous Black Skull (KNM-WT 17000) (stained blue-black by manganiferous minerals), dated to 2.5–2.4 Ma, that is the principal specimen of Paranthropus aethiopicus. • the skull and partial skeleton of the “Nariokotome (or Turkana) boy” (KNM-WT 15000), dated to 1.6 Ma, that has uniquely illuminated the biology of Homo ergaster. An isolated lower irst molar (KNM-WT 42718), dated to roughly 2.3 Ma and assigned to early Homo, supplements the sparse sample of fossils (table 4.11) that document the emergence of Homo 2.5–2.4 Ma. A cranial fragment from the Kalachoro Mb that was once once thought to represent Homo habilis is now thought to be recent and has been reassigned to H. sapiens. Oldowan artifacts have been reported from multiple sites. he oldest (in the Kalochoro Member) occur at Lokalalei 1, where they date from about 2.35 Ma and at Lokalalei 2C, where they are probably about 100 ky younger. he youngest (in the lower Kaitio Mb) come from Kokiselei 1, 2, 3, and 5 and Naiyena Engol 1 and 2, where they are dated to 1.8–1.7 Ma. An extension at Lokalalei 1 provided the molar of early Homo sp.

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TABLE 4.5. Hominin taxa (and artifact traditions) represented in the Nachukui Formation, northern Kenya, the geologic units from which they originate, and their geologic ages, estimated mainly from K/Ar and 40Ar/39Ar determinations. Figure 4.14 presents the same information graphically for the entire Turkana Basin.

Hominin Taxon (and Artifact Traditions, if any)

Geologic Unit

Estimated Age

Kenyanthropus platyops

Kataboi Mb (skull KNM-WT 40000)

3.5 Ma

Kenyanthropus platyops

lower Lomekwi Mb (a partial let maxilla and other fragmentary specimens)

3.3 Ma

cf. Australopithecus afarensis

lower Lomekwi Mb (two fragmentary mandibles and some isolated teeth)

3.3 Ma

Paranthropus aethiopicus

base of the Lokalalei Mb (the “Black Skull” and a mandible)

2.5–2.4 Ma

early Homo (Oldowan)

base of the Kalochoro Mb (an isolated lower right M1)

2.3 Ma

Paranthropus boisei (Oldowan)

Kalochoro and Kaitio Mbs (fragmentary fossils)

between 2.3 and 1.6 Ma

Homo ergaster (= African H. erectus) (Acheulean)

Natoo Mb (principally the “Nariokotome Boy”)

1.6 Ma

referred to above, and Kokiselei 1 has provided eight teeth or tooth fragments attributed to Paranthropus boisei. he Lokalalei Oldowan occurrences are equaled or exceeded in age only by occurrences in Shungura Formation Member F (Omo Valley), southern Ethiopia and at Hadar and Gona, north-central Ethiopia. Acheulean bifaces and associated artifacts occur at Kokiselei 4 (upper Kaitio Member), where they are dated to roughly 1.65 Ma. Together with Acheulean sites of similar antiquity in the Gona region (upper Busidima Formation), Ethiopia, Kokiselei 4 suggests that H. ergaster (or early African H. erectus) and the Acheulean emerged at roughly the same time, 1.8–1.7 Ma. SOURCES: tuf correlations among Turkana Basin Formations (Brown et al. 1985b; Feibel et al. 1989; Harris et al. 1988a); Nachukui Fm fauna (Brugal et al. 2003; Harris et al. 1988a, 1988b; Leakey et al. 2001); K. platyops (Leakey et al. 2001); the Black Skull (Leakey and Walker 1988; Walker et al. 1986a); Turkana Boy (Brown et al. 1985a; Walker 1993); KNM-WT 42718 early Homo (Prat et al. 2005); Kalachoro Fm cranial fragment (Harris et al. 1994); W. Turkana Oldowan artifacts (Kibunjia 1994; Kibunjia et al. 1992; Roche et al. 2003); dating of Lokalalei archaeological sites (Brown and Gathogo 2002); Oldowan in Shungura Mb F (Howell et al. 1987) and at Hadar and Gona (Kimbel et al. 1996; Semaw 2000; Semaw et al. 1997, 2003); Kokiselei 4 Acheulean (Roche 1995; Roche et al. 2003; Roche and Kibunjia 1994); Gona early Acheulean (Quade et al. 2004)

Fejej

he Fejej Plain lies roughly halfway between the mouth of the Omo River and Chew Bahir (former Lake Stephanie) in extreme southern Ethiopia. he sequence comprises luviolacustrine deposits and interbedded

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volcanics that span much of the later Cenozoic. Seven isolated hominin teeth have been found at a locality (FJ-4) that is probably between 4.2 and 4 my old, based on 40Ar/39Ar dating and paleomagnetism. he teeth closely resemble those of Australopithecus afarensis but they are perhaps more likely to represent A. anamensis, given their geologic age. At a second locality (FJ-1), Oldowan artifacts occur in sediments that dated to about 1.96 Ma.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Fejej sediments (Asfaw et al. 1991), antiquity of hominin teeth (Kappelman et al. 1996); assignment of teeth to A. afarensis (Fleagle et al. 1991) or A. anamensis (Ward et al. 1999a, 2001); Oldowan artifacts (Asfaw et al. 1991, 1993; de Lumley and Beyene 2004)

Konso (Formerly Konso-Gardula)

he Konso locality is in the Main Ethiopian Rit Valley, roughly 180 km northeast of the Lake Turkana Basin. he fossiliferous sediments formed mainly in an ancient lake and along streams that lowed into it. hey reach a thickness of more than 180 m and they include more than 30 interbedded volcanic tufs. 40Ar/39Ar dating on the tufs and geochemical correspondences between them and dated tufs in the Turkana Basin bracket the Konso Formation between about 1.95 and 1.3 Ma. his makes it broadly coeval with the upper Burgi, KBS, and Okote Members of the Koobi Fora Formation at East Turkana, the Kaitio, Natoo, and Nariokotome Members of the Nachukui Formation at West Turkana, Members H, J, K, and L of the Shungura Formation in the lower Omo Valley, and Olduvai Gorge Beds I and II in northern Tanzania. Fieldwork at Konso has produced nearly 8,000 identiiable mammalian fossils, mainly from horizons dated to 1.9 and 1.5–1.4 Ma. More than sixty-eight species are represented, including two hominins—Homo ergaster (or African H. erectus) (eight specimens) and Paranthropus boisei (nine specimens)—both from deposits dated to 1.5–1.4 Ma. he fossils of P. boisei presently mark the northern limit of the species and they include a nearly complete skull (KGA10–525). Like contemporaneous east African faunas, the Konso fauna implies a shit to grassier and probably drier conditions between 1.8 and 1.6 Ma. Two Konso sites dated to about 1.9 Ma have provided Oldowan artifacts, while sites dated to 1.6–1.5 Ma and later contain abundant Acheulean bifaces that are among the oldest known. SOURCES: Konso—location, fossils, and hominins (Asfaw et al. 1992, 1995; Suwa et al. 1997, 2003, 2007a), dating (Katoh et al. 2000; Nagaoka et al. 2005)

Hadar and Dikika

he Hadar site is in the Awash River Valley within the Afar Depression, about 300 km northeast of Addis Ababa, Ethiopia. he fossiliferous sediments at Hadar have been grouped into the Hadar Formation, which has

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four members (from bottom to top): the Basal Member, the Sidi Hakoma Member, the Denen Dora Member, and the Kada Hadar Member (ig. 4.15). Streams deposited most of the he Hadar Formation sediments in a basin that was also periodically inundated by a large lake. Interbedded volcanic tufs and a basalt layer provide materials for radiometric dating, and the tufs allow regional and interregional correlations. he Dikika area immediately south of Hadar is now distinguished from Hadar for collection purposes, but presents the same sedimentary sequence and is considered with Hadar here. he abbreviation AL (Afar Locality), followed by a site number and a serial number, denotes Hadar fossils in the narrow sense. Dik replaces AL for newly discovered fossils from Dikika. Altogether, the four members of the Hadar Formation have furnished more than 360 specimens of the australopith species Australopithecus afarensis, also represented at the nearby Middle Awash locality, at Laetoli in northern Tanzania, and more equivocally at other sites in Ethiopia and Kenya. High-precision single-crystal laser fusion 40Ar/39Ar dates now bracket the Hadar A. afarensis fossils between slightly more than 3.4 Ma and 2.9 Ma (ig. 4.15). Most derive from the Denen Dora Member, in which a single site (AL 333) has produced about 250 specimens from at least nine adults and four juveniles (the “irst family”). Other particularly notable inds include a 40%-complete skeleton (“Lucy,” or AL 288-1), a nearly complete skull (AL 444-2), a fragmentary mandible (Dik-2-1), which is the oldest known fossil of A. afarensis at Hadar, and the skull and partial skeleton of a child (Dik-1-1) who died at about age three, assuming an African ape rate of dental development. he adult skeleton comes from a diminutive individual, presumed to be female and the nearly complete skull from a much larger one, presumed to be male. Comparable size variation marks the rest of the Hadar sample, leading some specialists to suggest that it comprises two (or more) species or that A. afarensis exhibited a high level of sexual dimorphism, comparable to the level in gorillas and orangutans. his position is accepted here, but as discussed below, the actual level of dimorphism is controversial. Sediments formerly assigned to the Kada Hadar Member of the Hadar Formation but now to the overlying Busidima Formation have provided a maxilla of Homo irmly dated to about 2.3 Ma by 40Ar/39Ar dating and associated fauna. Only four other sites, listed in table 4.11, have provided equally old or older fossils of Homo. he Hadar maxilla has been attributed to H. habilis, while the remaining fossils have been assigned to H. rudolfensis. he oldest Hadar artifacts date to about 2.3 Ma, and they include an assemblage associated with the maxilla of Homo. Together with artifacts dated to 2.6–2.5 Ma at nearby Gona, they suggest that stone artifact

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BKT-3 2.33 + 0.07 my ago ✽ AL 666

p p p 160 p p T

140

Homo sp. and artifacts

AST-1

Disconformity BKT-2 2.92 + 0.03 my ago GMT

p

Kada Hadar

✽ AL 444 ✽ AL 438

p p Tp

120 BKT-1

Hadar Formation

100

Denen Dora

p

80

60

Confetti clay ✽ AL 288 (“Lucy”) KHT 3.18 + 0.01 my ago DD-3 Sand

✽ AL 333 (“First Family”)

TT-4 3.22 + 0.01 my ago

T 40

Sidi Hakoma

Pink marl KMB 3.28 + 0.04 my ago SH-3 Sand

p T p

T

Basal

KMT 20

0

SH-2 Sand

✽ AL 417

Australopithecus afarensis

Busidima Formation

meters

177 FIGURE 4.15. schematic stratigraphy of the hadar formation (redrawn after Kimbel et al. [1996]). The constituent members are listed on the extreme left. T’s in the adjacent column mark deposits that accumulated in a large (“transgressive”) lake and p’s mark the positions of paleosols, which formed when deposition was relatively quiescent. 40ar/39ar dating provided all the ages to the right of the stratigraphic column. asterisks indicate the stratigraphic horizons of particularly important fossil sites (afar or dikika localities, abbreviated respectively by AL or Dik and an appropriate serial number). other abbreviations refer to key geologic marker horizons.

✽ DIK-1-1 (juvenile) ✽ AL 137 SHT 3.40 + 0.03 my ago

-10 -20

✽ DIK-2-1

manufacture and Homo emerged simultaneously. Both the Hadar and the Gona artifacts closely resemble those from sites dated between about 2.35 and 1.7 Ma in the Lake Turkana Basin and at Olduvai Gorge, and they have been assigned to the same Oldowan Industrial Complex. he next youngest artifacts reported from the Hadar region are Acheulean bifaces and associated pieces from luvial deposits that unconformably overlie the Hadar Formation and that are clearly much younger (early or middle Quaternary). Fossil pollen shows that Hadar was generally much cooler and moister when A. afarensis was present, between roughly 3.4 and 2.9 Ma. Pollen and animal fossils indicate a vegetation mosaic in which forest, wet or dry grassland, and woodland dominated at diferent times. he vegetational luctuations do not seem to have afected A. afarensis, which

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persisted through them. By comparison to much earlier times, Hadar appears to have been signiicantly grassier when the maxilla of Homo and associated Oldowan artifacts accumulated about 2.3 Ma.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Hadar Formation sediments—origin (Chavaillon 1982; Johanson et al. 1982b; Kimbel et al. 1996) and 40Ar/39Ar dating (Kimbel et al. 1994; Wynn et al. 2006); the “irst family” (Johanson et al. 1982b; White and Johanson 1989); “Lucy” (Johanson et al. 1982a); AL 444-2 (Kimbel et al. 2004); Dik-2-1 (Alemseged et al. 2005); Dik-1-1 (Alemseged et al. 2006); Busidima Homo maxilla (Kimbel et al. 1996, 1997); early Oldowan artifacts from Hadar and Gona (Hovers 2003); Hadar Acheulean (Corvinus 1975, 1976) and pollen (Bonneille et al. 2004); open vegetation associated with Homo and oldest Hadar artifacts (Kimbel et al. 1996)

Gona (Kada Gona)

he Gona locality lanks the northern Awash River, immediately southwest of Hadar. Hadar and Gona have been investigated mainly by diferent teams, but they contain broadly similar geologic sequences, including the Hadar Formation as just described. Gona also preserves two formations that have not been recorded at Hadar. hese are the Sagantole Formation, which formed between roughly 5.6 and 3.9 Ma and precedes the Hadar Formation, and the Busidima Formation, which accumulated between about 2.7 and 1000 cc) and laterally expanded sidewalls (Senut et al. 2000).

Sites (and Dates of Discovery)

Proposed Age (and Basis for Estimate)

Fossils

References

Morocco Jebel Irhoud (Ighoud) (1961, 1963, 1968, 1969)

A nearly complete skull (Irhoud 1), a skullcap (Irhoud 2), a juvenile mandible (Irhoud 3), a juvenile humerus shat (Irhoud 4), and a partial adult mandible

between 190 and 90 ka (ESR)

(Amani & Geraads 1993; Ennouchi 1962; Ennouchi 1963; Ennouchi 1968; Ennouchi 1969; Grün & Stringer 1991; Hublin 1993; Hublin 2000; Hublin et al. 1987)

Kébibat (Mifsud-Giudice Quarry, Rabat) (1933)

Mandible, let maxilla, and occipital fragments of an adolescent

between 200 and 130 ka (associated mammals and U-series on an overlying marine shell)

(Debénath 2000; Hublin 1985; Saban 1977; Stearns & hurber 1965)

Sidi Abderrahman (Littorina Cave) (1954)

Two fragments of a mandible

?400 ka (faunal and artifactual associations; and OSL)

(Arambourg & Biberson 1956; Biberson 1964; Debénath 2000; Geraads 1980; Geraads et al. 1980; Howell 1960; Jaeger 1975; Raynal et al. 1995b; Rhodes et al. 2006)

homas Quarries I and III Caves (1969, 1972, 1996) (homas III = Oulad Hamida 1)

he let side of a mandible, cranial fragments, and 3 isolated teeth

between 600 and 400 ka (Bräuer 1984b; Debénath (faunal associations, ESR, 2000; Geraads 1980; Gerand OSL) aads et al. 1980; Geraads et al. 2004; Howell 1978a; Hublin 1985; Raynal et al. 1995b; Raynal et al. 2001; Rhodes et al. 1994; Rhodes et al. 2006; Rightmire 1980)

Salé (1971)

Fragmentary skull

?400 ka (probable faunal associations and ESR)

(Debénath 2000; Hublin 1985; Jaeger 1975)

Djibouti Wadi Dagadlé (1983)

Let maxilla

Between ?400 and ?250 ka (faunal associations and TL)

(Bonis et al. 1984; Bonis et al. 1988)

Sudan Singa (1924)

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292

170–150 ka (ESR and Useries, associated fauna)

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TABLE 5.4. (continued)

Sites (and Dates of Discovery)

Proposed Age (and Basis for Estimate)

Fossils

References

Ethiopia Bodo, Middle Awash (1976, 1981, 1990)

Partial skull with face, parietal of a second skull, and a distal humerus shat

Between 640 and 550 ka (40Ar/39Ar)

(Asfaw 1983; Clark et al. 1984a; Clark et al. 1994; Conroy 1997; Conroy et al. 1978; Conroy et al. 2000b; Kalb et al. 1982a; Renne 2000; Rightmire 1996)

Garba III (Melka Kunturé) (1977)

Cranial fragments

?400–?250 ka (faunal and artifact associations)

(Chavaillon 1979; Chavaillon 1982; Chavaillon et al. 1974)

Kenya Kapthurin (Baringo) sites GnJh-01 and GnJh-19 (1966, 1983)

Two mandibles (KNM-BK- Between 543 and 509 ka 67 and 8518), a phalanx (40Ar/39Ar) (KNM-BK-64), a right metatarsal (KNM-BK63), and a right ulna (KNM-BK-65)

(Cornelissen et al. 1990; Day 1986b; Deino & McBrearty 2002; Leakey et al. 1969a; McBrearty 1999; McBrearty et al. 1996; McBrearty & Jablonski 2005; Solan & Day 1992; van Noten 1983; Wood & van Noten 1986)

Lainyamok (1976, 1984)

hree associated maxillary teeth and a femur shat fragment

between 392 and 330 ka (40Ar/39Ar)

(Potts & Deino 1995; Potts et al. 1988; Shipman et al. 1983)

Lake Ndutu (1973)

Skull

?500–?300 ka (artifactual associations)

(Clarke 1976; Clarke 1990; Leakey & Hay 1982; Mturi 1976; Rightmire 1983)

Eyasi (1935, 1938, 1993)

A fragmentary skull (Eyasi I), a partial occipital (Eyasi II), a second, separate occipital fragment (Eyasi IV), and small cranial fragments and associated teeth (Eyasi III)

?400–?200 ka (faunal and artifactual associations)

(Bräuer & Mabullah 1996; Mehlman 1984; Mehlman 1987; Trinkaus 2004)

A nearly complete skull, a cranial fragment, a right maxilla, a fragmentary humerus, two pelvises, six femur fragments, and a tibia fragment

?600–?400 ka (probable faunal associations)

(Clark 1959a; Clark et al. 1968; Clark et al. 1950; Kennedy 1984b; Klein 1973a; Pycrat et al. 1928; Singer 1958; Stringer 1986a; Woodward 1921)

Tanzania

Zambia Kabwe (= Broken Hill) (1921–1925)

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TABLE 5.4. (continued)

Site (and Dates of Discovery)

Geologic Age (and Basis for Estimate)

Fossils

References

Namibia Berg Aukas (Grootfontein, Otavi Mountains) (1965)

Proximal femur

?500–?200 ka (degree of mineralization)

(Grine et al. 1995)

Cave of Hearths, Limpopo Province (1947)

Right mandible fragment with three teeth; possibly also a radius of uncertain stratigraphic origin

?400–200 ka (faunal and artifactual associations)

(Latham & Herries 2004; Mason 1962; Mason 1988b; Pearson & Grine 1997; Tobias 1971)

Florisbad, Free State Province (1932)

Partial skull and an isolated upper third molar

ca. 260 ka (ESR)

(Brink 1987; Clarke 1985; Douglas 2006; Dreyer 1935; Grün et al. 1996; Scott & Rossouw 2005)

South Africa

Elandsfontein (= Hopeield Skullcap and mandible = Saldanha), Western fragment Cape Province (1953, 1954)

?600 ka (faunal and artifac- (Drennan 1953; Klein et tual associations) al. 2007; Klein & CruzUribe 1991; Singer 1958; Singer & Wymer 1968)

TABLE 5.5. European sites with fossils of early Homo neanderthalensis. Italics and boldface designate sites where the fossils might be alternatively assigned to H. heidelbergensis.

Site (and Dates of Discovery)

Geologic Age (and Basis for Estimate)

Fossils

References

United Kingdom Swanscombe (Upper Middle Gravel) (1935, 1936, 1955)

Occipital and right and let parietals of a single skull

ca. 400 ka (geomorphic assessment, associated fossils, and climate correlation [OIS 11])

(Barton & Clark 1993; Conway et al. 1996; Marston 1937; Oakley 1952; Oakley 1957; Roberts et al. 1995; Stringer 1996b; Stringer & Hublin 1999; Stringer et al. 1984; Wymer 1955; Wymer 1964)

Boxgrove, West Sussex (1993, 1995)

Tibia shat, two isolated incisors

ca. 500 ka (mammalian fauna and climate correlation [OIS 13])

(Roberts 1986; Roberts et al. 1994a; Streeter et al. 2001; Stringer 1996b; Stringer 2006, pp. 63–67; Stringer et al. 1998; Trinkaus et al. 1999b)

Pontnewydd, Wales (1978–1985)

Seven isolated teeth and a right maxillary fragment with two teeth

ca. 200 ka (U-series, TL, and climate correlation [OIS 7])

(Green 1981; Green et al. 1989; Green et al. 1981)

A nearly complete mandible, and some isolated teeth

?150–130 ka (climate correlation [OIS 6])

(Billy 1982; Billy & Vallois 1977; Stringer et al. 1984)

France Montmaurin, Haute Garonne (1949)

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TABLE 5.5. (continued)

Site (and Dates of Discovery)

Geologic Age (and Basis for Estimate)

Fossils

References

La Caune de l’Arago (Tautavel), Eastern Pyrenees (1964–2001)

More than 100 specimens, ca. 425–400 ka (ESR, including a facial U-series, and climate skeleton (Arago 21) and correlation [OIS 11]) a parietal (Arago 47) possibly from the same individual, a mandible (Arago 2), a half mandible (Arago 13), a left pelvis (Arago 44), and 60 isolated teeth

Day 1982; Day 1984; de Lumley 1979; de Lumley et al. 1984; Falguères et al. (2004; Sigmon 1982; Stringer et al. 1984)

Orgnac 3, Ardeche (1962, 1968–71)

Seven, mostly deciduous isolated teeth

ca. 310–300 ka (ESR, U-series, and climate correlation [OIS 9])

(Combier 1971; Cook et al. 1982; Moncel et al. 2005)

Le Lazaret Cave, Nice (1953–64)

A juvenile right parietal, a deciduous upper incisor, an adult lower canine, and seven additional fragments

170–130 ka (ESR, Useries, and climate correlation [OIS 6])

(de Lumley 1969b; de Lumley 1975; de Lumley et al. 2004; Falguères et al. 1992; Stringer et al. 1984; Valensi 2000)

La Chaise-de-Vouthon (Bourgeois-Delaunay and Suard Caves), Charente (1960’s and 1970’s)

Approximately eighty 185–100 ka (TL and Uspecimens, mainly series [OIS 6]) isolated teeth and cranial fragments, and a few fragmentary postcranial bones; at least 3 children are represented by cranial fragments

(Blackwell et al. 1983; Debénath 1976; Debénath 1977; Debénath 1988; Matilla 2005; Piveteau & Condemi 1988; Stringer et al. 1984; Teilhol 2003)

Fontéchevade Cave, Charente (1947, 1957)

A frontal fragment (Fontéchevade I), a skullcap (II), and a parietal

?150 ka (associated fauna and climate correlation [OIS 6])

(Henri-Martin 1965; Stringer et al. 1984; Vandermeersch et al. 1976)

Biache-Saint-Vaast, Pas-de-Calais (1976)

he rear half of a cranium, a fragmentary maxilla with all six molars, and ive isolated teeth

175 ka (TL and associated fauna [OIS 6])

(Aitken et al. 1986; Stringer et al. 1984; Tufreau 1989; Tufreau et al. 1978; Tufreau et al. 1982)

Bau de l’Aubesier, Vaucluse (1994–2000)

A partial mandible, an isolated I2 and an isolated M1 or M2

ca. 200 ka (associated fauna and TL [OIS 7])

(Lebel 2002; Lebel et al. 2001)

A let mandibular canine, and a let maxillary M3

>= 240 ka (associated fauna and a composite ESR/U-series reading [?OIS 9])

(Trinkaus et al. 2003a)

hree nearly complete craniums, six partial craniums, and more than 5500 other cranial and

between 600 and 530 ka (U-series, associated fauna, and climate correlation [OIS 15])

Portugal Galeria Pesada, Almonda Karstic System, Torres Novas (2001–2002) Spain Atapuerca SH (= Sima de los Huesos [“Pit of the Bones”] = Cueva Mayor/ Ibeas) (1976 to present)

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TABLE 5.5. (continued)

Site (and Dates of Discovery)

Geologic Age (and Basis for Estimate)

Fossils

postcranial bones representing at least twentyeight individuals

References

1994; Arsuaga et al. 2003; Arsuaga et al. 1991; Arsuaga et al. 1996; Arsuaga et al. 1999a; Arsuaga et al. 1993; Arsuaga et al. 1997a; Arsuaga et al. 1997b; Bermúdez de Castro 1988; Bermúdez de Castro 1993; Bermúdez de Castro 1996; Bermúdez de Castro et al. 1997b; Bermúdez de Castro et al. 2004; Bermúdez de Castro & Nicolás 1995; Bermúdez de Castro & Nicolás 1997; Bermudez de Castro et al. 2001; Bischof et al. 1997a; Bischof et al. 2003; Bischof et al. 2007; Carretero et al. 1997; CuencaBescós et al. 1997; García & Arsuaga 1999; García et al. 1997; Martínez & Arsuaga 1997; Rosas 1995; Rosas 1997; Rosas et al. 1991; Stringer et al. 1996)

Italy Venosa-Notarchirico, Basilicata (1986)

?400 ka (U-series, TL, faunal associations, and climate correlation [OIS 11])

(Belli et al. 1991; Condemi 1991; Mussi 1995; Piperno et al. 1990; Villa 2001)

Fontana Ranuccio, Anagni, Four isolated teeth Latium (1970’s and 1980’s)

ca. 400 ka (radiopotassium, associated fauna, and climate correlation [OIS 11])

(Bidduttu et al. 1979; Bidduttu & Celletti 2001; Condemi 1991; Mussi 1995; Segre & Ascenzi 1982)

Castel di Guido, Latium (1979–1990)

Fragmentary right and let parietals, a right temporal, an occipital fragment, a right maxillary fragment without teeth, and two femoral shats

ca. 315 ka (associated fauna and climate correlation [OIS 9])

(Boschian & Radmilli 1995; Condemi 1991; Mallegni et al. 1983; Mallegni & Radmilli 1988; MarianiCostantini et al. 2001; Mussi 1995)

Rebibbia-Casal de’Pazzi, Latium (1983)

Parietal fragment

?200 ka (associated fauna and climate correlation [OIS 7])

(Anzidei & Cerilli 2001; Bietti 1985; Manzi et al. 1990)

Cava Pompi, Foi, Latium (1961, 1974)

Cranium fragments, a proximal ulna fragment, and a tibia shat

?400 ka (associated fauna and climate correlation [OIS 11])

(Bidduttu & Celletti 2001; Condemi 1991; Cook et al. 1982)

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TABLE 5.5. (continued)

Site (and Dates of Discovery)

Geologic Age (and Basis for Estimate)

Fossils

Visogliano Shelter and Breccia, Trieste (1983, 1985, 1987, 1992, 1993, 1996)

References

A mandible fragment, 5 isolated teeth, and 2 dental fragments

386–321 ka (combined (Abbazzi et al. 2000; BosESR/U-series, associated chian et al. 1995; Cattani fauna, and climate coret al. 1991; Condemi relation [OIS 11]) 1991; Falguères 2003)

Grafenrain Sand Quarry, Mauer, Heidelberg (1907)

A mandible

Ca, 500 ka (associated fauna and climate correlation [OIS 13])

(Bosinski 1995b; Brunnacker 1975; Howell 1960; Koenigswald 1973; Roebroeks & van Kolfschoten 1994; Rosas & Bermúdez de Castro 1998; Schoetensack 1994 (1908); Stringer et al. 1984)

Reilingen, BadenWurttemberg (1978)

A skull with both parietals, a nearly complete right temporal, and the upper 70% of the occipital

?200 ka (associated fauna and climate correlation [?OIS 7])

(Czarnetski 1991; Dean et al. 1998; Dean et al. 1994; Ziegler & Dean 1998)

Steinheim an der Murr, Wurttemberg (1933)

a nearly complete skull

?300 ka (associated fauna and climate correlation [?OIS 9])

(Adam 1985; Bosinski 1995b; Howell 1960; Stringer et al. 1984)

Bilzingsleben (Steinrinne Quarry), huringia (1972–1993)

Approximately twentythree skull fragments from two or three adult individuals; a mandible, one deciduous tooth and six isolated permanent teeth

412–320 ka (U-series, associated fauna, and climate correlation [OIS 11])

(Bridgland et al. 2004; Harmon et al. 1980; Mania 1986; Mania 1998; Mania et al. 1994; Mania & Vlček 1981; Schwarcz et al. 1988a; Stringer 1996b; Stringer et al. 1984; Vlček et al. 2000)

Weimar-Ehringsdorf (1908–1925)

An adult skullcap, adult mandible, a juvenile mandible, a partial child’s skeleton, and other fragmentary cranial and postcranial bones from perhaps nine individuals

ca. 225 ka (U-series, ESR, associated fossils, and climate correlation [OIS 7])

(Blackwell & Schwarcz 1986; Bridgland et al. 2004; Cook et al. 1982; Mallick & Frank 2002; Smith 1984)

An adult occipital bone, two isolated deciduous teeth, and two deciduous dental fragments

?400 ka (U series, associated fauna, and climate correlation [?OIS11])

(Dobosi 1988; Kretzoi & Dobosi 1990; Schwarcz & Latham 1984; homa 1972; homa 1978; Valoch 1995b; Wolpof 1977)

Germany

Hungary Vértesszöllös (1964–1965)

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TABLE 5.5. (continued)

Site (and Dates of Discovery)

Geologic Age (and Basis for Estimate)

Fossils

References

Greece Skull

Petralona Cave, Halkidiki (1960)

?400–?200 ka (ESR, U-series, possible mammalian associations)

(Bonis & Melentis 1991; Cook et al. 1982; Grün 1996; Hennig et al. 1982; Kurtén 1983; Latham & Schwartz 1992; Papamarinopoulos et al. 1987; Poulianos 1989; Stringer 1983; Stringer et al. 1979; Wintle & Jacobs 1982; Wolpof 1980a)

“primitive H. sapiens” from Florisbad, South Africa, discussed below. In short, the names attached to the three lineages in igure 5.1 may violate accepted taxonomic practice, but for the moment, there is no obvious, less contentious alternative nomenclature. Tables 5.3–5.5 list the dates of discovery and other summary information for fossils of Homo ergaster (early African H. erectus), later African fossils assigned to H. sapiens, H. heidelbergensis, or both, and European fossils assigned to H. neanderthalensis or H. heidelbergensis. Boldface in tables 5.4 and 5.5 distinguishes putative specimens of H. heidelbergensis. SOURCES (not including those in table 5.1): Eugène Dubois—the discovery of H. erectus (Dubois [1892] 1994; Howells 1990; Oakley 1964; Shipman 2001; Shipman and Storm 2002; heunissen 1989; heunissen et al. 1990) and of Wajak (Dubois 1922); the Mojokerto skull—indspot (Hufman et al. 2006), age at death estimated to 0.5–1.5 years (Coqueugniot et al. 2004) or 4–6 years (Antón 1997); discovery of Beijing Man (Boaz et al. 2004; Howells 1991; Jia and Huang 1990; Shapiro 1971, 1974); 1939 comparison of Javan and Chinese fossils (von Koenigswald and Weidenreich 1939); loss of the original Beijing Man fossils (Janus 1975); sinking of Sinanthropus into Pithecanthropus (Le Gros Clark 1955) and of Pithecanthropus into Homo (Le Gros Clark 1964); early acceptance of H. erectus (Howell 1960; Howells 1966); Longgudong and Juyuandong fossils (Zhang 1985b); overlap of Gigantopithecus and H. erectus in China and Vietnam (Ciochon et al. 1996; Louys et al. 2007); H. helmei (Dreyer 1935)

Geologic Antiquity Most fossils of primitive Homo are imprecisely dated, either because their stratigraphic position and associations were not carefully recorded, they were not accompanied by materials that are amenable to radiometric methods, or both. he diiculty of placing many key fossils in time, combined with the relatively small size of the fossil samples, explains why reasonable authorities can disagree on the course of human evolution between 1.8 Ma and 130 ka. he discussion here addresses geologic age by species, which means in efect by region, starting with Homo erectus

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in the Far East. Figure 5.5 summarizes the estimated ages of the principal fossils, regardless of species. Earliest Homo erectus in China

he Chinese Homo erectus fossils are ostensibly the best dated, thanks primarily to more carefully documented stratigraphic provenance, to paleomagnetic analyses, and to occasional radiometric dates. he radiometric dates are based mainly on uranium-series analyses of bones and teeth and on electron spin resonance (ESR) analyses of teeth. Some dates have also been obtained from uranium-series analysis of cave lowstones. he results based on bones and teeth are questionable because they rely on unveriiable assumptions about the rate at which the bones or teeth gained and perhaps lost uranium in the ground. he dates on lowstone require fewer questionable assumptions, and they are to be preferred where they contradict dates on bones or teeth. Some deeply stratiied Chinese sites, above all Zhoukoudian Locality 1, record climatic luctuations that can be provisionally correlated with the dated oxygen-isotope stages of the global marine stratigraphy, and the correlations can then supplement paleomagnetic and radiometric readings. he results from paleomagnetism, uranium-series and ESR, and climatic correlation are probably most reliable for the cave sites, especially Zhoukoudian Locality 1, and least reliable for the open-air sites, especially Gongwangling, Chenjiawo, and Yuanmou, where uncertainty about the stratigraphic origin of the H. erectus fossils adds an additional complication. Still, with all the caveats in mind, table 5.2 presents age estimates that are probably broadly correct. he oldest Chinese H. erectus fossil may be the Gongwangling (Lantian) skull, whose age has been variously estimated at 1.15–1 Ma or 800–750 ka. he other principal Chinese sites—Zhoukoudian, Chenjiawo, Hexian, Nanjing, and Yunxian, together with ham Khuyen Cave in Vietnam—all appear to postdate 800 ka. hus, based on H. erectus fossils alone, it could be argued that people arrived in China only ater 1 Ma. However, two Chinese sites or site complexes that lack diagnostic H. erectus fossils—Longgupo Cave and the Nihewan Basin—suggest a much earlier arrival, as long ago as 1.9 Ma. Each site thus requires more detailed consideration, for if the dates from either are valid, they could mean that H. erectus evolved in China from a yet more primitive hominin species and then spread back to Africa. Longgupo Cave. At Longgupo Cave, paleomagnetic analysis places an

isolated human upper lateral incisor, a mandible fragment with the fourth premolar and irst molar, and two possible stone artifacts within the Olduvai Normal Subchron, between 1.96 and 1.77 Ma. he result is intriguing but problematic, for two reasons. First, the cave ill is highly complex and ill-suited for paleomagnetic analysis, and the actual age of

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EUROPE

AFRICA

Ma

EASTERN ASIA

Ma

0.1 0.2

0.1 0.2

Singa, Eliye Springs, Guomde

Dagadlé

0.3 0.4 0.5 0.6 0.7

Sidi Abderrahman

Salé, Garba III, Lainyamok, Ndutu, Kapthurin, Eyasi, Broken Hill, Cave of Hearths

Swanscombe, Steinheim, Reilingen, Arago & Bizlingsleben

0.3 0.4 Tham Khuyen Cave

0.5

Thomas Quarries Bodo Elandsfontein

H. sapiens/ H. heidelbergensis

H. neanderthalensis/ H. heidelbergensis

0.6 Yunxian

H. ergaster

Tighenif

Chenjiawo

?

0.8

0.7 0.8

Ceprano

0.9 1.0

Daka

0.9 Olorgesailie

Buia

Gomboré II

1.1 1.2

Olduvai Hominid 9

? Swartkrans

Nariokotome III

?

1.4 1.5

Garba IV

H. erectus

Koobi Fora

1.7 1.8

1.2 1.3

Konso

1.4

1.6

1.1

Homo sp.

1.3

1.5

1.0

1.6 1.7

H. ergaster

1.8

H. habilis

FIGURE 5.5. approximate ages of the main sites providing fossils of Homo ergaster, H. erectus, early H. neanderthalensis, and early H. sapiens. The shaded vertical lines indicate time ranges that possibly include the fossils. The datings are based variously on faunal correlations; u-series, esr, or Tl determinations; paleomagnatism; and presumed correspondences between the sequences of glacial-interglacial events recorded at the sites and the global oxygen-isotope stratigraphy. (Atapeurca TD 6 = atapuerca Gran dolina layer 6; Atapuerca TE 9 = atapuerca sima del elefanted layer 9.)

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the human fragments may be closer to an ESR estimate that brackets a deer tooth from a higher-lying unit between 1.02 and 0.75 Ma. Second, the morphology of the mandible fragment suggests it came from a late representative of the small fossil ape, Lufengpithecus, while the incisor is identical to incisors of living east Asians. his raises the possibility that it was an undetected intrusion from a higher part of the deposit. Nihewan Basin. In the Nihewan Basin, paleomagnetic readings and

estimated sedimentation rates at the neighboring sites of Donggutuo, Xiaochangliang, and Majuangou ix successive artifact-bearing surfaces between 1.66 and 1.07 Ma, in the interval following the Olduvai Normal Subchron. he deposits that buried the surfaces formed mainly within ancient lakes, and they are ideal for paleomagnetic analysis. he oldest surface, with an estimated age of 1.66 Ma, occurs at Majuangou and has provided 443 laked stones and 1,014 fragmentary bones, mainly from elephant (Elephas sp.). Animal trampling has afected both the ancient land surface and many of the bones, and it could arguably have produced the crude core forms that are common among the artifacts. However, it could not have created the lakes with striking platforms that are also present, and only human transport could have introduced the stone particles into an otherwise ine-grained sedimentary deposit. Fresh paleomagnetic results from the Nihewan Basin, published after the text of this book was completed, illuminate the importance of assumed sedimentation rates in estimating local paleomagnetic ages, and the new results suggest that the oldest Nihewan artifacts, including those from Majuangou, may date closer to 1.2 Ma than to 1.6 Ma. he diference is important, but even at 1.2 Ma, the Nihewan artifacts would be the oldest found so far in eastern Asia. If an estimate of 1.2 Ma (or more) is valid, archaeologists should eventually encounter equally early artifacts in southern and Southeastern Asia and also along the probable route from Africa, perhaps above all in the deep sequences of windblown silts (loesses) that blanket parts of central Asia. So far, the oldest reported site in central Asia is at Kuldara, Tadjikistan, where paleomagnetism brackets deposits with about forty crudely laked pebbles and stone fragments between the Jaramillo Normal Subchron at 1.07 Ma and the Brunhes/Matuyama boundary at 780 ka. No other central Asian site clearly antedates 600 ka. SOURCES (not including those in table 5.2): paleomagnetic dating of the oldest Nihewan artifacts to ca 1.2 Ma (Li et al. 2008); Kuldara (Ranov et al. 1995); lack of sites >600 ka in central Asia (Vishnyatsky 1999)

Latest Homo erectus in China

he upper age limit for Chinese Homo erectus is partly a matter of deinition. Virtually all specialists agree that Hexian and especially Zhoukoudian

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extend the span to ater 500 ka. Fossils putatively dated between 300 and 100 ka, including especially those from Jinniushan, Dali, and Maba, are commonly regarded as more advanced. As discussed below, if they had been found in Europe or Africa, they might have been assigned to “archaic” H. sapiens or to H. heidelbergensis. Arguably however, they differ from the Zhoukoudian or Hexian representatives of H. erectus little more than the younger Zhoukoudian and Hexian representatives difer from the older Gongwangling form, and they may represent a further development along the same lineage. If this is accepted, then Chinese H. erectus, broadly understood, survived to 100 ka or later, until it was eventually replaced by or perhaps admixed with immigrant H. sapiens from Africa. A late persistence for Chinese H. erectus is tentatively accepted here, pending a denser, more securely dated Chinese fossil record that could demonstrate otherwise.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES (not including those in table 5.2): (Chen et al. 1994; Etler 1996; Wu and Poirier 1995)

Earliest Homo erectus in Java

he Javan specimens of Homo erectus remain the most poorly dated. In the Sangiran-Trinil area of central Java, they are conventionally said to come from two rock-stratigraphic units, each with its own distinctive fauna. he older unit, which provided relatively few H. erectus remains, comprises the Pucangan (or Putjangan) Beds with the Djetis (Jetis) fauna; the younger unit, which has provided most of the H. erectus fossils, comprises the Kabuh Beds with the Trinil fauna. In fact, both the lithostratigraphy and the biostratigraphy are probably far more complex, and the exact stratigraphic provenance of most of the H. erectus fossils is uncertain in any case. Fluorine analysis has been used to narrow the provenance possibilities, but it is still diicult to relate the fossils to paleomagnetic determinations or to radiopotassium and ission-track dates that have been obtained on volcanic tufs in the Pucangan Beds and on tufs and pumices in the Kabuh Beds. Pucangan dates range from 2 Ma to 570 ka, while Kabuh dates vary between 1.6 Ma and 470 ka. he patent inconsistency is further underscored by contradictions between some of the dates and associated paleomagnetic readings or between the dates and the regional sequence of mammalian faunal change. he older Pucangan dates are also inconsistent with 40Ar/39Ar determinations on underlying volcanic mudlows (lahars) that mark the irst full emergence of Java from the sea. hese determinations are supported by paleomagnetic readings, and they imply that people and other large mammals could not have occupied Java before about 1.7 Ma. Explanations for the dating discrepancies include erroneous stratigraphic assessment in the ield or the selection of samples that were reworked from older sediments.

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he bottom line is that the antiquity of Javan H. erectus remains controversial. he best age estimates may come from a systematic stratigraphic and dating program conducted in the southeastern quadrant of the Sangiran region or dome. he sediments in this sector have been minimally deformed or distorted by subsequent tectonic movement, thereby reducing the likelihood of stratigraphic confusion. he Pucangan and Kabuh Beds are known here alternatively as the Sangiran and Bapang Formations, respectively, and they have provided nearly eighty H. erectus fossils. he large majority are thought to come from the Bapang Formation, but none were found during the kind of carefully monitored ieldwork that has commonly accompanied fossil recovery in eastern Africa. With this caveat in mind, it may still be meaningful that a stratigraphically consistent suite of 40Ar/39Ar determinations places the fossiliferous Bapang sediments mainly between about 1.5 and 1 Ma. he underlying Sangiran sediments must then antedate 1.5 Ma, and this supports a widely publicized 40Ar/39Ar date of 1.66 Ma on volcanic pumice overlying a central Sangiran layer that provided an H. erectus maxilla and skullcap in 1978. he researchers who obtained the Bapang dates argue, however, that the Sangiran Formation contains no pumice, and others have suggested that the dated pumice came from an underlying volcanic mudlow that was locally uplited so that it appeared to overlie the H. erectus indspot. his possibility underscores the continuing diiculty of estimating the antiquity of Javan H. erectus, and there is the further problem that tektites have been reported from deep within the Bapang Formation. hese are small glassy objects that rained down widely over Southeastern Asia following a large asteroid impact about 800 ka, and their presence would imply that much of the Bapang Formation, including perhaps the majority of H. erectus fossils, could be no older than 800 ka. Still, a 1.5–1 Ma age estimate is broadly consistent with the 1.6–1.1 Ma estimate for human presence in the Nihewan Basin of north China, and the sum may imply that H. ergaster/H. erectus reached the Far East not long ater it emerged in Africa. Passage to Java could have occurred during one of the early Pleistocene glaciations when sea level dropped by more than 60 m and a broad land bridge connected Java to mainland Southeast Asia. he fossil mammals associated with Javan H. erectus testify to repeated connections under relatively dry, glacial conditions. Glacial intervals probably promoted regional H. erectus populations because forest cover shrunk and more food became available at ground level. SOURCES (not including those in table 5.2): rock-stratigraphic units in the Sangiran-Trinil region (Bartstra 1983; Bouteaux et al. 2007; Leinders et al. 1985; heunissen et al. 1990; von Koenigswald 1962); geologic complexity of the Sangiran-Trinal region (Hertler and Rizal 2005; Sondaar 1984); luorine analysis of Sangiran fossils (Matsu’ura 1986; Matsu’ura et al. 2000); paleomagnetism and absolute

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dates for the Pucangan and Kabuh Beds (de Vos et al. 1994; Howell 1986; Larick et al. 2001; Pope 1995; Sémah 1984; Swisher et al. 1994); contradictions between dates and associated paleomagnetic readings (Hyodo et al. 2002) and between dates and fauna (Hertler and Rizal 2005); time when Java fully emerged form the sea (Sémah et al. 2000); 40Ar/39Ar dates on Bapang sediments (Larick et al. 2001); 40Ar/39Ar determination on pumice overlying 1978 H. erectus fossils (Swisher et al. 1994) and possible displacement of the pumice (Sémah et al. 2000); Bapang Formation tektites (Langbroek and Roebroeks 2000) and their dating (Hou et al. 2000); mammals as indications of connections between Java and the Southeast Asian mainland (Storm 2001)

Latest Homo erectus in Java

he upper age limit of Javan Homo erectus is also debatable. If as morphology argues, the Ngandong and Sambungmacan specimens are included, then H. erectus persisted in Java until at least 300–250 ka and perhaps until sometime between 53 and 27 ka. he older age is based on faunal associations, a uranium-series date on one of Ngandong skulls, and a ission-track date from the Notopuro Beds that have sometimes been equated with the fossiliferous sediments at Ngandong. he younger date is based on ESR and uranium-series analysis of bovid teeth that accompanied the human specimens at both Ngandong and Sambungmacan. If the younger date is correct, it would indirectly support the local replacement of H. erectus by H. sapiens spreading from Africa ater 50 ka. However, even the older date supports Chinese evidence (mainly from Zhoukoudian and Hexian) that H. erectus persisted in eastern Asia ater 500 ka when other kinds of people had appeared in Africa and Europe. he Africans are referred here to the lineage that culminated in living H. sapiens, while the Europeans are referred to the antecedents of classic H. neanderthalensis. SOURCES: latest H. erectus in Java—300–250 ka (Jacob 1978; Sémah 1984; Storm et al. 2005) or 53–27 ka (Grün and horne 1997; Swisher et al. 1996)

The Antiquity of Homo ergaster

he separation of Homo ergaster from H. erectus is controversial, and even advocates commonly restrict H. ergaster to a handful of relatively complete African fossils dated to about 1.8–1.4 Ma. To these, the same specialists would probably add human fossils from Dmanisi, Georgia, that may be roughly as old. As discussed below, some or all of the Dmanisi specimens may actually have been deposited closer to 1.2 Ma, but even in this instance, they could be the oldest evidence for people far outside of Africa and they would document the persistence of H. ergaster ater 1.4 Ma. Virtually all specialists assign later African specimens, dated between perhaps 1.2 and 0.6 Ma to H. erectus, implying either that H. erectus evolved from H. ergaster in Africa and then spread to eastern Asia or

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that H. erectus evolved in Asia and then spread back to Africa. he position taken here is that H. erectus developed exclusively in eastern Asia, beginning 1 Ma or before, and that H. ergaster persisted in Africa and perhaps in neighboring western Asia until the emergence of H. sapiens or H. heidelbergensis 600–500 ka. he issue can be decided only by the recovery of a much larger number of diagnostic, well-dated fossils, however, and readers should be aware that the present use of H. ergaster is not commonplace, particularly for fossils that postdate 1.2 Ma. In recognition of this, in what follows H. ergaster is oten designated alternatively as “African H. erectus.” Homo ergaster is far more irmly dated than east Asian H. erectus because the stratigraphic positions of key H. ergaster fossils are wellestablished, and the fossils oten come from deposits that are amenable to 40Ar/39Ar and other radiometric dating methods and to paleomagnetic analysis. hey are also usually associated with fossils of other mammalian species whose timespans have been determined by radiometric techniques. Table 5.3 presents the ages that radiometric dating, paleomagnetism, and associated mammals imply, alone or in combination, for fossils assigned here to H. ergaster. A large pelvic fragment (KNM-ER 3228) and a partial occipital bone (KNM-ER 2598) might indicate that H. ergaster was present as early as 1.95 or 1.89 Ma, respectively, but the least equivocal specimens—skulls and partial skeletons from Koobi Fora, East Turkana, and Nariokotome, West Turkana—are irmly dated between 1.8–1.7 Ma and about 1.4 Ma. Skulls from Daka-Bouri, Buia, and Olorgesailie extend the range to 1 Ma or later, and Terniine takes it to ater 780 ka, assuming the fragmentary Terniine specimens actually derive from H. ergaster. he most problematic fossils in the table are probably those from Olduvai Gorge (Bed II, Beds III/IV, and the Masek Beds), partly because paleomagnetism provides the main basis for dating, and two contradictory paleomagnetic schemes exist. he table accepts one that places all the relevant deposits within the Matuyama Reversed Paleomagnetic Chron, before 780 ka (ig. 4.9 in the previous chapter). here is also the problem that among all the African fossils considered here, the skullcap designated Olduvai Hominid (OH) 9 most blurs the distinction between H. ergaster and (east Asian) H. erectus. he antiquity of OH 9 is speculative because it was found on the surface, and if the distinction between H. ergaster and H. erectus is maintained, OH 9 could mean that H. ergaster and H. erectus (of east Asian origin) once coexisted in Africa. Pending fresh discoveries that could resolve such uncertainties, the most economic reading of the table is that H. ergaster existed alone in Africa from 1.8 Ma to at least 1 Ma and perhaps to 600 ka.

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The Antiquity of Early Homo sapiens

In the view that is tentatively accepted here, Homo sapiens is a strictly African ofshoot of H. ergaster. he earliest specimens are distinguished from H. ergaster (and H. erectus) by several advanced features, including larger endocranial volume (table 5.6), a more rounded (less angulated) occipital, expanded parietals, and a broader frontal with more arched (vs. more shellike) browridges. hese features are not expressed equally in every specimen, perhaps in part because the specimens vary in age. Among the early H. sapiens fossils in table 5.4, the principal ones that exhibit one or more advanced features are the skulls or partial skulls from Salé, Kébibat, and Jebel Irhoud in Morocco, Singa in Sudan, Bodo in Ethiopia, Lake Ndutu and Eyasi in Tanzania, Kabwe in Zambia, and Elandsfontein and Florisbad in South Africa. he table includes the remaining fossils on the admittedly circular grounds that they fall in the same time range as the diagnostic specimens. he number of diagnostic early H. sapiens skulls is small, and among them, only the principal Bodo skull comes from a setting where 40Ar/39Ar can be applied. Table 5.4 shows that the other diagnostic skulls have have been dated by ESR, by their mammalian and artifactual associations, or by both. ESR ages can almost always be questioned for reasons discussed in chapter 2, while associated mammals or artifacts usually allow age estimates only within broad limits. Still, with these caveats in mind, the estimated ages for early H. sapiens all fall in the interval between about 600 and 130 ka. he Bodo skull has been assigned to the H. sapiens category because its relatively broad frontal resembles the frontals of Kabwe and Elandsfontein. Its face, however, is remarkably massive, particularly in the midregion, and it might therefore be expected to be the oldest fossil in the group. he age estimates in table 5.4 suggest that it could be. he Elandsfontein skull might be equally old, but whatever its precise age, the rich associated fauna demonstrates that it is signiicantly older than a much more modern-looking skull from Florisbad. Eighteen of the fortyive Elandsfontein mammal species are extinct, and nine occur in extinct genera. At Florisbad, only ive of the twenty-six mammalian species are extinct, and only two are in extinct genera. he next chapter (on the Neanderthals and their contemporaries) reconsiders the fossils from Florisbad, Irhoud, and Singa because they surely overlap in age with European fossils that exhibit derived Neanderthal features. he Bodo and Elandsfontein skulls show that early H. sapiens had appeared in Africa by 600 ka, but they are similar to their probable European contemporaries. he Florisbad, Irhoud, and

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TABLE 5.6. he mean and standard deviation of endocranial capacity in

Homo ergaster and later fossil representatives of Homo. he means for early African later H. ergaster, early H. sapiens, and early H. neanderthalensis should be regarded only as rough guides, since the fossils represent lineages in which average endocranial capacity may have been increasing with time. (KNM = Kenya National Museum; ER = East Turkana (Rudolf ); WT = West Turkana; OH = Olduvai Hominid.) Taxa (and Estimated Age)

Estimated Endocranial Capacity (cc)

Source

H. ergaster (= early African H. erectus) core (1.8–1.4 Ma)

794.6 + 75.8 (N = 5)

KNM-ER 3732

750

(Holloway et al. 2004)

KNM-ER 3733

848

(Holloway et al. 2004)

KNM-ER 3883

804

(Holloway et al. 2004)

KNM-ER 47200

691

(Spoor et al. 2007)

KNM-WT 15000

880

(Begun & Walker 1993)

Proposed African later H. ergaster (1.4–0.6 Ma)

872.8 ± 149 (N = 5)

Buia

775

(Abbate et al. 1998)

Daka

995

(Asfaw et al. 2002)

Olorgesailie

800

(Potts et al. 2004)

OH 9

1067

(Holloway et al. 2004)

OH 12

727

(Holloway et al. 2004)

Dmanisi H. ergaster (?1.7–1.6 Ma)

685 ± 83.2 (N = 3)

D2280

650

(Gabunia et al. 2000b)

D2282

780

(Gabunia et al. 2000b)

D3444

625

(Lordkipanidze et al. 2006)

classic Trinil/Sangiran Indonesian H. erectus (“Pithecanthropus”) (?1–0.5 Ma)

933 ± 84 (N = 7)

(Holloway et al. 2004)

Zhoukoudian H. erectus (?670–400 ka)

1043 ± 113 (N = 5)

(Weidenreich 1943)

Sambungmacan and Ngawi later Indonesian H. erectus (?500–300 ka)

955 ± 75 (N = 4)

(Baba et al. 2003) and (Zeitoun & Widianto 2001)

Ngandong late Indonesian H. erectus (?300–50 ka)

1148 ± 89 (N = 6)

(Holloway et al. 2004)

Dali

1120

(Wu & Poirier 1995)

Jinniushan (Yinkou)

1390

(Wu & Poirier 1995)

“early” H. sapiens (Africa) (600–130 ka)

1201 ± 281 (N = 8)

Jebel Irhoud 1

1305

(Holloway et al. 2004)

Jebel Irhoud 2

1400

(Holloway et al. 2004)

Salé

880

(Holloway et al. 2004)

Chinese “late H. erectus” (between 300 and 100 ka)

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TABLE 5.6. (continued)

Taxa (and estimated age)

Estimated Endocranial Capacity (cc)

Source

Singa

1340

(Stringer et al. 1985)

Bodo

1250

(Conroy et al. 2000b)

Lake Ndutu

1100

(Rightmire 1983)

Kabwe

1325

(Holloway et al. 2004)

Elandsfontein

1225

(Drennan 1953)

west Asian (Skhul-Qafzeh) “nearmodern” humans (120–90 ka)

1545 ± 27 (N = 5)

(Trinkaus 1983b)

early Upper Paleolithic (fully modern) humans (35–24 ka)

1577 ± 135 (N = 11)

(Trinkaus 1983b)

“early” H. neanderthalensis (Europe) (530–130 ka)

1248 ± 1048 (N = 9)

Swanscombe

1325

(Holloway et al. 2004)

Arago 21/47

1166

(Holloway et al. 2004)

Biache-Saint-Vaast

1200

(Stringer et al. 1984)

Atapuerca SH 4

1390

(Arsuaga et al. 1997b)

Atapuerca SH 5

1125

(Arsuaga et al. 1997b)

Atapuerca SH 6

1220

(Lorenzo et al. 1998)

Reilingen

1430

(Holloway et al. 2004)

Steinheim

1150

(Howell 1960)

Petralona

1230

(Stringer et al. 1979)

European and West Asian classic Neanderthals (130–40 ka)

1,435 ± 184 cc (N = 15)

(Holloway et al. 2004)

Singa skulls difer conspicuously from their European counterparts, and they show that H. sapiens had diverged conspicuously from H. neanderthalensis by 250–200 ka. SOURCES: table 5.4.

The Antiquity of Early Homo neanderthalensis

As conceived here, Homo neanderthalensis includes the classic Neanderthals, who occupied Europe and southwestern Asia between roughly 130 and 50–40 ka, and their antecedents, who appear to have been strictly European. Some of the pre- (ante-) Neanderthal fossils have been dated by ESR on associated animal teeth, TL on heated artifacts, and U-series disequilibrium on stratigraphically associated lowstones. Many of the specimens can be ordered in time relative to others by associated mammalian fossils. In addition, the mammals, sometimes supplemented by

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5.5

δO18 (o/oo)

5.0

4.5

2

4.0

3.5

1

3

NEANDERTHAL ENGIS KRAPINA SACCOPASTORE MONTMAURIN BIACHE PONTNEWYDD L’AUBESIER EHRINGSDORF

4

Neanderthals

5

0.1

6 0.2

some key European sites

7 8

STEINHEIM

9

PreCASTEL DI GUIDO Neanderthals

millions of years ago

10 0.4

SWANSCOMBE ARAGO BILZINGSLEBEN

11 12

BOXGROVE MAUER VISOGLIANO

13

0.5

BRUNHES

0.3

14 15

0.6

ATAPUERCA SH

16 17

0.7

309 FIGURE 5.6. Tentative correlation of global marine oxygen-isotope stages and key european human fossil sites (modified after Hublin [1998a], 577). Odd numbered stages correspond to continental interglacials, even-numbered stages to glaciations. The lower layers of the Atapuerca GD (Gran Dolina) and Atapuerca SE (Sima del Elephante), Spain, contain the oldest well-documented artifacts and human remains in Europe. More or less continuous occupation of Europe is clear only from (interglacial) stage 15, and in northern Europe, human presence seems to have confined mainly to interglacials prior to (glacial) stage 6.

18

0.9

21

22

23

24

25

26 1

colder

First ATAPUERCA GD Europeans { ?CEPRANO

19

20

MATUYAMA

0.8

28

(LE VALLONET)

27

warmer

(LES BATTANTS) (ORCE) ATAPUERCA SE

remains of other vertebrates, plants, freshwater mollusks, and even beetles, oten reveal whether the climate was glacial or interglacial. In combination with numeric dates and mammalian fossils, paleoclimate can then be used to correlate sites with the oxygen-isotope stages of the global marine stratigraphy. he correlations are usually tentative, particularly when numeric dates are lacking or when they’re based on questionable assumptions or poor target material, and the correlation problem is particularly acute for sites that antedate oxygen-isotope stage (OIS) 7 (245–187 ka). Yet older sites formed mostly under interglacial conditions, and it is especially diicult to distinguish ones that formed during OIS 9 (334–301 ka) from ones that formed during OIS 11 (427–364 ka). Still, with these diiculties in mind, table 5.5 implies that as a group, the pre-Neanderthals date between about 600 and 130 ka. Figure 5.6 places key sites from table 5.5 in relation to the global marine isotope stratigraphy. he igure also includes six sites older than

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600 ka that are addressed in the section below on the earliest occupation of Europe. hree—Orce, Blassac-Les Battants, and Le Vallonet—present only dubious evidence for human presence, but the others—Atapuerca GD (Gran Dolina), Atapuerca SE (Sima del Elephante), and Ceprano— have provided human fossils. he Atapuerca fossils are closely associated with artifacts and broken-up animal bones, and they are more irmly dated. However, neither the Atapuerca nor the Ceprano specimens are considered in this subsection because they don’t anticipate the Neanderthals to the same degree as many fossils that postdate 600 ka. hey may record sporadic and ultimately unsuccessful attempts to colonize Europe earlier on, and it is only the “pre-Neanderthals,” ater 600 ka, who appear to have established a more or less permanent foothold. he most securely dated sites in table 5.5 are the youngest, assigned to OIS 7 and 6, between 245 and 130 ka, followed by the oldest, assigned to OIS 15, between 621 and 568 ka. he oldest—Atapuerca SH (the Sima de los Huesos)—is also by far the richest, and the human fossils occur below a speleothem (lowstone) fragment that formed at least 530 ka, based on high-resolution U-series analysis. he fragment is ideal for the U-series method, and the only question is whether it grew in place or whether it might have originated from elsewhere in the site. In the latter case, it could be older than the human fossils by an unknown amount. he SH deposits have produced remarkably few animal bones, and the only ones that could provide an independent age check come from lion, Panthera leo, and red-backed vole, Clethrionomys acrorhiza. Both species imply an age of 600 ka or less, based on dated specimens elsewhere in Europe. he only SH artifact—a classic later Acheulean hand ax—supports the same lower limit, since hand axes seem to have appeared about 600 ka in Europe. In sum, if the dated speleothem fragment formed in place, the SH fossils date from between 600 and 530 ka. If the fragment did not form in place, they could be younger, and the issue is crucial for establishing the antiquity of the Neanderthal lineage and the course of its evolution. As discussed in the next section, the SH dating also bears directly on the validity of Homo heidelbergensis. Cranial or dental remains from Pontnewydd, Biache-Saint-Vaast, Montmaurin, La Chaise-de-Vouthon, l’Aubesier, and Ehringsdorf show that the Neanderthal lineage was in place during OIS stages 7 and 6, between 245 and 130 ka. here are no like-aged European fossils that are clearly non-Neanderthal, though some, like those from the Fontéchevade and Le Lazaret Caves are too incomplete for secure characterization. Yet older European fossils are more variable. Among those that are suiciently complete for diagnosis, some, like those from Atapuerca SH, Swanscombe, Arago, Reilingen, and Petralona Cave, anticipate the Neanderthals in one or more derived traits, variably including a tendency

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to midfacial projection, an elliptical area of roughened bone (the suprainiac fossa) overlying the occipital torus, or a dentition in which the rear teeth are signiicantly diminished in size relative to the front teeth. Others, including especially the fossils from Bilzingsleben and Vértesszöllös, show no conspicuous Neanderthal specializations and are variously characterized instead by a shellike browridge, an angulated occipital with a strong, centrally developed torus, very thick cranial walls, or yet other features that closely recall Homo erectus. A chronologic sequence might be implied, if it could be shown that the more Neanderthal-like fossils postdated the others, but current dating uncertainties permit us to only to bracket both groups between 600 and 250 ka. In sum, dating and morphology show that the Neanderthals developed in Europe, beginning as much as 600 ka, but the available fossils could also be used to argue that more primitive populations with few or no Neanderthal features persisted locally until 250 ka. Alternatively, and perhaps more likely, the dating and morphological variability imply that Neanderthal features accumulated piecemeal, mainly as a consequence of genetic drit in small isolated populations, and that their frequency varied strongly from place to place until ater 250 ka. A denser, far better dated fossil record will be necessary to choose between the alternatives.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: table 5.5.

The Antiquity of Homo heidelbergensis

Homo heidelbergensis is diicult to date, in large part because it is dificult to deine. he nomenclatural type is a massive, chinless human mandible recovered in 1907 from the Mauer sand quarry, about 10 km southeast of Heidelberg, Germany. Associated microfauna implies that the deposits may have formed about 500 ka (references in table 5.5). However, the description of H. heidelbergensis depends mainly on skulls from Petralona (Greece), Arago (France), Bodo (Ethiopia), Lake Ndutu (Tanzania), Kabwe (Zambia), and Elandsfontein (South Africa). Like the Mauer mandible, they could all date from between 600 and 400 ka, and they are suiciently similar and morphologically generalized to represent a common ancestor for Neanderthals and modern humans. If the age of H. heidelbergensis has been accurately assessed, it would approximate the inferred age for the last shared mitochondrial DNA ancestor of Neanderthals and living humans (discussion in chap. 7) and for the Africans who brought later Acheulean hand axes to Europe (discussion below). he convergence of fossil, genetic, and archaeological observations adds conidence to each, but there is the problem that the main European representatives of H. heidelbergensis—from Petralona, Arago, and Mauer—may already anticipate the Neanderthals in some

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features, and the Atapuerca SH skulls certainly do. he morphological indications are reviewed below, and if the Atapuerca SH fossils approach 600 ka in age, H. sapiens and H. neanderthalensis could not have shared a common ancestor aterward. In addition, according to the rules of zoological nomenclature, if the Mauer mandible were added to the preNeanderthal group, the name H. heidelbergensis could be no more than an informal designation for early H. neanderthalensis, and some specialists already use it that way. However, before such a usage is accepted, the SH indications for an especially early emergence of Neanderthal morphology need to be replicated elsewhere. If H. heidelbergensis is retained, it could be represented in western Asia by a partial skullcap from Kocabaş, Turkey, that was discovered in a block of travertine (a variety of limestone) around 2005 (reference in table 5.12), and its easternmost representative could be a skullcap found at Hathnora in the Narmada Valley, north-central India, in 1982 (references in table 5.14). he Narmada locality later produced a human clavicle. he Kocabaş skullcap lacks artifactual associations, but mammalian fossils and thermoluminescence dates from nearby travertines tentatively place it near 500 ka. In its frontal and browridge morphology, it broadly resembles the skull from Kabwe, Zambia, that is central to the notion of H. heidelbergensis. he Narmada skullcap was associated with mammalian fossils and Acheulean artifacts that suggest a possible age between 600 and 400 ka. Like European and African fossils that some assign to H. heidelbergensis, the Narmada skullcap combines primitive features (including a massive forwardly projecting supraorbital torus, thick cranial walls, and great basal breadth) with derived traits (including an endocranial capacity of more than 1,200 cc, a relatively steep [nonreceding] frontal, and a relatively rounded occipital). Fossil skulls from Jinniushan, Xujiayao, Dali, and Maba, China, broadly resemble the Narmada skull in their combination of primitive and derived characters, and on this basis, they could be added to H. heidelbergensis, but they are almost certainly much younger than 400 ky old. heir inclusion would thus imply that west-to-east gene low intensiied ater 400 ka or that the features that deine H. heidelbergensis (particularly brain expansion) evolved later on the east than on the west. If the Chinese fossils are included in H. heidelbergensis and they acquired their diagnostic features independently, H. heidelbergensis would become a grade concept similar to “archaic Homo sapiens,” with no phylogenetic utility.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES (not including those in table 5.5): H. heidelbergensis as the last shared ancestor of H. sapiens and H. neanderthalensis (Rightmire 1996; Stringer 1995, 1996a) or as an informal designation for early H. neanderthalensis (Rosas and Bermúdez de Castro 1998); Chinese fossils that could be included in H. heidelbergensis (Groves and Lahr 1994; Rightmire 1996, 1998)

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Morphology Morphologically, Homo ergaster, H. erectus, and the early representatives of H. sapiens and H. neanderthalensis difered in relatively subtle cranial characteristics, and their separation arguably turns as much on geography and dating as it does on morphology. hey closely resembled living humans in postcranial form and function and in the fundamental coniguration of the semicircular canals within the inner ear. Together, these features imply that they were comparably committed to terrestrial bipedalism. In this, they difered from H. habilis (or one of its variants) and the australopiths, all of which probably depended more on a mix of bipedalism and tree climbing. Many of the novelties that separated H. ergaster and its successors from earlier species would also have facilitated the uniquely human capacity for endurance (sustained long-distance) running, and natural selection for endurance running could especially explain their development. In their limb bones, individuals of early Homo difered from living humans primarily in the greater constriction (stenosis) of the medullary (marrow) cavities and in the concomitant thickness of the surrounding (cortical) walls. hese features served to confer extraordinary structural strength and resistance to strain. he ruggedness of the lower limbs (known mainly from the pelvis and the femur) implies a high level of recurrent stress during extended bouts of heightened muscular exertion, while the powerful muscle markings on the arm bones indicate phenomenal upper body strength (muscular hypertrophy). Together with artifacts described below, greater bodily robusticity argues that compared to modern humans, early Homo relied far less on technology for survival. Postcranial bones show also that unlike H. habilis and the australopiths, which tended to be small by modern standards, H. ergaster, H. erectus, and the early representatives of H. sapiens and H. neanderthalensis generally approximated living people in size or, perhaps more precisely, that they varied in size to about the same extent as living people. hus, like living humans in sunny, tropical climates, H. ergaster in tropical latitudes tended to be relatively tall and long limbed. A largely complete skeleton from Nariokotome, Kenya, suggests that adult males sometimes reached 180 cm (6´1˝) and femurs in a mixed-sex sample of six individuals collected at other Kenyan localities imply that average adult height was about 170 cm (5´7˝). Two partial skeletons of H. ergaster from Dmanisi, Georgia, suggest slightly shorter average adult stature, while femurs of H. erectus from Zhoukoudian, northern China, indicate that adult males probably stood no more than 160 cm (5´3˝). he geographic diferences probably relect the adaptive advantage of long, linear bodies in sunny, tropical climes where dissipating heat was critical, as opposed to the advantage of shorter, squatter bodies in cooler settings where

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ChaP Ter f ive TABLE 5.7. he minimum numbers of postcranial elements recovered from Atapuerca SH (Sima de los Huesos) through 1995 (ater (Arsuaga et al. 1997a, p. 122)) compared to the total numbers from like-aged (mid-Quaternary) sites elsewhere in the world. he SH sample has increased signiicantly in the meanwhile, creating an even greater disparity.

Anatomical Element

Atapuerca SH

Rest of the World

Sternums

1

0

Vertebrae

54

9

Clavicles

16

2

Ribs

28

13

Scapulae

14

0

Humeri

18

7

Radii

14

1

Ulnae

15

3

Carpals

61

10

Metacarpals

23

2

Manual phalanges

162

10

Pelvises

18

8

Sacra

7

1

Femurs

29

27

Patellae

14

1

Tibiae

20

6

Fibulae

15

1

Tarsals

65

11

Metatarsals

26

3

Pedal phalanges

113

13

Total

713

126

retaining heat was more important. Since such ecogeographic variation in body size and proportions is observable within living humans (and other widespread mammalian species), it does not by itself indicate species distinctions, and it would be surprising if it had not developed once populations of primitive Homo had colonized temperate latitudes. In sum, postcraniums not only fail to diferentiate H. ergaster, H. erectus, and the early representatives of H. sapiens and H. neanderthalensis, they also imply similar terrestrial lifestyles. Archaeology shows that these lifestyles centered on a shared, if rudimentary, reliance on technology and carnivory. It follows that species divergence, if it is accepted, may have resulted less from natural selection than from random genetic drit in small widely separated populations. In these circumstances, the

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species would tend to difer more quantitatively than qualitatively, and they would overlap extensively in morphology, particularly early on. In short, the diferences among them would be relatively subtle, and compelling species recognition would depend on large samples from which both means and ranges of variation could be established. he unusually large sample from Atapuerca SH, Spain, illustrates the point. he SH assemblage comprises more than 5,500 meticulously excavated specimens that have been provisionally dated between 600 and 530 ka. he sample is more than ive times larger than the combined samples from all other sites of comparable (mid-Quaternary) age and it includes multiple elements from nearly every part of the postcranial skeleton (table 5.7), together with three well-preserved skulls and fragments of more than six others (references in table 5.5). Both the postcranial bones and the skulls are remarkable because they combine primitive features, including ones that are retained in Homo sapiens, with derived features that uniquely distinguish H. neanderthalensis, and because they vary in the extent to which the primitive-derived mix is expressed. Neither the combination nor its variability could have been anticipated in advance, and it suggests that Neanderthal specializations did not evolve as an integrated complex, but perhaps by accretion, as might be expected if genetic drit played a larger role than natural selection. Unfortunately, the Atapuerca SH sample is as unique as it is informative, and even if multiple species within primitive Homo are accepted, small sample size impedes their description. he logical species to start with is Homo erectus, since it was the irst to be described, it is the most widely accepted, and it remains the species into which many specialists would sink one or more of the others. It thus provides the standard against which the validity of the others must be judged. SOURCES (not including those in table 5.5): postcranial similarities between H. ergaster and living humans (Ruf 2008); traits in H. ergaster and later species that imply a commitment to terrestrial bipedalism (Spoor et al. 1994) and a capacity for endurance running (Bramble and Lieberman 2004); limb bone stenosis and wall thickness in early Homo (Ruf et al. 1993); implications of early Homo limb bones for strength and resistance to stress (Trinkaus 1987a); stature in east African H. ergaster (Ruf and Walker 1993), in Dmanisi H. ergaster (Lieberman 2007a; Lordkipanidze et al. 2007) and in Chinese H. erectus (Antón 2003); evolution of Neanderthal features by accretion (Dean et al. 1998; Hublin 1998b)

Homo erectus

he description here relies on the type Javan and Chinese fossils, which are the only ones that all specialists assign to H. erectus. It depends heavily on characterizations on the sources at the end of this section. It deines a rough average, and it downplays temporal and geographic variation that some specialists stress over the average. Figures 5.7 and 5.8 illustrate the cranial and facial features to which it refers.

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FIGURE 5.7. franz Weidenreich’s restorations of Homo erectus skulls from sangiran in Java and Zhoukoudian locality 1 in China (redrawn by Kathryn Cruz-uribe partly after originals by Janis Cirulis in howells [1967], 156, 169). The features that unite the sangiran and Zhoukoudian skulls in H. erectus include a thick, forwardly projecting, shelflike supraorbital torus; a large face with pronounced alveolar prognathism; a long, low, flat braincase; sagittal keeling; and a highly angulated occiput with a prominent occipital torus. The similarities have long been taken to outweigh differences like the tendency for the Zhoukoudian skulls to exhibit a more conspicuous supratoral sulcus, a more steeply rising frontal, greater breadth across the ear apertures, and a narrower occiput. However, later Javan skulls, from Sambungmacan and Ngandong, maintain the basic suite of H. erectus features, while later Chinese skulls, from Dali, Yinkou, and Mapa, do not. The implication may be that Indonesian and Chinese H. erectus were on separate evolutionary trajectories and should be separated at the species level.

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ChaP Ter f ive

parasagittal depression

sagittal keel

flat, receding frontal

supraorbital torus

occipital squama

projecting nasal bones (external nose)

occipital torus nuchal squama zygomatic arch

alveolar prognathism (forwardly projecting jaws)

(no chin)

cm

Indonesian Homo erectus (Weidenreich reconstruction) sagittal keel

supratoral sulcus

angular torus

occipital squama

supraorbital torus

occipital torus nuchal squama

alveolar prognathism (forwardly projecting jaws)

supramastoid crest mastoid mastoid crest process cm (no chin)

Zhoukoudian Homo erectus (Weidenreich reconstruction) Braincase. Long, low vaulted, and thick walled; average cranial ca-

pacity slightly more than 1,000 cc, probably increasing through time, particularly if the sample is expanded to include the later Quaternary specimens from Sambungmacan/Ngawi and Ngandong, Java (ig. 5.9); greatest breadth near the base, oten coincident with biauricular breadth (breadth between the ear apertures); frontal bone low and receding, with a thick supraorbital torus (browridge) that tends to be straight when viewed from the front or above and that is equally well-developed in the middle and at the sides; variable development of a supratoral sulcus (an inlection in the frontal proile just behind the torus) and also of sinuses within the torus; pronounced postorbital constriction. Conspicuous midline keeling on the frontal and along the sagittal suture atop the braincase, associated with (parasagittal) depressions on either side of the midline and a thickening of the bone at and near bregma along the coronal suture; occipital bone mostly sharply angled in proile, with the upper (occipital) plate (scale or squama) usually smaller

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occipital (upper) squama

receding frontal squama

supraorbital torus occipital torus

nuchal (lower) squama ear aperture

strong alveolar prognathism

317 FIGURE 5.8. skulls of Javan Homo erectus and a robust modern person, illustrating key differences, including the more prominent supraorbital torus, lower, more receding frontal, more highly angulated occipital, and greater alveolar prognathism of H. erectus (redrawn after le Gros Clark [1964], 98).

Homo erectus (Java)

receding mental symphysis (no chin) more vertical frontal squama

occipital (upper) squama supraorbital torus

nuchal (lower) squama

external occipital protuberance (occipital torus absent)

Homo sapiens (robust modern) cm

vertical mental symphysis (chin)

than the lower (nuchal) one; at the juncture of the occipital and nuchal plates, a conspicuous mound or bar of bone (the transverse occipital torus), usually projecting farthest near the midline, where it can form a blunt triangular eminence; frequent development of a bony ridge or mound (the angular torus) along the posterior length of the temporal line; relatively small mastoid processes (downwardly projecting bony protuberances on the temporal bone below and behind the ear apertures); conspicuous mastoid and supramastoid crests marking muscle attachments on the temporal bone, sometimes separated by a gully or sulcus and sometimes merged; basicranium variably lexed or arched between the hard palate and foramen magnum. he cranial similarities between the Indonesian and Chinese H. erectus skulls are usually considered to outweigh the diferences. However, the classic Chinese skulls from Zhoukoudian, Nanjing, and Hexian

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FIGURE 5.9. Top: front and side views of skull Xi from ngandong (solo), Java (redrawn by Kathryn Cruzuribe mainly after originals by Janis Cirulis in howells [1967], 160). Bottom: front and side views of skull 17 (Pithecanthropus viii) from sangiran, Java (drawn by Kathryn Cruz-uribe from photos). The sangiran skull illustrates classic indonesian Homo erectus, and it probably antedates the ngandong skull by several hundred thousand years, yet the two skulls share many conspicuous morphological features, including thick cranial walls, a low, flat frontal, a sagittal keel, a prominent occipital torus, an angulated (flexed) occipital, and the tendency for the skull walls to slope inward from a broad base. The Ngandong skulls exceed those of classic Javan H. erectus in endocranial capacity (table 5.6), their browridges are thinner in the central part, and they exhibit less postorbital constriction, but the differences are small compared with the similarities. The Ngandong people are thus often regarded as a late variety of H. erectus that occupied Indonesia long after H. sapiens and H. neanderthalensis had appeared in the West.

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ChaP Ter f ive

sagittal keel shelflike supraorbital torus

flat, receding frontal

occipital squama of the occipital flat, receding frontal

nuchal squama of the occipital

cm

Ngandong XI (late Indonesian Homo erectus) sagittal keel flat, receding frontal

shelflike supraorbital torus

occipital squama of the occipital flat, receding frontal

highly angulated occipital nuchal squama of the occipital

Sangiran 17 (classic Indonesian Homo erectus) difer from both the earlier (Trinil/Sangiran) and later (Sambungmacan/Ngawi/Ngandong) Indonesian skulls in several respects, including (1) a supraorbital torus that is generally less massive and that tends to be straighter when viewed from above, projecting no more at the midline than at the sides (ig. 5.10); (2) a more continuous supratoral sulcus associated with greater frontal convexity; (3) generally greater biauricular breadth; (4) more inwardly sloping temporal walls; and (5) relatively narrower frontal and occipital bones. he contrast must relate to geography rather than to time because even when all the dating uncertainties are considered, the oldest and youngest Indonesian specimens certainly bracket the principal Chinese skulls in age. he diferences might be regarded as phylogenetically irrelevant, but later Chinese skulls, from Jinniushan, Maba, and Dali, described below, difer much more from the Indonesian skulls, and if classic Chinese H. erectus was ancestral to the later Chinese populations, then the Chinese lineage diverged increas-

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Ngandong 5

forward projection of the browridge along the midline

Sangiran 17

Zhoukoudian XI

cm

conspicuous supratoral sulcus

Sangiran 17

Ngandong 6

Zhoukoudian XI

319 FIGURE 5.10. browridge structure form in indonesian and Chinese Homo erectus (drawn by Kathryn Cruz-uribe from photos in antón [2002], fig. 6). The top three illustrations provide superior views, the bottom three sagittal (or side) views. The browridges are continuous on all Homo erectus skulls, but they project further forward along the midline on Indonesian specimens (represented in the figure by Ngandong 5, Ngandong 6, and Sangiran 17) than on Chinese ones (represented in the figure by Zhoukoudian XI). On Indonesian skulls, the frontal squama (forehead) slopes steeply backward, while on Chinese ones it rises more vertically. The difference produces a marked inflection point or supratoral sulcus on the Chinese skulls, which is lacking on the Indonesian ones. The Sangiran and Ngandong skulls bracket the Zhoukoudian skulls in age, which implies that the browridge differences reflect geography not time.

ingly from the Indonesian one through time. In the future, paleoanthropologists may decide that the Chinese specimens should be removed from H. erectus to a separate species, for which the name H. pekinensis would be most appropriate. Face. Short (from top to bottom), but massive and relatively wide, with

the nasal aperture projecting forward relative to the adjacent maxillary and zygomatic regions; pronounced alveolar prognathism (forward projection of the jaws); mandible robust and backward sloping below the incisors, resulting in the absence of a chin (“mental eminence”); a tendency for multiple mental foramina (passages for blood vessels and nerves on the outer surface of the mandible, generally below P4 or M1) (ig. 5.11). Dentition. Cheek teeth large relative to those of modern people, but gen-

erally reduced compared with those of Homo habilis or the australopiths and possibly reducing progressively with time; sporadic occurrence (in Javan specimens) of a diastema (gap) between the upper canine and

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ChaP Ter f ive

anterior teeth (canines and incisors)

FIGURE 5.11. mandibles of indonesian Homo erectus and a robust modern person (redrawn after le Gros Clark [1964], 94). unlike modern mandibles, those of H. erectus lacked a chin, and unlike mandibles of most hominins, they often had multiple mental foramina. These are perforations for the passage of blood vessels and nerves that occur on the outer surface of the mandible generally below P4 or m1.

cheek (= posterior) teeth (premolars and molars)

ascending ramus (broken)

Homo erectus

(no chin) mandibular corpus mental foramina or body

mental foramen

Homo sapiens

chin 0

5 cm

lateral incisor, associated with a projecting, though spatulate, upper canine; third molar generally smaller than the second, as in like-aged or later species of Homo but not in the australopiths; incisors large; upper incisors oten spatulate or shoveled (curled inward at the lateral edges). Postcranium. Known mainly from the femur and the pelvis, which are

generally similar to those of modern people. However, the pelvis was remarkably robust and distinguished especially by a thick buttress (or pillar) of cortical (outer) bone arising vertically above the acetabulum (the socket for the femur). he femur was equally robust, with extraordinarily thick external (cortical) bone, a relatively narrow internal (medullary or marrow) canal, and pronounced muscle markings. he femoral shat difered from that of later people in its greater fore to at compression (greater platymeria, resulting in a more oval, less round shat circumference), in the absence of a longitudinal bony ridge or pilaster on the rear surface, and in the more distal position of minimal shat breadth. SOURCES (not including those in tables 5.1 and 5.2): fundamental characterization of H. erectus (Antón 2003; Howell 1978a; Howells 1980; Hublin 1986; Kaifu et al. 2005; Le Gros Clark 1964; Rightmire 1981, 1984a, 1985, 1988, 1990); variation that may imply H. erectus comprises multiple species (Schwartz 2004; Schwartz and Tattersall 2003); basicranial lexion in H. erectus (Baba et al. 2003; Laitman 1985); diferences between skulls of Chinese and Indonesian H. erectus (Antón et al. 2002; Baba et al. 2003; Santa Luca 1980; Weidenreich 1943); H. erectus postcranium (Day 1984, 1995); diferences between H. erectus and later people in the femur (Kennedy 1983, 1984b; Weidenreich 1941)

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frontal bone supratoral sulcus occipital squama

supraorbital torus elevated nasal bones

occipital torus nuchal squama

maxilla zygomatic ear arch aperture

alveolar part of the maxilla

0

5 cm

temporal line

supraorbital torus

321 FIGURE 5.12. skull Knmer 3733 from Koobi fora, east Turkana (formerly east rudolf), northern Kenya (redrawn by Kathryn Cruzuribe after howell [1978a], fig. 10.10). KNM-ER = Kenya National Museum–East Rudolf. The skull exhibits several features that distinguish it from preceding Homo habilis, including a large, forwardly projecting supraorbital torus, a supratoral sulcus separating the torus from a low, receding frontal, a highly angulated occipital bone with a pronounced occipital torus, and an expanded braincase (with an estimated endocranial capacity of 848 cc). It is one of three north Kenyan fossils documenting the emergence of Homo ergaster about 1.7–1.6 Ma.

zygomatic arch nasal aperture

maxilla

KNM-ER 3733 (Homo ergaster)

postorbital constriction

Homo ergaster

In the scheme that is profered here, H. ergaster is a mainly African species that existed between 1.8–1.7 Ma and perhaps 600 ka (table 5.3). As indicated in the section above on geologic age, it is founded mainly on fossils dated between 1.8 and 1.4 Ma in the Lake Turkana Basin of northern Kenya. he core specimens are two skulls (KNM-ER 3733 and 3883) (ig. 5.12) and a partial skeleton (KNM-ER 1808) from Koobi Fora, East Turkana, and a skull and associated skeleton (KNM-WT 15000) (igs. 5.13 and 5.14) from Nariokotome III, West Turkana. To these can be added a skull (SK 847) (ig. 5.15) from Swartkrans Cave, South Africa, that is morphologically similar to the Turkana Basin specimens and that probably also dates between 1.8 and 1.5 Ma. he only compelling extra-African specimens are skulls from Dmanisi, Georgia, that also closely recall East and West Turkana skulls (references in table 5.12) (igs. 5.41 and 5.42).

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FIGURE 5.13. skull of Knm-WT 15000 from nariokotome iii, West Turkana, northern Kenya (drawn by Kathryn Cruz-uribe from photos in Walker and leakey [1993c]). The skull came from a subadult male who died roughly 1.5 ma, and it exhibits the same features that distinguish other skulls of Homo ergaster (or early african H. erectus) from those of H. habilis, including a distinct supraorbital torus, an enlarged braincase (with an endocranial capacity of 880 cc), and a reduction in the postcanine dentition. KNM-WT = Kenya national museum–West Turkana.

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supraorbital torus

flat, receding frontal

long, low vault

highly prognathic face and jaws

cm

receding mandibular symphysis (no chin)

KNM-WT 15000 (Nariokotome III) As noted below, they could date from as much as 1.7 Ma, in which case they would show that H. ergaster expanded from Africa almost as soon it evolved. Alternatively, their age could be closer to 1 Ma, in which case they could join the 1-my-old skulls from Buia, Eritrea, and Daka, Ethiopia (ig. 5.16) to document the persistence of H. ergaster morphology for 800–700 ky. he partial cranium (KNM-OL 45500) from Olorgesailie, Kenya, shows that some H. ergaster individuals retained small endocranial capacities (around 800 cc) as recently as 970–900 ka. H. ergaster was ancestral to all later species of Homo, including H. erectus, so it is not surprising that it resembled H. erectus in many key features. Particularly notable is a suite of shared, derived similarities that separate H. ergaster and H. erectus jointly from preceding H. habilis (broadly understood). he shared novelties include an increase in the size of the brain (from a mean of roughly 630 cc in H. habilis to about 800 cc in H. ergaster to more than 1,000 cc in H. erectus); a decrease in the postcanine dentition (and in the robusticity of the associated jawbones); a reduction to one in the number of roots on the upper (maxillary) premolars; a dental eruption schedule (inferred from KNM-ER 820 and WT 15000) that closely approximated the schedule in living people; a vertical shortening of the face; the formation of a pronounced supraorbital torus with supratoral sulcus; the development of an occipital torus; the forward projection of the nasal aperture; a reduction in relative arm length leading to arm/leg proportions like those in living humans; an increase in average body size to roughly that of living humans; and a signiicant reduction in sexual dimorphism compared to the australopiths and perhaps H. habilis. In addition, the nearly complete skeleton of specimen KNM-WT 15000 (ig. 5.14) implies that H. ergaster was further distinguished from

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0

Homo ergaster (KNM-WT 15000)

50 cm Australopithecus afarensis Enlarged A. afarensis (AL 288-1) (AL 288-1)

the australopiths and possibly H. habilis by a narrow hip region (the pelvis was remarkably narrow between the crests of the iliac blades) by a barrel-shaped chest (rib cage), and by larger hind limb and vertebral joint surfaces. In contrast, the australopiths (as known especially from AL 288–1, the famous Hadar skeleton of “Lucy”) had relatively much broader hips (pelvises), their rib cages were shaped like cones or inverted funnels as in chimpanzees, and their hind limb and vertebral joint surfaces were closer in size to those of chimpanzees. Homo ergaster may have evolved from H. habilis in a burst of rapid evolution (in the mode predicted by punctuated equilibrium) or it may have emerged more gradually. A inal decision probably depends on whether H. habilis is retained as one species or split between two, and this in turn depends on the accumulation of many new, well-dated fossils. An abrupt increase in aridity and seasonality across eastern Africa 1.8–1.7 Ma could explain an abrupt species origin, since it produced the kind of natural selective pulse that is presumed to underlie most punctuated evolutionary events. he conclusion to the previous chapter briely summarized the terrestrial and ofshore geologic indications of increased aridity and seasonality. he essentially modern sequence of dental eruption in H. ergaster might imply a modern rate of growth and maturation, but as discussed in the section below titled “Maturation Rates and Longevity,” H. ergaster (and H. erectus) developed their dental crowns more rapidly than do living people, and the rate was probably about the same as in H. habilis and Australopithecus. More rapid crown formation implies short childhoods

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323 FIGURE 5.14. Left: skeleton of Homo ergaster from nariokotome, West Turkana, northern Kenya; Middle: skeleton of Australopithecus afarensis (al 288-1, “lucy”) from hadar, ethiopia; Right: The A. afarensis skeleton scaled to the height of the H. ergaster skeleton (redrawn after ruff [1993], 55). H. ergaster was the first hominin species to achieve the stature and bodily proportions of living humans. The H. ergaster individual whose skeleton is pictured was a boy whose age at death has been variously estimated between eight and twelve years. Yet he stood about 1.60 m tall (5'3“) and if he had experienced an adolescent growth spurt, he might have reached 1.85 m (6'1“) at adulthood (Ruff and Walker 1993). His body was long and linear like the bodies of living humans who inhabit similar hot, dry savanna environments. KNM-WT = Kenya National Museum–West Turkana; AL = Hadar.

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ChaP Ter f ive

supratoral sulcus

FIGURE 5.15. fossils of Homo ergaster (or early african H. erectus) from swartkrans Cave (SK), south africa (drawn by Kathryn Cruz-uribe from photos in Clarke [1994c], 187). faunal associations indicate that the fossils date from between 1.8 and 1.5 ma, and they may join similar specimens from the lake Turkana basin, Kenya, in the core sample of H. ergaster. The partial skull labeled sK 847 actually comprises three separately numbered pieces that were reassembled by r. J. Clarke. The fragmentary mandible sK 45 may represent the same individual. all the fossils are readily distinguishable from those of the robust australopith, Paranthropus robustus, which occur in the same deposit.

(thick) supraorbital torus prominent nasal bones

prognathic upper jaw

SK 847 cm

large canine alveolus (socket) compared with that of Paranthropus

much narrower molars than those of Paranthropus M1

M2

M3

SK 15

SK 45 M1 M2

by modern standards, and since childhood (time to sexual maturity) and potential lifespan are closely linked in mammals, H. ergaster, H. erectus, and their predecessors probably also had relatively short lifespans, perhaps more apelike than humanlike. Analyses of crown formation times summarized in the next chapter imply that prolonged childhoods and lifespans like those of living humans may have appeared only ater 200 ka in the Neanderthals and more certainly in their modern (or nearmodern) African contemporaries. he increased body size of H. ergaster could account for most of its brain enlargement, and when its larger mass is considered, H. ergaster was only slightly bigger brained (more encephalized) than H. habilis.

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sagittal keel

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large, double-arched supraorbital torus

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Daka calvaria (Bouri, Middle Awash, Ethiopia)

maximum breadth near the base

Substantially greater encephalization occurred only ater 600 ka in people who resembled H. ergaster in size but had much larger endocranial volumes (generally exceeding 1,200 cc). Limited encephalization in H. ergaster and a relatively short childhood were probably linked, and they both imply a limited capacity to acquire new knowledge and behaviors. Still, the invention of hand axes and other Acheulean artifacts 1.7–1.6 ka strongly suggests a cognitive advance. More sophisticated brains and more reined technology in turn could help explain how H. ergaster expanded into arid, highly seasonal settings in eastern and southern Africa and how it became the irst hominin species to colonize Eurasia. Range expansion implies population growth, and population growth in turn implies that the novel features of H. ergaster signiicantly enhanced human ability to survive and reproduce. In this sense, the emergence of H. ergaster signals genuine evolutionary as opposed to strictly historical change. Adaptation to aridity may explain the change in the position of the nasal aperture, which relects the appearance of the typically human external nose with downwardly facing nostrils (vs. the relatively lat nose, with more forwardly facing nostrils, of the apes and of hominins before H. ergaster). he key point is that the external nose is usually cooler than the central body, and during periods of heightened activity, it thus tends to condense and recycle moisture that would otherwise be exhaled.

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325 FIGURE 5.16. fossil skullcap from the daka (dakanihylo) member of the bouri formation, middle awash valley, ethiopia (drawn by Kathryn Cruz-uribe from photos in asfaw et al. [2002], fig. 2). Like skulls of broadly contemporaneous Far Eastern Homo erectus, the Daka skull has its maximum breadth near the base, a keel along the midline at the top, and a massive browridge. Its endocranial volume of 995 cc closely approximates the Far Eastern mean. It differs from classic Far Eastern H. erectus in the arching of the browridge over the orbits, the height of the vault relative to the breadth, the absence of a well-defined occipital torus, and the relative thinness of the vault bones. In the absence of a distinct occipital torus and vault bone thinness, it recalls earlier African H. ergaster, and it is taken here to indicate the persistence of H. ergaster in Africa after H. erectus had emerged in the Far East.

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Adaptation to hot, arid environments must have required yet other mechanisms for cooling the body and especially for preventing the brain from overheating. First and foremost may have been an enhanced capacity to sweat, which would have been efected most eiciently by a substantial reduction in body hair. H. ergaster may thus have been the irst hominin species to possess a largely hairless, naked skin. If so, it was probably also the irst to have dark skin. he irst hominins probably had light skin under dark hair (like the chimpanzees and gorilla). However, once the hair was lost in populations occupying hot, sunny environments, natural selection would have rapidly favored dark skins to reduce the damaging efects of ultraviolet radiation on sweat glands and on folate, a form of vitamin B9. Photoprotection of folate in the blood may have been particularly important, since folate deiciency is known to cause spontaneous abortions and birth defects. Reduced arm length in H. ergaster probably signals the inal abandonment of any apelike dependence on trees for feeding or refuge and, conversely, the development of the more thoroughly terrestrial adaptation to savanna life that characterized historic, tropical hunter-gatherers. A more exclusively terrestrial lifestyle and a greater reliance on bipedalism probably also drove the narrowing of the pelvis and the change from a cone-shaped to barrel-shaped chest. he narrower pelvis increased the eiciency of muscles that operate the legs during bipedal locomotion, and it would have forced the lower part of the rib cage to narrow in concert. he result would have been the slimmer waist that sharply distinguishes people from chimpanzees, and combined with shorter arms, longer legs, and expanded hind limb and lumbar vertebral joint surfaces, the slimmer waist would particularly have facilitated endurance running. To maintain chest volume (pulmonary or lung function) as the lower rib cage contracted, the upper part would have had to expand, and the modern barrel shape would follow. he narrowing of the pelvis also constricted the birth canal, and this must have forced a reduction in the proportion of brain growth that occurred before birth. he implication is a longer period of infant dependency, foreshadowing the remarkably long period that distinguishes living humans from other species. Pelvic narrowing probably also reduced the volume of the intestines (gut), but this could occur only if food quality improved simultaneously. Circumstantially then, pelvic narrowing might imply increased consumption of meat and marrow, more efective processing of tubers and other vegetal foods, or both. It might especially mean the invention of cooking, but as indicated below, the oldest reasonably secure evidence for control over ire dates from ater 800 ka. A more complete adaptation to terrestrial life probably also selected for increased body size, and it could even explain the decrease in sex-

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ual dimorphism. In extant higher primates, strong sexual dimorphism tends to relect polygynous mating systems in which males compete vigorously for females, while reduced sexual dimorphism is associated with monogamous mating systems in which males and females pair for long periods. Decreased dimorphism in H. ergaster may thus signal the beginnings of the distinctively human pattern of sharing and cooperation between the sexes that promoted historic hunter-gatherer survival in savanna environments. Reduced cheek tooth size in H. ergaster/erectus probably relects a greater reliance on technology (artifacts) to process food. here may have been a simultaneous increase in the use of the incisor teeth for biting, gripping, and tearing, culminating in the heavily shoveled upper incisors of H. erectus. Shoveling greatly increases the area of the occlusal surface, and it could thus prolong incisor life under heavy use. If increased emphasis on the front teeth is accepted, it could further mean that the forwardly projecting supraorbital torus evolved as a structure to resist the stress that repeated incision (biting between the front teeth) would produce on the frontal bone. However, gauges attached to the frontal above the orbits of living macaque monkeys and baboons show that incision does not signiicantly strain the frontal bone, and the supraorbital torus may have developed simply to connect the braincase and face in fossil people whose faces grew entirely in front of the brain (rather than partly below it as in living humans). he derived features that H. ergaster and H. erectus shared are striking, and they explain why many authorities prefer to regard H. ergaster simply as early H. erectus. Separation depends on the argument that H. erectus was also derived relative to H. ergaster in some key features, including the tendency for H. erectus to have a lower, less domed cranium; thicker cranial walls; a strong sagittal keel (an elongated mound of bone or torus starting on the frontal and continuing between the parietals along the top of the skull); a much more conspicuous occipital torus (bony bar) around the rear of the skull; a more projecting supraorbital torus (browridge) that is as thick laterally as it is in the middle; a thickened tympanic plate (the bony enclosure of the middle ear that supports the eardrum and is fused to the temporal bone at the base of the skull); a signiicantly reduced mound of bone (postglenoid process or eminence) behind the articular depression (fossa) for the mandibular condyle on the base of the temporal bone; and a more massive face. he diferences between H. ergaster and H. erectus are admittedly more subtle than the similarities, particularly since they are mainly quantitative, and they must be assessed on a relatively small number of specimens. If the diferences are accepted, however, H. erectus is a plausible descendant of H. ergaster, and it a less likely ancestor for

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FIGURE 5.17. skullcap of olduvai hominid 9 from upper bed ii, olduvai Gorge (drawn by Kathryn Cruz-uribe from a cast and slides). The skullcap is thought to date from about 1.2 ma. it is assigned here to Homo ergaster, but among all african fossil skulls, it is the most difficult to separate from skulls of classic Far Eastern H. erectus. Conspicuous features it shares with H. erectus include a massive, forwardly projecting supraorbital torus, a low, receding frontal bone, a highly angulated occipital, and thick cranial walls.

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supratoral sulcus

(receding) frontal squama

supraorbital torus

angulated occipital

0

5 cm

Olduvai Hominid 9

ear aperture

H. sapiens and H. neanderthalensis, both of which tend to be derived away from H. ergaster in features that also diferentiate them from H. erectus. hese features include a supraorbital torus that tends to be thicker in the middle than at the sides, parietals that usually lack a midline keel and that tend to bulge (or boss) outward, and occipitals that are relatively rounded. Primitive morphology relative to the other three species and the geologic antiquity of H. ergaster together explain its basal position in igure 5.1. On strictly morphological grounds, the strongest objection to H. ergaster as advocated here is probably OH 9 (ig. 5.17), a previously mentioned skullcap from Olduvai Gorge. It was found in pieces on the loor of a gully that cut through Upper Bed II, Bed III, and Bed IV, but it is generally assumed to have come from Upper Bed II. If so, it probably dates from about 1.2 Ma, and it is distinguished from the 1.8–1.5-myold Turkana skulls by some of the same features that have been said to distinguish H. erectus from H. ergaster, namely, thicker cranial walls, a massive supraorbital torus, and a more angulated occipital. Its estimated endocranial capacity of 1,067 cc also exceeds that of all other supposed H. ergaster skulls (table 5.6) by an average of 37%, and even strong proponents of H. ergaster in eastern Africa have placed OH 9 in H. erectus. Still, like the Turkana specimens of H. ergaster and also H. sapiens, OH 9 lacks the interparietal keel that marks classic (Far Eastern) H. erectus, and it resembles both Turkana H. ergaster and H. sapiens in details of the tympanic plate. At least tentatively then, it can be assigned to H. ergaster, and it illustrates the diiculty of separating closely related species on quantitative characters alone when the mean conditions and variation around the means are poorly established. A special irony is that if OH 9 is included in H. ergaster, H. ergaster should perhaps be renamed H. leakeyi, since H. leakeyi was applied to OH 9 before H. ergaster was proposed for Turkana Basin fossils.

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In sum, H. ergaster and H. erectus resembled each other closely, and reasonable specialists can disagree on whether they should be separated. At the moment, the strongest argument for separation may be that the conspicuous evolutionary divergence between west (Africa and western Eurasia) and east (eastern Asia) ater 1 Ma must have deeper roots. A compelling morphological argument will depend on a much larger number of well-dated fossils, but in their absence, specialist disagreement should not be allowed to obscure the fundamental evolutionary implications of the key African fossils. Recall that Homo habilis, or the constituents into which it may ultimately be divided, was variable in the extent of its departure from the australopiths. Some specimens retained large cheek teeth and an australopith-like chewing apparatus, while others were not especially large brained and some also retained postcranial features suggesting an australopith-like combination of bipedalism and tree climbing. In contrast, H. ergaster or African H. erectus was uniformly advanced in brain enlargement, reduced cheek tooth size, and a postcranial structure that relects an exclusive reliance on bipedalism. Archaeology also shows that H. ergaster or African H. erectus was the irst hominid species to colonize arid, seasonal environments like those in which historic hunter-gatherers are well-known. If H. habilis may be seen as transitional between the australopiths and later humans, then H. ergaster may be regarded as the irst true human species. SOURCES (not including those in table 5.3): persistence of H. ergaster morphology for 800–700 ky (Manzi 2004); dental crown development and eruption schedule in H. ergaster (Moggi-Cecchi 2001a); description of KNM-WT 15000 (Walker and Leakey1993a, 1993b), including the hip region (Walker and Ruf 1993) and the rib cage (Jellema et al. 1993); increased aridity and rainfall seasonality about 1.7 Ma inferred from dust in deep-sea deposits of Africa (deMenocal 1995, 2004; deMenocal and Bloemendal 1995), from soil carbonates in eastern Africa (Cerling 1992; Wynn 2004), from a lowstone in Bufalo Cave, South Africa (Hopley et al. 2007), and from plant and animal fossils (Bonneille 1994, 1995); dental crown formation and maturation rate in H. ergaster (Dean et al. 2001; Moggi-Cecchi 2001b); pace of enamel growth in Irhoud early-modern H. sapiens (Smith et al. 2007a); fast enamel growth in H. heidelbergensis and H. neanderthalensis (Ramirez Rozzi 1993b and Bermúdez de Castro 2004); slow enamel growth in the near-modern African contemporaries of the Neanderthals (Smith et al. 2006) and in Upper Paleolithic/Mesolithic Europeans (Ramirez Rozzi and Bermúdez de Castro 2004); encephalization in H. ergaster (McHenry 1994a, 1994b) and increased encephalization ater 600 ka (McHenry 1994b; Walker and Leakey1993b); the external nose—emergence (Franciscus and Trinkaus 1988) and function (Trinkaus 1987a); natural selection and skin color (Jablonski and Chaplin 2000); need for mechanisms to cool the brain in H. ergaster (Falk 1990); explanation for narrowing of the waist and development of a barrel-shaped chest (Jellema et al. 1993); structural specializations for endurance running in H. ergaster (Bramble and Lieberman 2004); narrowed waist and the birth canal in H. ergaster (Walker and Leakey1993b); relation between pelvic narrowing, gut reduction, and food quality (Aiello and Wheeler 1995); possible increased use of the incisors as tools in early Homo (Wolpof 1980b, 1985b); biting between the front teeth and the development of the supraorbital torus—pro (Russell 1985) and con (Hylander and Johnson 1992); the supraorbital torus as a connection between the braincase and a forwardly positioned face (Lieberman 2000); H. ergaster as an early form of H. erectus (Bräuer 1994; Bräuer and Mbua 1992; Rightmire 1990; Walker and Leakey1993b) or as separate, less specialized species (Andrews 1984; Clarke 1994c; Groves 1989; Groves and Lahr 1994; Manzi 2004; Stringer 1984; Wood 1984); OH 9—discovery (Leakey 1961; 1971, 229), geologic age (Leakey and Hay 1982), similarities to H. erectus (Groves 1989; Wood 1991) and to H. ergaster/H.

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sapiens (Clarke 1990), endocranial capacity (Holloway et al. 2004, 298), and assignment to H. leakeyi (Heberer 1963); the initial proposal for H. ergaster (Groves and Mazák 1975)

Homo sapiens, H. neanderthalensis, and H. heidelbergensis

Like Homo ergaster, the early representatives of H. sapiens (Africa) and H. neanderthalensis (Europe) shared many features with H. erectus, including large browridges; a low, lattened frontal bone; a relatively broad cranial base; thick cranial walls; relatively massive, chinless mandibles with large teeth; and powerfully constructed postcraniums. However, they diverged from both H. ergaster and H. erectus in average endocranial volumes that signiicantly exceeded the average (of just over 1,000 cc) for the H. erectus core (table 5.6), and in a shared tendency to more rounded occipitals, expanded parietals, and broader frontals with more arched (vs. more shellike) browridges. he similarities among early H. sapiens, early H. neanderthalensis, and H. erectus are probably primitive retentions from a shared H. ergaster ancestor, whereas the traits that separate H. sapiens and H. neanderthalensis from H. erectus are derived characters. hey indicate that the line or lines leading to H. sapiens and H. neanderthalensis had diverged from the line that produced H. erectus by 1 Ma or before. As noted previously, H. sapiens in Africa and H. neanderthalensis in Europe may have inherited their shared, derived traits from a common ancestor that lived roughly 600–400 ka and that is represented by the skulls from Bodo, Ndutu, Kabwe, and Elandsfontein in Africa, by those from Petralona and Arago in Europe, and perhaps by the like-aged Kocabaş skull from Turkey and the Narmada skull from north-central India (ig. 5.18). Each of these skulls tends to exhibit roughly the same mix of primitive features that are shared with H. erectus and of more advanced features that separate both later H. sapiens and H. neanderthalensis from H. erectus. To the extent that the African and European skulls cannot be separated morphologically, they are reasonably joined in a common species for which the name Homo heidelbergensis has been most commonly proposed. Artifactual contrasts discussed below underscore the divergence of eastern Asia from Africa and Europe by 1 Ma, while artifactual similarities indicate the likelihood of a population dispersal from Africa to Europe around 600 ka. As already noted, however, the concept of H. heidelbergensis is problematic, partly because it includes fossils that may vary greatly in age, partly because the putative European specimens may anticipate the classic Neanderthals (ater 200–130 ka) in derived features, and partly because the Atapuerca SH fossils certainly do. he SH specimens are not commonly included in H. heidelbergensis, but they may be as old as or older than the fossils that are, and if so, the last shared

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superior temporal line

parietal (receding) frontal

upper plate of the occipital

lower plate of the occipital

supraorbital torus

331 FIGURE 5.18. skullcap of archaic Homo from the narmada valley, northcentral india (redrawn by Kathryn Cruz-uribe after de lumley and sonakia [1985b], 26, 30). associated animal bones and “upper” acheulean artifacts imply the skull dates from between 600 and 400 ka. it combines primitive features like a thick, forwardly projecting supraorbital torus, thick cranial walls, and great basal breadth with advanced features like expanded parietals, a less angulated (flexed) occipital, and a relatively large endocranial capacity (>1,200 cc). In its mix of primitive and derived characters, it recalls like-aged African and European specimens that are variably assigned to early Homo sapiens, early H. neanderthalensis, or H. heidelbergensis.

superior temporal line

Narmada Skull

0

5 cm

ancestor of the Neanderthals and modern humans must have existed before H. heidelbergensis. he name would then have to be abandoned except perhaps as an informal designation for the early representatives of H. neanderthalensis. he issue can be resolved only by fresh fossils and fresh dates, but even if H. heidelbergensis is retained, its morphology can be adequately understood from a consideration of H. sapiens and H. neanderthalensis, and it will not be considered further here. he case for separate H. sapiens and H. neanderthalensis lineages rests primarily on European fossils that date between roughly 600 and 200 ka and that combine primitive features of the genus Homo with evolutionary novelties that uniquely mark the classic Neanderthals who occupied Europe ater 130 ka. In contrast, African fossils dated between 600 and 200 ka exhibit no Neanderthal specializations, but they plausibly anticipate the modern or near-modern people who inhabited Africa

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prelambdoidal flattening

parietal

lambda

lambdoidal suture

asterion

Swanscombe FIGURE 5.19. fossil skull from swanscombe, england (drawn by Kathryn Cruzuribe partly after originals by Janis Cirulis in howells [1967], 218; and photos in stringer et al. [1984]). The skull is probably between 400 and 250 ka old. in its thick cranial walls and maximum breadth near the base, it recalls skulls of Homo erectus. however, it anticipates skulls of the classic neanderthals in the flattening of the parietal region just in front of lambda (a point on the rear of the skull where the lambdoid and sagittal sutures join) and especially in the presence of an incipient suprainiac fossa (an elliptical, depressed area of roughened bone) over the occipital torus. It is assigned here to early H. neanderthalensis.

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occipital torus

thick cranial walls

cm

incipient suprainiac fossa

and the adjacent southwestern corner of Asia ater 130 ka. he relevant European fossils include the skulls from Swanscombe (England) and Reilingen (Germany) and above all, the extraordinary sample of human remains from Atapuerca SH (Spain) (references in table 5.5). he African fossils include the skulls from Salé and Jebel Irhoud (Morocco), Singa (Sudan), Bodo (Ethiopia), Lake Ndutu (Tanzania), Kabwe (Zambia), and Elandsfontein and Florisbad (South Africa) (references in table 5.4). he skulls from Swanscombe (ig. 5.19) and Reilingen both lack faces, but they are similar in the position of maximum breadth near the base of the skull and in occipital morphology, including a torus that is better developed at the sides than at the center and that is surmounted by an elliptical depression of roughened bone (the suprainiac fossa, illustrated in igs. 6.5 and 6.6 in the next chapter). Maximum breadth near the cranial base is a primitive feature that is not retained in Neanderthal skulls, whose maximum breadth is at midparietal level (ig. 5.20). he Reilingen skull is also primitive in the large size of its mastoid process, which tends to be signiicantly smaller in Neanderthals (the mastoid region is not preserved on the Swanscombe skull). In contrast, the occipital morphology that Reilingen and Swanscombe share is a derived feature that is expressed more fully in the classic Neanderthals. he Atapuerca SH sample includes three nearly complete skulls, large fragments from at least six other skulls, numerous small craniofacial fragments, more than forty-one complete or partial mandibles and numerous isolated teeth from at least twenty-eight individuals, and hundreds of postcranial bones. To an even greater extent than the Reilingen and Swanscombe skulls, the SH sample is remarkable for its combina-

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low

pentagonal

? intermediate interparietal (sagittal) keel

interparietal (sagittal) keel

Homo erectus (Zhoukoudian, China)

Homo erectus (Ngandong, Indonesia)

high pentagonal parietal boss

early Homo sapiens (Salé, Morocco)

modern Homo sapiens

cm interparietal (sagittal) keel

globular (“en bombe”)

intermediate pentagonal incipient suprainiac fossa

suprainiac fossa

large mastoid process

small mastoid process

early Homo neanderthalensis (Atapuerca SH 5, Spain)

early Homo neanderthalensis (Swanscombe, England)

classic Homo neanderthalensis (La Chapelle, France)

FIGURE 5.20. skulls of Homo erectus, early H. sapiens, modern H. sapiens, early H. neanderthalensis, and classic H. neanderthalensis viewed in occipital (rear) view (redrawn after hublin [1996], 41; and arsuaga et al. [1996], photo on 224). from this perspective, skulls of H. erectus are pentagonal with sidewalls that slope sharply inward from near the base (“low pentagonal”); skulls of early H. sapiens and early H. neanderthalensis are pentagonal with walls that tend to rise more vertically; skulls of classic H. neanderthalensis are globular with walls that bulge outward at midparietal level; and skulls of modern H. sapiens are pentagonal with walls that rise vertically to a point high on the parietals before sloping inward (“high pentagonal”). skulls of modern H. sapiens also tend to show a boss or bulge at the point where the parietals turn inward. arguably the early H. sapiens skull from salé, morocco, shows an incipient boss.

tion of primitive features also seen in H. erectus or H. sapiens, derived features that are shared between H. sapiens and H. neanderthalensis, and derived features that are unique to H. neanderthalensis (table 5.8). he most conspicuous Neanderthal specializations in the SH skulls are a face that projects far forward along the midline and an oval area of roughened or porous bone overlying the occipital torus (ig. 5.21). Midfacial projection and other Neanderthal facial features are particularly impressive in the most complete skull (SH 5) (ig. 5.22). Some other facial fragments are less obviously Neanderthal-like, and one large piece (AT [Atapuerca] 404) exhibits a distinctly non-Neanderthal canine fossa (delation or hollowing of the bone above the canine tooth between the nasal aperture and the zygomatic arch). Such a fossa also characterizes the somewhat younger skull from Steinheim, Germany, which is arguably Neanderthal-like in its relatively rounded occipital. he roughened oval area on the SH occipitals varies from slightly less depressed than on the

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Characters that human fossils from Atapuerca SH (Sima de los Huesos), Spain, share with fossils of Homo erectus (including H. ergaster), the classic Neanderthals (ater 130 ka), and H. sapiens (modiied ater (Stringer 1993, p. 502)). he features shared with H. erectus are primitive features for the genus Homo, whereas those shared exclusively with H. sapiens probably characterized a common ancestor of H. sapiens and the Neanderthals. he features shared only with the Neanderthals are uniquely derived, which suggests that the SH population should be placed on or near the line leading to the Neanderthals. TABLE 5.8.

Homo Erectus

Classic Neanderthals

Homo Sapiens

Vault broadest near the base

X

Substantial, overall facial projection (prognathism)

X

Laterally thick supraorbital torus

X

Relatively lat occipital squama

X

Interparietal (sagittal) torus

X

Mandibular robusticity

X

X

Lower limb robusticity

X

X

Cranial capacity range

X

X

X

Double-arched supraorbital torus

X

X

Incipient suprainiac fossa

X

Substantial midfacial projection

X

Anterior teeth large relative to posterior ones

X

Large retromolar space, an asymmetric sigmoid notch, and a large, posteriorly projecting coronoid process on the mandible

X

Lateral occipital proile relatively rounded

X

X

Relatively thin tympanic bone

X

X

High cranial vault

X

X

Temporal squama high and rounded

X

X

Rear parietal proile

X

Adult mastoid process large

X

Swanscombe or Reilingen skulls to lat or even convex, but it still foreshadows the classic Neanderthal suprainiac fossa. he numerous primitive features on the SH skulls include a maximum breadth near the base (vs. a maximum breadth at midparietal level in Neanderthals); a horizontal occipital torus that is well-developed centrally (versus the doublearched torus of Neanderthals, which is better developed laterally); a tendency for the rearmost point on the cranium to lie on the occipital torus (rather than on the occipital plate above the torus as in Neanderthals); a supraorbital torus that is double arched (as in Neanderthals and early H. sapiens) but that is very thick laterally (as in H. erectus); a large mastoid process; and an endocranial capacity that not only fell below the

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Cranium 4

Cranium 5

Cranium 6 midfacial projection

large mastoid process

cm

Atapuerca Sima de los Huesos interparietal (sagittal) keel

incipient suprainiac fossa

Neanderthal average of roughly 1,450 cc but also encompassed the entire range observed in other broadly contemporaneous European specimens (table 5.6). Two skulls also exhibit an upraised mound or keel of bone between the parietals that is otherwise common only in H. erectus. he SH mandibles and dentitions display a similar amalgam of primitive and derived features, but the mandibles and teeth are highly variable in size and shape. Like adult Neanderthal mandibles, the SH specimens all exhibit a marked gap (the retromolar space) between the rear edge of the third molar and the leading edge of the ascending ramus, but they vary in the extent to which they show the highly asymmetric sigmoid notch and large, posteriorly projecting coronoid process that mark most Neanderthal mandibles (ig. 5.23). he SH dentitions resemble those of the Neanderthals (and of other broadly coeval Europeans, insofar as these are known), but not those of H. erectus or H. sapiens in

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335 FIGURE 5.21. skulls 4–6 from the sima de los huesos (sh), atapuerca, spain (drawn from photos in arsuaga et al. [1997b], figs. 2, 4, 6). The skulls probably date from between 600 and 530 ka, and they combine characters that are primitive for the genus Homo with derived features that indicate a special relation to the classic Neanderthals, who inhabited Europe after 130 ka. The primitive features include a large mastoid process and a pentagonal shape in rear view (vs. the typical Neanderthal condition, in which the mastoid is very small and the skull is globular in rear view). The derived Neanderthal features include the pronounced forward projection of the face along the midline and the presence of an oval area of roughened or porous bone overlying the occipital torus. The roughened area anticipates the suprainiac fossa of the classic Neanderthals. Skull 6 contrasts somewhat with skulls 4 and 5 in its greater occipital rounding and with skull 5 in its more flattened face, but this is probably mainly because skull 6 came from a juvenile, whereas skulls 4 and 5 came from adults. However, the larger SH sample exhibits substantial variability in the expression of Neanderthal facial and occipital morphology. The sum suggests that Neanderthal features evolved piecemeal rather than as an integrated complex. This may mean that Neanderthal evolution owes more to gene drift than to natural selection.

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ChaP Ter f ive

double-arched supraorbital torus

FIGURE 5.22. facial view of skull 5 from atapuerca sh, spain (drawn by Kathryn Cruz-uribe from a photograph). sh 5 anticipates classic neanderthal skulls in its double-arched browridge and strong projection of the face along the midline (midfacial prognathism).

forward projection of the face along the midline

cm

Atapuerca SH 5 the large size of the irst incisor through the third premolar (I1 through P3) relative to the small size of the fourth premolar through the third molar (P4 through M3). In addition, the SH molars tend toward the taurodontism (enlarged pulp chamber and root fusion) that is common in the Neanderthals. However, the SH P4 through M3 are unique for their small average size, in which they resemble only recent humans. Twenty of the SH premolars and molars, representing at least ive individuals, exhibit interproximal grooves at the base of the crown that probably resulted from tooth picking. Similar grooves have been observed occasionally in all species of Homo, including living humans, but they may be particularly common in the Neanderthals. heir high frequency at Atapuerca SH may thus be added to other features in which the SH people anticipate the Neanderthals. Finally, the SH postcranial bones are notable mostly for the extreme robusticity that marks primitive Homo everywhere. he postcranium lacked obviously derived Neanderthal characters, except perhaps for the scapula, which resembled Neanderthal specimens in the tendency for a dorsal (as opposed to a ventral) sulcus or groove on the axillary margin (illustrated in ig. 6.15 in the next chapter). he pelvis, which is known from a nearly complete specimen and numerous fragments, lacked Neanderthal novelties (particularly the platelike pubic ramus described in the next chapter), and in its overall robusticity, including a thick buttress or pillar of cortical (outer compact) bone above the acetabulum, it more

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very large, posteriorly projecting coronoid process large, posteriorly projecting coronoid process

asymmetric sigmoid notch

retromolar space

highly asymmetric sigmoid notch

very long retromolar space

more vertical ascending ramus receding symphysis (no chin)

AT 505

receding symphysis (no chin)

AT 605

closely recalls pelves of much older H. ergaster (or African H. erectus) from Koobi Fora (East Turkana) and Olduvai Gorge—the same is true of a pelvic fragment from Arago, France, that may only slightly postdate Atapuerca SH, and fundamental ruggedness and a lack of speciic Neanderthal features mark other European lower limb bones from the same time frame, including multiple femoral fragments from SH, three femoral fragments from Arago, a femoral shat from Notarchirico, Italy, and a tibia shat from Boxgrove, England (references in table 5.5). he femora in particular have straight, lat shats, unlike Neanderthal femora, in which the shats tend to be rounded and strongly bowed from front to back (ig. 6.14 in the next chapter). he Atapuerca SH limb bones imply adult body masses up to 95 kg (209 lbs), beyond the known range of the Neanderthals or any other fossil people for whom body mass has been reasonably estimated. Coupled with the great breadth of the trunk (inferred from the breadth of the pelvis), large body mass may have been a physiological adaptation to cold. If so, in a general sense, if not in the speciics, the SH people may have anticipated the Neanderthals, for whom likely bodily adaptation to cold is described the next chapter. he large body mass of the SH people was not matched by especially large endocranial capacity (table 5.6), which might mean they were cognitively limited compared to their smallerbodied and larger-brained Neanderthal successors. In their mix of primitive features and Neanderthal novelties and in the variable expression of the mix, the SH skulls recall not only Reilingen and Swanscombe but also other well-known European fossils that probably date between 600 and 200 ka. he others encompass some skulls or skull fragments that are remarkably primitive in occipital or frontal form (especially at Bilzingsleben and Vértesszöllös) (ig. 5.24), some that are arguably Neanderthal-like in facial or occipital features (but not both)

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FIGURE 5.23. mandibles aT 505 and aT 605 from the sima de los huesos, atapuerca, spain (drawn by Kathryn Cruz-uribe from photos in rosas [1997], 329). The mandibles are both adult, but they differ conspicuously in size. They both show typical neanderthal features, including a prominent retromolar space, a large posteriorly projecting coronoid process, and an asymmetric sigmoid notch, but the larger mandible expresses these features more strongly. it also has a somewhat steeper mental symphysis (chin region) and a more vertical ascending ramus. The difference in size presumably reflects sexual dimorphism, and it shows that sex (size) may also influence the expression of phylogenetically diagnostic morphological features. AT = Atapuerca.

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ChaP Ter f ive Vértesszöllös

FIGURE 5.24. midline (midsagittal) sections through the occipital bones of some key fossils assigned here to Homo erectus (Pithecanthropus iv, Pithecanthropus ii, and Sinanthropus ii), early H. neanderthalensis (bilzingsleben, vértesszöllös,swanscombe, and steinheim), and later H. neanderthalensis (ehringsdorf, la Chapelle, la ferrassie, and Circeo 1) (redrawn after vl ek [1978], 244). note that with respect to occipital flexion (angulation between the occipital and nuchal planes), the Vértesszöllös and especially the Bilzingsleben fossils are more like H. erectus than they are like the Swanscombe, Steinheim, and later Neanderthal fossils.

Sinanthropus II Pithecanthropus II Pithecanthropus IV

Bilzingsleben occipital plane 0

inion = opithsocranion nuchal plane

Homo erectus Swanscombe

Steinheim

Ehringsdorf

?early Homo neanderthalensis La Chapelle

La Ferrassie

Circeo 1

O O

5 cm

O

O

O

opisthocranion

inion

early Homo neanderthalensis

Homo neanderthalensis

(at Arago, Petralona, and Steinheim) (ig. 5.25), and some that clearly anticipate the Neanderthals in key respects (especially at Swanscombe and Reilingen). Like-aged mandibles and teeth are similarly variable, and a Neanderthal-like retromolar space, for example, is apparent on one mandible from Arago (no. 2), absent in a second (no. 13), and only tenuously (incipiently) expressed on yet a third from the site of Montmaurin, France (ig. 5.26). he Mauer mandible adds to the variation, since it lacks a retromolar space in the strict sense, but it would have had one if the extraordinary breadth of the ascending ramus did not obscure a large Neanderthal-like gap behind the third molar. he SH fossils surely represent a single population, and similar variability in the broader set of European fossils might thus mean that Neanderthal morphology evolved as independent features ixed by random genetic drit rather than as an integrated complex driven by natural selection. Drit is particularly likely to afect small populations, and even under optimal conditions, pre-Neanderthal numbers were probably small. Repeated cold snaps periodically reduced them even more, and local extinction was probably commonplace, particularly where environmental change was most adverse. In this situation, groups that survived by luck, because they happened to occupy a favorable locale, would see their features increase in frequency by chance alone. A succession of fortuitous increases between 600 and 130 ka could have culminated in classic Neanderthal morphology, largely independent of natural selection.

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evoluTion of The Genus HO MO expanded parietal supraorbital torus

rounded occipital

Steinheim

canine fossa cm

expanded parietal

flat, receding frontal massive supraorbital torus

Arago 21 and 47 flat, receding frontal

upper (occipital) plate of the occipital

occipital torus

strong alveolar prognathism

massive supraorbital torus

lower (nuchal) plate of the occipital

incipient midfacial prognathism (no canine fossa)

Petralona

An accretional process, driven mainly by chance, could explain the lowfrequency persistence of primitive character states alongside their derived Neanderthal counterparts even ater 130 ka. Infrequent persistence of the primitive condition is known particularly for inner ear (semicircular canal) form and lower fourth premolar morphology. he demographic history of the Neanderthal lineage may always be sketchy, but the limited variability of classic Neanderthal mitochondrial DNA, addressed in chapter 7, points to a population bottleneck 300–250 ka, corresponding to (glacial) oxygen-isotope stage 8. Genetics could not detect bottlenecks before the last one, and the archaeology of the Neanderthal lineage is too sparse and weakly dated to provide

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339 FIGURE 5.25. fossil skulls from steinheim, Germany, arago, france, and Petralona, Greece (steinheim drawn by Kathryn Cruzuribe from originals by Janis Cirulis in howells [1967], 218; arago from casts and from a photo in day [1986b], 48; and Petralona from photos in stringer [1983], 73). none of the skulls is reliably dated, but each is probably between 500 and 250 ky old. The arago skull is a composite based on a face (arago 21) and a large parietal fragment (arago 47) that may not come from the same individual. if the combination is accepted, however, the arago braincase exhibited parietal expansion that might anticipate such expansion in the classic neanderthals, after 130 ka. in other respects, including its flat, receding frontal, massive supraorbital torus, and strong alveolar prognathism, the Arago skull more closely recalls those of Homo erectus. The Steinheim and Petralona skulls also foreshadow Neanderthal skulls, but in different, contrasting ways. Thus in Steinheim the resemblance is primarily in the expansion and rounding of the braincase. The face is flat with a canine fossa more in the manner of H. sapiens than of H. neanderthalensis. In Petralona the resemblance is primarily in the face, which anticipates those of Neanderthals in its incipient forward projection along the midline. The braincase is more like that of H. erectus, particularly in the angulation of the occipital. All three skulls are tentatively assigned here to the Neanderthal lineage.

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ChaP Ter f ive

FIGURE 5.26. fossil human mandible from montmaurin, france (redrawn by Kathryn Cruz-uribe after billy and vallois [1977], 280). associated animal remains and pollen suggest that it is between 186 and 130 ky old. it combines large size, the absence of a chin, and other features that are primitive for the genus Homo with molar taurodontism and an incipient retromolar space that probably imply a special relation to the classic neanderthals, after 130 ka.

ascending ramus

mental foramen

mandibular condyle

mandibular corpus or body mandibular symphysis

(incipient retromolar space)

0

5 cm

Montmaurin

mandibular symphysis (incipient retromolar space)

ascending ramus

mandibular corpus or body

(no chin)

independent conirmation. However, it has yet to reveal a Neanderthal or proto-Neanderthal site that unquestionably formed during a glacial maximum, only sites that formed under relatively mild conditions, particularly prior to stage 8 (discussion below; data in table 5.11). Full glacial impact on European environments was dramatic, and even the culturally advanced Upper Paleolithic hunter-gatherers who succeeded the Neanderthals were driven from much of Europe during the last glacial maximum, about 20 ka. heir numbers were reduced, and they survived mainly in refuges on the southeast and southwest. Determining whether drit truly dominated Neanderthal evolution will require a much larger sample of well-dated fossils scattered through time. However, if it should turn out that specimens like those from Arago, Vértesszöllös, and Bilzingsleben are as old or older than those from Atapuerca SH and that these in turn are older than those from Swanscombe, Steinheim, and Reilingen, one could argue that Neanderthal upper facial

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frontal keel broad, flattened, receding frontal thick, double-arched supraoribtal torus

thick cranial walls

Bodo

massive, forwardly projecting face

and mandibular morphology evolved before the typically Neanderthal occipital region and that the occipital region evolved before the characteristically large and globular Neanderthal vault. his became more or less universal only ater 130 ka. he more ancient African skulls—from Bodo (ig. 5.27), Lake Ndutu (ig. 5.28), Kabwe (ig. 5.29), and Elandsfontein (ig. 5.30)—exhibit no uniquely derived features of modern humans, but they equally lack any Neanderthal specializations. he younger skulls—from Irhoud (ig. 5.31), Singa (ig. 5.32), and Florisbad (ig. 5.33)—also lack Neanderthal novelties, and they anticipate the derived modern human tendency for the face to be tucked below the forepart of the brain (rather than mounted in front of it). Arguably the skull from Salé, which could be chronologically intermediate, is also morphologically transitional in its outwardly bulging (bossed) parietals (ig. 5.20) and in its weakly expressed occipital torus, but the occipital shows signs of pathology and its phylogenetic signiicance is therefore dubious. he Salé skull also exhibits an interparietal (sagittal) keel. An isolated right maxilla from Kabwe is much less robust than the maxilla on the famous complete skull, and unlike the maxilla on the complete skull, it exhibits a canine fossa. his could imply that the Kabwe people were as variable as those from the Sima de los Huesos, or it could record change through time toward the more modern condition. he stratigraphic relationships of the Kabwe bones are uncertain, but the specimens are known to have come from diferent parts of the cave. he small number of African limb bones generally resemble their European counterparts in their great robusticity. Only an ulna from Kapthurin, Kenya, is described as gracile in the manner of modern specimens. In contrast, a pelvis from Kabwe, Zambia, preserves the same

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FIGURE 5.27. The principal fossil skull from bodo, middle awash, ethiopia (drawn by Kathryn Cruz-uribe from photographs in rightmire [1996], 24–25). radiometric readings and associated fauna indicate it is probably about 600 ky old. in its massive, forwardly projecting face, thick supraorbital torus, flat, receding frontal, frontal keel, and thick cranial walls, the skull recalls those of Homo erectus. However, it is distinguished from H. erectus by its great frontal breadth, the double arching of its supraorbital torus, and its relatively large braincase. It is assigned here to early H. sapiens.

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ChaP Ter f ive

expanded parietals broad (receding) frontal

thick cranial walls

enlarged, vertical occipital scale of the occipital

massive face relatively small occipital torus

Ndutu

cm

FIGURE 5.28. fossil skull from deposits of seasonal lake ndutu near the western end of the main olduvai Gorge, northern Tanzania (drawn by Kathryn Cruz-uribe from photos of a reconstruction in Clarke [1990], 705, 713). The skull is thought to be about 400 ky old. like the Kabwe and elandsfontein skulls (in figs. 5.29 and 5.30), it combines primitive features, including a large, projecting supraorbital torus, a low receding frontal, and thick cranial walls, with more advanced features, including expanded parietals and a nonangulated (rounded) occipital. The occipital is also notable for the large size of the upper (occipital scale), which arises vertically (instead of sloping forward as in Homo erectus). Ndutu differs from the Kabwe and Elandsfontein skulls primarily in its smaller size (with an endocranial capacity of about 1,100 cc) and its weaker occipital torus. If Ndutu represents a female and the others represent males, the difference could reflect sexual dimorphism in the same fundamental population. All three skulls are tentatively assigned here to early Homo sapiens.

cortical buttress over the acetabulum found at Atapuerca SH and Arago and in specimens from much earlier Homo. Femoral fragments from Kabwe, a proximal femur from Berg Aukas, northern Namibia, and a femoral shat from Lainyamok, Kenya, are variable in morphological detail, but they tend to have very thick cortical bone by modern standards. A tibia from Kabwe also has thick cortical bone, but it is very long, implying that the distal leg (between the knee and ankle) was long, as in most living tropical or subtropical Africans. In contrast, the Neanderthals had short distal legs, like those of modern arctic peoples, whose short limbs help to reduce heat loss. he Cro-Magnons who succeeded the Neanderthals in Europe had tropical limb proportions, and this reinforces cranial evidence that they evolved in Africa. In general, body form says more about ecology than about phylogeny, but a larger limb bone sample from both Europe and Africa may nonetheless help to illuminate the divergence of early H. neanderthalensis from contemporaneous H. sapiens. In sum, the African fossil record is sparse between 600 ka and 200 ka, but the contrast with the European record is still clear. Together, the African and European fossils document an evolutionary separation that

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upper (occipital) plate of the occipital

receding frontal

Kabwe (Broken Hill) supraorbital torus

partially healed puncture

occipital torus lower (nuchal) plate of the occipital dental caries and abscessing

(no canine fossa)

dental caries and abscessing

by 130 ka produced the classic Neanderthals in Europe and near-modern or modern people in Africa. SOURCES (not including those in tables 5.4, 5.5, and 5.8): random genetic drit to explain the development of the Neanderthal lineage (Dean et al. 1998; Hublin 1998b, 2000; Stringer and Hublin 1999); persistence in Neanderthals of primitive inner ear form (Ponce de León and Zollikofer 1999) and lower fourth premolar morphology (Bailey 2002; Bailey and Lynch 2005); limited variability in Neanderthal mitochondrial DNA (Lalueza-Fox et al. 2005); contraction of European populations during the Last Glacial Maximum (Terberger and Street 2002); hypothetical chronological sequence for the appearance of Neanderthal features (Hublin 1996, 1998b); face positioned below the front of the braincase as a modern human novelty (Lieberman 1995)

Late Homo erectus or H. heidelbergensis in China? he next two chapters argue that fully modern Homo sapiens spread from Africa 50–40 ka to replace H. neanderthalensis in western Eurasia and late surviving H. erectus in the Far East. However, the replacement hypothesis is truly compelling only for western Eurasia with its relatively dense and well-dated fossil and archaeological records. It is much harder to sustain for the Far East, where the corresponding fossil record is much poorer and where the associated archaeological record is almost nonexistent. Within the Far East, replacement is most plausible for Southeastern Asia, or more precisely Java, where classic H. erectus fossils (from the Pucangan and Kabuh Beds) and later ones (from Sambungmacan and Ngandong) document a single evolving lineage that may have survived to 50 ka or later. It is much less persuasive for China, where the evolutionary

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FIGURE 5.29. fossil skull from Kabwe (formerly, broken hill), Zambia (drawn by Kathryn Cruz-uribe from a cast and photographs). Possibly associated animal remains suggest the skull may be about 400 ky old. in its massive face, thick browridges, flat, receding frontal, and relatively great basal breadth, the skull recalls those of Homo erectus. However, it is readily distinguished from H. erectus skulls by its large endocranial capacity (about 1,325 cc) and by the vertical orientation of the upper or occipital plate of the occipital bone above the torus. It is assigned here to early H. sapiens.

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ChaP Ter f ive parietal

FIGURE 5.30. fossil skullcap from elandsfontein (saldanha, or hopefield), South Africa (drawn by Kathryn Cruz-Uribe from a cast and photos). Faunal associations suggest an antiquity of about 600 ka. The skull has a massive supraorbital torus, a low, receding frontal, thick cranial walls, and other features that tend to characterize primitive Homo everywhere, together with expanded parietals, a relatively rounded occipital, and a broad frontal that suggest it is derived in the direction of H. sapiens. It is placed here in early Homo sapiens.

(receding) frontal

supratoral sulcus

occipital (upper) plate of the occipital

supraorbital torus

0 nuchal (lower) plate of the occipital

5 cm

Saldanha skull (broad) frontal

occipital parietal

supraorbital torus

connection between classic H. erectus fossils (especially from Lantian, Zhoukoudian, and Hexian) and later fossils is more equivocal. Table 5.9 lists the most informative later Chinese fossils, and igures 5.34–5.36 illustrate the most complete skulls, from Dali, Jinniushan, and Maba, respectively. Dating depends mostly on U-series analysis of associated animal bones or teeth, but the reliability of the results is questionable because the stratigraphic relationship to the human fossils is not always clear and because the dated specimens may have exchanged uranium with their burial environment. he most secure dates are probably those for Jinniushan, where a combination of U-series and ESR determinations on animal teeth place the skull and associated postcranial bones near 200 ka. If the dates are taken at face value, the fossils are all between roughly 230 and 100 ky old. Recall that African fossils of this antiquity foreshadow living humans, while European ones clearly anticipate the Neanderthals. he skulls in table 5.9 variably combine massive, uninterrupted browridges, keeled, lat, receding frontal bones, low vault heights or other primitive features that characterize H. erectus with larger, more rounded braincases, less massive faces, and other advanced features that mark

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long, low vault

FIGURE 5.31. right lateral and facial views of Jebel irhoud skull 1, morocco (drawn by Kathryn Cruzuribe from photographs). The skull is estimated to be between 190 and 90 ka old, yet it anticipates skulls of living people in its relatively convex frontal (forehead) and in its relatively short, flat face tucked beneath the forepart of the brain. It is less conspicuously modern in the thickness and projection of the supraorbital region and in the low height of the braincase relative to length and breadth. Conceivably, it represents a population that was near the ancestry of the nearmodern or early-modern people from Skhul and Qafzeh Caves, Israel.

moderately convex frontal

projecting supraorbital torus

cm

345

broad, flat non-protruding face, tucked in below the forepart of the brain

Jebel Irhoud 1

near-vertical frontal

Singa supraorbital torus thick both centrally and laterally

face below the forepart of the brain

cm

FIGURE 5.32. Left lateral view of the skull from Singa, Sudan (drawn by Kathryn Cruz-Uribe from a photograph in Stringer [1979], 77). The skull is estimated to date from 170–150 ka, yet it resembles skulls of living people in its highly convex frontal, in its high, short vault, and in the placement of the face beneath the forepart of the brain. Arguably only the thickness of the browridge, particularly at the sides, is a distinctly archaic feature.

H. sapiens. hey difer from both their African and their European contemporaries, but in their mix of archaic and derived features, they recall the older African and European fossils that are sometimes grouped in H. heidelbergensis. his could imply that H. heidelbergensis extended eastward to China (but not to Java), or it could imply that Chinese H. erectus

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ChaP Ter f ive

puncture from carnivore tooth relatively convex frontal

thick supraorbital region that grades imperceptibly into the frontal squama above

thick cranial walls

broad, massive, but relatiely flat face

cm

Florisbad

FIGURE 5.33. right lateral and facial views of the skull from florisbad, south africa (drawn by Kathryn Cruz-uribe from photos in Clarke [1985], 303). esr analysis of an associated molar suggests the skull is about 260 ky old. like most modern human skulls, florisbad lacks a true supraorbital torus (there is no supratoral sulcus or inflection point separating the supraorbital region from the frontal squama immediately above), and the frontal is relatively convex. However, the supraorbital region is heavily thickened, particularly at the sides, the face is very broad, and the cranial walls are very thick. Arguably when all features are considered, Florisbad is morphologically intermediate between older, more archaic African skulls like those from Bodo, Ethiopia (fig. 5.27), and Kabwe, Zambia (fig. 5.29), on the one hand, and later, more modern–looking skulls like the one from Singa (fig. 5.32) and other sites considered in the next chapter.

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and H. heidelbergensis evolved similar characters in parallel. Parallel evolution is tentatively favored here, partly because it is more parsimonious and partly because the Chinese archaeological record so far reveals no evidence for a population intrusion. In addition, the later Chinese fossils tend to resemble Chinese H. erectus in a handful of features, including a short maxilla, lat, horizontally oriented cheekbones, a very broad nasal bridge, strong shoveling of the upper incisors, and small third molars (M3s). If parallel evolution is accepted, then Chinese and Javan H. erectus followed divergent evolutionary pathways, and Chinese H. erectus might have to be removed to a separate species. Many new fossils, dates, and archaeological inds would be necessary to establish this, and fresh fossils and dates are also required to determine whether the Out-of-Africa scenario applies to China in the same way it does to Europe. SOURCES (not including those in table 5.9): survival of H. erectus on Java to 50 ka or later (Swisher et al. 1996); problematic U-series dates from Chinese sites (Chen and Yuan 1988); dates for Jinniushan (Chen et al. 1994); features of H. heidelbergensis on Chinese fossil skulls (Rightmire 1998)

Geographic Distribution Sites with artifacts that early people produced far outnumber sites that also contain their bones, and this section thus relies mainly on relevant archaeological sites to infer the geographic distribution of Homo ergaster and its immediate descendants. he next section examines the artifacts in detail, but it is important here to emphasize the broad distinction between ancient sites that contain hand axes and sites that don’t. Hand axes and related bifacial tools deine the Acheulean Industrial Tradition, which originated in Africa 1.7–1.6 Ma. H. ergaster almost certainly invented the Acheulean Tradition, and early H. sapiens and H. neanderthalensis continued it. H. erectus did not, and the next section centers on the evo-

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massive supraorbital torus

long, low vault

canine fossa

broad, short flat face

Dali

cm

mastoid process

lutionary implications of this basic artifactual divide. here is the complication, however, that not all populations of H. ergaster, early H. sapiens, and early H. neanderthalensis produced Acheulean tools, and the explanation remains speculative. he discussion examines diferent regions in rough order of their initial human colonization, beginning with Africa and proceeding to western Asia, eastern Asia, and Europe. Africa

Archaeological sites show that, unlike the australopiths or even Homo habilis, H. ergaster (or early H. erectus) was distributed throughout Africa, excepting only the extreme deserts of the north and south and the lowland rainforest of the Congo Basin. Archaeological discoveries indicate that about 1.5 Ma, shortly ater the emergence of H. ergaster, people more intensively occupied the drier peripheries of sedimentary basins on the loor of the eastern Rit Valley, and they colonized the Ethiopian high plateau (at 2,300–2,400 m) for the irst time. By 1 Ma, they had extended their range to the far northern and southern margins of Africa. Range extension northward inevitably led to dispersal out of Africa, probably mainly around the eastern end of the Mediterranean Sea and thence eastward to China and Indonesia and westward to Europe. If it is accepted that Eurasia was occupied before 1 Ma, then northern Africa must have been too, but the actual antiquity of north African occupation remains debatable. he oldest claim is for Aïn Hanech, Algeria, where large mammals and paleomagnetism suggest that an artifact assemblage that lacks hand axes or other bifacial artifacts accumulated within the Olduvai Paleomagnetic Subchron, between 1.95 and 1.77 Ma (references in table

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angulated occipital with a prominent transverse torus

FIGURE 5.34. The skull from dali County, shaanxi Province, northern China (drawn by Kathryn Cruzuribe from photos in Wu and Poirier [1995], 116). u-series analyses of associated animal bones indicate that the skull may be between 230 and 180 ky old. it shares many primitive features with Homo erectus, including a massive supraorbital torus, a flat, receding frontal, an angulated occipital with a prominent transverse torus, thick cranial walls, and a relatively small endocranial capacity (1,120 cc). However, it departs from classic H. erectus in its more expanded parietals and much more limited postorbital constriction. The face is flat with a canine fossa and without the midfacial projection that marks the Neanderthals. The skull is tentatively assigned here to late Chinese H. erectus.

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FIGURE 5.35. The skull from Jinniushan, near the city of Yinkou, liaoning Province, northeastern China (drawn by Kathryn Cruz-uribe from a photo in Wu and Poirier [1995], 121). esr and u-series dates on associated animal teeth suggest the skull may date from around 200 ka. it combines primitive features, including a large supraorbital torus, a flat, receding frontal, and a prominent occipital torus, with advanced features, including a relatively rounded braincase that is broadest well above the base, thin cranial walls, and a large endocranial capacity (1,390 cc). In its primitive-advanced mix, it broadly recalls 600–400-kyold African and European skulls that are sometimes assigned to Homo heidelbergensis. However, it is tentatively assigned here to late H. erectus. Part of the postcranial skeleton was associated, and it implies relatively large body mass, a broad trunk, and relatively short forearms that together suggest bodily adaptation to cold (Rosenberg et al. 2006).

ChaP Ter f ive

flat, receding frontal pronounced supraorbital torus broad nasal bridge

large, relatively well rounded braincase

broad, flat face

occipital torus shoveled incisor

cm

reduced M3

Jinniushan (Yinkou) 5.13). An alternative reading of the paleomagnetism and the mammals places the artifacts near 1.2 Ma. Mansourah (Constantine), also Algeria, has provided a broadly similar, non–hand ax assemblage that could be equally old or older, but fresh ieldwork is necessary to establish its stratigraphic relation to possibly associated mammalian fossils. Elsewhere in northern Africa, the oldest documented artifact occurrence is in homas Quarry 1 Level L near Casablanca, Atlantic Morocco, where reversed magnetism, tentative OSL readings, and mammal species suggest an age between 1.5 and 1 Ma for an assemblage that includes crude bifaces. SOURCES (not including those in table 5.13): early human colonization of the drier peripheries of the eastern Rit (Harris 1983), the Ethiopian high plateau (Clark and Kurashina 1979), far northern Africa (Raynal et al. 1995b, 2001), and far southern Africa (Klein 1994); Aïn Hanech at 1.95–1.77 Ma (Sahnouni et al. 2002, 2004) or 1.2 Ma (Geraads et al. 2004); Mansourah (Chaid-Saoudi et al. 2006)

Western Asia

If we assume that early human expansion from Africa occurred over land rather than water, adjacent western Asia should provide the oldest irm indication for an African exodus. Erk-el-Ahmar and ‘Ubeidiya in Israel and especially Dmanisi in the Republic of Georgia broadly conirm this expectation (references in table 5.12; locations in igs. 5.37 and 5.53.) Erk-el-Ahmar. Erk-el-Ahmar is the least secure early west Asian site be-

cause it has provided only a few Oldowan-style core tools and lakes, and the details of their recovery have not been described. hey lay in

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receding frontal massive supraorbital torus

broad nasal bridge

cm

Maba luvial deposits near the top of the Erk-el-Ahmar Formation, and paleomagnetic readings and the sedimentation rate calculated from a locality about 1.5 km away suggest the artifacts could date from more than 1.71 Ma. An age of this order is in keeping with fossil mollusks that place the Erk-el-Ahmar sediments before those at ‘Ubeidiya, discussed immediately below. If the dating is accepted and the Oldowan-like artifacts were in place, Erk-el-Ahmar might be the oldest known archaeological site outside of Africa, and it could even antedate the emergence of Homo ergaster. However, details on the stratigraphic context of the artifacts are necessary to evaluate their true antiquity. ‘Ubeidiya. ‘Ubeidiya is far more compelling than Erk-el-Ahmar, since it

was subject to large-scale, well-described excavations between 1959 and 1999. hese produced more than 3,400 early Acheulean or Developed Oldowan artifacts that come from eighty distinct archaeological horizons within luvial and lake-edge sediments of the ‘Ubeidiya Formation. Associated mammalian fossils imply the deposits formed in the interval between 1.4 and 1 Ma. his age estimate is consistent with the reversed magnetism of the sediments and radiopotassium determinations of about 0.79 Ma on a superimposed basalt. As at most other early Paleolithic openair sites, tool-marked bones are rare and the artifact-bone association may mean only that standing water attracted people and other animals, mostly on separate occasions. Human remains comprise three cranial fragments and three isolated teeth, but only one specimen (a heavily worn lower incisor) was unquestionably sealed in place. It has been tentatively assigned to Homo ergaster. he other mammals are mainly Eurasian (Palearctic), but some are African (Ethiopian), and the artifacts, including crude bifaces, closely resemble broadly contemporaneous pieces from Upper Bed II at Olduvai Gorge. ‘Ubeidiya may thus relect a slight ecological enlargement of Africa more than a true human dispersal. Israel is only technically outside Africa today, and it was repeatedly invaded by African species during the past 1–2 my. he invasions occurred mainly during

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FIGURE 5.36. a partial skull found in a limestone cave near the village of maba, Guandong Province, southern China (drawn by Kathryn Cruz-uribe from photos in Wu and Poirier [1995], 139). u-series analyses of associated animal bones indicate the skull may be between 140 and 119 ky old. it exhibits more postorbital constriction than either the dali or the Jinniushan skulls (figs. 5.34 and 5.35), but it broadly resembles them in its large supraorbital torus and receding frontal. With them, it is tentatively assigned here to late Chinese Homo erectus.

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ChaP Ter f ive

TABLE 5.9. Sites with fossils that could document late Homo erectus or H. heidelbergensis in China (Brown 2001; Wu & Poirier 1995; Wu & Wu 1985). Figure 5.4 locates the sites. he list excludes possibly contemporaneous fossils from Changyang, Dingcun, Miaohoushan, and Tongzi because they are isolated teeth or jaw fragments that are diicult to diagnose (Wu & Poirier 1995). It should arguably include a handful of fossils from the Korean Peninsula, but so far, these are too sparsely reported for independent evaluation (Norton 2000).

Geologic Age (and Basis for Estimate)

Site (and Date of Discovery)

Fossils

Dali (Jiefang Village), Shanxi Province (1978)

A partially distorted cranium with face

Jinniushan (Yinkou), Liaoning Province (1984)

A partial skeleton with skull, one ca. 200 ka (combined Ucervical vertebra, 5 thoracic vertebrae, series and ESR on associtwo let ribs, a let ulna, a complete let ated animal teeth) pelvic bone, and numerous articulated bones of both hands and feet

230–180 ka (U-series analysis of possibly associated animal teeth)

Maba (Mapa), Shizishan Cave, Guandong fragments of the frontal, parietals, and Province (1958) upper face of a single skull

135–129 ka (U-series analysis of associated animal teeth)

Xujiayao, Shanxi Province (1976–1979)

partial juvenile maxilla, 12 parietal fragments, 2 occipital fragments, a temporal bone, 2 isolated upper molars, and an isolated lower molar

125–104 ka (U-series analysis of associated animal teeth)

Chaoxian (Chaohu), Anhui Province (1982–83)

a fragmentary occipital and a partial maxilla

200–160 ka (U-series analysis of associated animal teeth)

interglacials, including the last one (between roughly 130 and 80 ka), when the African invaders included early-modern or near-modern humans. SOURCES (not including those in table 5.12): ancient expansions of African mammals to Israel (Tcher-

nov 1992)

Dmanisi. At Dmanisi, multiyear excavations into ancient luvial depos-

its have produced ive skulls (one unpublished), four mandibles (three linked to skulls), and at least sixteen isolated teeth and twenty-four postcranial bones. Some analysts believe the specimens represent a novel species, for which they have suggested the name Homo georgicus, but most specialists include them in Homo ergaster (or “African Homo erectus”). he excavations have also provided more than 1,000 Oldowan-like pebble tools and lakes and more than 3,000 animal bones that may antedate those at ‘Ubeidiya. Human expansion from Africa is indisputable, since Dmanisi is at 41°N, between the Greater and Lesser Caucasus Mountain Chains, roughly 1,500 km north of ‘Ubeidiya and Erk-el-Ahmar, and its fossil fauna is overwhelmingly Eurasian. he fossiliferous deposits overlie a basalt that has been dated by 40Ar/39Ar to about 1.85 Ma, within the Olduvai Normal Paleomagnetic Subchron. he Olduvai Subchron interrupted the Matuyama Reversed Chron between roughly 1.95 and 1.77 Ma, and the Dmanisi basalt exhibits the expected normal polarity.

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Swanscombe Boxgrove

Schöningen Bilzingsleben Mauer Vertesszöllös 0.6 1.1 Steinheim Dmanisi

St. Acheul Arago Atapuerca

Zhoukoudian Ceprano

Ambrona

Kuldara

1.7-1.2

Petralona

Ternifine Casablanca

Kocaba Mansourah & Aïn Hanech

Lantian

Gesher Benot Ya’aqov ‘Ubeidya

1.6-1.1

Nanjing

Hexian

1.6-1.1

Buia

1.8

Nihewan

Daka & Bodo Koobi Fora Nariokotome Olorgesailie Olduvai & L. Ndutu

Kabwe

Sangiran Mojokerto

Elandsfontein 0

2000 km

FIGURE 5.37. The approximate locations of key early archaeological or human fossil sites in africa, western asia, and europe, the most probable human dispersal routes from africa, and the dates in millions of years before present (ma) when people first inhabited different parts of Europe and Asia. The artifacts from ‘Ubeidiya, Israel, show that people had occupied southwestern Asia between 1.4 and 1 Ma. Human fossils and artifacts from Dmanisi, Republic of Georgia, are probably as old or older. Paleomagnetic readings on artifact-bearing deposits put people in north China between 1.66 and 1.1 Ma, and some human fossils in Indonesia may be equally as old. A human skull from Ceprano, Italy, and human fossils and artifacts from Atapuerca SE (layer TE 9), northern Spain, show that people penetrated Mediterranean Europe by 1.1 Ma, but they may have appeared to the north only 600–500 ka, and it was only after this time that they maintained a continuous presence. Dispersal into Europe probably occurred across the Dardanelles or Bosporus, the narrow straits that now separate Asian and European Turkey.

he surface of the basalt is fresh, indicating that it was covered by luvial deposits soon ater it formed, and the luvial deposits also show normal magnetization. his means that they probably also accumulated during the Olduvai Subchron, before 1.77 Ma. he Dmanisi H. ergaster fossils could thus be as old as any in Africa. However, the fossils and those of other animals occur in large hollows eroded within the normally magnetized deposits, and the hollows are illed with deposits that exhibit reversed magnetism (ig. 5.38). he fossils must then be younger than 1.77 Ma, and they could be as young as 1.19 Ma, when the Matuyama Reversed Chron was interrupted by the Cobb Mountain + Jaramillo Normal

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FIGURE 5.38. a schematic profile through the Dmanisi fossil and archaeological site, Republic of Georgia (adapted from Gabunia et al. [2001], fig. 3). 40Ar/39Ar dating and normal magnetism place a basal basalt and overlying alluvial deposits within the Olduvai Normal Subchron, between 1.95 and 1.77 Ma. Fossils and artifacts occur in deposits filling lens- or tunnel-like structures within the alluvium, and the reversed magnetism implies that the fills formed within the later part of the Matuyama Reversed Chron, between 1.77 and roughly 1.2 Ma. The fossils include three published human skulls that closely resemble specimens assigned to Homo ergaster (early African Homo erectus) and one that is more H. habilis–like. Accompanying mammal species may indicate an age nearer to 1.77 than to 1.2 Ma, but even at 1.2 Ma, the human fossils and artifacts would be among the oldest known outside Africa. Tooth marks, abundant carnivore bones, and coprolites suggest that hyenas or other carnivores probably accumulated most of the bones. People are unlikely to have played a major role since few if any of the bones exhibit unequivocal stonetool marks.

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ChaP Ter f ive

B2 (-) calcareous soil B2 (-)

no artifacts or bones

2m

B1 (-)

co

numerous artifacts & few bones (no human fossils)

calcium carbonate layer

(-) 1

ium

v llu

lens- and tunnel-like structures with occasional artifacts & numerous bones (including human ones)

A2 (+)

-

-

(-)

m viu

(-)

u all A1 (+)

0

Masavera Basalt (+) (1.85±0.01 mya) + normal magnetism - reversed magnetism

DMANISI SCHEMATIC PROFILE Subchrons, or even as young as 780 ka, when the Matuyama Chron gave way to the Brunhes Normal Chron. he Dmanisi fauna is argued to favor the earlier date, near 1.77 Ma, but it comprises a unique mix of taxa whose regional time ranges are poorly ixed, and an age near 1.19 Ma thus remains possible. he artifacts lack hand axes, which could mean that they antedate the appearance of hand axes in Africa at 1.7–1.6 Ma or that like many other Africans (and Europeans) ater 1.7–1.6 Ma, the Dmanisi people either did not make hand axes or did not leave them at every site they occupied. At the moment, the skull labeled Dmanisi (D) 2700/D2735 (ig. 5.39) probably presents the best case for an age near 1.77 Ma, since in its small endocranial capacity (about 600 cc), its relatively thin supraorbital torus, its well-rounded occipital (with no trace of an occipital torus), its diminutive face, and its facial proile (lacking an external nose), it recalls skull KNM-ER 1813 from northern Kenya. his is usually assigned to Homo habilis, which on present evidence is mainly, if not entirely, older than 1.65 Ma. A mandible (D211) difers from the type mandible of H. ergaster in some details (ig. 5.40), but the associated skull (D2282) and two other described skulls (D2280 and D3444/D3900) (igs. 5.41 and 5.42) closely resemble H. ergaster (or early H. erectus) counterparts from eastern Africa, and on this basis, they could all date to 1.7 Ma. However, they could also be younger, since the morphology that deines H. ergaster persisted in Africa until 1.4 Ma and perhaps later.

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0

10 cm

thin supraorbital torus well-rounded occiput lacking an occipital torus

facial profile lacking bony indications for an external nose

Dmanisi 2700 & 2735 Arguably, D2700 appears H. habilis–like in part because it represents a subadult, and the face in particular might have looked less primitive in adulthood. However, the face contrasts conspicuously with that of KNM-WT 15000, an H. ergaster juvenile who was dentally even younger, and the mix of morphologies at Dmanisi thus appears real. Whatever geologic date is preferred, Dmanisi then implies either that two human species expanded from Africa early on or that the deinition of H. ergaster must be enlarged to include skulls that might otherwise be assigned to H. habilis. Adult skullcap KNM-ER 42700, discovered at Ileret (Koobi Fora), northern Kenya, in 2000, may favor an enlarged deinition. KNMER 42700 dates to about 1.55 Ma, and it resembles east Asian H. erectus in its frontal and parietal keeling but recalls D2700 in its small endocranial capacity (691 cc) and weakly developed supraorbital torus. he inclusion of D2700 and KNM-ER 42700 in H. ergaster would buttress the argument that H. ergaster is more primitive than east Asian H. erectus and

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353 FIGURE 5.39. skull 2700 and mandible 2735 from dmanisi, Georgia (drawn by Kathryn Cruz-uribe from photographs). The skull and mandible were found separately but almost certainly represent the same individual. Ten isolated teeth from the maxilla and mandible occurred nearby. The third molars were in the process of erupting, indicating that the individual was not fully adult. in its small endocranial capacity (approximately 600 cc), its relatively thin supraorbital torus, its well-rounded occipital (lacking an occipital torus), its diminutive face, and its facial profile (lacking an external nose), the skull closely resembles skull KNM-ER 1813 (fig. 4.46), which is usually assigned to Homo habilis narrowly understood. The other described skulls from Dmanisi (pictured in figs. 5.41 and 5.42) more closely resemble those of Homo ergaster. The implication is either that two human species are represented at Dmanisi or that the definition of H. ergaster must be expanded to incorporate H. habilis in whole or in part.

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ChaP Ter f ive molars do not decrease in size from front to back

FIGURE 5.40. mandible Knm-er 922 from east Turkana, Kenya (drawn by Kathryn Cruz-uribe from photos in Wood [1991], 148) and mandible d211 from dmanisi, Georgia (redrawn after Gabunia and vekua [1995], 511). Knm-er 922 is reliably dated to about 1.6 ma and is the nomenclatural type for H. ergaster (Groves and mazák 1975). d211 may be as much as 1.77 ma old and it is linked to a skull (d2282) (fig. 5.41) that closely resembles skulls assigned to H. ergaster (or early H. erectus) in eastern Africa. The two mandibles differ subtly in the relative occlusal sizes of the molars and in the degree to which the mandibular symphysis is rounded, but if their shared assignment to H. ergaster (or H. erectus) is accepted, the differences have no taxonomic significance.

molars decrease in size from front to back

cm

more receding symphyseal contour

KNM-ER 992 (East Turkana)

more rounded symphyseal contour

D211 (Dmanisi)

mandibles of Homo ergaster (or early H. erectus) thus sensibly separated from it. Even if D2700 and KNM-ER 42700 are excluded from H. ergaster, the similarities between them further imply that D2700 antedates 1.5 Ma. SOURCES (not including those in table 5.12): Homo georgicus (de Lumley et al. 2006; de Lumley and Lordkipanidze 2006); possibility that D2700 would appear more H. ergaster–like if it came from an adult (Antón 2003); KNM-ER 42700 (Leakey et al. 2003; Spoor et al. 2007)

Eastern Asia

he timing of human arrival in eastern Asia is controversial and may long remain so, since a lack or rarity of evidence can never prove human absence. In addition, specialists disagree on what acceptable evidence is, and no one can deny that an indisputable site of very great antiquity might await discovery. As noted in the section above on dating, some authorities believe that people reached both China and Indonesia 1.8–1.7 Ma. Such an ancient arrival is appealing in part because it could explain why Far Eastern artifact assemblages always lacked hand axes and other

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supraorbital torus

flat, receding frontal

long, low braincase

forwardly projecting face and jaw

great breadth across the base of the skull

Dmanisi 2282

FIGURE 5.41. skulls d2280 and d2282 from dmanisi, Georgia (drawn by Kathryn Cruz-uribe from photographs in Gabunia et al. [2000b], figs. 2 and 3). The skulls resemble those of east Asian (classic) Homo erectus in their well-developed browridges overlying large orbits, in the tendency for the side walls to slope inward from a broad base, and in the long, low profile of the braincase. They fall within the range of Asian H. erectus in every measurable respect, but they are at the low end in the thickness of the skull walls and the browridges (both relatively thin) and in endocranial capacity (780 cc in 2282 and about 650 cc in 2280, compared to a mean of around 1,000 cc for classic Indonesian and Chinese H. erectus). The Dmanisi braincases also tend to be unusually narrow and high domed by H. erectus standards. In every respect in which the Dmanisi skulls depart from the classic east Asian norm, they recall east African skulls that have been dated between 1.8 and 1.5 Ma and that have been variously assigned to Homo ergaster or “African Homo erectus.” The Dmanisi skulls could be as much as 1.77 Ma old, in which case, they would imply that H. ergaster (or “African H. erectus”) spread from Africa only shortly after it appeared.

10 cm

0

Dmanisi 2280

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0

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5 cm

Dmanisi 3444/3900

FIGURE 5.42. dmanisi skull 3444 and associated mandible 3900 (drawn by Kathryn Cruz-uribe from photos in lordkipanidze et al. [2005], fig. 1; and [2006], figs. 4 and 5). The individual had lost all teeth before death, except for the left mandibular canine, and the sockets were totally resorbed, except those for the mandibular canines. Remodeling following the loss of the lower incisors probably explains the forward projection of the mandibular symphysis (“chin”). The skull is the oldest on record to show nearly complete tooth loss, and it may imply food provisioning and preparation by healthier members of a group. However, free-ranging great apes are known to survive substantial antemortem tooth loss without such help (DeGusta 2002, 2003).

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markers of the Acheulean Industrial Tradition, as discussed below. he Acheulean developed in Africa from the preceding Oldowan Tradition only 1.7–1.6 Ma, and if people penetrated eastern Asia at 1.7 Ma or before, they would have arrived without Acheulean tools. Human arrival in eastern Asia by 1.8 Ma might mean that the pioneer species was not Homo ergaster but its immediate ancestor. In this event, H. ergaster could even have evolved in eastern Asia and subsequently spread to Africa. his might explain why no African fossils unambiguously anticipate H. ergaster and why its emergence 1.8–1.7 Ma seems so abrupt. However, the lack of patent African predecessors could equally relect the relative poverty of the east African fossil record between 2.5 and 1.8 Ma. In addition, it is possible, perhaps even likely, that H. ergaster evolved in a sudden spurt when east African climate turned sharply drier and more seasonal about 1.8 Ma. he change promoted the initial development of typical African savanna grasslands to which H. ergaster seems to have been well-adapted. Finally, the problem remains that, so far, no Asian site unequivocally antedates the emergence of H. ergaster. he paleomagnetically dated sites in the Nihewan Basin, north China, discussed above and again below (references in table 5.2), come the closest, with estimated ages between 1.66 and 1.1 Ma, but even the younger limit will remain questionable until compelling sites of comparable antiquity are found elsewhere in eastern, southern, or central Asia. “Compelling” in this context means a site where the quality of the fossil or archaeological evidence and of the dating matches the standard that has been set at many African sites that date from at least 1.1 Ma.

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TABLE 5.10. European sites that have provided doubtful evidence for human presence before 600 ka. Figure 5.43 locates the sites approximately. For more extended discussion, see (Roebroeks 2001; Roebroeks & van Kolfschoten 1994; Villa 2001). At most of the sites, the key issue is the stratigraphic or cultural origin of the artifacts. In general, they are few. and in some cases, they were selected from a much larger number of nonartifactual stones. In these conditions, they could be geofacts or artifact mimics that natural rock impacts will occasionally produce (Bosinski 1995a). Such natural laking is now generally believed to explain artifacts of the “Kafuan” tradition that was once thought to precede the Oldowan in Africa (Clark 1958). It must also account for the occasional laked pieces known as “eoliths” (“dawn stones”) from early Tertiary deposits in Britain (Oakley 1959; Warren 1920).

Site

References

Le Vallonet (= Le Vallonnet) Cave, Nice, France

(de Lumley 1975; de Lumley 1976b; de Lumley et al. 1988; Eschassoux 1995; Terradillos Bernal & Moncel 2004)

Blassac-Les Battants, Massif Central, southeastern France

(Raynal et al. 1995a)

Chilac III, Massif Central, southeastern France

(Chavaillon 1991; Guth & Chavaillon 1985; Raynal et al. 1995a; Villa 1991)

Kärlich A and B alluvial sites, Neuwied Basin, west-central Germany

(Bosinski 1995b; Würges 1986)

Ca’Belvedere de Monte Poggiolo alluvial (or possibly colluvial) site, northeastern Italy

(Falguères 2003; Mussi 1995; Peretto 2001; Terradillos Bernal & Moncel 2004; Villa 2001)

Monte Peglia karst issure site, north-central Italy

(Mussi 1995; Piperno 1972; van der Meulen 1973)

Pirro Nord (Cava Pirro = Cava Dell’Erba) karst issures, southern Italy

(Arzarello et al. 2007)

Přezletice lakeside site, Prague, Czech Republic

(Fejfar 1976b; Fridrich 1976; Fridrich 1989; Fridrich 1991; Valoch 1976; Valoch 1982b; Valoch 1986; Valoch 1995b)

Stránská Skála cave sites, Brno, Czech Republic

(Fridrich 1976; Musil & Valoch 1969; Valoch 1976; Valoch 1986; Valoch 1991; Valoch 1995a)

Korolevo streamside site, southwestern Ukraine

(Gladiline & Sitlivy 1991; Valoch 1995b)

Whenever humans became established in eastern Asia, they were initially limited to middle and lower latitudes, even during interglacials, and archaeological sites at Majuangou, Donggutuo, Xiaochangliang, Maliang, and Cenjiawan in the Nihewan Basin (approximately 41°N) are the northernmost well-established occurrences. hey are only slightly north of Zhoukoudian (approximately 39.50°N), where fossil pollen suggests that Homo erectus was present only under interglacial conditions. North Asian (Siberian) artifacts that might be attributed to H. erectus either are not clearly artifactual or are almost certainly too young. SOURCES (not including those in table 5.2): possibility that H. ergaster evolved in eastern Asia (Dennell and Roebroeks 2005); drier, more seasonal climate in eastern (Trauth et al. 2005; Wynn 2004) and southeastern Africa (Hopley et al. 2007) beginning 1.8–1.7 Ma; pollen indicating that H. erectus occupied Zhoukoudian only during interglacial intervals (Aigner 1986); absence of evidence for H. erectus in Siberia (Yi and Clark 1983; and the summary of early Siberian prehistory in chap. 7)

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FIGURE 5.43. The approximate locations of european sites with early human fossils or artifacts. most of the sites postdate 600–500 ka, but some are possibly or probably much older. They include Korolevo in ukraine; stránská skála and P ezletice in the Czech republic; the lower horizons at Kärlich in Germany; Pakefield in England; Le Vallonet, Blassac-Les Battants, and Chilhac in France; Ceprano in Italy; and Atapuerca and Orce in Spain. However, only Atapuerca (the Gran Dolina and Sima del Elefante) provide compelling evidence for human presence before 600–500 ka. Parentheses mark sites older than 600 ka where evidence for human presence is either lacking or doubtful.

0o

10o

20o

30o

NOR TH SEA 50o

A Schöningen Bilzingsleben Weimar Ehringsdorf & (Süssenborn) (Prezletice) Mauer (Untermassfeld & Voigtstedt) (Les Battants, ATLANTIC Steinheim (Korolevo) (Stránská Skála) Chilhac & OCEAN Cannstatt (Deutsch-Altenburg) Fontéchevade Soleihac) (Sénèze) La Chaise Vértesszöllös Visogliano Combe Grenal Orgnac Montmaurin B l’Aubesier Atapuerca Arago Torralba & Ambrona Galeria Pesada Aridos Petralona (Orce) (Mosbach)

40o

MEDITERRANE

AN SEA

Pontnewydd A (West Runton) Grotte du Prince B High Lodge, Barnham Lazaret (Monte Poggiolo) & Beeches Pit (Le Vallonet) Pakefield Stanton-Harcourt Campitello & (Val d’Arno) Hoxne Terra Amata (Monte Peglia) Southfleet & Swanscombe Clacton Miesenheim Torre in Pietra, Saccopastore Baker’s Hole La Polledrara, (Tegelen) Fontana Ranuccio Kent’s Cavern Boxgrove Mesvin Castel di Guido, (Pirro Nord) Malagrotta, La Cotte Isernia Biache Casal de’Pazzi Abbeville, St. Altamura Cava Pompi Kärlich Acheul & Cagny Kartstein Menez-Dregan MaastrichtBelvédère Ariendorf & Tönchesberg

Ceprano Venosa-Notarchirico, Venosa-Loreto

Europe

It has long been assumed that people reached Europe as early as they reached the Far East, and Dmanisi, at the “gates of Europe” could be regarded as conirmation, even if it dates to no more 1 Ma. he irst edition of this book accepted an initial European penetration between 1 Ma and 780 ka, based on sites like those listed in table 5.10. Each such site had been dated by paleomagnetism, radiopotassium, ESR or luminescence determinations, associated mammal fossils, or some combination of these. However, critical assessment of these sites and most others that were supposed to be equally old subsequently suggested that few if any were valid. he problem was sometimes the dating, but more oten it was that the proposed artifacts could be naturally laked stones (especially at Blassac-Les Battants, Chilhac, Le Vallonet, Kärlich A, Přezletice, and Stránská Skála). he same review suggested that the oldest unquestion-

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able archaeological sites in Europe dated to no more than 500–600 ka. he oldest European human fossils appeared to be about the same age. hey included a tibia shat and two isolated lower incisors from Boxgrove, England, a mandible fragment and isolated teeth from Visogliano, Italy, and the mandible from Mauer, Germany, that is the nomenclatural type for Homo heidelbergensis (references in table 5.5). Rich well-known European paleontological sites like the Tegelen pits in the Netherlands, Untermassfeld, Voigtstedt, and Süssenborn in Germany, West Runton in England, Sénèze in France, Deutsch Altenburg in Austria, and the upper Val d’Arno basin in Italy, all of which formed between 1.5 Ma and 600–500 ka appeared to conirm that people were absent before 600–500 ka, since none provided artifacts or tool-marked bones. In contrast, paleontological sites that postdate 600–500 ka commonly provide both, and as discussed in the section on subsistence below, the issue then is not whether people were present but whether they killed or butchered the animals. he contrast between the evidence for people in Europe before and ater 600 ka recalled the contrast in evidence for human occupation of the Americas before and ater 12 ka. he rarity and poverty of American evidence older than 12 ka has convinced many archaeologists that people reached the Americas only about 12 ka (discussion in chap. 7). here is the diference, however, that known geographic or ecological obstacles could have prevented human expansion to the Americas before 12 ka, whereas it is hard to imagine what circumstances could have excluded people from Europe for 500 ky or more ater they had reached north China. One possibility is that before 600 ka people lacked the means to adapt to the dense deciduous forests that covered temperate Europe during interglacial peaks and to the grassy steppes or steppe-tundras that replaced the forests during glaciations. Another is that before 600–500 ka, people depended heavily on scavenging and they could not compete efectively with the lion-sized, short-faced hyena, Pachycrocuta brevirostris, that inhabited Europe. P. brevirostris had massive bone-crunching teeth, and it probably relied heavily on the large number of intact bones that saber-toothed cats let at kills. It coexisted with two saber-tooths— the jaguar-sized, dirk-toothed cat, Megantereon cultridens, and the lionsized, scimitar-toothed cat, Homotherium latidens—and it disappeared when the last of these did, about 500 ka. A third possibility is that people actually colonized Europe before 600 ka, but only sporadically, and the evidence will thus be sparse. Five sites currently support this case—Ceprano, Orce-Venta Micena, Atapuerca GD, Atapuerca SE, and Pakeield (references in table 5.11; locations in ig. 5.43). Ceprano, Orce-Venta Micena, Atapuerca GD, and Atapuerca SE are in southern Europe (south of the Alps and the Pyrenees), and they

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might mean that previous evidence for absence before 600 ka was really for absence only to the north, where archaeological and paleontological investigations have been most intense. However, Pakeield is in southeastern England, and it raises the possibility that people appeared equally early north and south. Each site is described in greater detail immediately below, but in advance, it can be said that only Atapuerca GD and SE are truly compelling. he other sites take their credibility partly from them, since they are unlikely to have been alone.

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Pakefield, Southeastern England.  At Pakeield, archaeologists have ex-

tracted thirty-one lint lakes (one crudely retouched) and a lint core from river channel deposits of the Cromer Forest-bed Formation. he Forest-bed Formation is exposed discontinuously along more than 80 km of the North Sea Coast, and since the mid-nineteenth century, it has yielded abundant mammalian bones, insect remains, and plant fossils that imply mild interglacial conditions. he artifact-bearing channel ill repeats the pattern, and its sediments, fauna, and lora together relect a warm Mediterranean-like climate. he Forest-bed Formation is now thought to span four-to-six successive temperate interglacial phases, but on stratigraphic grounds, the Pakeield channel ill probably accumulated during an earlier one, either global oxygen-isotope stage 17, between about 712 and 659 ka, or perhaps stage 19, between 787 and 760 ka. Depending on the choice, the Pakeield site would be 150–300 ky older than the oldest widely accepted north European archaeological sites, all of which are assigned to isotope stage 13 (table 5.11). Water vole bones underscore the greater antiquity of Pakeield, since it has supplied only Mimomys (with rooted molars), while stage 13 sites provide only the descendant genus Arvicola (with rootless molars). he Mimomys-Arvicola transition is well-ixed across Europe between 600 and 500 ka. he cultural origin of the Pakeield artifacts and their association with Mimomys have convinced a leading skeptic that humans penetrated northwestern Europe before 600 ka, at least on those occasions when climatic conditions were particularly favorable. However, the artifacts do not originate from a single horizon or from a paleosurface like those represented at later open-air sites such as Boxgrove (England) or Ambrona and Torralba (Spain) that postdate 600–500 ka, or even from a single locality, and the circumstances of their extraction have not been speciied. he problem also remains that other broadly contemporaneous Cromer Forest-bed sites, especially West Runton, have failed to demonstrate human presence, despite many decades of investigation. In that sense, the Cromer Forest-bed Formation as a whole continues to recall broadly contemporaneous paleontological sites elsewhere in northern Europe. he Pakeield artifacts may now stimulate searches that will uncover a Boxgrove- or Ambrona-like occurrence where artifacts and bones are

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he main sites that document human presence in Europe before global isotope Stage 5. Boldface and italics mark sites with human remains. Figure 5.43 locates the sites approximately. he Orce-Venta Micena sites are included for the sake of convenience, although the text suggests they provide only questionable evidence for human presence.

TABLE 5.11.

Site

References

STAGE 6 (glacial) (186–130 ka) La Cotte de St. Brelade, Jersey

(Callow 1986; Callow & Cornford 1986; Scott 1980; Scott 1986a; Scott 1986b; Scott 1989)

Biache-Saint-Vaast, France

(Aitken et al. 1986; Sommé et al. 1986; Tufreau 1988; Tufreau 1989)

Fontéchevade, France

(Henri-Martin 1965; Stringer et al. 1984; Vandermeersch et al. 1976)

La Chaise-du-Vouthon (l’Abri Suard), France (Blackwell et al. 1983; Debénath 1976; Debénath 1977; Debénath 1988; Matilla 2005; Piveteau & Condemi 1988; Stringer et al. 1984; Teilhol 2003) Combe Grenal layers 63–56, France

(Bordes 1972; Delpech & Prat 1995)

Montmaurin (La Niche), France

(Billy 1982; Billy & Vallois 1977; Cook et al. 1982; Stringer et al. 1984)

Lazaret Cave, France

(de Lumley 1969b; de Lumley 1975; de Lumley & Boone 1976; de Lumley et al. 2004; Michel et al. 1997; Michel et al. 2000; Rousseau et al. 2005; Valensi 2000; Valensi & Psathi 2004)

Ariendorf 2, Germany

(Bosinski 1986; Conard & Prindiville 2000; Gaudzinski et al. 2005; Turner 1986)

Tönchesberg 1A, Germany

(Conard 1992; Conard & Prindiville 2000)

STAGE 7 (interglacial) (245–186 ka) Torre in Pietra, Italy (upper horizons)

(Caloi et al. 1998; Malatesta et al. 1978; Mussi 1995; Mussi 2002)

Rebibbia-Casal de’Pazzi, Italy

(Anzidei & Cerilli 2001; Bietti 1985; Manzi et al. 1990)

Pontnewydd Cave, Wales

(Green 1981; Green 1984; Green et al. 1989; Green et al. 1981)

Stanton-Harcourt, England

(Buckingham et al. 1996; Scott & Buckingham 2001; Zhou et al. 1997)

Baker’s Hole, England

(Bridgland et al. 2004; Roe 1981; Wymer 1968)

Bau de l’Aubesier, France

(Lebel 2002; Lebel et al. 2001)

Maastricht-Belvédère, Netherlands

(Roebroeks 1986; Roebroeks et al. 1992b; Roebroeks et al. 1993)

Weimar-Ehringsdorf, Germany

(Blackwell et al. 1983; Bridgland et al. 2004; Cook et al. 1982; Mallick & Frank 2002)

Stuttgart-Bad Cannstatt, Germany

(Bosinski 1995b; Vollbrecht 1995; Wagner 1984; Wagner 1990)

STAGE 8 (glacial) (301–242 ka) Ariendorf 1, Germany

(Bosinski 1986; Turner 1986)

Mesvin IV, Belgium

(Cahen & Michel 1986)

STAGE 9 (interglacial) (334–301 ka) Atapuerca TD (Trinchera Galeria), Spain

(Carbonell et al. 2001; Falguères et al. 2001)

Torre in Pietra, Italy (lower beds)

(Caloi et al. 1998; Malatesta et al. 1978; Mussi 1995; Mussi 2002)

La Polledrara di Cecanibbio, Italy

(Anzidei 1995; Anzidei & Cerilli 2001; Caloi et al. 1998; Mussi 1995)

Castel di Guido, Italy

(Boschian & Radmilli 1995; Caloi et al. 1998; Mallegni & Radmilli 1988; Mariani-Costantini et al. 2001; Mussi 1995; Mussi 2002)

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TABLE 5.11. (continued)

Site

References

Malagrotta, Italy

(Caloi et al. 1998; Mussi 2002; Villa 1991; Villa & Soressi 2000)

Cagny-l’Épinette, France

(Bahain et al. 2007; Dibble et al. 1997; Tufreau & Antoine 1995; Tufreau et al. 1997)

Orgnac 3, France

(Combier 1971; Cook et al. 1982; Moncel et al. 2005)

Terra Amata, France

(de Lumley 1969a; Villa 1983; Villa 1991)

Hoxne, England

(Singer et al. 1993; Singer & Wymer 1976)

Steinheim, Germany

(Adam 1985; Bosinski 1995b; Howell 1960; Stringer et al. 1984)

STAGE 11 (interglacial) (427–364 ka) Aridos, Spain

(Santonja et al. 2001; Santonja & Villa 1990; Villa 1990)

Ambrona and Torralba, Spain

(Butzer 1965; Falguères et al. 2006; Freeman 1994; Howell 1966; Howell et al. 1996 (1991); Santonja et al. 1997; Santonja & Villa 2006; Soto et al. 2001; Villa et al. 2005b)

Visogliano, Italy

(Abbazzi et al. 2000; Cattani et al. 1991; Falguères 2003)

Fontana Ranuccio, Italy

(Bidduttu et al. 1979; Bidduttu & Celletti 2001; Mussi 1995; Segre & Ascenzi 1982)

Cava Pompi, Italy

(Bidduttu & Celletti 2001; Mussi 1995)

Venosa-Notarchirico, Italy

(Belli et al. 1991; Condemi 1991; Mussi 1995; Villa 2001)

Menez-Dregan 1, France

(Mercier et al. 2004; Monnier et al. 1994)

Rue de Cagny (Saint-Acheul), Cagnyla-Garenne, and Cagny-Cimitière, France

(Bahain et al. 2007; Bourdier 1976; Commont 1908; Haesaerts & Dupuis 1986; Tufreau 1978; Tufreau & Antoine 1995; Tufreau et al. 1997; Villa 1991)

La Caune de l’Arago (Tautavel), France

(de Lumley 1975; de Lumley 1979; de Lumley et al. 1984; Falguères et al. 2004; Stringer et al. 1984)

Swanscombe, England

(Conway et al. 1996; Roberts et al. 1995; Roe 1981; Stringer 1996b; Stringer & Hublin 1999; Stringer et al. 1984; Waechter 1973; Wymer 1964; Wymer 1988)

Beeches Pit, England

(Preece et al. 2006; Preece et al. 2007; Roebroeks 2007)

Barnham, England

(Ashton et al. 1994; Preece & Penkman 2005; Roberts et al. 1995)

Clacton-on-Sea, England

(Bridgeland et al. 1999; Collins 1969; Oakley et al. 1977; Ohel 1977; Ohel 1979; Singer et al. 1973; Warren 1911; Wymer 1968; Wymer 1988)

Kärlich-Seeufer, Germany

(Bosinksi 2006; Bosinski 1995b; Gaudzinski et al. 1996)

Bilzingsleben, Germany

(Bridgland et al. 2004; Mania 1986; Mania 1991; Mania 1998; Mania et al. 1994; Mania & Vlček 1981; Mania 1995)

Schöningen, Germany

(Dennell 1997; Roebroeks 2001; hieme 1997; hieme 1998)

Vértesszöllös, Hungary

(Dobosi 1988; Kretzoi & Dobosi 1990; Schwarcz & Latham 1984; Valoch 1995b; Vértes 1965; Vértes 1975)

STAGE 13 (interglacial) (528–474 ka)

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TABLE 5.11. (continued)

Site

References

Kent’s Cavern, England

(Campbell & Sampson 1971; Proctor et al. 2005; Roberts et al. 1995; Roe 1981)

Boxgrove, England

(Roberts 1986; Roberts et al. 1995; Roberts & Paritt 1999; Roberts et al. 1994a; Stringer et al. 1998)

High Lodge, England

(Ashton et al. 1992; Cook & Ashton 1991; Roberts et al. 1995)

Mauer, Germany

(Bosinski 1995b; Cook et al. 1982; Howell 1960; Roebroeks & van Kolfschoten 1994; Schoetensack 1994 (1908))

Kärlich G and Miesenheim 1, Germany

(Bosinksi 2006; Bosinski 1995b; Turner 1995b; van Kolfschoten 1990)

Kartstein, Germany

(Bosinski 1995b)

STAGE 15 (interglacial) (621–568 ka) Abbeville, Carpentier Quarry

(Breuil 1939; Howell 1966; Tufreau & Antoine 1995)

Isernia La Pineta, Italy

(Anconetani et al. 1994; Coltorti et al. 1982; Coltorti et al. 2005; Coltorti et al. 1983; Peretto 1991; Peretto 1994; Sardella et al. 2006; Villa 1996; Villa 2001)

Venosa-Loreto, Italy

(Mussi 1995)

Atapuerca SH (= Sima de los Huesos), Spain

(Arsuaga et al. 1996; Arsuaga et al. 1997a; Bischof et al. 2003; Bischof et al. 2007; Cuenca-Bescós et al. 1997; García & Arsuaga 1999; García et al. 1997)

Atapuerca GD (= Gran Dolina) layer TD-8, Spain

(Bischof et al. 2007; Falguères 2003; Falguères et al. 1999)

STAGE 17 (interglacial) (712–659 ka) Pakeield, England

(Paritt et al. 2005; Roebroeks 2005; Roebroeks 2006; Stringer 2006, pp. 46–51)

STAGE 19 (interglacial) (787–760 ka) Atapuerca GD (= Gran Dolina) layers TD4TD6, Spain

(Arsuaga et al. 1999b; Bermúdez de Castro et al. 1997a; Bermúdez de Castro et al. 1999; Bermúdez de Castro et al. 2004; Bermúdez de Castro et al. 2003; Carbonell et al. 2005; Carbonell et al. 1995; Carbonell et al. 1999; Carbonell & Rodríguez 1994; Cuenca-Bescós et al. 1999; Falguères 2003; Falguères et al. 1999; Lorenzo et al. 1999; Parés & Pérez-González 1995; Parés & Pérez-González 1999; Terradillos Bernal & Moncel 2004)

Ceprano, Italy

(Ascenzi et al. 1995; Ascenzi et al. 1996; Ascenzi et al. 2000; Bruner & Manzi 2005; Bruner & Manzi 2007; Clarke 2000; Mallegni et al. 2003; Manzi 2004; Manzi et al. 2001)

STAGE UNCERTAIN, OLDER THAN 1 Ma Orce-Venta Micena, Spain

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(Gibert et al. 1994a; Gibert et al. 1998; Gibert et al. 1991; Gibert et al. 1989; Gibert et al. 1994b; Gibert et al. 2002; Martínez Navarro et al. 1997; Oms et al. 2000; Roe 1995; Santonja & Villa 2006; Terradillos Bernal & Moncel 2004; Tixier et al. 1995; Turq et al. 1996)

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flat, receding frontal (”forehead”)

relatively small braincase with thick walls

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Ceprano

occiput sharply angulated (not rounded)

FIGURE 5.44. lateral and anterior views of the reconstructed fossil skullcap from Ceprano, italy (drawn by Kathryn Cruz-uribe from photos in Clarke [2000], figs. 2 and 10). Compared to skullcaps of classic east Asian Homo erectus, the Ceprano specimen is relatively large, broad, and short, and it lacks a sagittal keel. However, it resembles skullcaps of H. erectus in its massive, shelflike browridge, its relatively thick walls, and its sharply angled occiput when viewed from the side. It has been provisionally dated to 900–800 ka, and if it had been found in Java, it might have been assigned to H. erectus. If the dating is correct, the anatomical contrast with roughly contemporaneous fossils from the Gran Dolina, Atapuerca, may indicate that there were at least two failed human attempts to colonize Europe before 600 ka.

massive, continuous brow ridge thickening above the orbit

0

5 cm

stratiied together. Alternatively, close examination may show that some of the Forest-bed large mammal bones are stone-tool-marked. In either case, the argument for 700-ky-old human penetration of northwestern Europe would be signiicantly strengthened. Ceprano, Central Italy. At Ceprano, bulldozing of ancient lake deposits

exposed a human skullcap. Artifacts and animal bones were lacking, but the deposits are bracketed between 1 Ma and 700 ka by 40Ar/39Ar readings on volcanic particles from underlying sands and by radiopotassium determinations on particles from an overlying sandy gravel. he nearest outcrop of the underlying sands is 25 km away, but if the stratigraphy has been accurately assessed, the skullcap probably dates from 900–800 ka. It resembles African and European skulls commonly assigned to Homo heidelbergensis and also skulls of east Asian H. erectus. Features shared with H. heidelbergensis include a relatively short, broad braincase, a relatively large endocranial capacity of about 1,150 cc, the absence of a sagittal keel, limited postorbital constriction, and the shape of the supraorbital torus, which thickens conspicuously above the midline of each orbit (ig. 5.44). H. erectus–like features include an occipital torus that is equally developed at the sides and in the center, a sharply angled occiput on which the lower (nuchal) squama or plane is oriented more to the back and less downward than in most European skulls postdating 600–500 ka, and remarkably thick cranial walls. he frontal and occipital bring to mind fragments of the same bones from Bilzingsleben, Germany. Among all the European fossils that postdate 500 ka, the Bilzingsleben specimens are perhaps the hardest to relate to the Neanderthal

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lineage. Based on its feature mix and proposed geologic age, the Ceprano skull might be viewed as a link between earlier H. ergaster (“African H. erectus”) and later African and European H. heidelbergensis. Orce-Venta Micena, Gaudix-Baza Basin, Southeastern Spain.  he Guadix-

Baza Basin near the city of Orce contains about 100 m of ancient lake sediments with three proposed paleoanthropological sites—Venta Micena, Fuente Nueva 3, and Barranco Léon. Based on paleomagnetism and fossil mammals, Venta Micena has been dated to about 1.6 Ma, while Fuente Nueva 3 and Barranco Léon have been bracketed between 1.3 and 1.2 Ma. he Venta Micena mammal bones come partly from African carnivores and ungulates, and they could signal a 1.6 Ma Out-of-Africa movement that included people. Four possible human bones—a cranial fragment and three partial long bone shats—provide support. However, to the extent the bones can be diagnosed, they are more likely to represent young ungulates. Since Venta Micena has also produced neither artifacts nor bones that were clearly damaged by artifacts, the site should probably join the list of rich paleontological localities that fail to indicate human presence in Europe before 500 ka. he mammal bones at Fuente Nueva 3 and Barranco Léon include no supposed human specimens, but they occur within and below deposits that contain possible stone artifacts. hese range from pieces that appear crudely, if humanly laked to more abundant ones on which the fracturing is less obviously artifactual. he published reports do not specify which pieces lay on or near the surface and which occurred at depth, and the origin and integrity of the assemblages are thus open to question. he assemblages may include the oldest known artifacts in Europe, or the portions that are truly old may not be artifactual. Atapuerca GD (Gran Dolina) and Atapuerca SE (Sima del Elefante), Burgos, Northern Spain.  At Atapuerca GD, a railway trench intersected a

sinkhole inilling in the early 1900s and exposed 18 m of sediment, divided among eleven layers. hese are numbered 1–11 from bottom to top, and the numbers are usually preceded by the letters TD, for Trinchera (trench) and Dolina (depression or hollow). he oldest artifacts occur in layer TD 4, but TD 6 is presently the most important. It has produced more than one hundred laked stones and roughly one hundred fragments of human bone from a subunit named the Aurora Stratum for a key excavator. he coarse sediments are not ideal for paleomagnetism, but widely accepted paleomagnetic readings place the transition from the Matuyama Reversed Chron to the Brunhes Normal Chron within TD 7, about 1 m above the Aurora Stratum. Constrained by the

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FIGURE 5.45. a juvenile maxilla (aTd6-69) from the Gran dolina site, atapuerca, spain (drawn by Kathryn Cruz-uribe from a photograph in bermúdez de Castro et al. [1997a]). The maxilla is readily distinguishable from those of juvenile neanderthals by the presence of a canine fossa and by the absence of strong forward projection along the midline. it has been dated to about 800 ka and it could represent the last shared ancestor of neanderthals and living humans. more securely, together with other human fossils from the same layer, it shows that people at least sporadically penetrated europe before 600–500 ka.

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canine fossa

0

5 cm

paleomagnetic result, ESR and U-series analyses on horse and rhinoceros teeth bracket TD 6 between 857 and 780 ka. Fossil pollen implies moist, mild climatic conditions that must then correspond either to oxygenisotope stage 21 or to stage 19. Considering all the evidence, the excavators propose an age of roughly 800 ka. Only the relatively evolved state of the ancestral water vole, Mimomys, may imply a somewhat younger age, perhaps closer to 600 ka. However, the association of Mimomys with human remains still makes Atapuerca GD the oldest fully documented human fossil site in Europe. he TD 6 artifacts that have been described so far comprise 268 crudely laked pebbles and simple nondescript lakes that closely resemble yet older artifacts in layer TD 4 below. he raw materials are mainly lint, quartzite, and sandstone that had to be transported to the site, and some pieces show signs of use in butchery or woodworking. Still, based strictly on form, they might be dismissed as geofacts (naturally laked stones), and it is the human remains that incontrovertibly conirm human presence. hey represent at least seven individuals, and the craniofacial fragments are striking for derived features that diferentiate them from H. ergaster–H. erectus and that may suggest a special relationship to H. sapiens. Particularly signiicant are a fragmentary juvenile maxilla and a young adult half-mandible. he maxilla (ATD6-69) (ig. 5.45) suggests that the midface did not protrude strongly forward (particularly along the midline as in Neanderthals) and it exhibits a deep hollowing (canine fossa) between the nasal aperture and the cheek bone (this region is inlated, not hollowed in Neanderthals). he same hollowing is conspicuous, if less marked on an adult maxillary fragment (ATD6-58). he mandible (ATD6-96) is remarkably small and slender, and like the

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maxillary fragments, it exhibits no features that anticipate H. neanderthalensis. Among other fossil mandibles to which it has been compared, it actually appears most like those of Zhoukoudian H. erectus. he TD 6 fossils have been assigned to a new species, Homo antecessor, on the grounds that they exhibit a unique combination of characters. If it is accepted that the facial features imply an especially close relationship to H. sapiens, H. antecessor must have become extinct by 600–500 ka when H. neanderthalensis (or H. heidelbergensis) appeared in Europe. Additional, more complete fossils are required to establish its actual phylogenetic relationships and also to determine if it might be represented at Ceprano. If not, TD 6 and Ceprano would provide evidence for two separate, ultimately unsuccessful early human movements from Africa to Europe. he same railway trench that exposes GD exposes a similar occurrence at Atapuerca SE, approximately 180 m to the southwest. he SE deposits are about 25 m thick, and since the text for this book was completed, the excavators have reported artifacts and a partial human mandible dated by paleomagnetism, associated mammalian fossils, and cosmogenic nuclides to 1.2–1.1 Ma. Atapuerca SE thus conirms that humans penetrated Europe at least occasionally long before 600 ka. SOURCES (also table 5.11): critical assessment of European sites older than 600–500 ka (Roebroeks 2001; Roebroeks and van Kolfschoten 1994; Villa 2001); possible early human inability to adapt to European environments (Roebroeks et al. 1992a) or to compete with the short-faced hyena (Palmqvist and Arribas 2001; Turner 1992; Turner and Anton 1997); Mimomys-Arvicola transition (van Kolfschoten 1993, 1998); acceptance of Pakeield by a skeptic (Roebroeks 2005); mammalian indications of a 1.6-my-old Out-of-Africa event (Martínez Navarro and Palmqvist 1995; Palmqvist 1997; Palmqvist and Arribas 2001); Venta Micena human bones as possible ungulate bones (Moyà-Solà and Köhler 1997; Palmqvist et al. 2005); Gran Dolina Mimomys (van Kolfschoten 1998); Sima del Elefante (Carbonell et al. 2008; Parés et al. 2006; Rosas et al. 2001)

The Route from Africa to Europe. he European archaeological sites that

probably or possibly antedate 600 ka, including Atapuerca GD, the Orce localities, and Pakeield, all lack hand axes or other markers of the Acheulean Tradition. heir artifact assemblages are sometimes described as Oldowan-like, but they are unlikely to derive directly from the Oldowan, which gave way to the Acheulean in Africa 800 ky to 1 my earlier. he source of the earliest European artifacts must still be African, but the artifacts themselves are too amorphous to infer a more speciic origin or route of dispersal. he origins of Acheulean artifacts in Europe, 700–600 ka, are less obscure. Acheulean assemblages in northern Africa and southern Europe are broadly similar, and this could imply dispersal across the Straits of Gibraltar or even the central Mediterranean from Tunisia to Sicily and thence to Italy. However, no other large mammal is known to have crossed the Mediterranean in the same interval, probably because

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substantial stretches of open water remained even when glacier growth depressed sea level. Successive large mammal faunas in Sicily show no connection to Africa. Instead, they imply repeated invasions from Italy when sea level was low, interspersed with long periods of isolation when sea level was higher. People may have occupied Sicily only ater 40 ka. he sum suggests that human movements from Africa to Europe occurred around the eastern shore of the Mediterranean to Anatolia and from there to southeastern Europe across the Dardanelles or Bosporus, which were oten dry during glacial periods. he lakeside Acheulean site at Gesher Benot Ya’aqov (GBY), Israel (references in table 5.12), may record a movement that paralleled the Mediterranean coast and that could have produced, irst, the still sparsely known Acheulean of Anatolia and, later, the much better known Acheulean of Europe. he GBY sequence is 34 m thick, and it records a shit from reversed to normal magnetic polarity. Together with U-series and ESR determinations on animal teeth, the paleomagnetism ixes GBY between roughly 800 and 700 ka. he animal remains include the oldest well-established Eurasian record of the straight-tusked woodland elephant, Elephas (Palaeoloxodon) antiquus, which evolved from E. (Palaeoloxodon) recki in Africa before 1 Ma. By 600 ka, E. antiquus or its descendants were distributed from Spain to China, and its bones consistently characterize the earliest European Acheulean sites, dated to between 600 and 300 ka. he GBY artifacts are striking for their similarity to contemporaneous African ones in form and in mode of manufacture. Like their African contemporaries, the GBY Acheuleans preferentially chose volcanic rock for biface manufacture, and the bifaces and associated tools they produced have a distinctively African lavor. Perhaps most telling are large basalt lakes that have a bulb of percussion on both surfaces. Similar pieces are otherwise common only in eastern and northern Africa, where they are oten called Kombewa lakes. At the African sites and at GBY, Kombewa lakes were oten used to produce the guillotine-like bifaces known as cleavers, and they place GBY irmly within the African Acheulean tradition. It remains possible, however, that GBY represents not so much an African exodus as another in a long series of episodes when Africa expanded ecologically to incorporate its southwest Asian periphery.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES (not including those in table 5.12): possible dispersal of the Acheulean across the Mediterranean (Alimen 1975; Santonja and Villa 2006); Pleistocene prehistory of Sicily (Villa 2001); origins and distribution if Elephas antiquus (Lister 2004)

Climate and the Early Human Occupation of Europe. Even ater people

successfully colonized Europe, continuous occupation appears to have been restricted to the warmer Mediterranean borderlands. North of the Alps and Pyrenees, where plant fossils, animal remains, or both occur,

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TABLE 5.12. Early west Asian archaeological and human fossil sites. Boldface and italics mark sites that have pro-

vided human fossils and the fossils are listed in parentheses. Figures 5.37 and 5.53 locate the sites. Estimated age (and Basis for Estimate)

Site

References

> = 600 ka GEORGIA Dmanisi (5 skulls [4 published], 4 mandibles (3 linked to skulls), and at least 16 isolated teeth and 24 postcranial bones)

= 1.7 Ma (paleomagnetism)

(Bar-Yosef 1995; Bar-Yosef 1998a; Ron & Levi 2001; Verosub & Tchernov 1991)

Gesher Benot Ya’aqov (= Gesher Benot Ya’akov = GBY = Jisr Banat Yacub) (two femoral shats)

800–700 ka (paleomagnetism)

(Bar-Yosef 1987; Bar-Yosef 1994c; Bar-Yosef & Belfer-Cohen 2001; Belitsky et al. 1991; Geraads & Tchernov 1983; Goren-Inbar 1994; Goren-Inbar et al. 1992; Goren-Inbar et al. 2000; Goren-Inbar et al. 1994; Goren-Inbar & Saragusti 1996; Goren-Inbar et al. 2002; Rink & Schwarcz 2005; Rosenfeld et al. 2004; Saragusti & Goren-Inbar 2001)

Evron Quarry

1–0.78 Ma (paleomagnetism)

(Bar-Yosef 1994c; Ron et al. 2003; Tchernov et al. 1994)

Bitzat Ruhama

990–850 ka (TL and paleomagnetism)

(Zaidner et al. 2003)

TURKEY Dursunlu

SYRIA Latamne ISRAEL

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TABLE 5.12. (continued)

Estimated age (and basis for estimate)

Site

References

300 ka (TL, faunal associations, and climatic correlation)

(Ljubin & Bosinski 1995)

Tsona Cave, South Osetia

?400–300 ka (faunal and artifactual associations)

(Ljubin & Bosinski 1995)

?500–?250 ka (faunal and artifact correlations)

(Ljubin & Bosinski 1995; Lyubin 1989)

AZERBAIJAN Azykh Cave (layers VI and V) (a mandible fragment) TURKEY Yarimburgaz Cave (hrace, Euro- 390–>160 ka (ESR, U-series, (Farrand & McMahon 1997; Kuhn 2002; Kuhn pean Turkey) and mammalian associaet al. 1996; Stiner et al. 1996) tions) Kaletepe Deresi 3 (archaeological layers IV and V)

between ?440 and > 160 ka (radiopotassium)

(Slimak et al. 2004)

Karain Cave E (Units A to E)

?440–300 ka (climate correlations)

(Kuhn 2002; Otte 1998; Otte et al. 1998)

Yabrud Cave 1

>= 225 ka (ESR and TL); ?400–300 ka (artifact correlations)

(Copeland 2000; Mercier & Valladas 1994; Porat et al. 2002; Rust 1950; Solecki & Solecki 1986)

Adlun (Zumofen Shelter)

?400–300 ka (artifact correlations)

(Copeland 2000; Garrod & Kirkbride 1961)

Zuttiyeh Cave (a fronto-facial fragment of “Galilee Man”)

>160 ka (U-series and artifact associations)

(Gisis & Bar-Yosef 1974; Schwarcz 1994; Sohn & Wolpof 1993; Vandermeersch 1989)

Berekhat Ram

between 470 and 232 ka (40Ar/39Ar)

(Bahn 1996; d’Errico & Nowell 2000; GorenInbar 1985; Goren-Inbar 1986; Goren-Inbar & Peltz 1995)

Misliya Cave

?400–>133 ka (OSL and artifact associations)

(Weinstein-Evron et al. 2003)

Tabun Cave (Layers G, F, and E) (a femoral shat and other fragments)

500–270 ka (TL, ESR, and U-series)

(Bar-Yosef 1994c; Bar-Yosef 1995; Bar-Yosef & Meignen 2001; Garrod 1956; Garrod & Bate 1937; Gilead 1970; Jelinek 1982a; Jelinek 1982b; Matskevich et al. 2001; Mercier & Valladas 2003; Mercier et al. 1995b; Rink et al. 2004; Shifroni & Ronen 2000)

Jamal Cave (Units 3 and 2)

?500–>220 ka (U-series)

(Weinstein-Evron et al. 1999)

Qesem Cave

>382–207 ka (U-series)

(Barkai et al. 2003; Lemorini et al. 2006; Verri et al. 2004; Verri et al. 2005)

LEBANON

ISRAEL

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Site

Estimated age (and basis for estimate)

Holon

>= 200 ka (TL and ESR)

Revadim Quarry

300–245 ka (climate correla- (Gvirtzman et al. 1999; Marder et al. 1999) tion and OSL)

References

(Chazan 2000; Chazan et al. 2001; Porat et al. 2002; Porat et al. 1999)

Uum Qatafa Cave Layers E and D >= 213 ka (ESR);500–270 ka (Bar-Yosef 1994c; Bar-Yosef 1995; Gilead 1970; (artifact correlations) Neuville 1931; Porat et al. 2002)

they indicate mild, mainly interglacial conditions at nearly all sites that antedate oxygen-isotope (glacial) stage 8, between about 301 and 242 ka. Table 5.11 lists the key European sites that relect the pattern. he attributions to particular interglacial stages are oten debatable, particularly prior to stage 8 because they depend mainly on the correlation of short, weakly dated, terrestrial sequences with the complete record of Quaternary glacial and interglacial alternations detected on the deep-sea loor. However, problems of precise correlation aside, sites with animal and plant fossils that imply full glacial climate become common only in stage 6, between about 186 and 130 ka. La Cotte de St. Brelade on the Channel Island of Jersey illustrates a stage 6 occupation especially well. he La Cotte fauna not only includes species like mammoth, woolly rhinoceros, and reindeer that signal glacial conditions, but the species could have existed on Jersey only if low (glacial) sea level had created a land bridge that joined Jersey to the European (French) mainland. To some extent the rarity or absence of glacial-age sites before stage 6 may relect erosional destruction during subsequent glacial intervals. he ice sheets themselves were particularly destructive, and glacial scouring would have removed virtually all traces of very early sites from Scandinavia and other ot-glaciated parts of northern Europe. However, in areas like northern France that the ice sheets never reached, erosion is unlikely to have singled out glacial (vs. interglacial) sites, and ancient, glacial-age deposits (especially, wind-borne dust or loess) oten persist but without traces of human activity. hus, the absence of very early, glacial-age sites in northern Europe probably means that the earliest Europeans had to abandon the region during peak glacial cold. In this connection it is also noteworthy that in the more oceanic parts of Europe, both glacial and interglacial sites older than stage 5, beginning 130 ka, occur only south and west of a line roughly through Schöningen and Vértesszöllös (ig. 5.43). he overall pattern suggests that like east Asian Homo erectus, the earliest Europeans could not cope with truly continental conditions, even during interglacials. he Neanderthals and their contemporaries ater 130 ka

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did scarcely better, and it was only modern Homo sapiens, ater 40 ka that successfully colonized the harshest environments Eurasia had to ofer.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES (not including those in table 5.11): climatic conditions implied at European sites older than OIS 8 (Bosinski 1995b; Dennell and Roebroeks 1996; Hublin 1998b; Roberts et al. 1995; Roebroeks et al. 1992a; Roebroeks and van Kolfschoten 1994, 1995; Tufreau and Antoine 1995; Villa 2001)

Artifact Traditions he previous chapter showed that the irst toolmakers, the Oldowan people, had mastered the mechanics of stone laking and were skilled at producing sharp-edged lakes that could slice through hide or strip lesh from bones. However, they appear to have made little or no efort to shape the core forms from which they struck lakes, and they may have used core forms mainly as hammers to crack bones for marrow. For this purpose, core shape didn’t matter very much. Homo ergaster, however, initiated a tradition in which core forms were oten deliberately, even meticulously shaped, and shape obviously mattered a lot. he characteristic artifact of the new tradition was the hand ax or biface—a lat cobble or large lake that was more or less completely laked over both surfaces (hence the term biface) to produce a sharp edge around the entire periphery (igs. 5.46, 5.48, and 5.49). Many hand axes narrow from a broad base or butt at one end to a rounded point at the other, like large teardrops. Ovals, triangles, and other shapes are also common, and in some places, hand ax makers produced pieces with a straight, sharp, guillotine-like edge opposite the base (ig. 5.50). Archaeologists oten call such pieces cleavers to distinguish them from hand axes, on which one end tends to be more pointed. Together, hand axes, cleavers, and similar large bifacial tools deine the Acheulean Industrial (or Cultural) Tradition, named in 1872 by the pioneer French prehistorian Gabriel de Mortillet, following the repeated discovery of hand axes in deposits of the River Somme at Saint-Acheul, northern France. he Acheulean Tradition lacked the art and architecture of much later archaeological cultures, but it was much more impressive for its duration and distribution, for as discussed below, it spanned more than a million years and three continents. his section stresses geographic and temporal patterning in artifacts, with particular attention to the relation between assemblage composition and human type. One of the main conclusions is that diferences in artifacts between Africa, western Asia, and Europe, on the one hand, and eastern Asia, on the other, underscore the probable divergence of east Asian Homo erectus from African and European populations by 1 Ma or before. Another is that the similarity of African and European

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quartz

373 FIGURE 5.46. bifaces from site TK in upper bed ii, olduvai Gorge (redrawn after leakey [1971], 190). The artifacts are probably about 1.2 my old, and they are among the oldest known markers of the acheulean industrial Tradition, which began in africa 1.7–1.6 ma.

lava

cm

quartz

lava

quartz

Olduvai Gorge Upper Bed II artifact assemblages at 600–500 ka adds to the probability that Africans and Europeans shared a common ancestor about this time. SOURCES: discovery of the Acheulean at St. Acheul (Bahain et al. 2007; de Mortillet 1867; Oakley

1964)

Africa

Table 5.13 lists the principal African archaeological sites where Homo ergaster (African H. erectus) or early H. sapiens probably produced the artifacts. Figures 5.37 and 5.47 locate the sites. At Terniine, Buia, GonaBSN12, Daka-Bouri, Melka Kunturé (Gomboré II and Garba IV), Konso, Olorgesailie, Olduvai Gorge Beds II and IV, and Swartkrans, the artifacts are associated with demonstrable or presumed fossils of H. ergaster. At Sidi Abderrahaman, the homas Quarry Caves, Bodo, Lake Ndutu, Kapthurin, the Cave of Hearths, and Elandsfontein, they accompany certain or probable fossils of early H. sapiens (or H. heidelbergensis). In each

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22o

FIGURE 5.47. The approximate locations of the main african acheulean sites

14o

36o

6o

Lac Karar

2o Aïn Hanech

10o

18o

26o

34o

42o

50o

Sidi Zin

36o

Ternifine 28o

Sidi Abderrahman & Thomas Quarries T ihodaïne

20o

12

28o

Kharga Dakhla Bir Tarfawi

20o

Bir Kiseiba Buia Gona Middle Awash Melka Kunturé Gadeb Karari W. T urkana Konso Kapthurin Kilombe Kanjera Kariandusi Nsongezi Olorgesailie Lainyamok Isenya Peninj Olduvai & L. Ndutu

o

4o o

32

1000 km

0o 4o

Kalambo Falls

12o

4o

4o

Isimila 12o

12o

20o

28o

18o

10o

2o

Cave of Hearths Olieboompoort Wonderwerk Namib IV Kathu Vaal Gravels Pan Swartkrans, Doornlaagte Sterkfontein & Rooidam & Kromdraai Montagu Cave Cornelia Elandsfontein Amanzi Duinefontein 6o 14o 22o 30o 38o

20o

28o

46o

54o

instance, the artifacts include characteristic Acheulean hand axes, cleavers, or other bifacial tools. Radiopotassium dates from Gona and West Turkana show that the Acheulean Tradition had emerged in eastern Africa by 1.65 Ma. Melka Kunturé, Konso, East Turkana (Karari Escarpment), Olduvai Gorge (site EF-HR), and Peninj demonstrate that it was widely established by 1.6–1.4 Ma. Associated mammals at Sterkfontein Cave (Member 5) and Swartkrans Cave (Members 2 and 3) place it in southern Africa at roughly the same time. Future research may show that it originated 1.8–1.7 Ma, more or less contemporaneously with H. ergaster. he Acheulean Tradition surely developed from the Oldowan, and archaeologists commonly lump the Oldowan and the Acheulean together in the Earlier Stone Age or ESA. Oldowan core forms and lakes continued on in the Acheulean, and in the earliest Acheulean, they sometimes outnumbered hand axes and other bifaces. At Olduvai Gorge and elsewhere in eastern Africa, such biface-poor assemblages have been assigned to the Developed Oldowan B, which is presumed to be an outgrowth of the Developed Oldowan A, a late facies (variant) of the Oldowan proper. However, the diferences between Developed Oldowan B assemblages and

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TABLE 5.13. Key African archaeological sites where Homo ergaster, H. heidelbergensis, or early H. sapiens probably made the artifacts. Figures 5.37 and 5.47 locate the sites. Boldface and italics mark sites that have provided bones of the likely tool makers. Excepting Aïn Hanech, all the sites belong to either the early or the late stage of the Acheulean Industrial Tradition. he transition between the stages is weakly dated, but at present, a time around 700–600 ka seems likely.

Site

References

early Acheulean MOROCCO homas Quarry I Level L

(Geraads et al. 2004; Raynal et al. 1995b; Raynal et al. 2001; Rhodes et al. 2006)

ALGERIA Aïn Hanech

(Geraads et al. 2004; Sahnouni & de Heinzelin 1998; Sahnouni et al. 2002; Sahnouni et al. 2004)

Terniine

(Balout et al. 1967; Geraads et al. 1986)

EGYPT Bir Kiseiba

(Haynes et al. 1997a; Haynes et al. 2001)

ERITREA (Abbate et al. 1998; Martini et al. 2004)

Buia ETHIOPIA BSN-12, OGS-5, and OGS-12, Gona

(Quade et al. 2004; Semaw 2000)

Daka-Bouri, Middle Awash

(Asfaw et al. 2002; Clark & Schick 2000; Gilbert et al. 2003; WoldeGabriel et al. 2004)

Melka Kunturé (Gomboré II, Garba II, Garba IV, Garba XII, and Simbirro III)

(Chavaillon 1979; D’Andrea et al. 2002)

Gadeb 2 and 8

(Clark 1987; Clark & Kurashina 1979; Haileab & Brown 1994; Williams et al. 1979)

Konso (ex-Konso-Gardula)

(Asfaw et al. 1992; Nagaoka et al. 2005; Suwa et al. 1997)

KENYA Kokiselei (KS) Complex 5, West Turkana

(Roche 1995; Roche et al. 2003; Roche & Kibunjia 1994)

Karari Escarpment (Koobi Fora), East Turkana

(Isaac & Harris 1978; Isaac & Isaac 1997)

Kilombe

(Crompton & Gowlett 1993; Gowlett 1978; Gowlett 1991)

Kariandusi

(Evernden & Curtis 1965; Gowlett & Crompton 1994; Kleindienst 1961; Leakey 1931)

Olorgesailie

(Isaac 1977; Koch 1989; Koch 1990; Potts et al. 2004; Shipman et al. 1981)

Kanjera

(Behrensmeyer et al. 1995; Ditchield et al. 1999; Leakey 1935; Plummer et al. 1994; Plummer & Potts 1995)

TANZANIA Peninj (Lake Natron)

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TABLE 5.13. (continued)

Site

References

Olduvai Gorge (Beds II and IV)

(Bunn 1986; Leakey 1971; Leakey 1975; Leakey 1977; Leakey 1979; Leakey & Roe 1994; Potts 1982; Potts 1988; Shipman 1986b; Shipman 1988)

SOUTH AFRICA Kromdraai A

(Kuman et al. 1997; Vrba & Panagos 1982)

Swartkrans Cave

(Clark 1993b; Leakey 1970; Volman 1984)

Sterkfontein Cave, Member 5

(Kuman 1994a; Kuman et al. 2000; Kuman 1994b; Kuman 1998; Kuman & Clarke 2000)

Cornelia

(Butzer 1974c; Clark 1974b; Cooke 1974)

late Acheulean MOROCCO Sidi Abderrahman (Littorina Cave)

(Arambourg & Biberson 1956; Biberson 1964; Geraads 1980; Geraads et al. 1980; Howell 1960; Jaeger 1975; Raynal et al. 1995b)

homas Quarries I and III Caves (homas III = Oulad-Hamida 1, including Rhinoceros Cave)

(Geraads 1980; Geraads et al. 1980; Geraads et al. 2004; Howell 1978a; Hublin 1985; Raynal et al. 1995b; Raynal et al. 2001; Rhodes et al. 1994; Rhodes et al. 2006)

ALGERIA Lac Karar

(Balout 1955)

Erg Tihodaïne

(homas 1979)

TUNISIA Sidi Zin

(Balout 1955; Gobert 1950)

EGYPT Dakhla (Dakhleh) Oasis

(Churcher et al. 1999)

Kharga Oasis

(Smith et al. 2004a)

Bir Tarfawi

(Wendorf et al. 1994b; Wendorf & Schild 1980)

ETHIOPIA Bodo, Middle Awash

(Clark 1987; Clark et al. 1984a; Clark et al. 1994; Clark & Schick 2000; Kalb et al. 1982a)

KENYA Isenya

(Brugal 1986; Roche et al. 1988; Roche et al. 1987)

Kapthurin (Baringo)

(Cornelissen et al. 1990; Deino & McBrearty 2002; Leakey et al. 1969a; McBrearty 1999; McBrearty et al. 1996; Tryon & McBrearty 2002; Tryon & McBrearty 2006)

UGANDA Nsongezi

(Cole 1967; Howell & Clark 1963)

TANZANIA Lake Ndutu

(Mturi 1976)

Isimila

(Cole 1963; Howell & Clark 1963; Howell et al. 1962; Howell et al. 1972)

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evoluTion of The Genus HO MO TABLE 5.13. (continued)

Site

References

ZAMBIA Kalambo Falls

(Clark 1969; Clark 2001; Sheppard & Kleindienst 1996)

NAMIBIA Namib IV

(Shackley 1980)

SOUTH AFRICA Cave of Hearths

(Latham & Herries 2004; Mason 1962; Mason 1988b; McNabb et al. 2004)

Olieboompoort Shelter

(Mason 1962)

Kathu Pan

(Beaumont et al. 1984; Butzer 1984a; Klein 1988a)

Wonderwerk Cave

(Beaumont 1982; Beaumont 1990b; Beaumont et al. 1984; Beaumont & Vogel 2006; Binneman & Beaumont 1992; Butzer 1984a; Malan & Cooke 1941; Malan & Wells 1943)

Vaal “Younger Gravels” = (the Rietputs Formation), including Power’s Site

(Butzer et al. 1973; Cooke 1949; Helgren 1977; Helgren 1978; Helgren 1979; Klein 1988a; Partridge & Brink 1967; Power 1955; Van Riet Lowe 1935; Van Riet Lowe 1952; Wells 1964)

Doornlaagte

(Butzer 1974a; Deacon & Deacon 1999; Mason 1966; Mason 1967)

Rooidam

(Butzer 1974a; Fock 1968; Szabo & Butzer 1979)

Montagu Cave

(Goodwin 1929b; Keller 1973)

Elandsfontein (= Saldanha = Hopeield)

(Hendey 1974; Klein 1978c; Klein et al. 2007; Klein & Cruz-Uribe 1991; Milo 1994; Singer 1957; Singer & Crawford 1958; Singer & Wymer 1968)

Duinefontein

(Cruz-Uribe et al. 2003; Feathers 2002; Klein et al. 1999a; Sampson 2003)

Amanzi Springs

(Deacon 1970)

contemporaneous Acheulean ones may simply relect diferences in the activities carried on by the same people at diferent sites or, in some cases, diferences in the local availability of raw material. Most authorities thus subsume the Developed Oldowan B within the Acheulean Tradition. Oldowan core forms broadly anticipate Acheulean bifaces, yet no Oldowan or Acheulean assemblage contains tools that are truly intermediate between the two, and the biface concept seems to have appeared abruptly, perhaps in a punctuational event like the one that may have produced H. ergaster. he earliest makers of bifaces made one other noteworthy discovery that was oten tied to biface manufacture: they learned how to strike large lakes, sometimes 30 cm or more in length, from large boulders, and it was from these that they oten made hand axes and cleavers. Ancient stone-tool assemblages that contain large lakes can be assigned to the Acheulean Tradition even on those occasions when, perhaps by happenstance, hand axes are absent. he end of the Acheulean Tradition is not irmly dated, but the ensuing Middle Stone Age (MSA), or Middle Paleolithic, was widely installed

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in Africa between 300 and 200 ka. (In northern Africa, archaeologists oten use the European labels Middle Paleolithic and Mousterian instead of MSA.) In the Kapthurin Formation, Kenya, geochemical characterization (“ingerprinting”) of volcanic tufs to correlate scattered sites suggests a complex transition in which Acheulean and MSA occupations coincided or alternated from perhaps 500 ka until the Acheulean inally disappeared about 285 ka. However, as emphasized below and in the next chapter, MSA artifact assemblages difered from late Acheulean ones mainly in the absence of large carefully trimmed hand axes and other bifaces. he Kapthurin artifact samples tend to be small, and they comprise a variable mix of excavated and surface-collected artifacts that may or may not be contemporaneous. Chance could play a strong role in the composition of such samples, and the Kapthurin results would be more compelling if they were mirrored at a single deeply stratiied Acheulean/ MSA locality. So far, wherever Acheulean and MSA (or Middle Paleolithic) assemblages are stratiied together in Africa, western Asia, or Europe, mostly in caves like Montagu, Wonderwerk, or the Cave of Hearths in South Africa, the Acheulean always underlies the MSA, with no indication for alternation or chronological overlap. he reason that MSA Africans and their west Eurasian contemporaries stopped making large bifaces is obscure, but a reasonable supposition is that they had invented a method for hating smaller tools on wooden handles or shats. he new composite tools would have been more portable, and they may have been quicker and easier to produce. In Africa, the technological advance may have coincided broadly with the appearance of more advanced people, like those represented at Florisbad in South Africa, Singa in Sudan, Omo-Kibish (individual no. 1) and Herto in Ethiopia, and Djebel Irhoud in Morocco. he fossils in each case surely or probably antedate 130 ka, and they clearly anticipate the nearmodern humans who inhabited Africa immediately aterward. However, in Europe, the people who made MSA-like (Middle Paleolithic or Mousterian) tools were Neanderthals and if it is accepted that they evolved in Europe, they either developed the new composite tool technology independently or it spread between continents entirely by difusion. he dates indicate that the Acheulean Tradition persisted for 1.5–1.4 my, and Acheulean assemblages separated by tens or hundreds of thousands of years commonly difer little if at all. In addition, for most of the long Acheulean timespan, assemblages separated by hundreds or thousands of miles, oten resemble each other closely, and local raw material quality, or a combination of raw material quality and the extent to which local people reshaped or refreshed used hand axes, could account for most observable geographic variability. It is oten thus said that Acheulean people were extraordinarily conservative in their behavior. However, numeric dates and mammalian associations imply there were

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early Acheulean bifaces (BOU-A1, Daka Mb of the Bouri Fm)

late Acheulean bifaces (BOU-A8, Herto Mb of the Bouri Fm)

actually two long periods of behavioral stasis: a more primitive early Acheulean stage and a signiicantly more reined late one. he available dates place the transition from early to late between 1 Ma and 500 ka, and future research may show that it occurred rapidly around 600 ka, perhaps simultaneously with the increase in brain size that distinguished early H. sapiens (or H. heidelbergensis) from H. ergaster. his increase in brain size was particularly notable because it increased encephalization, meaning that it occurred in the absence of an increase in body size. In the early Acheulean, the bifaces tended to be much thicker, less extensively trimmed, and less symmetrical (igs. 5.48–5.50). hey were commonly shaped by a small number of lake removals, oten less than ten, and the lake scars are usually deep. Modern experiments indicate that such scars result from the use of hard (meaning stone) hammers. Late Acheulean hand axes are oten equally crude, but many are remarkably thin and extensively trimmed, and they are highly symmetric not just when viewed in plan form but also when observed edge-on. he inal lake scars are shallow and lat, and replication eforts indicate that they were probably produced with sot (meaning wooden or bone) hammers. Late Acheulean people also manufactured a wider range of lake tools that foreshadowed those of their MSA and Middle Paleolithic successors (ig. 5.51), and they learned how to fashion a core that would provide a lake of predetermined size and shape. Archaeologists call such deliberate core preparation the Levallois technique, named for a western suburb of Paris

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379 FIGURE 5.48. earlier and later acheulean bifaces from the bouri formation, bouri Peninsula, middle awash, ethiopia (redrawn by Kathryn Cruz-uribe after de heinzelin et al. [2000], 148 and 165). The earlier acheulean pieces are from site bou-a1 in the daka member of the bouri formation, where they have been dated to about 1 ma. The later acheulean pieces are from site bou-a8 in the herto formation, where they are more tentatively dated to about 400 ka. like all bifaces from before 700–600 ka, those from bou-a1 were finished by hard-hammer percussion, which produces deep, bold flake scars. The number of flake scars is small and the finished pieces are relatively asymmetrical in both plan and side view. Similarly crude bifaces also occur in Acheulean sites after 600 ka, but they are often accompanied then by more sophisticated ones like those from BOU-A8. These were probably finished by soft-hammer percussion, which produced much shallower flake scars, the scars are much more abundant (reflecting greater effort in shaping), and the final products are more symmetrical in both plan and side view. The raw materials from which the illustrated bifaces were made—fine-grained volcanic rock at BOU-A1 and silicified limestone at BOUA8—flake about equally well, implying that a difference in human behavior determined the difference in the final products.

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FIGURE 5.49. an earlier acheulean hand ax (in chert) from sterkfontein Cave (redrawn after Kuman [1994b], fig. 6), compared to a Later Acheulean hand ax (in banded ironstone) from Kathu Pan (drawn from the original by Kathryn Cruz-Uribe). Mammalian associations suggest that the Sterkfontein hand ax dates from roughly 1.5 Ma, while the one from Kathu Pan is about 600 ky old. Like other earlier Acheulean hand axes, the one from Sterkfontein exhibits relatively few flake scars, and the scars are deep, implying that the maker used only a “hard” (stone) hammer. Later Acheulean hand axes are sometimes equally crude, but many like the one from Kathu Pan are remarkably thin and extensively flaked, and the final flaking probably required a “soft” (wooden or bone) hammer.

0

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early Acheulean hand axe (Sterkfontein Mb 5)

late Acheulean hand axe (Kathu Pan)

where the lakes that reveal such preparation were found and recognized in the 1860s. As discussed in the next chapter, specialists now identify several variants of the Levallois technique, and from the late Acheulean onward, some people employed one or more variants extensively, while other people hardly used them at all. Much of the diference probably relects diferences in the availability of suitable stone raw material. Finally, in distinction from early Acheulean assemblages, late ones varied more obviously through time and space, and the inal Acheulean anticipated the MSA in its extent of regional diferentiation. he behavioral advances relected in late stage artifacts could help explain the spread of late Acheulean people to Europe about 600 ka, and the advances may have been more signiicant than the changes that ushered in the MSA 300–200 ka, since judged strictly on artifact typology and technology, the late Acheulean and the MSA resembled each other much more closely than either resembled the Later Stone Age (LSA). he LSA succeeded the MSA in Africa roughly 50 ka, and chapter 7 argues that

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381 FIGURE 5.50. an earlier acheulean cleaver (in quartzite) from sterkfontein member 5 (redrawn after Kuman [1994b], fig. 6), compared to a later Acheulean specimen (in silcrete) from Elandsfontein Cutting 10. Note the much more extensive shaping on the later Acheulean specimen.

early Acheulean cleaver (Sterkfontein Member 5)

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late Acheulean cleaver (Elandsfontein Cutting 10)

the behavioral advances it implies help explain how the descendants of early LSA people rapidly replaced other kinds of people in Eurasia and eventually colonized every habitable corner of the globe. SOURCES: Developed Oldowan B—deinition (Leakey 1971, 1975), the possibility that only activity or raw material diferences diferentiate it from the Acheulean (Jones 1994), and its common incorporation in the Acheulean (Gowlett 1988; Stiles 1991); dating of the Acheulean/Middle Stone Age interface (Beaumont and Vogel 2006; Clark 1994; Evernden and Curtis 1965; Kuman et al. 1999; McBrearty et al. 1996; hackeray 1992a; Volman 1984; Wendorf et al. 1975, 1994a); interingering of the Acheulean and MSA in the Kapthurin Fm (Deino and McBrearty 2002; Tryon and McBrearty 2006); hating of smaller tools to explain the disappearance of hand axes (Clark 1993a; Schick and Toth 1993); raw material quality and rejuvenating to explain geographic variability in hand ax form (McPherron 2000); division of the Acheulean between early and late stages (Clark and Schick 2000; Isaac 1975; Schick and Clark 2000; Wynn 2002); increase in encephalization ca. 600 ka (Ruf et al. 1997); discovery of the Levallois technique (de Mortillet 1883); variants of the Levallois technique (Boëda 1993)

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FIGURE 5.51. later acheulean artifacts from elandsfontein Cutting 10 (redrawn after originals provided by T. P. volman). all were made from silcrete, a fine-grained, locally available siliceous rock. Elandsfontein has furnished numerous typical Acheulean large bifacial tools, exemplified here by a hand ax and a partial biface or bifacial knife. It has also provided small, thin flakes that were probably produced during biface manufacture or trimming; larger, thicker flakes that were struck for use as tools; and cores from which the larger flakes were struck. Some of the flakes exhibit the radial pattern of dorsal scars that is a characteristic feature of the Levallois technique for determining flake size and shape in advance. Later Acheuleans, after roughly 600 ka, were the first to apply the Levallois technique routinely, and they often produced flake tools that anticipate those of the succeeding Middle Stone Age in their mode of manufacture, secondary modification (retouch), and variety. At Elandsfontein, the flake edges were mostly unretouched or only lightly retouched, perhaps partly from use, but some were deliberately serrated or denticulated, perhaps for sawing or shredding.

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denticulate

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flake core partial biface or bifacial knife

Elandsfontein Cutting 10 Eastern Asia

No artifacts have been reported in direct association with Homo erectus fossils in Java, nor are they unquestionably known from the same stratigraphic units. It has long been assumed that classic H. erectus or the succeeding Solo (Ngandong) people made the choppers, chopping tools, lakes, and occasional hand ax–like tools assigned to the Pacitanian (Patjitanian) Industry in south-central Java. However, it is now known that Pacitanian tools in the type area come from alluvial terrace ills that probably postdate H. erectus, and the Pacitanian Industry may be coeval with the typologically similar end-Pleistocene/early Holocene Hoabinhian Industry of mainland Southeastern Asia. A lake and a chopper from alluvial deposits near Sambungmacan probably also date from the late Quaternary, rather than from the mid-Quaternary as irst suggested. At present, the oldest known tools in Java may be some small, relatively amorphous cores and lakes recovered from alluvial sediments at Ngebung near Sangiran in the 1930s and again more recently. In both cases the actual age of the artifacts is uncertain, and they could have been

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made either by classic H. erectus or by the succeeding Sambungmachan/ Ngandong (Solo) people. he absence of archaeological sites that are unambiguously associated with H. erectus on Java may relect a shortage of ancient lake basins or other low-energy sediment traps, together with limited archaeological survey. Fractured basalt, andesite, and chert stones from alluvial deposits in the Soa Basin on the island of Flores, east of Java, may illustrate what Java could one day produce. he most proliic indspot is at Mata Menge with 507 mostly basalt lakes and fractured pebbles. At this locality and others, the fractured stones occur in the same deposits as bones of the extinct elephant-like species, Stegodon trigonocephalus, which apparently arrived in Java about the same time as H. erectus. Paleomagnetic analysis suggests that the fossiliferous deposits accumulated shortly ater the shit from the Matuyama Reversed Chron to the Brunhes Normal Chron at roughly 780 ka. he magnetism is inconsistent with issiontrack dates that suggest an age of 800–880 ka, but either dating would imply that the maker was classic H. erectus. he possibility exists, however, that the Mata Menge fractured stones are not artifacts since they recall the amorphous laked pieces that once identiied the Kafuan Pebble Culture in Uganda and other parts of sub-Saharan Africa. Based on geomorphic context, the Kafuan was thought to antedate the Oldowan, but subsequent research showed that Kafuan laking could have resulted when pebbles or cobbles collided naturally, and its human origin is no longer accepted. he Mata Menge laked stones were selected from an unspeciied number of unmodiied pebbles or cobbles in the same deposits, and 12% were heavily abraded, implying that stream transport was important in their history. he issue of whether they are artifacts is especially consequential, since unlike Java and other Indonesian islands further to the west (west of Bali), Flores was never linked to the Southeast Asian mainland by dry land. Access to Flores, even during periods of lower sea level, required an openwater crossing of at least 19 km, and few terrestrial vertebrates made it. he exceptions include a rodent that could have crossed on natural rats and Stegodon that could have swum across, if it was like living elephants. People probably always required boats, and before Mata Menge was publicized, it was thought that only H. sapiens ater 60–50 ka could produce suitable ones (chap. 7). hus, if the crudely laked Mata Menge stones are artifacts, they reveal a totally unexpected degree of cultural sophistication. Elephants and other large mammals have repeatedly evolved dwarf forms when they became isolated on islands, and Flores Stegodon provides a prime example. H. erectus may provide another, since diminutive human bones found on Flores have been assigned to a dwarf form,

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Homo loresiensis, that is argued to derive from H. erectus. he alternative is that H. loresiensis is not a valid species and that its bones actually come from one or more modern humans alicted by dwarism and microcephaly (discussion in the addendum to chap. 7). Artifacts that are demonstrably as old as the oldest H. erectus are unknown on the southeastern or southern Asian mainland, with two possible exceptions. he oldest and potentially the most important case concerns three laked quartzite cobbles and two quartzite lakes from Riwat, near Rawalpindi in northeastern Pakistan. he Riwat deposit has been dated to at least 1.9 Ma by a combination of paleomagnetism and stratigraphic relation to a ission track–dated layer in the same region. Unfortunately, however, the deposit is a consolidated gravel (conglomerate) in which it is diicult to separate laking produced by natural collisions of cobbles from laking produced by humans, and the presumed artifacts have been distinguished from other, apparently natural lakes or laked cobbles mainly by quantitative criteria, including more (but still scant) lake scars, a smaller amount of remaining cobble cortex, and clearer bulbs of percussion. If the supposed artifacts are genuine, their near-2 My age has profound implications for the history and nature of human dispersal to Eurasia, but it remains possible that they represent simply one extreme along a continuum of naturally laked pieces. he remaining exception is less momentous. It involves three laked pebbles from alluvial deposits near Ban Mae ha, northern hailand. he alluvium is overlain by a paleomagnetically reversed basalt that has been radiopotassium-dated to 800 ± 300 and 600 ± 200 ka. Together the paleomagnetic and radiopotassium readings imply an age near the end of the Matuyama Reversed Polarity Chron, approximately 780 ka. If the stratigraphic position of the pebbles has been properly assessed, and if they are genuinely artifactual, they would be among the oldest welldated artifacts in eastern Asia. In southern Asia, the next oldest, welldated pieces are probably Acheulean hand axes and other artifacts from alluvial gravels near Dina, northeastern Pakistan, and at Bori, Maharashtra State, west-central India. At Dina, paleomagnetism and stratigraphic correlation with radiometrically dated sediments elsewhere bracket the artifacts between 780 and 400 ka. At Bori, 40Ar/39Ar determinations place the artifacts at roughly 670 ka. he Narmada skullcap, from Hathnora, north-central India, may represent the people who made the Dina and Bori hand axes. Acheulean hand axes occurred nearby, and the associated mammal fauna suggests broad contemporaneity with Dina and Bori. Morphologically, the Narmada skull recalls like-aged European and African specimens that are sometimes lumped in Homo heidelbergensis, and it may strengthen the case for an African exodus roughly 600–500 ka that brought Acheulean hand ax makers to both Europe and southern Asia. Alternatively, Bori

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may imply that Acheulean people penetrated southern Asia irst, perhaps because it was more like Africa environmentally. In China, artifacts are said to be stratigraphically associated with H. erectus at Lantian (the Gongwangling and Chenjiawo localities), Yunxian, Yuanmou, and Zhoukoudian Locality 1. he association at Zhoukoudian is by far the best documented and least equivocal. Artifacts have also been found at sites where geological or paleontological context ingers H. erectus as the artifact maker, although H. erectus fossils are absent. As noted previously, the oldest unquestionable artifacts in sealed stratigraphic context come from Donggutuo, Xiaochangliang, and Majuangou in the Nihewan Basin, 150 km west-northwest of Beijing. Paleomagnetic analysis of ine-grained sedimentary sequences at both sites brackets the artifact horizons between 1.66 and 1.1 Ma. he next oldest artifacts come from the Bose Basin, where they have been dated to about 800 ka, from the Lantian sites and Yunxian, where they are arguably between 800 and 600–500 ky old. he 800 ka age for the Bose Basin artifacts follows from their stratigraphic association with tektites whose age has been well-established. he artifact assemblages produced by Chinese H. erectus share many features that are particularly well-illustrated by the large and relatively well-described sample from Zhoukoudian Locality 1. he Locality 1 collection comprises perhaps 100,000 pieces, but it lacks hand axes and other well-made bifacial tools that characterize many contemporaneous assemblages in Africa and Europe. It has long been taken as the type assemblage for the so-called Choukoutienian (Zhoukoudianian) chopper– chopping tool industry, but it actually contains relatively few choppers and chopping tools (bifacial choppers). Instead, it is heavily dominated by lakes, a small proportion of which are modiied by retouch (ig. 5.52). In early systematic overviews of the east Asian Paleolithic, the Harvard archaeologist, Hallam L. Movius, emphasized the absence of hand axes in the Choukoutienian and likened it to the Pacitanian of Java, the Anyathian of Burma, the Tampanian of Malaysia, and the Soan (or Soanian) of Pakistan, in all of which hand axes are rare or absent. Movius assumed that these industries were all broadly contemporaneous with each other and with industries in Africa and Europe where hand axes abound, and he concluded that there were two great early-Paleolithic culture areas— the hand ax (Acheulean) tract from peninsular India westward across southern Asia into Europe and Africa (ig. 5.2) and the chopper–chopping tool tract eastward from northern India through eastern and Southeastern Asia. Movius’s formulation is questionable today on three counts. First, the east Asian industries assigned to the chopper–chopping tool complex are not necessarily contemporaneous either with each other or with hand ax industries to the west. Some, like the Pacitanian, are clearly

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FIGURE 5.52. sandstone and quartz artifacts associated with Homo erectus at Zhoukoudian locality 1 (redrawn after movius [1949], figs. 22, 23). The Zhoukoudian assemblage lacks hand axes and other typical Acheulean tools found at many sites in Africa, western Asia, and Europe, and it probably represents a totally distinct Acheulean-age artifact tradition that was widespread in eastern Asia.

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10 cm

choppers and chopping tools

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retouched or utilized flakes

Zhoukoudian Locality 1 younger. Second, many well-known early Paleolithic assemblages in the west also lack hand axes. Important examples include the assemblages from Atapuerca TD 6 in Spain; Isernia La Pineta and La Polledrara in Italy; Bilzingsleben and Schöningen in Germany; Vértesszöllös in Hungary; and Clacton and the lower horizons at Swanscombe in England (references in table 5.11). hird, hand ax–like bifacial artifacts have now been recorded at several east Asian early Paleolithic sites, including Lantian, Dingcun, Kehe, and the Bose Basin in China and Chon-Gok-Ni and other sites in the Imjin/Hantan River Basins of South Korea. Some large core tools from Dingcun, Kehe, and Chon-Gok-Ni broadly it the deinition of a hand ax, but they tend to be thicker and more crudely made than typical African or European hand axes, and they oten recall the less formal, three-sided (trihedral) core tools or “picks” that mark some post-Acheulean (Sangoan) assemblages in central Africa. In addition, the Dingcun and Chon-Gok-Ni sites are likely to date to 200 ka or later, and if so, they postdate the Acheulean in the west.

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In contrast, the Bose Basin artifacts more closely resemble true Acheulean pieces, and they have been dated to 800 ka, irmly within the long Acheulean interval. Like contemporaneous Acheulean artifacts in Africa, the Bose Basin tools were oten made on large lakes (exceeding 10 cm in diameter), they were frequently trimmed and thinned by extensive laking, and some are roughly symmetrical in plan form with a cutting edge that extends around much of the periphery. However, unlike classic Acheulean hand axes, they tend to be mainly unifacial (worked on one surface only), and few if any exhibit the ovoid plan form and biconvex cross-section that is common for Acheulean bifaces. In sum, they do not demonstrate that the Acheulean Tradition penetrated China, but they do show that Chinese H. erectus could produce equally sophisticated stone artifacts. Conceivably, comparably reined artifacts are lacking at Zhoukoudian and other east Asian sites because the inhabitants lacked access to stone raw material of suitable size and quality. Arguably, the Bose Basin artifact makers produced large extensively trimmed pieces only for a brief period, ater a rain of molten tektites ired the local vegetation and exposed large cobbles. he fundamental point remains that true Acheulean hand axes and related tools are still unknown north and east of peninsular India, and the reason is not a paucity of excavated archaeological sites, since commercial quarrying has repeatedly produced hand axes on the west for nearly two centuries. In addition even without quarrying or excavation, hand axes and related tools have oten been found on the surface in parts of Africa, western Europe, southwestern Asia, and India, where covering deposits are currently eroding or where they never formed. hus, ity years ater the “Movius line” was irst formulated, it continues to imply a fundamental culture-geographic divide dating from 1 Ma or before. Flake-and-chopper assemblages of Acheulean age from Kuldara and other sites in Tadzhikistan and Kirgizstan may mean that the divide extended westward across south-central Asia, and it may even have reached Europe (ig. 5.2). Here, it may have separated a non-Acheulean region in the east and center from well-documented Acheulean areas in the south and west. In China, people like those represented at Dali, Mapa, and Jinniushan ater 300–200 ka produced artifact assemblages that closely resembled those of classic H. erectus before them. he principal tools remained lakes, accompanied by choppers and chopping tools. Arguably, stone-artifact assemblages changed little throughout most of China and adjacent Southeast Asia even ater the appearance of fully modern people like those represented at Zhoukoudian Upper Cave. Radiocarbon tentatively brackets the Upper Cave deposits between perhaps 30 and 11 ka. However, the Upper Cave people produced formal bone artifacts and personal ornaments that broadly resemble like-aged items in the Far West,

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and future research may show that a major artifactual transformation occurred throughout the Far East 50–40 ka, just as it did in Africa and Europe.

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SOURCES (not including those in table 5.14): Kafuan Pebble Culture (Clark 1958, 1959b, 113–114); H. loresiensis (Brown et al. 2004; Morwood et al. 2004, 2005); analysis of the east Asian Paleolithic by Movius (1944, 1948, 1949, 1955); post-Acheulean age of some east Asian chopper–chopping tool assemblages (Hutterer 1985); bifacial tools in Korea (Norton 2000; Norton et al. 2006a); absence of true Acheulean bifaces north and east of peninsular India (Clark 1994; Corvinus 2004); non-Acheulean region in Europe (Bosinski 1995a); Chinese artifacts postdating 300–200 ka (Qiu 1985, 1992; Schick 1994); dating of Zhoukoudian Upper Cave (Kamminga 1992; Kamminga and Wright 1988)

Western Asia

he archaeological record of primitive Homo in western Asia broadly parallels the record in Africa (references in table 5.12; site locations in ig. 5.53). he oldest west Asian artifact assemblages—from Erk-el-Ahmar (Israel) and Dmanisi (Georgia)—date from 1.7 Ma or before, and they have been assigned to the Oldowan Tradition. Hand axes and related tools from ‘Ubeidiya (Israel) show that the Acheulean Tradition appeared in western Asia by 1.4 Ma, and as in Africa, it can be divided between an early stage before about 700–600 ka and a later stage aterward. Hand axes and related tools tend to be smaller and more inely crated in the later stage, and they are accompanied by a wider range of lake tools, sometimes on Levallois lakes. Besides ‘Ubeidiya, prominent early Acheulean sites include Latamne in Syria and Evron Quarry and Gesher Benot Ya’aqov (GBY) in Israel. GBY is the most informative for its large artifact and faunal samples, the remarkable resemblance of its Acheulean artifacts to like-aged ones in Africa (possibly implying an Out-of-Africa dispersal); the preservation of fruits, seeds, and bark, some from edible plants; fragments of wood, including a “plank” that the human inhabitants appear to have polished; and the oldest reasonably persuasive evidence for human control of ire. he most proliic later Acheulean sites are caves including Yabrud 1, Adlun (Zumofen Shelter), Tabun, Misliya, and Qesem in the east Mediterranean coastal zone, oten referred to as the Levant. Assemblages that date between ≥400 ka and 250–200 ka in these caves and others are now usually assigned to the Acheuleo-Yabrudian or sometimes, the Mugharan Tradition. Small well-made Acheulean hand axes are characteristic, particularly near the bottom and top of deeply stratiied sequences, but the most common tools are thick lakes on which one or more edges have been modiied (retouched) to produce the tools that archaeologists call sidescrapers. Blades (lakes that are at least twice as long as wide) are also common, especially in the middle or upper parts of thick Acheuleo-Yabrudian sequences, and the blade-rich assemblages are usually assigned to the Amudian variant of the Acheuleo-Yabrudian.

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TABLE 5.14. he principal east and central Asian early archaeological sites addressed in the text. Dina, Hathnora,

and Bori have all provided typical Acheulean bifaces. he remaining sites lack Acheulean tools, and they have oten been assigned to the “East Asian Chopper-Chopping Tool Tradition,” although this term masks substantial variation between assemblages. At some sites (perhaps above all Riwat and Mata Menge), there is the additional problem that the laked stones may not be artifacts. he listed Chinese sites are mainly ones that have also provided remains of Homo erectus, but artifacts also occur alone at sites where H. erectus was the likely tool maker, based on geological or paleontological context (Aigner 1978a; Aigner 1978b; Howell 1986; Jia 1985; Olsen & Miller-Antonio 1992; Yi & Clark 1983; Zhang 1985a). he most signiicant examples are in the Nihewan and Bose Basins. Site

References

Java (Indonesia) Pacitan, Baksoka River (Pacitanian Industry)

(Bartstra 1984; Movius 1948)

Sambungmacan vicinity (a lake and a chopper) (Bartstra 1985; Jacob et al. 1978) Ngebung Hill (Sangiran)

(Bartstra 1985; Bartstra 1982a; Sémah et al. 1992)

Flores (Indonesia) Mata Menge and other localities in the Soa Basin

(Brumm et al. 2006; Corvinus 2004; Moore & Brumm 2007; Morwood et al. 1999; Morwood et al. 1998; O’Sullivan et al. 2001; Sondaar et al. 1994; van den Bergh et al. 1995)

Pakistan Riwat

(Dennell 1998; Dennell et al. 1988)

Dina

(Rendell & Dennell 1985)

India Bori

(Corvinus 2004; Mishra et al. 1995)

Hathnora (Narmada)

(de Lumley & Sonakia 1985a; de Lumley & Sonakia 1985b; Kennedy et al. 1991; Sankhyan 1997; Sonakia 1985; Sonakia 1992)

hailand Ban Mae ha

(Pope et al. 1986)

China Lantian (Gongwangling and Chenjiawo)

(An & Ho 1989; Brown 2001; Etler 1996; Hyodo et al. 2002; Zhang 1985a; Zhu et al. 2003)

Yunxian

(Chen et al. 1997; Vialet et al. 2005)

Yuanmou

(Brown 2001; Hyodo et al. 2002; Jia 1985; Olsen & Miller-Antonio 1992; Zhu et al. 2003)

Zhoukoudian (ex-Choukoutien) Localities 1 and 15

(Pei & Zhang 1985; Wu & Lin 1983; Zhang 1985a)

Nihewan (ex-Nihowan) Basin (Donggutuo, Xiaochangliang, and Majuangou)

(Pei 1939a; Pope 1998; Schick & Dong 1993; Schick et al. 1991; Zhu et al. 2003; Zhu et al. 2001; Zhu et al. 2004)

Bose Basin

(Hou et al. 2000; Koeberl et al. 2000; Olsen & Miller-Antonio 1992)

Dingcun and Kehe (ex-K’oho)

(Aigner 1978a; Qiu 1985; Schick 1994; Yi & Clark 1983)

South Korea Chon-Gok-Ni

(Norton 2000; Norton et al. 2006a; Schick 1994; Yi & Clark 1983)

Tadzhikistan and Kirgizstan Kuldara (and other sites)

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Ceprano

Kuldaro

Tsona Akhalkalaki

Yarimburgaz Petralona

Dmanisi Dursunlu Denizli Kalatepe Deresi 3 Karaïn

Latamne 100 km

Azykh

Kharga Dakhla

Yabrud Adlun Zuttiyeh Evron Berekhat Ram Misliya, Gesher Benot Bir Tarfawi Tabun & Yaʼaqov Jamal ʻUbeidiya Qesem ʻErk-el-Ahmar Holon Uum Qatafa Revadim Ruhama 1000 km FIGURE 5.53. locations of the main excavated, early Paleolithic sites in western asia. Where space permits, the map also locates important early Paleolithic sites in adjacent northeastern africa and southeastern europe. The concentration of excavated sites in israel reflects especially propitious circumstances for site formation, environmental conditions that have commonly favored human presence, and a relatively large number of interested archaeologists. Excepting Erk-el-Ahmar and Bitzat Ruhama, the Israeli sites all contain hand axes and other Acheulean tools. Excepting Kalatepe Deresi 3, the small number of excavated sites to the north and east do not, but hand axes are common on the surface throughout western Asia (Bar-Yosef 1994c), particularly in Turkey (Kuhn 2002). In Saudi Arabia, Iraq, and Iran, they demonstrate at least sporadic Acheulean presence even where sealed sites are rare or absent, and they underscore the validity of the “Movius line” separating the Acheulean culture zone west and south of northern India from the non-Acheulean zone to the east and north.

Amudian assemblages were initially referred to as Proto-Aurignacian, on the assumption that they anticipated the early Upper Paleolithic Aurignacian culture that appeared in Europe and western Asia ater 45–40 ka. he resemblances are purely supericial, however, and no one today infers a historic connection from the shared abundance of blades. At an age of >200–250 ka, Amudian assemblages are more appropriately compared to some Acheulean assemblages with blades that have been reported from the Kapthurin Formation of Kenya and from Wonderwerk Cave and other sites in the South African interior. Sometime between 250 and 200 ka, depending on how the available dates are read, the Acheuleo-Yabrudian was replaced by the Middle Paleolithic or Mousterian Tradition, described in the next chapter. hroughout western Asia, Mousterian assemblages are distinguished from Acheulean ones mainly by the absence of hand axes, and if the as-

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semblages occurred in Africa, they could be comfortably assigned to the Middle Stone Age. As between the two possible Oldowan sites, Dmanisi is the more compelling because the dating is irmer and the artifact assemblage is much larger. Still, it remains possible that Dmanisi postdates the advent of the Acheulean Tradition in Africa and that hand axes are missing for reasons other than age. Even sites that clearly postdate the spread of the Acheulean across western Asia sometimes lack hand axes and related tools, and the reasons are generally unclear. he most conspicuous example is probably Bitzat Ruhama (Israel), where deposits irmly dated to about 0.9 Ma have produced a large artifact assemblage that comprises only small lakes and cores. he Dmanisi human fossils (described above) do not help to identify the associated artifacts, since they comprise one skull that could be classiied as Homo habilis in the narrow sense and three others that closely resemble skulls of H. ergaster (or African H. erectus). It is generally assumed that H. habilis produced Oldowan tools in Africa, while H. ergaster invented the succeeding Acheulean. Turkey further illustrates the problem of understanding assemblage variability within the Acheulean timespan, since hand axes and related bifacial tools have been widely reported on the surface, but so far, there are only four excavated sites (table 5.12) where, based on the age of the deposits, hand axes could be expected in place, and only one site—Kaletepe Deresi 3—has provided them. heir absence at the other sites—Dursunlu, Yarimburgaz Cave, and Karaïn Cave—may relect small sample size or a lack of suitable stone nearby. In addition, Yarimburgaz and Karaïn were probably occupied by carnivores as much as or more than by humans. he case is particularly clear for Yarimburgaz, where bones from at least forty-two individual cave bears comprise 93% of the faunal sample. Bears or other carnivores also appear to have been the principal occupants of the Kudaro and Tsona Caves in Georgia and of Azykh Cave in Azerbaijan, and in each case, the implication may be that Acheulean people were relatively rare nearby. So far, in western Asia, only the Acheuleo-Yabrudian caves imply relatively intense human occupation during the long Acheulean timespan and perhaps then only under optimal climatic conditions. Isolated human fossils (table 5.12) occur with early Acheulean artifacts at ‘Ubeidya and Gesher Benot Ya’aqov and with late Acheulean artifacts at Kudaro 1, Azykh Cave, and Tabun Cave (layer E), but none are diagnostic to species. hey could represent H. ergaster before 600 ka, and early H. sapiens, H. heidelbergensis, or early H. neanderthalensis aterward. A fronto-facial fragment that may have been associated with Acheuleo-Yabrudian artifacts at Zuttiyeh Cave has been variously assigned to early H. sapiens or early H. neanderthalensis. As discussed in the next chapter, human remains are well-known from west Asian sites that

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postdate 130 ka, and the people appear to have been irst near-modern humans (H. sapiens) and later Neanderthals (H. neanderthalensis).

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES (not including those in table 5.12): age of the Acheuleo-Yabrudian/Mousterian transition (Porat et al. 2002); Acheulean blades from the Kapthurin Fm (McBrearty 1999) and from sites in the South African interior (Beaumont and Vogel 2006); surface Acheulean artifacts in Turkey (Kuhn 2002)

Europe

he oldest proposed archaeological assemblages in Europe lack hand axes and other Acheulean markers, and they instead comprise Oldowanlike choppers, chopping tools, and lakes. Ot-noted early nonbiface sites include Le Vallonet Cave and Chillac III in France that date to 1 Ma or before and the lower layers at Atapuerca GD and Atapuerca SE, Spain, the lower horizons at Kärlich, Germany, and the sites of Stránská Skála and Přezletice, Czech Republic, which date variously between 1.1 Ma and 500 ka. In most cases, however, there is the problem that the laked stones may not be artifactual, and human remains to conirm an artifactual origin occur only at Atapuerca GD and Atapuerca SE. If the validity of all the sites is accepted, however, the earliest Europeans either did not bring hand axes from Africa or, for reasons that remain obscure, they developed a separate Oldowan-like tradition that persisted for perhaps 500 ky. Alternatively, it is possible that the earliest Europeans sometimes made hand axes but failed to leave them at the known sites, that the environs of the known sites lacked suitable raw material for hand ax manufacture, or that hand axes are absent solely by chance. Chance could perhaps explain the lack of hand axes in the lowermost horizons of Atapuerca GD, particularly in layer TD 6, where the total number of artifacts is less than two hundred. Late Acheulean artifacts that broadly resemble African ones (igs. 5.54, 5.55) occur at numerous European sites that probably formed between 600 and 400 ka, including Atapuerca SH (Sima de los Huesos), Torralba, and Ambrona in Spain; Fontana Ranuccio and Venosa-Notarchirico in Italy; Abbeville (Carpentier Quarry), Saint-Acheul (rue de Cagny), and Cagny-la-Garenne in France; and Boxgrove, Barnham, and Swanscombe (middle or upper horizons) in England. Acheulean assemblages that probably date from between 400 and 300 ka are known from Atapuerca TG (Trinchera Galeria) in Spain; Castel di Guido, Malagrotta, and Torre in Pietra in Italy; Cagny l’Epinette and Terra Amata in France; and Hoxne in England (references in table 5.11; locations in ig. 5.42). he abundant human fossils from Atapuerca SH and a partial skull from Swanscombe imply that the hand ax makers included early members of the Neanderthal lineage. he Acheulean undoubtedly intruded Europe from Africa, and its abrupt appearance roughly 600 ka underscores the

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393 FIGURE 5.54. acheulean bifaces from Torralba, spain (redrawn from originals provided by f. C. howell and l. G. freeman). The artifacts probably date from sometime between 600 and 400 ka, and they were stratigraphically associated with bones of elephants, horses, and other large animals. as at most sites in the same time range, few of the animal bones show stone-tool marks, and the artifact/bone association need not mean that people hunted or butchered many of the animals.

0

10 cm

Torralba bifaces likelihood that H. neanderthalensis and H. sapiens shared an ancestor at about the same time. Although most European assemblages that date between roughly 600 and 250–200 ka contain hand axes or other large bifaces, there are also assemblages where the tools comprise a variable mix of lakes and laked pebbles (choppers and chopping tools) without bifaces (ig. 5.56). Prominent nonbiface sites that fall squarely within the Acheulean timespan include Clacton in England; most layers at Arago Cave in France; Bilzingsleben, Bad Cannstatt, and Schöningen in Germany; Vértesszöllös in Hungary; and La Polledrara, Isernia La Pineta, and some layers at Venosa-Notarchirico in Italy (references in table 5.11; locations in ig. 5.43). he nonbiface assemblages have oten been assigned to independent traditions like the Clactonian, Tayacian, proto-Mousterian, or Taubachian that are presumed to have paralleled the Acheulean and to have been produced by diferent people. Alternatively, in some cases the absence of large bifaces could simply relect the local absence of suitable raw material, the conduct of activities that did not require bifaces, or some combination of these factors. Raw material or activity diferences

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FIGURE 5.55. late acheulean artifacts from southern england (redrawn after Wymer [1968], 147). finely crafted hand axes of various shapes and sizes are the hallmark of the late acheulean in europe, but biased collection in the late nineteenth and early twentieth centuries probably artificially raised their frequency in many assemblages. Fine hand axes tend to be much less abundant in samples that have been carefully excavated in more recent decades.

ovate hand ax cordate hand ax

sidescraper

subcordate hand ax

cm

irregular hand ax ovate hand ax

are perhaps particularly likely to explain the absence of large bifaces at sites within regions like southern France or central Italy, where Acheulean and non-Acheulean assemblages formed more or less simulaneously. hey are less likely to explain the absence or near absence of hand axes over larger regions like east-central Europe, where only lakeand-chopper assemblages occur. Arguably, then, east-central European lake-and-chopper sites like Vértesszöllös in Hungary and Bilzingsleben in Germany relect a separate non-Acheulean tradition with roots in eastern or central Asia. he implication could be for two human migrations to Europe roughly 500 ka: one from Africa that introduced hand axes to western and west-central Europe and a second from Asia that introduced the lake-and-chopper tradition farther east. Such a scenario

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bifacial choppers

Tayac points

double convexconcave sidescraper

Clactonian notch

395 FIGURE 5.56. Tayacian (non-acheulean) artifacts from la Caune de l’arago, southern france (redrawn after de lumley [1975], figs. 7, 8). The term Tayacian has been applied to a wide range of European assemblages that lack hand axes but that were broadly coeval with Acheulean assemblages between roughly 500 ka and 250–200 ka. Among competing explanations for the difference, the most popular are that different people produced Tayacian and Acheulean assemblages or that Acheulean people produced both, but that they left Tayacian artifacts where they didn’t use hand axes or where they could not find suitable raw material nearby. Both explanations may be valid, depending on the site or assemblage.

denticulate 0

5 cm

denticulate chunk

might seem implausible on its face, but it could further explain why the human remains from Vértesszöllös and Bilzingsleben are the most H. erectus–like of all the fossils that are tentatively assigned here to the Neanderthal lineage. As in Africa and western Asia, the Acheulean in Europe was succeeded 250–200 ka by (Middle Paleolithic or Mousterian) lake industries that lack large bifaces. Diagnostic human remains are rare at early Middle Paleolithic sites, but where they occur—mainly at Biache-Saint-Vaast and l’Aubesier, France, and Ehringsdorf, Germany (table 5.5)—they contrast sharply with contemporaneous (early Middle Paleolithic/Middle Stone Age) human remains in Africa. Whereas the African fossils anticipate

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modern people, the European ones anticipate the classic Neanderthals who occupied Europe ater 130 ka. he contrast in human types could not have been anticipated from the associated artifacts, which are remarkably similar between Europe and Africa. A conspicuous artifactual diference between the two continents appears only ater 50 ka with the emergence and spread of modern humans from Africa. he next chapter argues that the similarity before 50 ka implies a shared primitive behavioral repertoire in which innovation was unusual, whereas the diference ater 50 ka implies a shared derived behavioral repertoire in which innovation was commonplace.

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SOURCES (not including those in tables 5.5 and 5.11): possibility that laked stones at many European sites older than 600–500 ka are not artifacts (Roebroeks and van Kolfschoten 1994); non-Acheulean industries coeval with the Acheulean in Europe (Bordes 1968; de Lumley 1975; Svoboda 1987; Valoch 1984) and the possibility they relect diferences in activities or in raw material availability (Villa 1983); possible origin of east-central European non-Acheulean industries from outside Europe (Rolland 1992); diferences between the Acheulean and Middle Paleolithic in Europe (Villa 1991)

Other Aspects of Behavior and Ecology In total, the fossil sample of Homo ergaster, H. erectus, early H. sapiens, and early H. neanderthalensis comes from perhaps 130 individuals, represented mainly by bits and pieces, and the behavior of the species must be inferred from fewer than ity reasonably well-excavated archaeological sites. he skeletal remains and sites are widely scattered in time and space, and the time dimension is poorly controlled, particularly outside Africa. Compounding these problems of small sample size and poor temporal control, the archaeological evidence for behavior is oten highly ambiguous, and nontrivial behavioral inferences are mostly tentative. Habitat Preference

Paleoenvironmental indicators indicate that, in general, Homo ergaster and its descendants favored open, nonwooded or lightly wooded settings and that in this preference, they difered from the chimpanzees. his would explain why human fossil and archaeological sites ordinarily lack chimpanzee fossils. he only known exception is an area within the Kapthurin Formation, Kenya, where a hominin mandible has been found at site GnJh-19 and chimpanzee fossils have been found at Locality 99 only one kilometer away within the same K3 geological unit. 40 Ar/39Ar dating and stratigraphic position suggests that each site formed between 540 and 500 ka. Table 5.4 lists the mandible and other hominin fossils from the same geological unit and assigns them to early Homo sapiens, based mainly on their presumed antiquity. he chimpanzee fossils comprise two incisors and two molars, and the incisors are particularly diagnostic. hey probably derive from the common chimpanzee, Pan

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troglodytes, and barring the possibility, considered in the last chapter, that Ardipithecus includes an early chimpanzee, they are the only chimpanzee fossils so far. he Kapthurin sites are about 100 km east of the present-day eastern limit of chimpanzee distribution in western Uganda and Tanzania. In the easternmost part of their range, chimpanzees commonly occupy semiarid woodland or savanna of the kind that human foragers are thought to favor, and vertebrate fossils and sediments imply that similar vegetation existed at Kapthurin 540–500 ka. his clearly provided acceptable, if perhaps marginal, habitat for chimpanzees in the Rit Valley, signiicantly east of their current range. Still, Kapthurin remains unique, and even if chimpanzee fossils are eventually found at other human fossil or archaeological sites, their rarity will still imply that the divergence of hominins and chimpanzees has long been grounded in a preference for contrasting environments. Environments favored by chimpanzees have occurred mostly west of the Rit Valley in the forested regions of central and western Africa, while those preferred by early hominins occurred mainly east and south of the Rit Valley, in the semiarid savannas and grasslands of eastern and southern Africa. SOURCES: chimpanzee fossils (McBrearty and Jablonski 2005); K3 vertebrate fauna (McBrearty

1999)

Site Location and Modification

African Acheulean sites are overwhelmingly associated with ancient stream or lake deposits, and this could mean that Acheulean people were closely tied to water. In contrast, succeeding Middle Stone Age (MSA) and Later Stone Age (LSA) sites occur more commonly in caves, and the connection to water is not so clear. he diference may mean that MSA and LSA people had developed technology that allowed them to roam farther from water, but it could also be a result of preservation, since most caves of Acheulean age collapsed long ago and their deposits were eroded away or are presently inconspicuous. Open-air, water-edge sites (vs. caves) also dominate the archaeological record in Eurasia before the Middle and Upper Paleolithic, and the reason is the same. Early humans almost surely required shelters of some kind, particularly ater they colonized Eurasia, but the surviving evidence is remarkably sparse and ambiguous. Cited examples include seemingly patterned arrangements of large, nonartifactual rocks at the proposed pre-Acheulean site of Soleihac, France, and at four Acheulean sites: Melka Kunturé (Ethiopia), Olorgesailie (Kenya), Latamne (Syria), and Terra Amata (France). People at each site might have arranged the rocks to make the foundations of huts or windbreaks, but in each case the

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travertine wall channels in marshy area slope to streambed

artifact workshops

concentrations of bones and stones

tusk hearth areas

artifact workshops

0

4m fissure

Bilzingsleben Excavation Plan

responsible agent could equally have been stream low, soil creep, or some other natural process. Many more sites contain clusters of artifacts, bones, and other debris that could mark hut bases or specialized activity areas. he evidence is more conjectural than compelling, but the most noteworthy examples perhaps include those from Bilzingsleben and Ariendorf 1 in Germany, and from Le Lazaret Cave in southern France (references in table 5.11). he sites vary in age from perhaps 350 ka at Bilzingsleben, to a time within global oxygen isotope stage (OIS) 8 (301–242 ka) at Ariendorf 1, and a time within OIS 6 (186–130 ka) at Le Lazaret.

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3m

N

rocks anvils pebble tools

399 FIGURE 5.57. Left: arrangement of bone and artifact concentrations dated to 400–300 ka at bilzingsleben, Germany (adapted from mania [1986], 244). Right: Close-up of part of the oval accumulation of stones and bones whose position is indicated to the left (adapted from mania [1986], 245). The accumulation may mark the base of a structure in which large stones and bones were used to build walls, though the spatial patterning is less convincing than it is in “ruins” at many sites that postdate 40 ka. The location of a second, similar accumulation is not shown on the plan (left) because it was excavated only after the plan was drawn.

animal bones charcoal charred wood and charcoal

At Ariendorf 1, several large quartz and quartzite blocks (measuring up to 60 × 30 × 30 cm) occur in a ine-grained, sandy-silty deposit, where they could only have been introduced by people. Conceivably, they mark the base of a structure, and they partly surround a scatter of artifacts and fragmentary bones. Artifacts that can be reitted to reconstruct the core from which they were struck and conjoinable bone fragments imply that many objects were deposited at more or less the same time. At Bilzingsleben, the ancient inhabitants camped alongside a stream lowing from a nearby spring to a small lake. hey also settled on parts of the streambed that were periodically dry. he evidence for structures

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FIGURE 5.58. Putative floor plan of an early Middle Paleolithic layer dated between 190 and 130 ka at Le Lazaret Cave, southern France (adapted from de Lumley and Boone [1976], 639). Stone artifacts and fragmentary animal bones were heavily concentrated between the cave wall and a line of large rocks. The rocks may have supported poles of a tent pitched against the wall. Artifacts and bones spilling out from between the rocks at two points might mark former doorways, while concentrations of small seashells could come from seaweed introduced as bedding. Two roughly circular concentrations of charcoal may define former fireplaces (hearths). The integrity of the plan is questionable, mainly because artifacts and bones within it frequently conjoin to counterparts in underlying and overlying layers (Villa 2004). The Lazaret example illustrates the weakness of the evidence for substantial structures before 50 ka.

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charcoal concentrations (hearths)

floor plan of Le Lazaret Cave indicating area enlarged to the left below

U T

possible internal division

S

R

N Q

P

boulder ?doorway

O

?doorway N 18

17

16

15

14

0

13

12

11

10

9

8

7

5m

concentrations of small shells possibly introduced on seaweed used as bedding

large rocks possibly used as weights to support the poles of a tent pitched against the cave wall

consists of one circular and two oval concentrations of artifacts and fragmentary animal bones (probable food debris) associated with large stones and bones that could have been used to build walls. he concentrations are 2–3 m in diameter, and each is immediately adjacent to a spread of charcoal interpreted as a ireplace. Also nearby are clusters of artifactual “waste” that may represent workshops (ig. 5.57). At Le Lazaret, the presence of a structure is suggested by an 11 × 3.5 m concentration of artifacts and fragmented animal bones bounded by a series of large rocks on one side and by the cave wall on the other (ig. 5.58). he area also contains two hearths, as well as numerous small marine shells and carnivore teeth that could derive, respectively, from seaweed and skins that were introduced as bedding. he rocks could have supported poles over which skins were draped to pitch a tent against the wall of the cave. he apparent pattern is intriguing, but its integrity is questionable, in large part because many of the items it includes conjoin to items that came from overlying and underlying layers. Broadly contemporaneous concentrations of artifacts and other cultural debris surrounded or accompanied by natural rocks in other French caves (especially La Baume Bonne and Orgnac) provide even less persuasive evidence for structural remnants. In sum, the evidence for housing before 130 ka is remarkably sparse and equivocal, and the following chapters show that it becomes abun-

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dant and unambiguous only ater the advent of fully modern humans between 50 and 40 ka. People before 50 ka must have built shelters, particularly at open-air sites in midlatitude Europe and Asia, but the structures were apparently too limsy to leave unmistakable archaeological traces. he more substantial structures that appeared ater 50–40 ka help explain how fully modern humans were able to colonize the most continental parts of Eurasia where no humans had lived before. SOURCES (not including those in table 5.11): putative ruins at Soleihac (Bonifay et al. 1976), Melka Kunturé (Chavaillon 1979), Olorgesailie (Isaac 1977), Latamne (Clark 1968b), and Terra Amata (de Lumley 1969a); possible ruins at Bilzingsleben (Mania et al. 1994), Ariendorf 1 (Turner 1986), Le Lazaret—pro (de Lumley 1975) and con (Villa 2004, 2–5), and La Baume Bonne and Orgnac (Villa 1976)

Artifact Function

Early Paleolithic people undoubtedly used stone artifacts for many purposes, including cutting, whittling, scraping, shredding, and butchering. Damage and wear polish on tool edges can sometimes reveal the mode of use and, more oten, the material to which an artifact was applied. Wear polish analyses show that H. ergaster, early H. sapiens, and early H. neanderthalensis applied tools to essentially the entire range of materials (bone, antler, meat, hide, wood, and nonwoody plant tissue) that the methodology can diferentiate. However, only a small proportion of tools preserve diagnostic use traces, and trace identiication is time consuming. Since there is also no obvious one-to-one relation between tool form and inferred use, the functional signiicance of whole assemblages remains obscure. Flakes and other light-duty tools are much more common than hand axes at some sites where animal bones suggest that butchering was important, which may mean that lakes, rather than hand axes, were the primary butchering tools. At the Campitello Quarry, central Italy, the only artifacts found among the bones of a straight-tusked elephant (Palaeoloxodon antiquus) were three unretouched lint lakes. Associated rodent species suggest that the site formed during OIS 6 (186–127 ka) or before, broadly within the Acheulean interval. Two of the lakes were partly covered with birch-bark tar, and if it is accepted that the tar fastened the lakes to wooden handles, the site provides the oldest known evidence for hating. Hated lakes may have been especially useful for slicing through skin and muscle. However, experiments show that hand axes or other large bifacial cutting tools are at least as eicient as lakes for butchering large animals, and neither the rarity nor the abundance of hand axes probably informs on butchery. he persistence of hand axes and related bifacial tools for more than a million years and their broad distribution across Africa, western Asia,

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and much of Europe must mean that hand axes were central to Acheulean culture, yet their functions remain conjectural, even mysterious. he term hand ax implies that each piece was handheld and used for chopping, but many were far too large and unwieldy for anything like this. he puzzle is heightened at Melka Kunturé in Ethiopia, Olorgesailie in Kenya, Isimila in Tanzania, Kalambo Falls in Zambia, and other sites in eastern and southern Africa, where hand axes occur by the hundreds, oten crowded close together and with no obvious signs of use. Such sites could imply that the hand ax was the Acheulean equivalent of a male peacock’s plumage—an impressive emblem for attracting mates. When a female saw a large, well-made hand ax or other biface in the hands of its maker, she might have concluded that he possessed just the determination, coordination, and strength needed to father successful ofspring. Having obtained a mate, a male might simply discard the badge of his success, alongside others that had already served their purpose. he mate selection hypothesis cannot be disproven, but sites with large concentrations of seemingly unused hand axes are less common than ones where hand axes are rarer and sometimes do show signs of use. Since the tools come in a wide variety of sizes and shapes, they were likely used in various ways. Some of the more carefully shaped, symmetric examples may have been hurled at game like a discus; other, more casually made, pieces may have served simply as portable sources of sharp-edged lakes; and yet others could have been used to chop or scrape wood. As already noted, experiments have also shown that hand axes make effective butchering tools, particularly for dismembering the carcasses of elephants or other large animals. he truth is that hand axes may have been used for every imaginable purpose, and the type probably had more in common with a Swiss Army knife than with a peacock’s tail.

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SOURCES: interpretation of stone-tool wear polish (Keeley 1977, 1980); lakes as the primary Acheulean butchery tools (Clark 1975; Clark and Haynes 1970); Campitello Quarry lakes (Mazza et al. 2006); utility of hand axes for butchering large animals (Jones 1980); hand axes as the Acheulean equivalent of a peacock’s plumage (Kohn and Mithen 1999); hand axes perhaps hurled like discuses (Calvin 2002) or used as cores (Jelinek 1977)

Raw Material Use

Early Paleolithic people preferentially selected locally available rock types with the best laking properties, and they commonly used diferent rock types to produce diferent kinds of tools. hus the occupants of many sites tended to make large heavy-duty core tools out of soter rocks like limestone and basalt, while they produced smaller, more fully shaped core tools and lakes out of harder rocks like lint. Stone artifacts dominate the Paleolithic record because of their durability, but early people certainly used other raw materials, including

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403 FIGURE 5.59. a flaked fragment of elephant bone from Bilzingsleben, Germany (after Mania [1995], fig. 4). Bilzingsleben probably dates to 400–300 ka, and it has provided more than 100 flaked bone artifacts associated with numerous stone artifacts and some fragmentary human remains. Together with flaked bones from other sites, especially in Italy, the Bilzingsleben objects show that early Paleolithic people sometimes flaked bone in the same way they flaked stone. However, only late Paleolithic people after 50 ka routinely carved or ground bone to produce formal artifacts.

bone and highly perishable substances like wood, reeds, and skin. Animal bones survive at many sites of primitive Homo in Africa, Europe, and Asia, and percussion or cut marks sometimes suggest that the people used individual bones as hammers, retouchers, anvils, or cutting boards. In addition, as discussed in the previous chapter, polished tips show that people in the vicinity of Swartkrans, Drimolen, and Sterkfontein Caves, South Africa, 1.8–1.5 Ma used bovid limb bone fragments and horncores to dig in rocky soil or to open termite nests. Also, as noted in the previous chapter, the people who lived at Olduvai Gorge between roughly 1.8 and 1 Ma sometimes laked bone by percussion, in the same way that they laked stone. A few European sites that date between roughly 600 and 300 ka have produced additional laked bones. he most striking examples come from the open-air sites of Fontana Ranuccio, Malagrotta, Castel di Guido, La Polledrara, and Rebibbia-Casal de’Pazzi near Rome, central Italy and from the broadly contemporaneous open-air site at Bilzingsleben, Germany (references in table 5.11). At Fontana Ranuccio, Malagrotta, and Castel di Guido, the laked bone artifacts include unequivocal elephantbone bifaces. At Bilzingsleben, they are also made of elephant bone, but they resemble scrapers or choppers (ig. 5.59). At both the Italian sites and Bilzingsleben, the people may have turned to elephant bone because they lacked access to stone of suitable size and quality. However, despite the variety of laked bone artifacts these sites have produced, neither they nor any other sites of similar or greater age contain bones that were whittled, carved, or polished to form points, awls, borers, and so forth. Such artifacts are common only in sites of fully modern humans ater

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50–40 ka, and chapter 7 argues that fully modern people were the irst to routinely shape bone by cutting, carving, and polishing. Artifacts in the most perishable raw materials are scarce almost by deinition. Bamboo makes edges that rival or exceed those of stone in sharpness and durability, and it is especially likely to have been used in eastern Asia, where it enjoys roughly the same distribution as the chopper–chopping tool tradition deined by Movius. An emphasis on bamboo could explain why east Asian stone-artifact assemblages oten appear typologically impoverished next to contemporaneous Acheulean ones from Africa, western Asia, and Europe, but actual bamboo artifacts remain unknown. Reed and leather artifacts are similarly lacking, and wooden artifacts have been found at only four African, west Asian, and European Acheulean or Acheulean-age sites: a possible club and other shaped objects at Kalambo Falls (Zambia); a wooden fragment apparently polished by use from Gesher Benot Ya’aqov (Israel); a wooden spear point from Clacton-on-Sea (England); and four complete spears and one partial spear from Schöningen (Germany) (references below and in tables 5.11– 5.13). Schöningen has also provided a shorter double-pointed wooden stick of uncertain function and three shaped and grooved wooden branches that may have been handles for stone tools. he sites all date from between 780 and 300 ka ago, and they are distinguished by unusually dense, anaerobic deposits that inhibited bacterial decay. In Africa, the artifact makers were probably early Homo sapiens; in western Asia, they could have been late H. ergaster; and in Europe, they were either H. heidelbergensis or early H. neanderthalensis. he wooden artifacts are mostly nondescript, and their functions are debatable. he Clacton spear point, for example, might actually have tipped a digging stick, a stake, or even a snow probe. Only the Schöningen spears are unequivocal (ig. 5.60). hey are over 2 m long, and they are heaviest and thickest near the pointed ends, like modern javelins. However, they probably could not have been thrown hard enough to penetrate an animal from a distance, and they are thus more likely to have been thrusting spears, employed at close quarters. hey are estimated to be 400 ka old, and they provide the oldest, most compelling evidence for human hunting.

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SOURCES (not including those in table 5.11): diferential early Paleolithic use of harder and soter rocks (Villa 1996); artiicially polished bone artifacts from Swartkrans Mbs 1–3 and Sterkfontein Mb 5 (Backwell and d’Errico 2001; Brain 1985a, 1988; Brain and Shipman 1993), and Drimolen (Keyser et al. 2000); laked bones from Olduvai Gorge (Leakey 1971); early European bone artifacts (Villa and d’Errico 2001); Italian bone bifaces (Gaudzinski et al. 2005; Villa 1991, 2001); possible use of bamboo at east Asian early Paleolithic sites (Bordes 1978; Leng and Shannon 2000; Pope 1993); wooden artifacts from Kalambo Falls (Clark 1969), Gesher Benot Ya’aqov (Belitsky et al. 1991), Clacton-on-Sea (Warren 1911), and Schöningen (hieme 1998); function of the Clacton spear point (Gamble 1986, 1987)

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FIGURE 5.60. multiple views of two of the five wooden throwing spears recovered from 400-ky-old lakeside deposits at Schöningen, Germany (redrawn after Thieme [1996], fig. 9). The spears are the oldest known unequivocal human hunting weapons. They were probably used to hunt horses, whose bones abound in the same deposits.

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throwing spear 1 (recovered in five parts)

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Geographic Variation in Artifact Form

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he only compelling artifactual divide before 250 ka is between the Acheulean biface tradition of Africa, western Asia, and Europe and the chopper–chopping tool (or chopper-chopping-tool-and-lake) tradition of eastern Asia. he diference between east and west endured from the time that people irst reached eastern Asia, more than 1 Ma until beyond 250 ka, and the reason for its persistence is obscure. One possibility, already alluded to above, is that the irst east Asians arrived nearly 2 Ma, before the biface tradition had been invented in Africa. Another is that early on east Asians began using bamboo tools for purposes that bifaces served elsewhere. A third, arguably more plausible, explanation is that migrants on their way to eastern Asia passed through a region that lacked suitable stone for hand ax manufacture, and when they emerged, they had lost the hand ax habit. hereater, isolation by distance maintained the long-standing diference between east and west. It was already noted that geographic diferences in artifact manufacture are diicult to discern within the expansive Acheulean tradition, and Acheulean assemblages tend to be remarkably similar over vast distances. Measurements that deine hand ax shape have shown that, on average, shape difers signiicantly among African, west Asian (Israeli), Indian, and European (English) specimens, but much of the apparent geographic variation might actually relate to diferences in mean geologic age between hand ax samples, to diferences in the size and quality of regionally available raw materials, or to diferences in the intensity of bifacial working (reduction) related to raw material quality. Continued working (rejuvenation or resharpening) tends to transform a hand ax that was initially large, long, pointed, and thick into one that is not only smaller overall but that is also relatively broader, more rounded, and thinner. Diferent degrees of reduction, conditioned by variation in raw material availability and the need for resharpening, will thus lead to signiicant modal diferences in hand ax shape. Whatever the reasons that hand axes difer in average shape among regions, the regions are too large to signal cultural (stylistic) traditions in the ethnographic sense, and the average tendencies probably span intervals that were many times longer than any known historic cultural tradition. Artifactual variation through time and space increased ater 600–500 ka and again ater 250 ka, but geographically and chronologically delineated artifactual units that can reasonably be equated with ethnic traditions or identity-conscious groups are conspicuous only ater 50 ka. SOURCES: bifaces absent in eastern Asia because hominins arrived before they were invented in Africa (Swisher et al. 1994), because east Asians used bamboo tools instead (Pope 1989, 1993), or because people lost the biface tradition on their way to eastern Asia (Schick and Toth 1993); shape diferences among African, west Asian, Indian, and European hand axes (Wynn and Tierson 1990); geographic variation in hand axes due to diferences in reduction intensity (McPherron 2000)

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Artifact Form and Cognition

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Modern novices who attempt to produce hand axes commonly fail because they remove too many lakes from the sides of the blank or preform before they begin to thin it, because they trim some parts of the periphery prematurely (producing an undesirable sinuous edge), because they overcorrect as they attempt to maintain a symmetrical outline (resulting in a highly asymmetrical one), because they cannot extend the peripheral edge through the thickest parts of the blank, or because they break a thinned biface in two by an overly strong blow. he same problems do not afect the production of simple lakes and choppers, and we may reasonably ask whether Acheulean people possessed cognitive abilities beyond those of their lake-and-chopper contemporaries. he answer is: not necessarily. To begin with, the principles involved in hand ax manufacture are easily transmitted and not particularly complex. hey can be learned entirely by imitation and repetition, and they are far simpler, for example, than the intricate and subtle grammatical rules than underlie all known languages. Second, and more important, stone artifacts surely formed only a small part of ancient material culture, and there is at present no basis for assessing the sophistication of the rest. he well-made wooden spears from Schöningen, discussed above, illustrate the point, since they are associated with a lake-and-chopper stone-tool assemblage, yet they relect at least as much forethought and control in manufacture as any hand ax. SOURCES: reasons that modern novices oten fail to produce hand axes (Schick 1994); rules for hand ax manufacture simpler than rules for language (Wynn 1991, 1995)

Art

It has already been noted that the late Acheulean people, who lived ater 700–600 ka, oten produced hand axes that appeal to the modern eye for their remarkable symmetry in both plan form and edge view. he makers shaped these hand axes extensively and meticulously, and there is the possibility that they were guided by an evolving aesthetic sensibility. Alternatively, they might have been driven by the need to produce a piece that was inely balanced, and the possibilities cannot be separated, since, as we have already seen, hand ax function remains largely conjectural. Among other possible indications of art from Acheulean or Acheuleanage sites, three examples stand out: (1) fragments of humanly introduced mineral pigment (ocher) associated with Acheulean artifacts and animal bones at Kapthurin (site GnJh-15), Kenya, at Kathu Pan, Wonderwerk Cave and Duinefontein 2, South Africa, and with like-aged non-Acheulean artifacts and bones at Twin Rivers, Zambia; (2) an elephant tibia shat fragment from Bilzingsleben, Germany, on which a fanlike set of lines has been incised, and (3) above all, a 35-mm long, modiied lava (tuf )

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incised lines

FIGURE 5.61. Left and center: an elephant tibia shaft fragment from bilzingsleben, Germany, with a partial fanlike pattern of incised lines. Right: a reconstruction of what the fanlike pattern might have looked like before the base of the shaft fragment was broken away (left, center, and right redrawn after mania and mania [1988], 93). The regular spacing of the incisions, their subequal lengths, and their v-like cross sections suggest that they were created at the same time with a single stone tool. The shaft fragment is 350–400 ky old, and it is one of only a handful of potential “art objects” from before 40 ka.

0

10 cm

Bilzingsleben pebble from Berekhat Ram on the Syrian/Israeli border that may represent a crude human igurine. At Kapthurin, Kathu Pan, and Duinefontein, artifacts and bones accumulated in the open air near ancient water sources, while at Wonderwerk and Twin Rivers they accumulated in caves. 40Ar/39Ar dating on a superimposed volcanic ash places the humanly collected pigment fragments at Kapthurin before 285 ka, while OSL dating of enclosing sands ixes the fragments from Duinefontein 2 near 270 ka. Mammalian remains and Acheulean artifacts together imply an age near 600 ka for the Kathu Pan pigment fragments. U-series disequilibrium determinations on lowstone suggest that pigment fragments began to accumulate at Wonderwerk by 500 ka and at Twin Rivers by 270 ka or perhaps before. he associated artifacts at Wonderwerk include late Acheulean hand axes, while at Twin Rivers, they have been assigned to a Middle Stone Age variant (the Lower Lupemban Industry) that is commonly thought to succeed the Acheulean in the Twin Rivers region. he Twin Rivers fragments are particularly notable for their number (196 separate pieces), for the variety of their colors, and for indications that some were modiied by grinding or rubbing. At all ive sites, the fragments anticipate those that are well-known at succeeding Middle Stone Age/Middle Paleolithic sites dated between 250 and 50 ka in Africa, western Asia, and Europe. As discussed in the next chapter, people before 50 ka may have employed pigments in hide tanning, medication, and perhaps especially

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deep groove setting off the putative head and neck

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409 FIGURE 5.62. Top: a possible female figurine from the Acheulean site of Berekhat Ram, Golan Heights (drawn by Kathryn Cruz-Uribe from photographs in d’Errico and Nowell [2000], fig. 1). Bottom: The well-known Upper Paleolithic (Gravettian) “Venus” figurine from Lespugue, France (drawn by Kathryn Cruz-Uribe from a cast). Note that the scales are different and that the putative Berekhat Ram figurine is tiny. It is a lava (tuff) pebble on which at least three grooves have been incised with a sharpedged stone tool. The contrast with the Lespugue Venus is stark, but if it is agreed that the grooves were intended to set off a head and arms, the pebble would be the oldest known example of representational art in the world.

5 cm

Lespugue (Upper Paleolithic Gravettian Culture)

as an ingredient in the mastic used to ix stone bits to wooden handles. However, in the absence of contrary evidence, body painting or some other broadly artistic behavior cannot be ruled out. Bilzingsleben is an ancient lakeside site that is well-known for fossil human skull fragments, but it has also produced numerous animal bones and a laked stone-artifact assemblage that lacks hand axes. ESR determinations and U-series dates suggest that the bones and artifacts accumulated sometime between 420 and 350 ka. Berekhat Ram is an ancient lakeside site that does not preserve bone but that has provided a classic late Acheulean artifact assemblage, including eight small hand axes and numerous well-made lake tools. 40Ar/39Ar dates on underlying and overlying basalts bracket the archaeological layer between 470 and 232 ka.

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Based on the stratigraphic position of the artifact-bearing deposits, the excavators favor an age between about 280 and 250 ka. he incised marks on the Bilzingsleben elephant shat fragment are noteworthy for their even spacing (ig. 5.61) and for the extent to which they replicate each other in length and especially in cross-section. he strong similarity in form implies that they were made in quick succession by a single stone tool, and neither their placement nor their form suggest butchery marks. hey could, however, relect repeated slicing of a sot substance that was laid over the shat fragment, and they may even have originated naturally when a large animal trampled the fragment on a sandy or gritty substrate. Elephant trampling of bones near modern African waterholes occasionally creates similar clusters of subparallel cut mark mimics. he Berekhat Ram pebble is remarkable for three distinct grooves— a deep one that encircles the narrower, more rounded end, setting of the putative head and neck, and two shallow, curved incisions that run down the sides and that could delineate the arms (ig. 5.62). he deep groove and, to a lesser extent, the shallower ones closely match experimental marks produced by sharp-edged lakes in the same material, and they are readily distinguishable from natural lines. he Berekhat Ram pebble was thus humanly modiied, even if its inal form only dimly anticipates the inely crated, aesthetically attractive human igurines that appear in Europe and elsewhere ater 40 ka. As discussed in the next chapter, objects that exhibit the same degree of possible artistic intent also occur sporadically in sites occupied by the Neanderthals and their near-modern African contemporaries between roughly 130 and 50 ka. However, objects whose artistic meaning is unequivocal become commonplace only ater 50 ka, when they are associated with the origins and spread of fully modern humans from Africa.

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SOURCES: mineral pigment at Kapthurin GnJh-15 (Tryon and McBrearty 2002), at Wonderwerk Cave and Kathu Pan (Beaumont and Vogel 2006), at Duinefontein 2 (Cruz-Uribe et al. 2003), and at Twin Rivers (Barham 2002); Bilzingsleben incised elephant bone (Mania and Mania 1988), and Berekhat Ram igurine (Goren-Inbar and Peltz 1995); pigment in Middle Stone Age/Middle Paleolithic sites in Africa (Henshilwood et al. 2002), western Asia (Hovers et al. 2003), and Europe (d’Errico 2003); Bilzingsleben incisions possibly produced by slicing of an overlying sot substance (Davidson 1990) or by trampling (d’Errico 1995); cut-mark mimics produced by elephant trampling (Haynes 1988, 1991); stone-tool marks on the Berekhat Ram pebble (d’Errico and Nowell 2000); possible art objects in sites of Neanderthals and their African contemporaries (d’Errico 2003)

Language

he complexity of late Acheulean hand ax manufacture and the sophistication of the Schöningen spears show that early Homo sapiens and H. neanderthalensis were cognitively far advanced over the australopiths, and they probably possessed a much more advanced form of commu-

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nication. But direct evidence for language is lacking, and it is diicult even to imagine what intermediate stages might have been like. Historically observed languages provide no clues, since they are all equally complex and equally far removed from the communication systems of other animals. he next two chapters employ archaeological evidence to argue that only fully modern humans ater 50 ka may have possessed fully modern language ability and that the development of this ability may even have underlain their emergence. By extrapolation, one could argue that the anatomical and behavioral advances that mark Homo ergaster as the irst truly human species imply that it was also the irst to possess rudimentary language. hree anatomical regions can be brought to bear on this. First is the brain or, more precisely, its size and coniguration. Size speaks mainly to potential, and on this basis H. ergaster is perhaps more likely to have had language than earlier species. Coniguration is arguably less ambiguous because two regions (Broca’s and Wernicke’s areas) on the external cerebral cortex are linked to motor control over speech, and both are detectable on endocranial casts. Endocast incompleteness or distortion oten hinders observation, and reliable detection is particularly diicult for Wernicke’s area. Wernicke’s area also appears to be well-developed in common chimpanzees, which of course totally lack the human ability to speak. Broca’s area is less problematic, and it appears to have been absent or relatively undeveloped in the australopiths and much better developed in H. habilis and H. ergaster. Some degree of humanlike speech may be implied. he remaining two lines of anatomical evidence are contradictory. On the negative side, there is the small diameter of the neural canals on the vertebrae of an adolescent skeleton (KNM-WT 15000) from Nariokotome, West Turkana, Kenya. he neural canals enclose the spinal cord with its nerve cells and nerve ibers, and small neural canal size in thoracic (chest) vertebrae implies limited nervous control over movements of the rib cage. Fine control is essential for articulate speech. On the positive side, there is the moderate degree of basicranial lexion (upward arching between the posterior edge of the palate and the anterior margin of the foramen magnum) that has been observed on H. ergaster skulls. In the degree of basicranial lexion, H. ergaster was apparently intermediate between the australopiths, who had very lat cranial bases (like those of extant apes and very young humans), and modern adult humans, who have highly lexed bases. In extant primates, the degree of lexion is related to the anatomy of the upper respiratory tract, including the position of the larynx, which rests much lower in the neck of modern human adults (with pronounced lexion) than in the neck of apes (with limited lexion). he low position of the larynx is crucial to the modern human ability to produce complex sound (phonemic)

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sequences in a single, short breath, and this in turn is crucial to fully modern speech. he moderate basicranial lexion in H. ergaster may thus suggest at least an incipient capacity for true speech. Since the basicranium is unknown in unequivocal specimens of H. habilis, H. ergaster is the earliest hominid for which basicranial lexion suggests a degree of articulate speech.

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SOURCES: role of Wernicke’s and Broca’s areas in speech (Cooper 2006; Lieberman 2006); Wernicke’s area—detection in fossils (Walker and Leakey1993b), presence in chimpanzees (Gannon et al. 1998); Broca’s area in fossils (Begun and Walker 1993; Tobias 1991); neural canal diameter in KNM-WT 15000 (MacLarnon 1993; Walker 1993); basicranial lexion in H. ergaster (Laitman 1985); laryngeal anatomy (Nishimura et al. 2006)

Fire

Homo ergaster (or H. erectus) may have had to master ire for warmth and cooking before it could colonize Eurasia, but direct archaeological support is tenuous. he oldest evidence (cited in the previous chapter) may be patches of baked earth in deposits dated to 1.5–1.4 Ma at Koobi Fora and Chesowanja in Kenya, but naturally ignited, smoldering vegetation might have produced the same results. Natural ires or ignition could also account for burning at most other early Paleolithic sites, including burned bones associated with Developed Oldowan or early Acheulean artifacts in Member 3 at Swartkrans Cave in South Africa, patches of burned earth at the Olorgesailie Acheulean site in Kenya, burned bat guano underlying the Acheulean layers at the Cave of Hearths in South Africa, dispersed ash in the Acheulean deposits at the Cave of Hearths and Montagu Cave, also South Africa, burned lint artifacts in the Acheulean site of Terra Amata in France, and dispersed charcoal in the Acheulean layers of Torralba and Ambrona in Spain and in the non-Acheulean site of Přezletice in the Czech Republic. he Přezletice charcoal may be irrelevant if, as noted above, the associated laked stones are not artifactual. At the other extreme, the Swartkrans burned bones may be particularly noteworthy because they are absent in older Swartkrans units where they might also be expected if they were produced by natural burning. If the Swartkrans bones relect human combustion, they would imply that H. ergaster possessed ire by 1.5 Ma, and the use of ire by its geographically scattered descendants might not require independent invention. Swartkrans Member 3 aside, the most persuasive evidence for early human use of ire comes from the ancient lakeside site of Gesher Benot Ya’aqov, Israel, and from the classic H. erectus cave of Zhoukoudian Locality 1, China. At Gesher Benot Ya’aqov, Acheulean horizons, ixed by paleomagnetism to the millennia following 780 ka, contain discrete clusters of burned lint chips surrounded by spreads of mainly unburned

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artifacts, bones, and even vegetal remains. A plausible explanation for the burned chip clusters is that they formed in and around Acheulean ireplaces. At Zhoukoudian Locality 1, where the ill is tentatively dated between 670 and 400 ka, layers 10 and 4 contain thin dark seams interpreted as fossil hearths together with numerous charred bones. Geochemical analysis has failed to identify wood ash or charcoal in the dark layers, but the burned bones have been conirmed, and the frequency of burned bones (about 12% of the total) recalls their frequency in much later (west Eurasian Mousterian) sites, where ireplaces are well-documented. As at later sites, the Zhoukoudian specimens are also directly associated with numerous artifacts, which increases the likelihood that deliberate ire produced the burning. In Europe, ire indications that may rival those of Zhoukoudian in age include concentrations of charred bones in shallow depressions at Vértesszöllös, Hungary; clusters of wood charcoal at Bilzingsleben, Germany; a reddened patch of sediment at Schöningen, Germany; burned artifacts, burned microvertebrate bones, and multiple patches of burned and baked sediment at Beeches Pit, England; and two “built ireplaces,” a charcoal illed pit, and burned quartz artifacts from Menez-Dregan, France. U-series determinations, ESR dates, TL readings (on burned lints), mammalian species, fossil pollen, amino-acid racemization of land snail shells, or some combination of these tentatively place each site at 400 ka, give or take a few tens of thousands of years (table 5.11). Like the much more ancient Kenyan sites with patches of baked earth, the European sites are all open air localities where natural burning cannot be excluded, but in the deeply stratiied cave ill at the Caune de l’Arago, France, where natural burning is less likely, burned objects appear only in layers provisionally dated to about 400 ka. hey are absent in layers dated to between 700 and 400 ka and also in broadly coeval deposits at Atapuerca GD, including the layer (TD-6) that has provided the oldest persuasive evidence for cannibalism, discussed below. he implication may be that Europeans began to control ire only about 400 ka, and that little or no control before this explains why they were apparently conined to southern Europe during glacial periods older than OIS 8, between 301 and 242 ka. he oldest conirmed full glacial occupations date from the OIS 6 (the Penultimate Glaciation), between roughly 186 and 130 ka, when ire use is implied by patches of ash and charcoal, abundant burned objects, or both at Le Lazaret, Pech de l’Azé, and perhaps other French caves and by abundant carbonized bones, less abundant charcoal, and numerous burned lint artifacts at the cave of La Cotte de St. Brelade (Jersey). Spreads of burned sediment, ash, and charcoal that almost certainly signal ireplaces are conspicuous in many sites occupied by the European Neanderthals and their near-modern African contemporaries ater

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130 ka, and it is generally assumed that people everywhere ater 130 ka could make ire when they needed it. he question then is when this ability evolved or, perhaps more precisely, whether a stage of full control followed on one where ire use was sporadic and opportunistic. he issue is diicult to address, since sites older than 130 ka are relatively rare and they are mostly open-air localities. Sites younger than 130 ka are not only more abundant, but they occur much more oten in caves. he diference relects the limited average lifespan of cave ills, noted previously, and it is crucial because caves are far more likely to preserve fossil ireplaces. he diiculty of separating natural from human burning at ancient open-air sites and the rarity of equally ancient caves means that reconstructing the history of human ire use may always depend more on speculation than on archaeological discovery.

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SOURCES: evidence for ire at Koobi Fora and Chesowanja (Clark 1955; Gowlett et al. 1981; Isaac 1982), Swartkrans Mb 3 (Brain 1993d; Brain and Sillen 1988; Sillen and Hoering 1993), Olorgesailie (Isaac 1977), Cave of Hearths (Butzer 1973a; Mason 1962), Montagu Cave (Butzer 1973a), Terra Amata (de Lumley 1969a; Villa 1983), Torralba and Ambrona (Howell 1966), Přezletice (Fridrich 1976), Gesher Benot Ya’aqov (Goren-Inbar et al. 2004), Zhoukoudian (Pei and Zhang 1985, Weiner et al. 1998; Zune 1985), Vértesszöllös (Dobosi 1988), Bilzingsleben (Mania et al. 1994); Schöningen (hieme 1997), MenezDregan (Monnier et al. 1994), Beeches Pit (Gowlett 2006), Arago (de Lumley 2006), Pech de l’Azé (Villa 1976), Le Lazaret (de Lumley et al. 2004); and La Cotte de St. Brelade, Jersey (Callow et al. 1986)

Diet and Ecology

Like ire, advances in the ability to obtain food could explain how Homo ergaster and its descendants managed to colonize new regions. However, the evidence for such advances is extremely limited. Both logic and observations on historic hunter-gatherers suggest that, in general, early people everywhere depended more on plants than on animals. Tubers and other underground storage organs may have been especially important since they were staples among historic low- and midlatitude huntergatherers. It may not be coincidental that H. ergaster emerged at a time 1.8–1.7 Ma when tubers had probably become more abundant, following a shit to yet drier, more seasonal climate over much of Africa. As indicated previously, key features of H. ergaster anatomy and behavior suggest it was especially adapted for foraging in arid, highly seasonal environments. he evolution of H. ergaster (or African H. erectus) has sometimes been tied to males’ enhanced ability to hunt, but it may actually have depended more on females’ enhanced ability to locate, excavate, and process tubers. Unfortunately, plant remains that could illuminate the importance of tubers or other potential plant foods rarely preserve in early human sites. In addition, in the few very early sites where remnants of edible plants do occur, they cannot be unequivocally linked to human activity. he sites include Locality 1 at Zhoukoudian in China; the Acheulean

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layers at Kalambo Falls in Zambia, Gesher Benot Ya’aqov (GBY) in Israel; and Kärlich-Seeufer in Germany; and the non-Acheulean deposits at Bilzingsleben in Germany and Vértesszöllös in Hungary (references below and in tables 5.11–5.13). he case for human consumption is strongest at GBY because the plant remains include specimens from seven diferent nut-bearing species, and the accompanying artifacts include pitted stones that could have been used to crack nutshells. A human origin for the pitting is indisputable, and if it resulted in whole or in part from nut cracking, similar pitted stones at other Acheulean sites may imply nut eating even though plant fossils are absent. Pitted stones like those GBY are especially abundant in Bed IV and the Masek Beds at Olduvai Gorge in Tanzania, and they are also known at other African sites, particularly Melka Kunturé and Gadeb in Ethiopia (references in table 5.13). Given the failure of plant tissues to preserve at the vast majority of sites, dietary reconstruction for early Homo depends on an unusual partial skeleton, KNM-ER 1808, from Koobi Fora, Kenya, on the stable-carbon isotope analysis of dental enamel, on molar microwear, and above all, on the animal bones found at many sites. KNM-ER 1808 is dated to roughly 1.7–1.6 Ma, and it comes from Homo ergaster. Its long bone shats are covered by a layer of abnormal, coarse-woven bone as much as 7 mm thick. A toxic excess of vitamin A (hypervitaminosis A) could be responsible, and if so, it might relect an overindulgenence on carnivore livers or on honeybee eggs, pupae, and larvae. Alternatively, it might mark the oldest known case of yaws, an infectious disease related to syphilis that induces similar bone growth in its inal stage. he discussion of Australopithecus africanus in the previous chapter outlined the principles behind stable carbon-isotope analysis. As applied to teeth of probable Homo ergaster from Swartkrans Cave, South Africa, it suggests a broad diet that could have incorporated seeds and rhizomes from grassy plants, together with fruits, nuts, and other edible parts of nongrassy plants. It might also have included the tissues of animals, including insects, that eat various kinds of plants. In its substantial dietary breadth, H. ergaster difered from various herbivores that tend to specialize on grasses or nongrasses, but not from A. africanus and Paranthropus robustus, both of which appear to have had at least equally varied diets. As noted in the previous chapter, the three species may have inherited their shared dietary eclecticism from a common ancestor ater 3 Ma, but this doesn’t mean they ate the same mix of foods. In particular, the reliance of H. ergaster on stone-artifact technology and its broad distribution through more open, arid environments suggests that it probably fed more on meat and marrow. he discussion of Australopithecus anamensis in the previous chapter summarized the principles behind microwear interpretation. In

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general, the molars of living primates that prefer nuts, seeds, or other hard, brittle items exhibit a higher frequency of large microwear pits, while the molars of primates that focus more on soter, tougher tissues, including leaves, tubers, and lesh, exhibit a greater number of striations (microscratches). hey also have fewer and smaller microwear pits. Microwear on fossil teeth is oten obliterated in the ground, and so far, it has been reliably observed on only nine molars of undoubted or probable Homo habilis (narrowly deined to exclude H. rudolfensis) and nine of certain or probable H. ergaster. he comparison suggests that H. ergaster consumed a higher proportion of tough, ibrous items, but microwear analysis cannot distinguish between meat and plant foods of similar toughness. Stone-artifact technology may imply that H. ergaster was more carnivorous, but based on microwear alone, it could also have relied heavily on the edible underground parts of plants, or on a combination of these and meat. Animal bones accompany the artifacts of primitive Homo at numerous sites, including Duinefontein, Elandsfontein, and Kathu Pan in South Africa; Olduvai Gorge and Isimila in Tanzania; Olorgesailie in Kenya; Terniine in Algeria; ‘Ubeidiya, GBY, Revadim, and Holon in Israel; Torralba, Ambrona, and Aridos in Spain; Venosa-Notarchirico, La Polledrara, Isernia la Pineta, and Rebibbia-Casal de’Pazzi in Italy; Bilzingsleben, Schöningen, Miesenheim 1, and Bad Cannstatt in Germany; and Boxgrove and Hoxne in England (references in tables 5.11–5.13). Until the 1970s, archaeologists commonly assumed that the stratigraphic association implied a functional relationship and that the artifacts were used to kill and butcher the animals. It followed that the people were accomplished big-game hunters, since the bones oten came from elephants, rhinoceroses, bufalo, and other formidable prey. In the 1970s, however, a growing number of archaeologists began to specialize in the analysis of animal bones, and ater close scrutiny it was soon realized that by itself, a stratigraphic association between artifacts and bones implies nothing about human ability to hunt and butcher. Most early archaeological sites, including all those listed above, formed in the open air near ancient springs, streams, or lakes that naturally attracted both people and animals. he animal bones at such sites could represent human kills, or they could represent carnivore kills or even natural deaths (from starvation, disease, etc.) that totally escaped human notice or that were subsequently scavenged by people (or carnivores). Separating the alternatives has required new approaches, above all the analysis of bone damage. he Elandsfontein Acheulean site, South Africa illustrates the point (references below and in table 5.13). Elandsfontein is especially well-known for a human skullcap assigned here to early Homo sapiens or H. heidelbergensis, but it has also provided more than 160 late Acheulean bifaces, thousands of associated

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“bushpig” (1.8%)

“warthog” (0.6%)

hippo (0.8%)

sivathere (1.1%) small bovids (7.2%)

white rhino (2%)

small-medium bovids (3.1%)

black rhino (1.8%) quagga (1.0%) Cape zebra (16.6%)

417 FIGURE 5.63. The representation of different kinds of large mammals at the elandsfontein acheulean site, based on the minimum numbers of individuals (mni) from which their bones must come. The percentages were calculated on a total mni of 712. figures 5.64 and 5.65 illustrate the proportional representation of bovids and carnivores, respectively.

large-medium bovids (29.1%)

elephant (2.0%)

carnivores (13.1%) gelada baboon (0.7%)

large bovids (10.0%) very large bovids (9.8%) pangolin (0.1%)

hare porcupine (0.7%) (0.3%)

Elandsfontein Main large mammals lake tools and laking debris, and nearly 13,000 iron-mineralized mammalian fossils. he bones come from forty-eight species (excluding the hominin), iteen of which have no historic descendants. Elandsfontein cannot be dated directly, but comparison of the fauna to radiometrically dated faunas in eastern Africa implies an age between 1 Ma and 600 ka. he morphology of the human skullcap (ig. 5.30) and the reined character of the Acheulean artifacts (ig. 5.51) suggest the true age is closer to 600 ka. he fauna includes a wide range of herbivores, from small antelopes to rhinoceroses and elephants, and a broad array of carnivores, from mongooses to hyenas and lions (igs. 5.63–5.65). he bones and artifacts accumulated on an ancient land surface that was subsequently covered by windblown sands that originated at the nearby coast. In 1906 or before, the local vegetation was disturbed or removed, and strong southerly winds whipped the sands into dunes over an area of about 1.6 × 3 km. he wind exposed the ancient land surface, together with its bones and

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FIGURE 5.64. The representation of bovid (antelope and buffalo) species at the elandsfontein acheulean site, based on the minimum number of individuals (mni) from which their bones must come. The percentages were calculated on a total bovid mni of 421.

“spiral horn” (1.7%)

long-horned buffalo (12.4%)

eland (12.8%)

greater kudu (1.9%) gemsbok (1.7%)

Cape grysbok (12.1%)

“giant hippotragine” (4.3%)

southern springbok (1.7%)

blue antelope (1.4%)

Reck’s springbok (1.0%)

reedbuck (12.4%)

gazelle (2.6%)

Arambourg’s hartebeest (10.9%) tsessebe-like antelope (4.5%) ?Damaliscus niro (3.3%)

“giant hartebeest” (1.2%) black wildebeest (10.9%)

?Parmularius sp. (1.2%)

?Damaliscus sp. nov. (2.1%)

Elandsfontein Main bovids artifacts, in the hollows or “bays” between the dunes. In one small area, excavation recovered objects still covered by intact sands, but about 90% of the bones and artifacts were collected from the exposed land surface. Collection occurred mainly in the 1950s and 1960s, and few records were kept of how the bones and artifacts were distributed. However, recently observed bone scatters tend to be dominated by axial parts, particularly skulls, vertebrae, and ribs, as opposed to appendicular or limb bones. he same pattern characterizes the broadly similar, somewhat younger Duinefontein 2 Acheulean site, about 45 km to the south, where wind erosion (delation) has been less severe, and the bones and artifacts remain mostly sealed in place. A reasonable inference at both sites is that the bone scatters represent the remains of carcasses from which the limb elements were largely removed. he abundance of reedbuck (Redunca arundinum) and hippopotamus (Hippopotamus amphibius) and the geochemical nature of the Elandsfontein sediments show that the bones and artifacts accumulated in and around a large marsh or pond. he herbivore bones come mostly from grazers (grass-eaters), especially a large zebra (Equus capensis), an extinct long-horned bufalo (Pelorovis antiquus), and seven diferent species of hartebeest/wildebeest-antelopes (Alcelaphini). Browsing species (ones that feed mainly on bushy plants) are less well-represented, but they include black rhinoceros (Diceros bicornis), kudu (Tragelaphus strep-

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leopard (3.7%)

419 FIGURE 5.65. The representation of carnivore species at the elandsfontein acheulean site, based on the minimum number of individuals (mni) from which their bones must come. The percentages were calculated on a total carnivore mni of 81.

dirk-toothed cat (2.5%)

lion (9.9%) black-backed jackal (22%)

caracal or serval (14.8%)

Cape fox (4.9%)

wildcat (2.5%)

hunting dog (2.5%) polecat (1.2%) honey badger (7.4%)

brown hyena (11.1%)

civet cat (1.2%)

spotted hyena (1.2%)

suricate (6.2%) water mongoose Egyptian mongoose (2.5%) (1.2%)

Elandsfontein Main carnivores siceros), and sivathere (Sivatherium maurusium, an extinct ox-bodied, short-necked relative of the girafe). he sum suggests a vegetation dominated by grass swards with smaller patches of bush. he abundance of bifaces and other artifacts among the bones might imply that Acheulean people routinely hunted or scavenged the large mammal species, and this could explain why axial parts outnumber limb bones. In general, limb bones carry more meat and marrow, and human hunters or scavengers might have taken them away to avoid competition with hyenas, lions, or other carnivores. However, lions and especially hyenas might have done the same, and numerous hyena coprolites (fossil feces) document hyena visitations. he analysis of bone damage ofers an opportunity to distinguish the alternatives. To assess damage, Milo (1994) microscopically examined the surfaces of all the available limb bones and larger tarsals (ankle bones) from wildebeest- and eland-sized antelopes. he samples comprised 1,092 and 516 specimens, respectively, for a total of 1,608, and the particular elements involved were precisely those on which either people or carnivores are most likely to have let telltale damage marks. Microscopic

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examination was critical because the naked eye may miss many of the relatively subtle marks that stone tools and carnivore teeth can produce. Milo found that 2% of the wildebeest-sized bones and 5% of the elandsized bones exhibited tooth marks from the African porcupine (Hystrix africaeaustralis), a large rodent that gnaws dried bones for their calcium and phosphates. he comparable igures for carnivore tooth marks were 1.6% and 1.4%, and for stone-tool marks only 0.7% and 0.2%. Tool marks were not only rare, but mostly ambiguous, and the conclusion is inescapable that despite the abundance of Acheulean tools, the toolmakers played little role in the Elandsfontein bone accumulation. It seems likely thus that the bone-artifact association originated mainly from natural mortality near a water source that independently attracted people and other large mammals. he rarity of tool-marked bones, further, may also mean that local Acheulean people obtained few of the available large mammals, whether by scavenging or hunting. It is diicult to determine if Elandsfontein represents an Acheulean rule or the exception because there are few similar “death” or “carcass” sites from which Acheulean people could have removed bones (to be distinguished from camp sites to which they might have brought them), and fewer yet have been analyzed for bone damage. At most sites for which damage numbers are available, including Duinefontein 2 (South Africa), Hoxne (England), Cagny l’Épinette (France), Torralba and Ambrona (Spain), and ‘Ubeidiya (Israel), stone tool–marked bones are no more common than at Elandsfontein, and a possible inference again is that local Acheulean people had little impact on the associated large mammal community. he overall incidence of tool-marked bones is also low at Boxgrove, England, but one unit (4b) produced numerous bones with cut marks and impact fractures, mostly from a single horse that was apparently butchered on the spot. Incompletely reported damage observations from the broadly contemporaneous (400ky-old), non-Acheulean locality at Schöningen, Germany, may imply horse butchery on a wider scale. he wooden spears from Schöningen, discussed above, demonstrate that humans actively hunted during the Acheulean interval, but they do not show how successful the hunters may have been. Bone damage aside, limited Acheulean ability to obtain large mammals might be inferred from the relatively low level of Acheulean technology and from the likely small size of Acheulean populations. he question then is whether the frequency of tool-marked bones at Acheulean or Acheulean-age death or carcass sites can provide conirmation. A compelling answer will require observations from a much larger number of sites to establish both the average frequency of tool-marked bones and the extent of deviation from the average. It will probably also require greater consensus on the recognition of tool marks. As noted in the

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previous chapter, experts, perhaps especially those who rely on microscopy, oten disagree on the identiication criteria, and their tool-mark estimates for the same bone assemblage can thus difer signiicantly. Part of the problem, emphasized for key European sites, is that cut marks can be diicult to distinguish from striations produced by trampling or sediment compaction, and impact fractures produced by stone tools can be diicult to distinguish from fractures produced by hyena teeth. he separation oten depends less on mark form than on sedimentary context, as at Boxgrove, where the most obviously tool-marked bones come from ine-grained, low-energy deposits in which natural abrasion is unlikely to have produced cut-mark mimics. If it is eventually conirmed that tool-marked bones tend to be rare at Acheulean or Acheulean-age death or carcass sites and if limited Acheulean ability to obtain large mammals is accepted as the probable reason, it would follow that tool-marked bones should, on average, be more common in comparable post-Acheulean death or carcass sites. Unfortunately, if such sites exist, they remain unreported, in contrast to well-known post-Acheulean cave occupations. Tool-marked bones usually abound in such caves, and their abundance probably implies active hunting. In itself, it does not mean that the people hunted more successfully than their Acheulean predecessors, but those who lived ater 50–40 ka almost certainly did, based on their more sophisticated technology and on indications of greater population density (discussion in chap. 7). Acheulean-age cave occupations are even rarer than carcass or death sites, mainly because caves themselves have a limited lifespan, and most of those that existed in Acheulean times have either collapsed or were long ago lushed of their deposits. he small number that survive intact, notably including Atapuerca GD in Spain, Arago Cave in France, and Zhoukoudian Locality 1 in China, contain animal bones that people probably introduced. Hyenas may have also have introduced some, particularly to Zhoukoudian 1, but stone-tool marks and fractures imply that people introduced all, or nearly all, to Atapuerca GD (discussed in the section titled “Cannibalism and Interpersonal Violence,” immediately below). Atapuerca GD conirms human interest, perhaps even dependence, on meat and marrow, but it does not necessarily imply substantial hunting or scavenging success. An Amudian (blade-rich Acheuleo-Yabrudian) layer at Qesem Cave, Israel, perhaps provides the best case for frequent hunting, followed by butchery. he layer formed sometime between 380 and 200 ka, and it has provided artifacts and bones that are unusually fresh looking (chemically and physically unaltered) for a cave occupation of such antiquity. he freshness of the artifacts is particularly remarkable, and it means that many exhibit unambiguous traces of wear from use. he next chapter discusses the wear traces on tools in much later Mousterian sites,

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dated ater 130 ka, and it points out that these result mainly from woodworking not butchery. At Qesem Cave, the reverse is true, and the abundance of butchering tools coincides with an unusually high proportion of tool-marked bones. hese are about twice as common as they are in any subsequent regional Mousterian or Upper Paleolithic site, and the artifacts and bones together suggest that Qesem Cave may have been largely dedicated to butchery, particularly of fallow deer (Dama sp.). Even if, as seems likely, plants dominated Acheulean diets almost everywhere, in part because Acheulean people rarely obtained carcasses of large mammals, Qesem Cave shows that meat was important in late Acheulean-age diets and it foreshadows abundant evidence for butchery by the Neanderthals and their African Middle Stone Age contemporaries ater 130 ka. As already indicated, the Neanderthals and their contemporaries commonly introduced large numbers of bones to cave sites, and numerous cut or percussion marks prove that the bones represent food debris. he ages of animals at time of death sometimes imply driving in Africa and in Europe, but authorities have disagreed vigorously on the relative importance of hunting versus scavenging. However, unlike the bones of early Homo, which rarely if ever retain protein (collagen), the bones of Neanderthals more commonly do, and stable-isotope analyses of residual protein, presented in the next chapter, imply that the Neanderthals relied heavily on meat. It is diicult to imagine that scavenging alone could have met their requirements, and the issue is then not so much whether they actively hunted, but how successful they were. Observations summarized in the next chapter imply that whatever means their southern African contemporaries used to obtain large mammals, they did not obtain them very oten. If it is fair to project backward, the more ancient people on which this chapter focuses were even less successful, and this supports the tentative conclusion from bone damage that they rarely fed on the bufalo, rhinoceroses, elephants, or other large animals whose bones accompany their artifacts at sites like Elandsfontein.

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SOURCES: possible relationship between the emergence of H. ergaster and an increase in underground storage organs (O’Connell et al. 1999); plant remains from Zhoukoudian (Chia 1975), Kalambo Falls (Clark 1969), Gesher Benot Ya’aqov (Goren-Inbar 1992), Kärlich-Seeufer (Gaudzinski et al. 1996), Bilzingsleben (Mania et al. 1994), and Vértesszöllös (Dobosi 1988); pitted stones with plant remains at Gesher Benot Ya’aqov (Goren-Inbar et al. 2002); KNM-ER 1808 skeletal pathology (Walker et al. 1982) possibly from yaws (Rothschild et al. 1995) or from hypervitaminois A (Skinner 1991); stable carbon–isotope analysis of early hominin teeth (Lee-horp et al. 2000; van der Merwe et al. 2003); microwear in H. ergaster and H. habilis (Ungar et al. 2006a, 2006b); Elandsfontein (Klein et al. 2007); damage on bone surfaces at Duinefontein 2 (Cruz-Uribe et al. 2003; Klein et al. 1999a), Hoxne (Stopp 1993), Cagny l’Épinette (Dibble et al. 1997), Torralba and Ambrona (Shipman and Rose 1983b; Villa et al. 2005b), ‘Ubeidiya (Gaudzinski 2004b, 2004c), Boxgrove (Paritt and Roberts 1999), and Schöningen (Roebroeks 2001; hieme 1997, 1998); discrepancies among observers in tool-mark frequencies (Lupo and O’Connell 2002); diicultly of distinguishing stone-tool marks from natural marks (Gaudzinski 1999b); probable hyena contribution at Zhoukoudian Locality 1 (Boaz et al. 2004); butchery indications at Qesem Cave (Lemorini et al. 2006) and in Mousterian/Middle Stone Age sites (Chase 1988; Marean and Kim 1998; Milo 1994, 1998; Stiner 1994); evidence for animal driving in the

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Middle Stone Age (Klein 1979) and Mousterian (Levine 1983); Mousterian scavenging (Binford 1985) vs. hunting (Chase 1988); stable-isotope evidence for Neanderthal diet (Bocherens et al. 1999, 2001, 2005; Fizet et al. 1995; Richards et al. 2000); relative ineiciency of Middle Stone Age hunters (Klein 1994)

Cannibalism and Interpersonal Violence

Cannibalism might be indicated by cut marks on the principal Bodo skull, Ethiopia (approximately 600 ky old), by arguably intentional removal of the rear part of the Arago skull, France (425–400 ky old), by possible cut-marks on skull fragments from Castel de Guido, Italy (325 ky old), and by possibly intentional removal of the cranial bases on the Ngandong (Solo) skulls, Java (200–50 ky old). However, deleshing without consumption is possible for Bodo, and postdepositional destructive forces could explain Arago, Castel de Guido, and Ngandong. For the moment, only Atapuerca GD layer TD 6 provides a compelling case for cannibalism among early Homo. Recall that TD 6 is dated to roughly 800 ka and that it ofers some of the most persuasive evidence for human presence in Europe before 600 ka (references in table 5.11). he TD 6 human fossils that have been described so far include eighteen skull fragments, ive partial jaws, fourteen isolated teeth, sixteen vertebrae, sixteen ribs, twenty bones of the hands and feet, two bones of the wrist, three collar bones, two radiuses, a femur, two patellas, and other fragments from a minimum of seven individuals. he human bones were broken and scattered more or less randomly among the animal bones within TD 6, and more than 25% of the human remains show one or more forms of humanly caused damage. hese include chop and cut marks where large muscles were severed or stripped away; roughened surfaces with parallel grooves or a ibrous texture that relects “peeling,” when a bone was partially broken by a blow and then bent across the break to separate the pieces; and percussion marks made when a bone was splintered for marrow extraction. Tool marks are about equally common on the nonhuman bones, and allowing for diferences in anatomy, the positioning of the marks on the human and nonhuman bones is broadly similar. he abundance and arrangement of the damage marks implies that the TD 6 people butchered other people for food and not for ritualistic purposes, and it is tempting to draw a parallel with the situation on Easter Island when Europeans irst arrived in the eighteenth century a.d. he Easter Islanders had severely degraded their environment, and their once-thriving population had shrunk by 80%. In desperation, the survivors had adopted a wide range of bizarre behaviors, including dietary cannibalism. In the short run, this helped some to carry on, but in the long term, it could only have hastened a slide toward population extinction. If cannibalism at TD 6 relects similar nutritional stress, it could

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explain why the TD 6 people ultimately failed in their attempt to colonize Europe. Neanderthal sites (discussed in the next chapter) have also occasionally provided evidence for cannibalism, but if the custom led to extinction, if afected only local populations. Still, as far as we know, the great apes do not turn to cannibalism when food is short, and the records from TD 6, the Neanderthals, Easter Island, and some late prehistoric sites in Europe and the American Southwest suggest that dietary cannibalism may be a specialized human tendency that the TD 6 people, the Neanderthals, and modern humans inherited from their last shared ancestor. Cannibalism by itself suggests interpersonal violence, and this may be further indicated by healed fractures and other possible predepositional damage to the Ngandong skulls. he Neanderthals also frequently broke their bones before death (discussion in the next chapter), but accidents rather than violence could be responsible, and the least equivocal inference is for a physically strenuous lifestyle. he main Kabwe skull (ig. 5.29), Zambia (?400 ky old), exhibits a partially healed perforation on the let temporal that could have been made by a pointed weapon. Arguably however, it is more consistent with a carnivore bite or with the lesion from a small tumor.

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SOURCES: evidence for cannibalism at Bodo (White 1985, 1986c), Arago (de Lumley 1975), Castel de Guido (Mariani-Costantini et al. 2001), and Atapuerca TD 6 (Fernández-Jalvo et al. 1999); postdepositional destruction at Ngandong (Dennell 2006); tool marks on nonhuman and human bones at Atapuerca TD 6 (Diez et al. 1999); Easter Island (Kirch 1984); cannibalism in late prehistoric Europe (Villa 1992) and the American Southwest (White 1992); antemortem damage to the Ngandong skulls (Coon 1962, 301–302); Kabwe skull perforation—from pointed weapon (Keith 1928), a carnivore bite (Tappen 1987), or a tumor (Montgomery et al. 1994; Price and Molleson 1974)

Maturation Rates and Longevity

Compared to the great apes, living humans have signiicantly longer childhoods (time from birth to puberty), substantially higher fertility (relected in shorter average interbirth intervals), and much greater longevity, which typically exceeds menopause by two-to-three decades (in chimpanzees menopause and longevity closely coincide). Two competing explanations have been ofered for the diference. he irst and more conventional idea is that prolonged childhoods and extended lifespans relect the need for children to accumulate culture and the value of older adults as cultural repositories. he second, known popularly as the “grandmother hypothesis,” argues that natural selection favored increased longevity at that point in human evolution when older women could enhance their reproductive itness more by provisioning their daughters’ or nieces’ children than by continuing to bear their own. In its most recent formulation, the grandmother hypothesis traces extended human lifespans to H. ergaster (or early African H. erectus).

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his is based in part on the previously noted possibility that H. ergaster emerged when climate in eastern Africa became drier and more seasonal about 1.8 Ma. he shit would have promoted greater dependence on tubers, rootstocks, or other underground plant organs, and in historic African hunter-gatherer societies, most notably the Hadza people of northern Tanzania, it is mainly the women who exhume these foods. Older (postmenopausal) women remain expert at it, but young children hardly do it at all because they lack suicient upper body strength. An increased reliance on underground plant organs could thus explain why H. ergaster females appear to have been so much larger than their predecessors, and if postmenopausal females signiicantly provisioned their grandchildren, their daughters could shorten their birth spacing and produce more ofspring. Maximum longevity and childhood length (time to sexual maturity) are tightly correlated across the mammals, so in theory, if either can be detected in the fossil record, the other can be inferred. Longevity is the more diicult to infer than childhood length, primarily because older adult age is hard to estimate from skeletal remains. It is especially diicult to assess age from the skeletons of living humans known to be more than forty-ive-to-ity years old, and this raises the issue of whether traditional aging methods can accurately determine longevity from fossils. Age at sexual maturity, or more precisely the rate of maturation, is more tractable, and the question to ask here then is whether Homo ergaster (and its descendants) matured at the same rate as living humans. To date, the most informative approach has been to examine growth increments in the enamel of fossil human teeth. he number and spacing of increments (perikymata) monitors the rate of dental crown formation, and this can be used to gauge the overall rate of skeletal maturation up to the age when the crowns are fully formed. he available analyses suggest that H. ergaster and H. erectus experienced remarkably rapid rates of enamel formation, more apelike than humanlike. As outlined in the next chapter, so far the oldest fossil to indicate a slow, basically modern rate of enamel formation is a mandible of early (or “near-modern”) Homo sapiens, dated to roughly 160 ka at Djebel Irhoud, Morocco. No publication so far has addressed the rate in H. heidelbergensis, and reports on the rate in H. neanderthalensis conlict. In advance, large brain and body size might favor a slow rate, comparable to that in H. sapiens. Relatively slow maturation and large adult brains mean that compared to the brains of other primates, human brains grow much more after birth. hus, on average, the brains of living human neonates average only about 25% of adult size, while in chimpanzees they average about 40%, and chimpanzees thereater approximate full adult brain size at a much earlier age. he degree of postnatal brain growth in early Homo is diicult to gauge directly because there are so few infant skulls for which

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ChaP Ter f ive

both age at death and endocranial capacity can be accurately assessed. So far, the main exception is the skull of a Homo erectus child from Mojokerto, Java. he reader will recall that this skull was found in 1936 and that its geologic age has been variously estimated between 1.8 and 30 ka (14C); between (Bräuer & Rimbach 1990; 128 and 40 ka? Debénath 1994; Debénath (Aterian artifact 2000; Debénath et al. 1986; associations) Debénath 1982; Ferembach 1976b; Hublin 1993)

Zouhrah Cave (El Harhoura 1)

A mandible and an isolated canine tooth

> 40 ka (14C and TL); between 128 and 40 ka? (Aterian artifact associations)

(Debénath 1980; Debénath 1994; Debénath 2000; Debénath 1982)

Between 128 and 40 ka? (Aterian artifact associations)

(Debénath 1994; Debénath 2000; Debénath 1982; Ferembach 1976a; Hublin 1993; Roche 1976; Roche & Texier 1976; Vallois & Roche 1958)

Ascending rami of two let mandibles

? between 130 and 50 ka (Mousterian artifact associations)

(Hublin 2000; McBurney 1967; McBurney 1975; Rak 1998; Tobias 1967b)

he partial, poorly preserved skeleton of an 8–10-year old child

Between 80.4 and 49.8 ka (Mousterian artifact associations and OSL on correlated eolian sands)

(Vermeersch et al. 1998)

A skull

between 170 and 150 ka (h/U, ESR, and associated fauna)

(Bate 1951; Clark 1988; Grün & Stringer 1991; McDermott et al. 1996; Spoor et al. 1998; Stringer 1979; Stringer et al. 1985)

Smugglers’ Cave Occipital, parietal, and frontal frag(Grotte des ments and a mandible, all probably Contrebandiers from the same individual [El Mnasra 2] á Témara) Libya Haua Fteah (Great Cave)

Egypt Taramsa Hill

Sudan Singa

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TABLE 6.2. (continued )

Site

Human Fossils

Age Estimate (basis)

Sources

Diré-Dawa (Porc-Epic)

A partial mandible

> 60 ka (obsidian hydration dating and associated Middle Stone Age artifacts)

(Assefa 2006; Clark 1982; Clark 1988; Clark et al. 1984b; Vallois 1951)

Herto, Middle Awash

A nearly complete adult skull; a partial juvenile skull; and 24 fragments from a second adult skull

Between 160 and 154 ka (40Ar/39Ar); Associated Acheulean and Middle Stone Age artifacts may suggest a broader age range, between perhaps 200 and 50 ka

(Clark et al. 2003; Faupl et al. 2003; White et al. 2003)

Aduma, Middle Awash

Fragments of 4 adult skulls

Between 105 and 70 ka (40Ar/39Ar, U-series, OSL, TL, and amino-acid racemization of molluscan shell)

(Haile-Selassie et al. 2004a; Yellen et al. 2005)

Omo-Kibish

A partial skull and associated postcranial bones (Omo 1); a second partial skull (Omo 2) and fragments of a third (Omo 3)

Between 198 and 104 ka, perhaps about 195 ka (40Ar/39Ar and correlations of depositional events in the Kibish Fm and Mediterranean deep sea sediments)

(Bräuer 1984a; Butzer et al. 1969; Day 1972; Day & Stringer 1982; Day & Stringer 1991; McDougall et al. 2005)

Eliye Springs (KNM-ES 11693) (West Turkana)

A skull

Between 300 and 200 (Bräuer et al. 2003; Bräuer & ka (possible stratiLeakey 1986) graphic origin near the top of the Koobi Fora Fm and cranial morphology)

Guomde (KNMER-3884 and ER-999) (Ileret, East Turkana)

A partial skull and a femur

Between 300 and 100 ka (geologic context, U-series, and cranial morphology)

(Bräuer 2001; Bräuer et al. 1992b; Bräuer et al. 1997; Feibel et al. 1989; Leakey & Leakey 1978; Trinkaus 1993a)

Ngaloba (Laetoli 18)

A skull

120 + 30 ka (U-series; geologic context; artifact and faunal associations)

(Day et al. 1980; Hay 1987; Leakey & Hay 1982; Magori & Day 1983; Rightmire 1984b)

Mumba Shelter

hree isolated molars

130–109 ka (U-series) (Bräuer 1984a; Bräuer & Mehlman 1988; Mehlman 1987)

Ethiopia

Kenya

Tanzania

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TABLE 6.2. (continued)

Site

Human Fossils

Age Estimate (basis)

Sources

Fragments of a radius

> 40 ka (MSA artifact associations)

(Barham 1995)

Equus Cave

A partial mandible and 10 isolated teeth

Between 75 and > 27 ka (14C; geologic context and associated fauna)

(Butzer 1984a; Butzer et al. 1978b; Grine & Klein 1985; Klein et al. 1991; Wadley 1993)

Florisbad

Facial, frontal, and parietal fragments of a single skull and a right upper third molar

259 + 35 ka (ESR, geologic context, and fauna)

(Brink 1988; Butzer 1984a; Butzer 1988; Grün 2006; Grün et al. 1996; Kuman & Clarke 1986)

Border Cave

An infant’s skeleton (BC3), an adult skull (BC1), two partial adult mandibles (BC2 and BC5), a right humerus shat (BC6), a right proximal ulna (BC7), and right metatarsals IV and V (BC8a & b)

Between perhaps 170 and 50 ka (14C, U-series, ESR, and isoleucine epimerization of ostrich eggshell)

(Beaumont 1980; Beaumont et al. 1978; Bird et al. 2003; Butzer et al. 1978a; Cooke et al. 1943; De Villiers 1973; De Villiers 1976; Grün 2006; Grün & Beaumont 2001; Grün et al. 1990a; Grün et al. 2003; Grün & Stringer 1991; Miller et al. 1999a; Miller et al. 1993; Morris 1992; Pearson & Grine 1996; Pfeifer & Zehr 1996; Rightmire 1979b; Rightmire 1984b; Sillen & Morris 1996)

Sea Harvest

An upper premolar and a phalanx

Between 128 and > 40 ka (C-14 and geologic inference)

(Grine & Klein 1993; Hendey 1974)

Hoedjies Punt

cranial fragments, isolated teeth, and postcranial bones

Possibly between (Berger & Parkington 1995; 300 and 200 ka Butzer 2004; Klein 1983; (Infrared StimuVolman 1978) lated Luminescence, hermoluminescence, stratigraphic context, associated fauna)

Die Kelders Cave 1

Twenty-three isolated teeth and two phalanges

Between 75 and 65 ka (ESR, geologic context; artifactual and faunal associations)

(Avery et al. 1997; Grine et al. 1991; Tankard & Schweitzer 1976)

Blombos Cave

Nine isolated teeth or tooth fragments

ca. 75 ka (OSL on sediments and TL on burned quartzites and silcretes)

(Grine & Henshilwood 2002; Grine et al. 2000; Henshilwood 2005; Henshilwood et al. 2001b; Jacobs et al. 2003b; Tribolo et al. 2006)

Zambia Mumbwa Cave South Africa

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Site

Human Fossils

Age Estimate (basis)

Sources

Klasies River Main

Five partial mandibles, two partial maxillas, fronto-nasal, zygomatic, temporal, and other cranial fragments, isolated teeth, an atlas, a lumbar vertebra, three metatarsals, a phalange, and portions of a clavicle, radius, and ulna

Between 115 and 60 ka (ESR, geologic correlation to the global marine stratigraphy; faunal associations)

(Bräuer et al. 1992a; Bräuer & Singer 1996a; Bräuer & Singer 1996b; Churchill et al. 1996; Deacon 1989; Deacon 1995; Deacon & Geleijnse 1988; Deacon & Shuurman 1992; Grine et al. 1998; Lam et al. 1996; Pearson & Grine 1997; Rightmire & Deacon 1991; Rightmire & Deacon 2001; Rightmire et al. 2006a; Singer & Wymer 1982)

Mugharet el 'Aliya M OR OC CO

Dar es Soltan, Zouhrah, & Smugglers' Caves Jebel Irhoud

TUNISIA Berard & Allobroges Taforalt Bir el Ater Rhafas El Guettar Météo

Karouba

Ain Fritissa Hassi Ouchtat Zaouia el Kebira

Tiouririne

LIBYA

RN

Taoudenni Basin

ES

TE

Esselesikine

W

Seggedim MAURITANIA

Nazlet Safaha Nazlet Khater & Taramsa Kharga Sodmein Cave Dungul Kubbaniya Bir Tarfawi & Bir Sahara East Sites 400, 415, Jebel Brinikol 420, 440, etc.

Adrar Bous MALI

BURKINA FASO

SUDAN Bilma

Tiguent SENEGAL

NIGER

Ed Debba CHAD

represent a terminal variant of the Acheulean Tradition that possessed many Middle Stone Age features, or it could mean that delation (wind erosion) concentrated bones and artifacts from Acheulean and overlying Middle Stone Age layers onto a single surface. In this case, depending on which kind of artifacts originally accompanied the human specimens, they could be as old as 200 ka or as young as 50 ka. Table 6.2 includes other fossils, particularly from Omo-Kibish, southwestern Ethiopia, and Border Cave, South Africa (discussed below), whose stratigraphic provenance and dating are problematic. It speciically excludes fully modern human cranial and postcranial bones from Kanjera, near Lake Victoria, western Kenya. L. S. B. Leakey (1935) believed the Kanjera bones were associated with animal fossils indicating great antiquity. From the time of Leakey’s irst report, however, other specialists argued that the

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N

Haua Fteah

EGYPT

Le Shati

HA

RA

Hajj Creiem

500 km

Wadi Gan

ALGERIA

SA

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FIGURE 6.20. approximate locations of key north african aterian and mousterian/msa archaeological sites (modiied after Wendorf and schild [1992], 44).

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470 FIGURE 6.21. approximate locations of the east african sites mentioned in the text.

40o

Sai Island 500

20o

1000 km REA

0

ERI T

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

ChaP Ter s iX

Adbur Reef

Singa SUDAN Herto & Aduma ETHIOPIA

Diré-Dawa

Gademotta Omo-Kibish Eliye Springs

Guomde

SOMALIA

KENYA Matupi Katanda

DEMOCRATIC REPUBLIC OF THE CONGO

UGANDA (Kanjera)

Kapthurin

Prospect Farm Enkapune Ya Muto Malawa Gorge Lukenya Nasera

0o

Ngaloba Mumba Kisese TANZANIA

associations were spurious and that the human bones were much younger. Trace-element analysis has supported the critics, and the human bones are now thought to come from shallow Holocene graves, postdating 10 ka. he table also excludes two human footprints found on the shore of Langebaan Lagoon, about 110 km north of Cape Town, South Africa. he prints were made on the slope of a moist dune and then buried by additional sand that was cemented into dune rock (eolianite). Luminescence and U-series dates on the dune rock vary between 228 and 103 ka, but none are considered reliable, and the footprints have been tentatively placed at 117 ka. his is the earliest time within the Last Interglacial when the enclosing dune would have been above sea level. However, the dune could have formed much later, perhaps during the Last Glaciation, between 71 and 11 ka, when signiicantly lower sea levels encouraged sand mobilization and dune formation on what is now the lagoon edge. he footprints lack diagnostic detail, and they are less compellingly human than nine prints in dune rock at Nahoon Point, about 800 km east of Cape Town. hree of the Nahoon Point prints show distinct toe and heel impressions that are lacking at Langebaan. Shell fragments and second-

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ary carbonates in the same layer at Nahoon Point have been radiocarbondated to 29 ka, but this is probably a minimum age, and the Nahoon prints could equal or exceed the Langebaan specimens in age. Not surprisingly, the Nahoon prints reveal feet that are unequivocally human in size and proportions. Among African fossils with secure provenance, the most questionable dating estimates are probably those for specimens associated with Aterian artifacts at Mugharet el ‘Aliya, Dar es Soltan Cave 2, Zouhrah Cave, and Smugglers’ Cave (Témara) in Morocco. Radiocarbon (14C) dates on Aterian layers vary from greater than 40 ka to less than 30 ka, and some authorities believe the Aterian persisted until 40 ka or later. However, minute amounts of undetectable younger carbon could make objects that are much older than 40 ka appear younger, and only the older, ininite Aterian 14C dates are probably reliable. Sedimentological context and faunal or loral remains oten imply that the Aterian existed under unusually moist conditions, especially in the Sahara, and there is growing evidence that northern Africa was signiicantly wetter mainly during interglacials. If this is accepted, then the Aterian could largely antedate the start of the Last Glaciation about 71 ka. Table 6.2, however, accepts the possibility that the Aterian persisted into the early part of the Last Glaciation. his is perhaps particularly likely in western Morocco, which has provided all the Aterian-associated human fossils. Among the fossils listed in table 6.2, the more complete skulls and jaws range from clearly archaic (especially Jebel Irhoud [ig. 6.23], OmoKibish 2, Eliye Springs, Smugglers’ Cave, and Florisbad) to only marginally archaic (Singa, Guomde, and Ngaloba [ig. 6.24]) to essentially modern (Zouhrah Cave, Dar es Soltan Cave 2 [ig. 6.25], Smugglers’ Cave, Haua Fteah, Taramsa Hill, Herto [ig. 6.26], Aduma, Omo-Kibish 1 [ig. 6.27], Border Cave, and Klasies River Main [ig. 6.28]). he context of the Omo-Kibish and Border Cave specimens requires special consideration because in both cases, it has long been controversial. he Omo-Kibish fossils come from Member I of the Omo-Kibish Formation, which was deposited primarily by the Omo River. OmoKibish 1 is a fragmented, but largely complete skeleton that was found eroding from the surface of Member I. Omo-Kibish 2 consists of a braincase (calvarium) found about 3.3 km away on the surface of Member I. Radiocarbon dating of shell from overlying Member III places the Member I human fossil horizon before 37 ka, while Ar40/Ar39 readings on underlying and overlying volcanic fragments (pumice clasts) bracket it between 196 and 104 ka. he Omo River originates in the same Ethiopian highlands as the Nile River, and proposed correlations between sequential depositional pulses recorded in the Omo-Kibish Formation and Nile pulses recorded in layers of organic mud (sapropels) in the eastern Mediterranean suggest that the Omo-Kibish human fossil horizon formed

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S 6o

D. R. CONGO

TANZANIA

Chambuage Mine

9o

Kalambo

ANGOLA 12o

ZAMBIA Kalemba

Mumbwa Humpata

15o

Twin Rivers

Malowa Zombepata

ZIMBABWE

Pfupi & Mtemwa Rocks

BOTSWANA Cave of Hearths

12o

15o

Mwulu’s Bushman Rock

Apollo 11 Kathu Border Cave Wonderwerk Florisbad Sehonghong Ha Soloja Holley Boegoeberg Orangia 1 Sibudu Melikane SOUTH AFRICA Umhlatuzana Grassridge Moshebi’s Strathalan Oakleigh

18o

Elands Bay Hoedjies Punt & Sea Harvest (Langebaan Lagoon) Ysterfontein Peers Cave, Tunnel Cave & Skildergatkop

21o

24o

Lion Cavern Sterkfontein Sibebe Witkrans Rose Cottage

Bremen Pockenbank

Oranjemund

Kalkbank

Olieboompoort

Zais Zebrarivier

BIQ

Tshangula

UE

Redcliff

18o

ZAM

NAMIBIA

White Paintings = Gi Pomongwe, Bambata & Nswatugi

MO

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

21o

24o

27o

30o

33o

36o

27o

30o 500 km 33o 39o

42o E

Klipfonteinrand, Klein Kliphuis Highlands & Hollow Rock Diepkloof Boomplaas Howieson’s Poort (Nahoon Kangkara Point) Buffelskloof Paardeberg Montagu Blombos Die Kelders Cape St. Blaize & Pinnacle Point

Klasies River Nelson Bay Herolds Bay

FIGURE 6.22. approximate locations of the principal middle stone age sites in southern africa (modiied after volman [1984], ig. 1). boldface marks deeply stratiied sites with well-preserved bone. Parentheses mark sites with possible middle stone age footprints.

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The nea nder Thal s and T heir ConTe mP or ar ies supratoral sulcus

lambdoidal flattening

receding frontal

occipital bun

supraorbital torus

mastoid crest or tuberosity

mastoid process

suprainiac depression

juxtamastoid crest

occipital torus

Spy 2 (Neanderthal) 0

mastoid process

juxtamastoid crest

5 cm

(no suprainiac depression)

473 FIGURE 6.23. a fossil skull from Jebel irhoud, morocco, compared to a neanderthal skull from spy Cave, belgium (redrawn after santa luca [1978], 624, 627). The irhoud skull has sometimes been labeled neanderthal, but it is probably 100 ky or more older than most known neanderthal skulls and it differs from them in several crucial features, including its shorter and latter face, its more rectangular orbits, its more parallel-sided (less globular) braincase, and its lack of a suprainiac fossa or depression. in most of these features, it anticipates anatomically modern people, and it could represent a population that lay on or near the line of modern ancestry.

occipital torus

Jebel Irhoud 1 (non-Neanderthal) about 195 ka. However, the two Omo-Kibish skulls contrast starkly in form (ig. 6.27), and this raises the possibility that one (or both) might be intrusive into Member 1. he likelihood is greater for Omo-Kibish 1 (more modern) because it is associated with a tight cluster of postcranial bones that by itself might suggest a grave, independent of geologic age. Erosion of the overlying deposits could have obscured the outlines of a burial pit, even if it were only a few thousand years old. A comparison of luorine, nitrogen, and uranium content between the Omo-Kibish 1 bones and animal remains from the surrounding deposits might resolve the issue. If intrusion is precluded, it follows that Omo-Kibish 1 and 2 must both be about 195 ky old and they would probably imply separate, overlapping human populations, only one of which was anatomically modern. Border Cave presents a comparable, if less momentous, dilemma since it is possible that some of the remarkably modern human fossils derive from graves dug into much older deposits. his is perhaps especially likely for BC1, an adult skull, and BC2, a mandible, that were found during nonscientiic excavations for guano (fertilizer). It also concerns BC3, an infant’s skeleton that was found by scientists but is much better

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474

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

ChaP Ter s iX

FIGURE 6.24. facial and right lateral views of the near-modern human skull from ngaloba, northern Tanzania (drawn by Kathryn Cruz-uribe from photos in magori and day [1983], 748–749). ngaloba shares the location of the australopithecine site of laetoli, and the skull is also known as laetoli hominid (LH) 18. The cranium combines large browridges, a lattened, receding frontal, a pronounced occipital torus, and other primitive features with a short, lat face, subparallel sidewalls, a well-rounded occipital, and other characters that ally it to modern humans. it has an estimated cranial capacity of 1,200 cc, below the modern average, but well within the modern range. it is tentatively dated to about 120 ka, and it supplements other african fossils which suggest that modern anatomy evolved in africa at a time when only neanderthals were present in europe.

browridge

Ngaloba (LH 18)

occipital torus 0

10 cm

preserved than animal bones from the same level. he contrast is striking since the animal bones were initially much more durable than the infant’s skeleton. Analysis of bone mineral crystallinity has now resolved the contradiction. Fresh bone is relatively noncrystalline, but crystallinity tends to increase ater burial and should be the same for bones from the same layer within a single site. Crystallinity analysis shows that the Border Cave infant’s bones must be signiicantly younger than the associated animal fossils, and it conirms great (Middle Stone Age) antiquity only for BC6 and BC7, fragments of a humerus and ulna respectively. An ESR age of 74 ± 5 ka on dental enamel implies comparable antiquity for BC5, a partial mandible. Whether or not the Omo 1 and Border Cave specimens are included, however, there are no African fossils with indisputably Neanderthal features. Some of the mandibles are large and rugged, but where the appropriate parts are preserved they lack retromolar spaces and they usually have distinct chins. Together with other facial bones, the mandibles indicate that Africans who lived between 250 and 40 ka tended to have sig-

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FIGURE 6.25. nearmodern human skull and mandible from the cave of dar es soltan 2, morocco (redrawn after debénath et al. [1986], 237). The braincase is higher (and probably shorter) than in neanderthals, and the face is distinctly shorter and less protruding, especially along the midline. except for the large projecting browridge, the skull exhibits no features that distinguish it signiicantly from most modern human skulls, yet the population it represents inhabited morocco at the same time that neanderthals occupied europe.

5 cm

0

475

Dar es Soltan 2

niicantly shorter, broader, and latter faces than did the Neanderthals. Similarly, some of the skulls (Florisbad, Jebel Irhoud, Omo 2, Herto, Singa, and Ngaloba) are ruggedly built, with large browridges, and in some cases (Jebel Irhoud, Omo-Kibish 2, and Ngaloba) relatively prominent transverse occipital tori, as well as pronounced bony crests or mounds in the occipitomastoid region (igs. 5.31–5.33 illustrate Jebel Irhoud 1, Singa, and Florisbad; ig. 6.26 illustrates Herto BOU-VP-16). In general, however, the African skulls tend to be shorter and higher than classic Neanderthal skulls, and some approach or equal modern skulls in basic vault shape. Where adult endocranial capacity can be reasonably estimated (for Guomde, Ngaloba, Omo-Kibish 1 and 2, Herto BOU-VP-16/1, Singa, Eliye Springs, Border Cave 1, and Jebel Irhoud 1 and 2), the African skulls vary from roughly 1,370 cc (Ngaloba) to 1,510 cc (Border Cave 1). he endocranial igures are all comfortably within the range of both the Neanderthals and anatomically modern people. In sum, the African fossil sample is small and dominated by fragments. It is less homogeneous than the Neanderthal sample, perhaps because it represents a larger time range and geographic area, perhaps because it represents populations undergoing more rapid change, or perhaps because it (erroneously) includes some specimens that actually date from ater 40 ka. Still, it is clear that even though most of the African fossils date from the same interval as the Neanderthals, none of them exhibit typical Neanderthal morphology. Instead they possess craniofacial features that are near-modern to modern, and as a group they strongly imply that anatomically modern people were developing in Africa at the

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braincase exceptionally long, but also high

bossed frontal rising relatively steeply above the supraorbital

thick supraorbital torus

sharply angulated occipital with a massive external protuberance (but no occipital torus)

short, flat face tucked in below the fore part of the braincase large, projecting mastoid process

cm

Herto BOU-VP-16/1 FIGURE 6.26. The principal adult skull (bou-vP-16/1) from herto, middle awash, ethiopia (drawn by Kathryn Cruz-uribe from photos in White et al. [2003], 743). it differs from neanderthal skulls in numerous features, including the short, lat face tucked in under the forepart of the braincase, the bossed (outward bulging), steeply rising frontal, the large projecting mastoid process, and the absence of an occipital torus. in all these features and others, it anticipates the skulls of living people from which it differs mainly in its great robusticity and exceptional size. Together with a second less complete adult skull and a partial juvenile skull from the same locality, it shows that essentially modern people lived in africa at a time when neanderthals (or their immediate ancestors) were the sole occupants of europe.

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same time that Neanderthals were the only people in Europe. he SaintCésaire face and partial braincase (ig. 6.29) show that even the latest Neanderthals, dated to near 40 ka, were as distinctive and nonmodern as the earliest, dated to 71 ka or before. Depending on how the African fossils are arranged in time, their variability could imply an evolutionary process in which “modern” craniofacial features accumulated piecemeal, mainly via random drit, in the same way that Classic Neanderthal craniofacial form may have evolved in Europe. he case for an African origin of modern human morphology becomes particularly strong if the near-modern human fossils from Qafzeh and Skhul Caves, Israel, are added to the African sample. As discussed below, TL and ESR dating bracket the Qafzeh-Skhul fossils between roughly 120 and 90 ka, when animal fossils suggest that Israel lay within a slightly expanded Africa. Like broadly contemporaneous people in Africa strictly deined, the Qafzeh-Skhul people were variable in the expression of chins, vertical foreheads, rounded occipitals, parietal bossing, and other modern features, and in some important respects, such as strongly developed browridges, large teeth, conspicuous buttressing on the inner side of the mandibular symphysis (chin), and a tendency to pronounced alveolar prognathism (ig. 6.30), some resembled more archaic humans. he variability and robusticity apparent in the Skhul craniums is so great that when they were irst monographically described, they were considered to represent the same population as the Neanderthals from nearby Tabun. Subsequently the Tabun and Skhul people together were sometimes lumped as “progressive Neanderthals,” versus “classic Neanderthals,” such as those from the caves of southwestern France. he Skhul people were clearly not Neanderthals, however, and their robust-

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upper (occipital) plate of the occipital flattened frontal

occipital torus

lower (nuchal) plate of the occipital

cm

Omo-Kibish 1

Omo-Kibish 2

FIGURE 6.27. skulls 1 and 2 from the omo-Kibish locality, lower omo river valley, ethiopia (drawn from photographs in day and stringer [1991]). skull 1 is partly reconstructed, particularly in the face. The skulls are commonly said to derive from the same deposits, now tentatively dated to about 195 ka, but they contrast morphologically. omo-Kibish 1 has a steeply rising frontal, inlated parietal regions, a rounded occipital with a relatively small nuchal plane, and other features that are unequivocally modern, whereas omo-Kibish 2 has a latter, more receding frontal, inwardly sloping sidewalls associated with a midsagittal keel, a highly angulated occipital with an extensive nuchal plane, and a strong occipital torus, all reminiscent of Homo erectus. however, it resembles skulls of Homo sapiens in its relatively small browridge and in its large internal (endocranial) capacity of about 1,400 cc. The strong contrast between the skulls is puzzling, and it may actually imply a signiicant difference in their geologic age.

icity and variable expression of modern traits is probably better encapsulated in the term near-modern. In overall appearance, they resembled living humans far more closely than the Neanderthals did, but they (and the Qafzeh people) difered signiicantly not only from living humans but also from the earliest universally accepted modern (Upper Paleolithic) inhabitants of both Europe and western Asia. Both individually and as a group, the African skulls contrast strongly with Neanderthal skulls, and it is pertinent to ask whether postcranial diferences were equally marked. Based on Skhul-Qafzeh, the answer is probably yes. he African fossil sample narrowly understood comprises the highly fragmentary skeleton of Omo-Kibish I and nine other postcranial bones that have been fully described. hese are a proximal ulna, a proximal radius, and three metatarsals from Klasies River Main, a proximal ulna and a humerus shat from Border Cave, a femur from Guomde (East Turkana), and a humerus from Jebel Irhoud, Morocco. In form and proportions, the Omo 1 postcranial skeleton closely resembles those of modern Sudanese to which it has been compared in detail. he similarity in limb proportions suggests a shared adaptation to hot, dry climatic conditions. he Klasies River Main (KRM) radius and ulna and the Border Cave ulna broadly resemble their Neanderthal counterparts in overall morphology and robusticity, but they are no further from

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KRM 41815 small

true chin (vertical manidibular symphysis

KRM 16424

chin region not preserved

tiny

0

true chin

5 cm

M1 M2

M3 missing, possibly congenitally absent, no retromolar space

chin region not preserved

KRM 41815 M1 M2

KRM 16424 M3

FIGURE 6.28. occlusal and buccal views of mandibles 41815 and 16424 from the Klasies river main (KRM) cave site, south africa (drawings by Kathryn Cruz-uribe). both mandibles were sealed in a stratigraphic unit that formed sometime between 110 and 90 ka, following the maximum high sea level of the last interglacial (global isotope substage 5e). most of the teeth from Krm 41815 were lost before excavation, but the sockets are present for all except the m3’s (third molars), which probably never developed. The available teeth are heavily worn, indicating advanced age, and there are signs of periodontal disease, including abscesses at the tips of the irst molar roots. The specimen is striking for its distinct chin (vertical mandibular symphysis) and for its short, broad dental arcade that lacked retromolar spaces (a gap between the rear edge of each m3, if it had been present, and the fore edge of the ascending ramus). The sum implies that Krm 41815 came from someone with a short, broad, lat face, more closely resembling the face of a living human than that of a neanderthal. based primarily on the absence of a retromolar space, Krm 16424 probably implies a similar face, but it is striking for its tiny size, even compared to Krm 41815, which lies on the small side of the modern human range. other specimens, including especially partial mandible Krm 13400 (not illustrated), are signiicantly larger. The size variation may relect sexual dimorphism, but if so, the degree was beyond the range found in later, fully modern populations. arguably, this means that the Klasies river main people should be considered “near-modern” rather than fully modern in their anatomy.

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receding frontal

supraorbital torus

midfacial prognathism

479 FIGURE 6.29. The reconstructed face and skull of a neanderthal associated with Châtelperronian artifacts at la roche à Pierrot, saint Césaire, france (drawn by Kathryn Cruzuribe from a photograph). The fossil is among the youngest known neanderthal specimens, but in all observable features, including the strong browridge, low frontal, and far forward protrusion of the face along the midline, it is no more modern than much more ancient neanderthal skulls. since it postdates modern or near-modern human remains elsewhere, it all but precludes the possibility that neanderthals evolved into modern humans.

retromolar space cm

Saint-Césaire

the modern mean than some terminal Pleistocene and recent African counterparts. One of the KRM metatarsals is morphologically unusual, but together, the three metatarsals exhibit no more than idiosyncratic separation from much later Africans, except that considered with other KRM fossils, especially mandibles, they imply an extraordinary degree of sexual dimorphism. Assuming that only one population is represented, the KRM people were probably more dimorphic than any known historic population. Added to the variable expression of modern craniofacial morphology at KRM and like-aged African sites, the dimorphism implies that the African contemporaries of the Neanderthals were anatomically near-modern rather than modern. he Border Cave humerus has slightly thicker cortical walls and a correspondingly narrower medullary (marrow) canal than most modern humeri, but it is morphologically unremarkable. he Guomde femur is robust but exhibits typically modern (and non-Neanderthal) morphology, including a distinct bony ridge (pilaster) on the posterior surface and a high neck-to-shat angle. he Jebel Irhoud humerus has a hyperrobust (hypertrophied) shat like those of the Neanderthals and earlier nonmodern humans, but it probably antedates the near-modern fossils from KRM, Border Cave, and Skhul-Qafzeh.

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FIGURE 6.30. Top: Top view of skull 9 from Qafzeh Cave, israel, partially reconstructed. Bottom: lateral view of the same skull, more fully reconstructed (adapted from vandermeersch [1981], igs. 9, 10, 43). The Qafzeh fossils are dated to roughly 90 ka, when neanderthals were the sole inhabitants of europe, and like the neanderthals, the Qafzeh people produced mousterian artifacts. unlike the neanderthals, however, they closely resembled modern people in key morphological respects, including a relatively high, short braincase that overlapped the face below. retraction of the face beneath the braincase is relected in the absence of a retromolar space and in the presence of a chin. The descendants of the Qafzeh people may have been displaced by neanderthals, who spread to israel after 80 ka.

steep forehead (frontal)

wellrounded occipital face retracted below the forepart of the braincase 0

5 cm

Qafzeh 9

no retromolar space chin

he Skhul-Qafzeh postcraniums are unequivocal. hey mark people who were robust, particularly in their legs, but who were essentially modern in form. his is particularly clear in the femur and pelvis, which difer from those of the Neanderthals in precisely the way that those of living humans do, but it is also patent in every other anatomical region for which there is evidence. If it is assumed that Neanderthal form largely relects a reliance on jaws and bodies to accomplish everyday tasks, the Skhul-Qafzeh postcraniums might imply that modern morphology arose when Africans began to rely more on tools (or culture). However, no fundamental behavioral diference is implied by toolkits, which are basically similar between the Neanderthals and their near-modern contemporaries everywhere. Artifactual change that could signal a major behavioral diference from the Neanderthals occurs only ater 50 ka, with the widespread appearance of fully modern humans. he implication is probably that the anatomical diferences between Neanderthals and

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near-moderns have more to do with climatic adaptation and genetic drit than with diferences in behavior. Archaeological indings reported below suggest that fully modern humans originated in eastern Africa only about 50 ka, and they may have replaced not only the Neanderthals but also more modern-appearing people like those at Klasies River Main, Dar es Soltan, and Skhul-Qafzeh. In this light, the known African and Israeli near-modern fossils are signiicant not because they represent the lineal ancestors of living people but because they point to Africa as the birthplace of modern human morphology. Similarities between the Florisbad and Omo-Kibish 2 skulls on the one hand and the older (mid-Quaternary) Kabwe and Elandsfontein skulls on the other and between the Qafzeh and Skhul skulls and the more ancient ones from Jebel Irhoud, Morocco, hint that it may one day be possible to trace the origin of modern humans through a succession of progressively less archaic African fossils. SOURCES: Taramsa Hill (Vermeersch et al. 1998); contemporaries of the Neanderthals or anteNeanderthals in Indonesia (Rightmire 1988; Santa Luca 1980) and China (Wu 1992; Wu and Bräuer 1993; Wu and Poirier 1995; Wu and Wu 1985); ESR dating of Ngandong fossils (Swisher et al. 1996); African contemporaries of the Neanderthals (table 6.2); Kanjera fossils—antiquity (Leakey 1977; Plummer and Potts 1989) and trace element analysis (Behrensmeyer et al. 1995; Plummer et al. 1994; Plummer and Potts 1995); Langebaan footprints (Gore 1997; Roberts and Berger 1997); Nahoon Point footprints (Mountain 1966) and dating (Deacon 1966a); Aterian—persistence to 40 ka or later (Debénath 1994, 2000; Debénath et al. 1986) or beyond the range of radiocarbon (Camps 1974, 1975; Close 2002; Cremaschi et al. 1998; Wendorf et al. 1990; Wendorf and Schild 1992; Wrinn and Rink 2003); interglacial humidity in northern Africa (Causse et al. 1988; Fontes and Gasse 1989; Wendorf et al. 1990; Wendorf and Schild 1992); Omo-Kibish fossils—dating by radiocarbon (Leakey et al. 1969b) and by Ar40/Ar39 (McDougall et al. 2005; Shea et al. 2007), morphology (Day and Stringer 1982); Border Cave fossils—crystallinity analysis (Sillen and Parkington 1996), conirmed MSA antiquity (Grün and Beaumont 2001; Sillen and Morris 1996), and ESR dating (Grün et al. 2003); endocranial capacity in African contemporaries of the Neanderthals (Bräuer 1984a; Holloway et al. 2004); possible accretional evolution of modern craniofacial form in Africa (Bräuer 2008; Pearson 2008); Qafzeh-Skhul fossils— dating (Tchernov 1992, 1994). morphological variability (Bräuer 1989; Corruccini 1992; Kidder et al. 1992; McCown and Keith 1939; Simmons et al. 1991; Stringer 1989, 1996a; Trinkaus 1991; Wolpof and Caspari 1990), and diferences from earliest modern Europeans (Howells 1989); postcranial bones— Omo-Kibish 1 (Day et al. 1991), Klasies River (Churchill et al. 1996; Pearson and Grine 1997; Rightmire et al. 2006a; Rightmire and Deacon 1991), Border Cave (Morris 1992; Pearson and Grine 1996; Pfeifer and Zehr 1996), Guomde (Trinkaus 1993a), and Irhoud (Hublin 1993; Hublin et al. 1987); similarities between skulls of “earlier” and “later” fossil H. sapiens in sub-Saharan Africa (Rightmire 1984b, 1987); similarities between skulls from Irhoud and Qafzeh-Skhul (Hublin 1993, 2000)

Artifact Industries Archaeologists commonly lump similar assemblages of like-aged artifacts into an “industry” or “industrial complex,” and the artifact assemblages the Neanderthals made are now usually assigned to the Mousterian Industrial Complex. his is named for the rockshelters at Le Moustier (southwestern France) where, in 1863, the pioneer prehistorians Edouard Lartet and Henry Christy initiated a long series of important excavations. In

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both Europe and western Asia, the term Middle Paleolithic is oten used as a synonym for Mousterian, though it has also been used more loosely for any artifact assemblages that are supposedly coeval with Mousterian ones regardless of the artifact types involved. It is in this relatively loose time sense that it has been applied in eastern Asia, where in general the artifact assemblages it refers to are still poorly known and poorly dated. In Africa the contemporaries of the Neanderthals produced artifacts that are very similar to Mousterian ones, and the term Mousterian has been applied directly to some north African assemblages, particularly in the Nile Valley, the eastern Sahara, Cyrenaica (northern Libya), and the Maghreb (northwestern Africa). Many other north African assemblages have been assigned to the Aterian Industry, named ater the site of Bir el Ater in northeastern Algeria. he Aterian is distinguished from the Mousterian primarily by the presence of stemmed or tanged pieces, but this diference is no greater than the diference among European industries assigned to diferent facies (variants) of the Mousterian, and the separation of the Aterian from the Mousterian owes less to its typology than to the now-abandoned idea that the Aterian postdated the European Mousterian. In fact, like the Mousterian, the Aterian almost surely dates to the earlier part of the late Pleistocene, at or beyond the 30–40-ky practical limit of 14C dating. In northern Africa, as in Europe and western Asia, the Mousterian (including the Aterian) was replaced by the Upper Paleolithic industrial complex. he Maghreb and the Sahara were sparsely populated or abandoned when the Upper Paleolithic appeared, but 14C dates from the Haua Fteah and Ed Dabba Cave in Cyrenaica, Libya, and Nazlet Khater in the Egyptian Nile Valley show that the Upper Paleolithic was present by 40–35 ka. In sub-Saharan Africa, the industries that are coeval with the Mousterian are conventionally assigned to the Middle Stone Age, or MSA. Like the Aterian, the MSA was once thought to postdate the Mousterian, but numerous 14C dates, obtained mainly since 1970, show that like the Mousterian, it terminated in the middle part of the Last Glaciation, before 40–30 ka. Artifactual variation within the MSA is as great or greater than between the MSA and the Mousterian, and as in the case of the Aterian, on strictly typological grounds, the various MSA industries could be regarded simply as facies of the Mousterian. Scholarly tradition and geographic distance are the principal reasons for separating the MSA and Mousterian. he time when the Mousterian/MSA began is not irmly established, partly because it lies in the period between 250 and 130 ka that is currently diicult to date radiometrically. here is also the problem that in both Africa and Europe the Mousterian/MSA difers from the preceding Acheulean mainly in the absence of large bifacial tools (hand axes and

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cleavers). However, people who made large bifaces need not have let them at every site they visited, and large bifaces do not occur at several African and European sites that are clearly contemporaneous with Acheulean sites in the same regions. In addition, at some sites where large bifaces do occur, they are rare, suggesting that chance (sampling error) could explain their absence elsewhere. Still, with the diiculty of distinguishing the MSA from the Acheulean in mind, radiometric dates at the sites in table 6.3 and the stratigraphic or paleontologic context of probable MSA artifacts at various sites in South Africa suggest the MSA was widely installed by 250–200 ka. he situation in Europe, western Asia, and northern Africa may be more complex since Mousterian industries may have replaced the Acheulean at diferent times in diferent places. However, there is growing evidence that the Mousterian appeared in most places by 250–200 ka, during the Penultimate Glaciation (oxygen-isotope stage 6) or the preceding interglacial (stage 7). In some regions, perhaps most, the Acheulean may not have terminated abruptly, Acheulean and MSA/Mousterian industries may have coexisted for thousands or even tens of thousands of years. he strongest case for such coexistence is in the Kapthurin Formation, Lake Baringo Basin, Kenya, where spatially discrete Acheulean and MSA assemblages are associated with dated volcanic tufs (ash horizons) that imply Acheulean/MSA overlap over a long interval before 285 ka. he Acheulean assemblages include ones that can be assigned to the Fauresmith variant (with inely made, highly symmetric hand axes) or the Sangoan variant (with coarser, heavy-duty “picks” and “core-axes”). In the Kapthurin region and elsewhere, the Fauresmith and Sangoan are usually regarded as late or inal Acheulean manifestations, perhaps diferentiated by adaptation to savanna (Fauresmith) and to woodland (Sangoan). However, in contrast to what the scattered Kapthurin occurrences imply, wherever Acheulean and MSA/Mousterian artifact assemblages have been found stratiied together in a single locality, the MSA/Mousterian layers consistently overlie the Acheulean ones, with no hint of interdigitation. his is true, for example, in deeply stratiied caves, such as Combe-Grenal, France, et Tabun, Israel, and Montagu, Wonderwerk, and the Cave of Hearths, South Africa. he Kapthurin Formation may appear diferent because the artifact assemblages tend to be small, and their assignment to the Acheulean or MSA is arbitrary. In addition, they were collected mainly on the surface, where their stratigraphic relationship to dated tufs is questionable. Finally, at most collection points, the tufs were not dated directly but by “inger printing” (geochemical similarity to dated tufs elsewhere), which introduces a further potential source of error. he various Mousterian/MSA industrial sequences almost certainly ended at diferent times, depending on the place. In sub-Saharan Africa

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TABLE 6.3. African sites where numeric dating indicates the Middle Stone Age began before 250–200 ka. Figures

6.21 and 6.22 locate the sites. Site

Dating Method

References

Sai Island (site 8-B-11), Sudan

OSL on sediments

(Van Peer et al. 2003)

Gademotta, Ethiopia

K/Ar on volcanic ash

(Clark 1988; Wendorf et al. 1994a; Wendorf et al. 1975)

Kapthurin Fm, Kenya

40

Malawa Gorge, Kenya

K/Ar on volcanic ash

(Evernden & Curtis 1965)

Twin Rivers Cave, Zambia

U-series on lowstone

(Barham & Smart 1996)

Ar/39Ar on volcanic ash

(McBrearty et al. 1996; Tryon & McBrearty 2002; Tryon & McBrearty 2006)

Lincoln Cave, Sterkfontein, South U-series on lowstone Africa

(Reynolds et al. 2003)

Wonderwerk Cave, South Africa

U-series on lowstone

(Beaumont & Vogel 2006)

Florisbad, South Africa

OSL on sediments

(Grün et al. 1996; Kuman et al. 1999)

they may have terminated between 50 and 45 ka, in northern Africa and western Asia between 45 and 40 ka, in eastern Europe between 43 and 39 ka, and in western Europe between 40 and 37 ka. At the southwestern corner of Europe (southern Spain and Gibraltar) some Mousterians may have persisted until 30 ka or later. (It should be recalled that for reasons presented in chapter 2, the stated ages are all in uncalibrated radiocarbon years.) Although it is heuristically useful to equate the Neanderthals and the Mousterian, it is important to stress that, strictly speaking, this is incorrect. he people who made Mousterian/MSA artifacts in Africa were clearly not Neanderthals, and at Saint-Césaire and especially Arcy-surCure (Grotte du Renne) in France, Neanderthal fossils are associated with the Châtelperronian artifacts, which some specialists assign to the early Upper Paleolithic. Conceivably the association implies that fully modern Châtelperronians sometimes accumulated Neanderthal bones, but the Châtelperronian is Mousterian-like in some respects, and most authorities therefore believe that the Saint-Césaire and Arcy Neanderthals represent its makers. Finally, and perhaps most important, the people who made Mousterian artifacts at the Skhul and Qafzeh Caves in Israel were near-modern people. he signiicance of these departures from the Neanderthal = Mousterian formula will be considered in the section below on the fate of the Neanderthals. SOURCES: Middle Paleolithic in eastern Asia (Olsen 1987; Qiu 1985, 1992); Mousterian—in the Nile Valley (Marks 1968b; van Peer 1991, 1998), the eastern Sahara (Close et al. 1990; Wendorf et al. 1993, 1994b; Wendorf and Schild 1980, 1992), Cyrenaica (McBurney 1967), and the Maghreb (Balout 1955; Camps 1974; Wengler 1986, 1990b); Aterian—typology (Bordes 1976–77; Debénath 1994; Ferring 1975) and dating (Bouzouggar et al. 2002; Camps 1974, 1975; Close 2002; Cremaschi et al. 1998; Wendorf et al. 1994a; Wendorf and Schild 1992); Upper Paleolithic dating in Cyrenaica (McBurney 1975) and the Nile Valley (Vermeersch et al. 1984a, 1984b, 1990); MSA—end (Clark 1988; hackeray 1992a; Vogel

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and Beaumont 1972; Volman 1984) and beginning (table 6.3 and Beaumont and Vogel 2006; hackeray 1992a; Volman 1984); beginning of the Mousterian in Europe, western Asia, and northern Africa (Bar-Yosef 1993, 1995; Bosinski 1982, 1986; Conard and Fischer 2000; Roebroeks et al. 1993; Santonja and Villa 2006; Tufreau 1978, 1979, 1982, 1992; Wendorf et al. 1993, 1994b; Wendorf and Schild 1992); Kapthurin Basin—Acheulean/MSA overlap (Tryon and McBrearty 2002) and nature of the Acheulean (McBrearty 2001; McBrearty et al. 1996); Fauresmith and Sangoan (Clark 1970, 1988); Acheulean to Mousterian/MSA at Combe Grenal (Bordes 1972), Tabun (Bar-Yosef 1995; Jelinek 1982b), and South African caves (Beaumont and Vogel 2006; Volman 1984); termination of Mousterian/MSA in subSaharan Africa (Ambrose 1998a), in northern Africa and western Asia (Bar-Yosef et al. 1996; Hole and Flannery 1967; Kuhn et al. 1999; Marks 1981a; McBurney 1967; Mercier and Valladas 1994; Schwarcz 1994; Solecki 1963), eastern Europe (Hofecker 2002; Kozlowski 1990b; Mellars 2004), and western Europe (Bischof et al. 1989, 1994; Cabrera Valdés and Bischof 1989; Straus 1989a; Straus 1993–94; Zilhão and d’Errico 1999); late persistence of Mousterian in southern Spain (Finlayson 2004; Hublin et al. 1995) and Gibraltar (Finlayson et al. 2006); fully modern humans as accumulators of Neanderthal bones (Bordes 1981a)

Mousterian/MSA Stone-artifact Technology Mousterian/MSA people were consummate stone knappers, who used a variety of sophisticated techniques to produce their tools. Beginning with the great French prehistorian François Bordes, other, mostly French, archaeologists have illuminated Mousterian technology by analyzing the chaîne opératoire. his term can be translated loosely as the core reduction sequence, or the sequence of actions between the selection of raw material for laking and the abandonment of exhausted tools. he intermediate steps comprise the primary production of sharp-edged blanks or preforms and their secondary modiication, use, or maintenance (refreshing). Broadly speaking then, the chaîne opératoire refers to the entire process of manufacturing and use, and the varied chaînes employed by Paleolithic people can be reconstructed by experimental replication or by reitting excavated pieces (ig. 6.31). he focus here is on Mousterian/MSA reduction of raw stone nodules or blocks to sharp-edged blanks. Blank modiication and use are discussed in the next section on Mousterian/MSA retouched tool typology. Traditionally, archaeologists have relied on retouched tool typology to reconstruct cultural relationships within the Mousterian/MSA, but the results are controversial, and similarities in the initial reduction sequences provide an alternative, arguably superior, basis. Reduction sequences have perhaps been used most fruitfully to isolate clusters of related Mousterian assemblages in southwestern Asia (the Near East). Even the earliest stone-tool makers preferentially selected the most desirable rock types at their disposal, and Mousterian/MSA people were particularly discerning. Generally speaking, the best rock types are relatively hard, fracture easily when struck, and have a smooth, homogeneous internal consistency. Among widespread rock types that best meet these criteria are ine-grained siliceous varieties such as lint and chert. Flint is common throughout Europe and southwestern Asia, where it

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FIGURE 6.31. Partial reconstitution of a lat, discoidal core by the reitting of 162 separately excavated lakes from mousterian site C, maastricht-belvédère, the netherlands (redrawn after roebroeks et al. [1993], ig. 7).

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was used so oten that the local Stone Age could almost be called the “lint age.” In Africa, where large nodules of lint or related rocks are uncommon in most regions, stone age people were oten compelled to use other materials, including volcanic rocks and quartzites. Some inegrained quartzites approach lint in quality, but among volcanic rocks only one—obsidian (volcanic glass)—is truly comparable. It is limited to parts of eastern Africa, where some MSA people used it intensively. Other more widespread volcanic rocks, such as basalt, are generally more diicult to work, and they may therefore yield cruder-looking artifacts. he reduction process begins when the knapper strikes a percussor against a raw stone nodule in order to remove a lake. he percussor can be made of stone or of soter material like wood, antler, or bone. Replication experiments suggest that Mousterian/MSA and earlier peoples mainly used stone, and their percussors are commonly called hammerstones. A nodule from which one or more lakes has been struck is called a core, and the resultant lakes are commonly said to have three main parts: the striking platform or butt, the ventral surface, and the dorsal surface. he striking platform is the part of the lake that was struck by the hammerstone when the lake was detached from the core. Part of the struck area remains on the core where it forms the striking platform of the core. A given core can have many striking platforms, but a lake will have only one (or very rarely two, formed when two blows hit a core at the same time; this can happen when a struck core bounces against another hard object). By convention, unless otherwise noted, lakes are illustrated with the striking platform down. he ventral surface is the one that was originally inside the core; the dorsal surface is the one that was outside. he ventral surface is usually smooth, though it always has a variably pronounced bulge or bulb of percussion immediately adjacent to the striking platform. his results from the rapid dissipation of the hammer blow through the interior of the core. Unlike the ventral surface, the dorsal one always exhibits the weathering rind (cortex) of the unlaked core, hollows and ridges (scars) from previous lake removals, or both.

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487 FIGURE 6.32. stages in the manufacture of a classic levallois core (redrawn after bordes [1961a], ig. 4). 1, raw nodule; 2, nodule with lakes struck off around the periphery; 3, nodule with lakes struck radially inward on one surface, by using the peripheral scars as striking platforms; 4, radial preparation completed; 5, inal hammer blow (indicated by the arrow); and 6, inal levallois core and lake.

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Once a knapper has removed one or more lakes from a core, the negative scars (or facets) let on the core can serve as striking platforms for additional lakes, and the process can be repeated until the core becomes too small for further reduction. A core that has been worked to this extent is said to be exhausted. Modern replication experiments show that Mousterian/MSA knappers oten shaped a core or modiied its striking platform to predetermine the size or shape of lakes. Platform modiication was commonly done to remove irregularities that would difuse or misdirect the force of the hammer blow. It results in lakes that have prepared or faceted striking platforms (or butts) as opposed to ones with unprepared or smooth platforms (or butts). A core that was extensively shaped to determine lake size and shape is called a Levallois core, ater a site in the Levallois suburb of Paris where Paleolithic examples have been known since the nineteenth century. he corresponding lakes are called Levallois lakes. In general, Mousterian/MSA knappers selected lattish nodules or blocks for Levallois reduction (ig. 6.32). hey irst struck downward to remove lakes around the entire periphery of the nodule and then used the peripheral lake scars to strike inward, removing lakes systematically from one surface of the core. When this surface was completely prepared, they again struck inward to remove one or more lakes whose shape and size was determined by the arrangement of previous lake scars on the core surface. Levallois lakes usually have faceted platforms, but the deining characteristic is the pattern of dorsal scars relecting deliberate preparation of the core surface. Well-made Levallois lakes and cores appear in some later Acheulean assemblages, dating between 500 and 200 ka, but they

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UNIPOLAR RECURRENT LEVALLOIS BLADE PRODUCTION

FIGURE 6.33. Two variants of the levallois technique for producing elongated lakes or blades (redrawn after boëda [1988], ig. 4). each method involves prior preparation of the core, indicated by small arrows in the lake preparation scars. each can produce multiple blades, in the irst case (the “unipolar” method) from a single striking platform and in the second (the “bipolar” method) from two opposed striking platforms. Continuing blade production requires continuing core preparation, but the process will eventually terminate when the core becomes too small for further working.

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become common only in Mousterian and MSA assemblages ater 200 ka. Not all Mousterian/MSA people produced Levallois lakes, but many did, and Levallois technology is sometimes regarded as a hallmark of the Mousterian/MSA. By replication and reitting, Eric Boëda has shown that Mousterian knappers actually employed a range of Levallois reduction techniques, variably designed to produce a single large lake (ig. 6.32), multiple smaller lakes, lakes that were naturally pointed at one end, or lakes that were exceptionally long (ig. 6.33). By convention, lake length is measured along a line that bisects the butt (or striking platform), and

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breadth is the maximum dimension perpendicular to this line. Flakes that are longer than they are wide are sometimes called lake blades, and ones that are at least twice as long as wide are called blades. Any extensive knapping session will produce some blades, but routine blade production requires special expertise in core preparation. Blades are usually considered a marker of the Upper Paleolithic culture complex that succeeded the Mousterian in Europe, western Asia, and northern Africa, but some Mousterian and MSA people produced numerous blades during the Last Interglacial or even before. As discussed below, blades or lake blades mark the opening phase of the LevalloisoMousterian complex in the Levant (the eastern Mediterranean coast and its hinterland). An even earlier Levantine industry has so many blades that it has been dubbed the “Pre-Aurignacian” for its rough resemblance to the very early Upper Paleolithic, Aurignacian Industry. Blades or lake blades also characterize many sub-Saharan MSA industries, and wellmade blades occur in late Acheulean assemblages at Kapthurin near Lake Baringo, Kenya, and at various sites in the South African interior. Blades tend to be rarer in the European Mousterian, but they are common at some sites, including Tönchesberg and Rheindahlen (Germany), Rocourt (Belgium), Seclin, (France), and other northwestern European sites that date to the end of the Last Interglacial or the early part of the Last Glaciation. In general, Mousterian/MSA people produced blades from one face of specially prepared Levallois cores. Like many Upper Paleolithic people, however, some Mousterian knappers may have struck blades from the top of a core around its entire periphery. Such a core comes to look like a prism, and prismatic blade production is sometimes taken as the pinnacle of knapping reinement since it maximizes the amount of cutting edge a single core can provide. As discussed below, many Mousterian/MSA people used Levallois technology almost exclusively, while others used it more rarely or not at all. he reasons for this variability are not well-understood, but the Levallois technique generally requires large nodules of high-quality raw material, and it was used most commonly in regions where suitable nodules were extremely abundant, such as parts of northern and southwestern France, and the Nile Valley. In advance, it might seem that Mousterian/MSA people who used non-Levallois technology were less sophisticated than their Levallois contemporaries, but this is not necessarily true. Analysis of the nonLevallois technology employed by the so-called Quina Mousterians of France shows that they reduced elongated lint cores systematically the way a picnicker would slice through a sausage (ig. 6.34). When they could, they selected appropriately shaped tubular nodules, but when they had to, they laked nodules or blocks of other shapes into tubular

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FIGURE 6.34. The technique that Quina mousterian people often used to “slice” lakes from an elongated core (redrawn after Turq [1992], ig. 6.5). The technique is less complex than the levallois method illustrated in igures 6.24 and 6.25, and it is also much less wasteful since it produces a very high percentage of immediately useful lakes.

raw nodule before the first flake is struck

top view after the first flake is struck

core before the second flake is struck

top view after the second flake is struck

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form. heir principal goal was to produce a series of lakes with roughly triangular cross sections on which a sharp edge was opposed by a very thick, dull one. he dull edge could have served as a inger rest during cutting or scraping, or it might have made hating easier, but whatever its purpose, Quina people showed great skill in producing lakes of the desired form. A distinct advantage of Quina-type reduction is that it generates relatively little waste (débitage), that is, lakes that are not intended for use. Experiments show that roughly 60%–75% of Quina lakes were usable fresh from the core, whereas Levallois reduction generally produces less than 30% immediately useful lakes. he bottom line is that Mousterian/MSA people not only had deep insight into the mechanics of stone laking but routinely performed lengthy and complex operations to transform raw stone nodules into desirable lakes and blades. A primary conclusion of this chapter is that they were more primitive than their Upper Paleolithic/Later Stone Age successors in many important behavioral respects, but their primitiveness clearly did not extend to the primary working of stone. In this respect they were as human as anyone, and they have never been surpassed. SOURCES: Mousterian stone reduction—François Bordes (Bordes 1947, 1961b) and other French archaeologists (Boëda 1988, 1991, 1993; Boëda et al. 1990; Geneste 1990; Turq 1992); the chaîne opératoire (Kuhn 1995; Mellars 1996; Schlanger 1994; Sellet 1993); reduction sequences to group Mousterian assemblages in SW Asia (Bar-Yosef and Pilbeam 2000); Levallois lakes in Acheulean assemblages (Beaumont and Vogel 2006; Moncel et al. 2005; Santonja and Villa 2006); varied Levallois techniques (Boëda 1988); blades in the early Levallois-Mousterian (Bar-Yosef 1995), in the MSA (hackeray 1992a), in the Kapthurin Acheulean (McBrearty et al. 1996), in South African late Acheulean sites (Beaumont and Vogel 2006), and in the European Mousterian (Kozlowski 1990a), especially in NW Europe during the Last Interglacial (Ameloot-Van der Heijden and Tufreau 1993; Conard 1990, 1992; Conard and Fischer 2000; Otte 1990a; Tufreau 1992); abundance of Levallois lakes in northern and southwestern France (Dibble and Rolland 1992; Mellars 1996; Rolland and Dibble 1990), Israel (Bar-

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Yosef 1992; Meignen and Bar-Yosef 1992), and in the Nile Valley (van Peer 1991); Quina Mousterian laking (Turq 1992)

Mousterian/MSA Retouched Tool Typology Edge damage and use wear show that many Mousterian/MSA lakes and blades were put to immediate use without further working. However, some lakes and blades were further reduced or modiied by removing small lakes or chips from one or more edges. Such modiication is called retouch, and the altered edges are said to be retouched. In most instances retouch chips were struck from the ventral surface, and the chip scars appear on the dorsal surface. Retouching blunts fresh edges, and it was thus done to alter the shape of an edge, to give it greater stability, or to resharpen it ater it had been dulled by use. Beginning with Gabriel de Mortillet and others in the nineteenth century, European archaeologists have emphasized the position and quality of retouch to deine diferent Mousterian tool types, and the pioneering French prehistorian François Bordes employed these criteria to formalize an inluential Mousterian/MSA typology (ig. 6.35a, b). He recognized a total of sixty-three discrete lake tool types, but his scheme centered on twenty-one types of sidescrapers (or racloirs, the French term, sometimes used in English), two or three types of retouched points, and three or four types of notched or denticulated pieces. He stressed these types because the underlying sidescraper, point, and denticulate classes dominate almost all Mousterian assemblages. Hand axes like those that mark the preceding Acheulean Industrial Complex tend to be rare in Mousterian/MSA sites, though small, usually triangular or cordiform (heart-shaped) ones characterize some Mousterian assemblages, mainly in France, and small pointed, bifacially retouched pieces that it the technical deinition of hand axes occur sporadically elsewhere, especially in central Europe. Endscrapers, burins, borers, backed pieces (lakes or blades with one edge intentionally dulled) and other so-called Upper Paleolithic/Later Stone Age (LSA) tool types also tend to be rare in Mousterian/MSA sites, and where they occur they could have been produced accidentally since they are usually atypical or poorly made. Numerous endscrapers and burins do mark some west Asian Mousterian assemblages, and well-made backed pieces typify localized Mousterian/MSA variants in both Africa and Europe, but most research on Mousterian/MSA retouched tool form and function has understandably focused on the far more common and widespread sidescrapers, points, and denticulates. Following long-standing tradition, Bordes deined a sidescraper as a lake on which one or more edges carry smooth continuous retouch. He distinguished diferent sidescraper types depending on how many edges

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FIGURE 6.35. basic mousterian and msa stone-tool types (as deined by bordes [1961b]). most assemblages are dominated by scrapers, points, and denticulates. The distinctions among the types are strictly formal rather than functional, and use wear studies show that there is no strong relation between type and function. Thus, pieces assigned to the same type often seem to have been used for different purposes, while different types often appear to have served broadly the same purpose.

typical Levallois flake

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Levallois point

retouched Levallois pseudo-Levallois point point

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simple, straight sidescraper simple, concave sidescraper

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simple, concave sidescraper

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were retouched, their shape (convex, concave, or straight) or a combination of shapes, their position with respect to the line bisecting the bulb of percussion, and so forth. He deined a point as a lake on which two continuously retouched edges converged directly opposite the striking platform (or butt). To qualify, the lake also had to be relatively thin; a thick, pointed lake became a “convergent sidescraper.” Finally, he deined a denticulate as a lake that was retouched to produce a ragged or serrate edge, comprising several adjacent indentations. He designated a denticulate with only a single indentation as a “notch.” Bordes believed that Mousterian people recognized essentially the same types or subtypes of sidescrapers, points, and denticulates as he did. However, this conclusion has been questioned, in part because the types frequently intergrade, and sorting retouched Mousterian/MSA pieces among types oten requires great tolerance for ambiguity and arbitrariness. he problem may be partly that some types or subtypes represent

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sidescraper on ventral surface

sidescraper with alternate retouch

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cm endscraper atypical endscraper

typical borer

typical burin atypical burin

Mousterian tranchet raclette atypical borer typical backed knife

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stages in the resharpening or refreshing of a tiny number of basic forms. Figure 6.36, for example, illustrates how reduction or rejuvenation could transform a relatively simple convex sidescraper (on which a single convex retouched edge is aligned perpendicular to the butt) into either a typical “convergent” form (on which two retouched edges converge to a point) or a classic “transverse” form (on which a single retouched edge lies opposite the butt). he logic behind igure 6.36 is appealing because resharpening undoubtedly occurred, and it could have altered sidescraper form in precisely the way the igure suggests. Indisputable rejuvenation lakes that removed the dulled edge of a retouched tool have been identiied at Combe-Grenal and La Micoque, southwestern France, at La Cotte de St. Brelade, Jersey, and at other Mousterian sites. Additionally, retouched lakes tend to be especially abundant in regions where high-quality raw material is relatively rare and rejuvenation of used lakes might have

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FIGURE 6.36. The manner in which progressive reduction can transform a simple, convex sidescraper into a double, biconvex form, and inally into a convex, convergent form (top) or into a more heavily retouched simple, convex form and inally into a convex, transverse form (bottom) (redrawn after dibble [1988a], igs. 1, 2). The sequences imply that some frequency variation in sidescraper types may relect only the extent to which mousterians at different sites refreshed or resharpened used scrapers.

simple, convex sidescraper

lightly retouched, simple, convex sidescraper

double, biconvex sidescraper

more heavily retouched, simple, convex sidescraper

convex, convergent sidescraper

convex, transverse sidescraper

been essential. But rejuvenation can explain only a portion of Mousterian retouched tool variability. To begin with, it fails to account for sites where sidescrapers or other reduced forms are numerous even though high-quality raw material for producing fresh lakes was readily available. It also neglects the existence of sidescraper-rich assemblages where the sidescrapers are relatively large when they should be small, assuming that an abundance of sidescrapers implies heavy reduction. Finally, it contradicts the observation that where transverse sidescrapers are most common—in the Quina variant of the French Mousterian (discussed below)—they rarely resulted from the reduction of other forms but were made on broad, short lakes that were specially struck to anticipate them. Short, wide lakes were also specially chosen for transverse sidescrapers at the Bir Tarfawi Mousterian site in the eastern Sahara, Egypt, and both in France and at Bir Tarfawi, transverse sidescrapers commonly lack the lateral, basal retouch they would have if they derived from simple, lateral forms. Whatever the precise reasons for variation in Mousterian/MSA tool form, Upper Paleolithic/LSA tool types tend to be much less variable (more standardized), and the result is that Upper Paleolithic/LSA tools are far easier to sort among a relatively large number of readily deined, discrete types. he contrast may relect a diference in intentionality;

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that is, Mousterian/MSA people may have tried less oten to impose a preconceived form on a raw lake or blade. Instead, unlike Upper Paleolithic/LSA people who were clearly concerned about the overall shape of inished tools, Mousterian/MSA people may have focused mainly on the character of an edge or the sharpness of a point. Another aspect of the diference may be that Mousterian/MSA people were mostly preoccupied with function, whereas Upper Paleolithic/LSA people were more concerned with style. In addition, as discussed below, Upper Paleolithic/ LSA tools may have served a wider variety of functions, including, for example, the working of bone, antler, ivory, and shell. he bottom line is that Upper Paleolithic tools are not only easier for a modern archaeologist to sort, they also tend to vary much more through time and space. In sum, unlike stone laking technology, which suggests no fundamental distinction between Mousterian/MSA people and their successors, tool typology implies a signiicant behavioral diference. Arguably this relects an underlying biological contrast in cognitive outlook. SOURCES: early deinition of Mousterian tool types (de Mortillet 1883); Bordes typology (Bordes 1961b); efect of resharpening on Mousterian types (Dibble 1988a, 1988b; Dibble and Rolland 1992; Rolland 1981; Rolland and Dibble 1990); rejuvenation lakes in Mousterian sites and the limitations of rejuvenation to explain tool variability (Mellars 1996); selection of broad, short lakes in the Quina Mousterian (Turq 1992) and at the Bir Tarfawi site (Close 1991); greater standardization of Upper Paleolithic tool types (Kuhn 1995; Mellars 1989b, 1996; Otte 1990a).

Mousterian/MSA Tool Function he names assigned to Mousterian/MSA stone-artifact types suggest that their functions are known, but this is mostly not the case. As outlined above, the types are deined on morphological grounds, and their functions are speculative, based on resemblances to historic tools of known function or on feasibility experiments with modern replicas. Even the conventional distinction between tools and manufacturing waste is strictly morphological. hus by convention, only retouched pieces have been classiied as tools, even though feasibility experiments show that many unretouched pieces could have been used efectively as knives, scrapers, and so forth. Additionally, as already noted, many unretouched pieces have macroscopic edge damage or microscopic wear polishes that formed during use. Polishes, striations, microfractures, edge rounding and other microscopic wear traces on both retouched and unretouched artifacts can show not only that an artifact was used but also how it was used (to cut, scrape, saw, etc.) and on what substance (wood, bone, hide, etc.). Ancient use is inferred from similarities to the wear patterns that form on modern replicas used in known ways. Microwear analysis has revealed the functions of many individual prehistoric tools, but it has some

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important limitations: it can be applied only to certain ine-grained rock types, it is time consuming, and it is only moderately accurate. It has not led to a reformulation of artifact typology along functional (vs. morphological) lines, nor is it likely to. his is partly because of its inherent practical and technical constraints and partly because of the loose relation between form and function. Analyses of French Mousterian tools illustrate both the power and the limitations of the microwear method. Microwear suggests that, as among the three major Mousterian tool classes, only two—notches and denticulates—were used mainly in just one way on just one material. he use was to shave, plane, or whittle, and the material was wood. Most notches and denticulates are ideally suited to wood working not only by their form but also by their tendency to have steeply retouched edges. Artifacts in the third major class—sidescrapers—are more variable in both form and edge angle, and it is therefore perhaps not surprising that they served a wider range of functions, not just to scrape or plane but also to cut or slice. hey were applied mostly to wood, followed by lesh, bone, and hide. Points (including convergent sidescrapers) were likewise used mostly on wood and less oten on bone or hide, and they difer from common sidescrapers mainly in stronger evidence that some were hated. he indications are polishes and striations not on the edges or the tip but on the ventral and dorsal surfaces below the tip, as if they formed by friction against a loosely attached wooden handle (ig. 6.37). At most Mousterian sites, wooden handles would long ago have vanished, and none are known from France. However, unusually dense (brown-coal) deposits at the Königsaue late Mousterian site, Germany, preserved two pieces of birch-bark pitch, one of which retained impressions from a wooden handle, a bifacial tool, and a human thumb. he pitch served as adhesive to anchor the tool to the handle, and it reinforces the more ancient indications for hating with birch-bark pitch from Campitello Quarry, Italy, noted in the previous chapter. Mousterian artifacts from Kebara and Qafzeh Caves, Israel, conform broadly to the French pattern. Most edge-worn Kebara and Qafzeh specimens were apparently used on wood, while smaller numbers were applied variously on lesh, hide, bone, and sot plant material. As in the French case, there was no obvious correspondence between artifact form and use mode. Unlike French points, however, some of the Israeli ones had symmetrically placed wear on their lower lateral edges (ig. 6.38), suggesting they were bound to wooden spear shats. he use of points as armatures could also explain the scars from hinge lakes or step lakes on the tips of some Israeli and Jordanian examples. Experiments show that such scars oten result when a point strikes bone or another hard object, including, for example, a rock near the actual target. Similar “impact fractures” have been observed on Mousterian points from La Cotte de

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upper limit of polish

0

5 cm

St. Brelade, Jersey. he likely Mousterian spear points were too large and heavy to arm long-distance missiles, and the spears to which they were attached must have been thrust or thrown close up. A broken Levallois point still embedded in the neck vertebra of a wild ass at Umm el Tlel, Syria shows that Mousterian hunters could throw or thrust with remarkable force. he Umm el Tlel point may have been ixed to a wooden shat with locally occurring bitumen (natural tar), traces of which have been observed on ive other Umm el Tlel tools and on one from the nearby site of Hummal. None of the tools with bitumen traces were points that are likely to have armed spears, and use-wear analysis implies that even most Levallois points at Umm el Tlel served as cutting tools not spear armatures. his underscores the loose connection between form and function in Mousterian tools. Microwear study of stone artifacts from Starosel’e Cave, Crimea (Ukraine), conirms that Mousterians oten used formally (typologically) similar pieces in diferent ways (or one piece in multiple ways) and that they mounted points, sidescrapers, and other formal types on wooden handles or shats. Some of the points that appear to have been hated exhibit impact fractures consistent with their use as spear armatures. Many Starosel’e artifacts preserve microscopic traces of plant and animal tissues that are argued to derive from human use (and not the burial environment), and the traces generally support the inferences from microwear.

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FIGURE 6.38. utilized levalloiso-mousterian artifacts from Kebara Cave, israel (redrawn after shea [1988], 447; [1989], 615). use wear studies indicate that the Kebara mousterians, like their european contemporaries, used their tools mainly on wood, though they also sometimes worked hide, bone, and other materials. unlike any known european mousterians, they may have also mounted points on wooden spear shafts with leather thongs.

projectile

projectile

hafting wear

hafting wear

hafting wear

hafting wear

retouched Levallois point

Levallois point

hide scraping wood scraping

Levallois blade

retouched Levallois blade

wood cutting

wood scraping

retouched Levallois flake (”sidescraper”)

bone scraping

bone scraping butchery

flake fragment truncated cortical flake

cm

hus, plant residues that could derive from mastic (adhesive) or binding ibers tend to concentrate on the same surfaces that were polished from rubbing against wooden handles or shats. Plant residues outnumber animal residues at Starosel’e, either because the people commonly used plant ibers and resins to bind tools to wooden handles, because they more oten processed plant foods or raw materials, including wood, or both. he use wear studies support three important generalizations. First, to the extent that worked material can be inferred, Mousterian tools were used primarily on wood. In contrast, Upper Paleolithic tools were used far more commonly on bone and lesh and especially on hide. he rarity of evidence for bone working in the Mousterian is in keeping with the rarity or absence of formal bone, antler, and ivory artifacts, which became common only in the Upper Paleolithic. Second, compared with Upper Paleolithic people, Mousterians more rarely mounted stone artifacts on handles or shats, and only Upper Paleolithic people produced stone

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points that were small, light, and aerodynamic enough to serve as longdistance projectile armatures. he contrast with the Mousterian is stark, especially since Upper Paleolithic people were also the irst to produce probable armatures in bone, antler, or ivory. he diference surely implies that Mousterian and Upper Paleolithic people difered fundamentally in hunting strategy and, more important, in hunting success. Finally, the absence of a strong one-to-one association between Mousterian tool form and function underscores the arbitrary nature of the conventional typology. In contrast, overall tool form and function appear to have been much more closely linked in the Upper Paleolithic. he diference once again suggests that form, as manifested in the eye of the modern archaeologist, was much less important to Mousterian toolmakers. MSA artifacts have not been extensively analyzed for function, but results from Sibudu Cave, South Africa, broadly recall those for the Mousterian. As at Starosel’e, the artifacts at Sibudu have been examined for microscopic organic residues, and these are especially common, owing to the persistently low moisture content of the deposits. All classes of Sibudu artifacts—including especially scrapers, points, and the backed, crescent-shaped objects referred to below as “segments”—retain residues. Plant residues dominate numerically, and on scrapers and points, they concentrate near the bases, where they probably derive from resin or iber used to ix tools on wooden handles or from friction with the handles. On segments, they concentrate on the backed (dulled) edges, where they probably represent the remnants of vegetal mastic (glue). As noted in the section “Pigment Collection and Utilization” below, mineral ocher, which was probably an emulsiier or hardening agent in the mastic, also concentrates near the bases of points and on the dulled edges of segments. Microwear and damage suggest that the Sibudu people used scrapers for a variety of purposes, independent of form, but impact fractures and microwear indicate that they oten ixed points and segments to wooden spear shats. In support of this inference, animal residues concentrate near the tips of points and on the fresh, unworked edges of segments. Experiments show that replicated segments function efectively as spear armatures. Like Mousterian points and backed elements, the Sibudu examples were generally too large and heavy to tip long-distance projectiles, and the Sibudu spears were probably thrust or thrown only short distances. Based on residue disposition alone, some of the points and segments might have been butchering tools rather than spear tips, and abundant cut marks on animal bones show that MSA people routinely butchered carcasses with stone tools, some of which could have been mounted. However, the MSA deposits at Klasies River Cave 1, South Africa, wellknown for its near-modern human fossils, provided the neck vertebra

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of a long-horned bufalo in which the tip of a stone point was still ixed. his conirms that like the Mousterians of Syria, the MSA inhabitants of South Africa hunted with stone-tipped weapons. In contrast to most MSA sites, ≠Gi, Botswana, and especially the Aduma complex, Ethiopia, have provided stone points that are small enough to have tipped projectiles, either darts propelled from a spearthrower or arrows shot from a bow. Both sites probably antedate 75–70 ka, and they might imply that the inhabitants possessed an advanced projectile technology, which is otherwise documented only ater 20–18 ka in Africa and Europe (discussion in the next chapter). If so, however, the technology remained bound to ≠Gi and Aduma, since other MSA sites, including Sibudu and other prominent South African examples with deposits that postdate 75–70 ka, contain few points or related artifacts that were small enough to tip darts or arrows. In this circumstance, it seems reasonable to propose that the ≠Gi and Aduma points are simply unusually small versions of the armatures that MSA people oten used to tip thrusting spears. Most authorities have long accepted that LSA/Upper Paleolithic people, ater 50–40 ka, originated projectile technology, and size-and-shape comparisons between LSA/Upper Paleolithic stone points and known stone dart points and arrow heads continue to support this conclusion. LSA/Upper Paleolithic people were also the irst to routinely produce bone artifacts that are likely to have been projectile points.

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SOURCES: microwear analysis of stone tools—general (Grace 1996; Jensen 1988; Keeley 1977, 1980; Semenov 1964; Shea 1992; van Gijn 1990), limitations (Jensen 1988; Moss 1987); microwear analysis of Mousterian tools from France (Anderson-Gerfaud 1990; Beyries 1987, 1988); Königsaue bark pitch (Grünberg 2002; Koller et al. 2001); microwear analysis of tools from Kebara and Qafzeh (Shea 1988, 1989); impact fractures on points from Israel and Jordan (Shea 1992) and La Cotte (Callow 1986); Uum el Tlel embedded point (Boëda et al. 1999) and bitumen (Boëda et al. 1996, 1998); Starosel’e use wear (Hardy et al. 2001); use wear contrast between Mousterian and Upper Paleolithic tools (Jensen 1988; Rots 2005); lack of projectile armatures in the Mousterian (Kuhn 1995; Shea 2006); greater link between tool form and function in the Upper Paleolithic (Jensen 1988); Sibudu Cave—organic residues on MSA tools (Williamson 2004) and microwear (Lombard 2004, 2005, 2008); experiments with replicated segments (Pargeter 2007); embedded stone tip at Klasies River (Milo 1998); ≠Gi and Aduma stone points {Brooks et al. 2006) and points from other MSA sites (Villa and Lenoir 2006); antiquity of projectile technology (Shea 2006)

Mousterian/MSA Interassemblage Variability As a whole, the Mousterian/MSA is remarkably uniform though time and space, and much supericial variability between assemblages from diferent regions or time periods can probably be ascribed to diferences in the size or laking quality of available stone raw materials. Flint of variable quality was available throughout much of Europe and western Asia but was generally absent in Africa, where quartzite and even volcanic rock types were widely used. Among the few tool types whose lo-

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calized distribution cannot be explained by diferences in raw material are the triangular and cordiform hand axes mentioned above (largely restricted to France, with extensions to Britain, Belgium, and Germany); small cleavers on lakes (cleaver lakes) found mainly in northern Spain; Keilmesser or “wedge knives” (elongated points that are usually blunt on one edge and bifacially retouched to a sharp edge on the other) found mainly in Germany and adjacent parts of central Europe; bifacially laked, leaf-shaped points (Blattspitzen) restricted mostly to central Europe and independently to an MSA variant in South Africa; backed elements, common in one of the Mousterian variants of southern France and, independently again, in an MSA variant in South Africa; and stemmed or tanged pieces (points, sidescrapers, etc.) restricted essentially to the western two-thirds of northern Africa. Except for these types and some others that are less strictly localized, most variability within the Mousterian/MSA is quantitative, that is, it consists of diferences in the percentage representation of the same widespread types of sidescrapers, points, and denticulates. SOURCES: raw material and Mousterian interassemblage variability (Dibble and Rolland 1992; Rolland and Dibble 1990)

Interassemblage Variability in Europe

Typological variability among Mousterian assemblages has been studied most thoroughly in Europe, above all in France, thanks to the descriptive artifact typology and analytic methodology developed by François Bordes. Working irst with data from such keystone French caves as Le Moustier, La Ferrassie, La Quina, Combe-Capelle, and Pech de l’Azé and later with material from his own meticulous, thirteen-year excavations at Combe-Grenal, Bordes recognized four basic Mousterian variants or facies: 1.

2.

3.

Mousterian of Acheulean Tradition (MAT). Characterized by numerous triangular or cordiform hand axes in addition to the sidescrapers, denticulates, and points that are ubiquitous in the Mousterian. In an earlier phase (MAT Type A), backed knives occur but are relatively rare. In a later phase (MAT Type B), they are much more numerous, essentially replacing small hand axes as the type implement of the MAT. Typical Mousterian. Characterized mainly by sidescrapers, with some denticulates and points. Hand axes and backed knives are rare or absent. Denticulate Mousterian. Characterized by the dominance of denticulates and notches, with much smaller numbers of sidescrapers, points, and such. Hand axes and backed knives are absent.

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4.

Quina-Ferrassie Mousterian (or Charentian). Characterized by a high proportion of sidescrapers; distinguished from the Typical Mousterian by abundant sidescrapers with distinctive steep, scalar retouch in which successive ranks of lake scars overlap like scales on a ish. Also diferentiated by the relative abundance of limaces—slug-shaped pieces that could be described as double convergent sidescrapers.

Within each variant, Bordes further distinguished between assemblages with many Levallois lakes and ones with few. He assigned Levallois-rich assemblages to the appropriate Levallois facies (for example, Typical Mousterian of Levallois Facies), except within the QuinaFerrassie variant, where he assigned them to the Ferrassie subvariant, as opposed to the Quina subvariant, in which Levallois lakes are rare. Apart from the Mousterian of Acheulean Tradition B, which was clearly later than Type A, Bordes argued that there was no chronological order to the facies and that any one of them could overlay any other within a site (ig. 6.39). He thought this was because the facies were produced by separate Mousterian tribes who moved from site to site, randomly replacing each other. In Bordes’s view, the separate tribes coexisted in France for tens of thousands of years, until one or perhaps two evolved into succeeding Upper Paleolithic tribes. He recognized that such prolonged social separation might produce discrete physical types, but human fossils so far do not suggest any diference. Specimens associated with the Typical Mousterian (at Le Moustier) and the Quina-Ferrassie Mousterian (at La Quina, La Chapelle-aux-Saints, La Ferrassie, Le Regourdou, Spy, and other sites) represent typical Neanderthals. Diagnostic human remains have not yet been found with the Denticulate Mousterian, but a child’s skull that was probably associated with the Mousterian of Acheulean Tradition Type B (MATB) at Pech de l’Azé Cave 1 is also characteristically Neanderthal. he Pech de l’Azé skull is particularly signiicant since Bordes thought the MATB may have anticipated the Upper Paleolithic Perigordian (or Gravettian), in which case the makers of MATB artifacts might have been anatomically modern humans. he Pech de l’Azé skull shows this was not the case. In addition, it probably dates from shortly before the time when the Upper Paleoltihic replaced the Mousterian, and it thus provides fossil evidence that the rupture was biological as well as cultural. Perhaps the most serious problem with Bordes’s tribal hypothesis is to imagine the social or cultural mechanisms that could have separated the tribes for tens of thousands of years. Another diiculty is that Bordes may have underestimated the amount of typological variability that is linked to time. hus when some questionably excavated or analyzed occurrences are ignored, there is a consistent sequence from Ferrassie Mousterian to Quina Mousterian to Mousterian of Acheulean Tradition within French

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Mousterian of Acheulean Tradition Mousterian

A A2-B2

Typical Mousterian Denticulate Mousterian Denticulate Mousterian Typical Mousterian

B3-B4 C D D2

Denticulate Mousterian Denticulate Mousterian Denticulate Mousterian Quina Mousterian Denticulate Mousterian

E1-2 F-H1 H I J

Quina Mousterian

K-M1

Ferrassie Mousterian Typical Mousterian

P Q-R1

Ferrassie Mousterian

U1-Y

Z Typical Mousterian Denticulate Mousterian α

Combe-Grenal (Dordogne)

caves. he principal obstacle to this chronological succession has been the sedimentologic/climatologic correlation of the sequences from Le Moustier (lower shelter) and Combe-Grenal, from which it appeared that the Mousterian of Acheulean Tradition at Le Moustier was as old as or older than the Ferrassie and Quina Mousterian at Combe-Grenal. However, TL dating of burned lints and ESR dating of animal teeth from Le Moustier, combined with climatic correlation of the Combe-Grenal sequence to the global oxygen-isotope stratigraphy, now suggests that the Mousterian succession at Combe-Grenal largely antedates that at Le Moustier (ig. 6.40), and the Ferrassie to Quina to Mousterian of Acheulean Tradition succession is thus supported. Moreover, some Mousterian assemblages excavated ater Bordes irst deined the facies clearly fall between these variants, suggesting that in part the facies variation is continuous rather than discrete. Partially in response to this problem, Bordes modiied and loosened the facies deinitions. Still, no one doubts there is signiicant quantitative variability among broadly contemporaneous Mousterian assemblages, not just in France but elsewhere in Europe where Bordes’s typological methods have been applied—for example, in Spain or in Ukraine and European Russia. Unlike Bordes and many French archaeologists, however, some prominent Anglo-American investigators believe the variation relects diferent tasks carried out by the same people (or their descendants) at diferent sites or at the same site at diferent times, perhaps at diferent seasons. hus denticulate-rich assemblages might indicate an emphasis on woodworking (bark stripping, whittling, or shaping), while sidescraper-rich

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FIGURE 6.39. schematic section through the deposits of Combe-Grenal Cave, southwestern france, showing the partially random interstratiication of mousterian facies (modiied after bordes [1961a], ig. 6). The apparently random pattern has been used to argue that the facies were produced by ethnically distinct mousterian groups who occupied france side by side for thousands of years.

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FIGURE 6.40. Correlation between the archaeological sequences at le moustier and Combe-Grenal, southwestern france, based on Tl dates for le moustier and climatic correlation of the Combe-Grenal sequence with the global marine stratigraphy (after mellars [1986]; mellars and Grün [1991]; valladas et al. [1986]). The sequence of ferrassie and Quina mousterian at Combe-Grenal is interrupted by a single level of denticulate mousterian in layer 20 and by three levels of either Typical or “attenuated” ferrassie mousterian in layers 28–30, but the correlated sequences nonetheless support the idea that the ferrassie mousterian, the Quina mousterian, and the mousterian of acheulean Tradition are temporally successive facies. if we assume that this succession is valid and has been properly dated at CombeGrenal, it permits some important southwestern french human fossils to be roughly ixed in time. it would also imply that much of the variability in the french mousterian relects temporal change within a single tradition rather than the coexistence of several separate traditions.

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Le Moustier

28

Combe-Grenal

Cro-Magnon

32 36 40 44

L

Aurignacian

K

Châtelperronian

J/I Typical/Denticulate H

48 52 56

G

Mousterian of Acheulean Tradition B

?

?

1-4

Mousterian of Acheulean Tradition A/B

5-16

Denticulate/ Typical

Mousterian of Acheulean Tradition A ? ?

60 64

Human Fossils

Saint-Césaire Le Moustier

La Chapelleaux-Saints

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4

68 72

27-35

Ferrassie

36-55

Typical/ Denticulate

La Ferrassie

76

5 115

Last Interglacial

504

Nonsequence >127 56-62

Acheulean

ones might relect a focus on food preparation or on hide processing (cutting and scraping). he activity variant hypothesis would clearly be strengthened if it could be shown that the principal tools that characterize each facies had a distinctive function. However, so far, ancient tool use-wear suggests just the opposite. As noted in the previous section, use-wear analysis implies that independent of tool type (denticulate or sidescraper), most tools in each facies were employed for wood working, whereas relatively few seem to have been used for butchering, hide working, or processing nonwoody plant materials. For reasons that may have to do with the geographic locations of the sites the tools come from, this contrasts with the results of use wear studies on (succeeding) Upper Paleolithic tools,

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most of which appear to have been used for working hides. However, it does not elucidate Mousterian facies variability. It is intriguing that a few Mousterian tools show wear from possible friction with a wooden hat or handle, but there is no indication yet that hating was more common in one Mousterian facies than in any other. Perhaps most important, there is no evidence for a close relation between Mousterian tool type and tool function. Since there is also no compelling evidence that diferent tool types or facies tend to associate with diferent kinds of bones or pollen, in the inal analysis the strongest argument for the activity variant model retains its plausibility. SOURCES: Mousterian facies in France (Bordes 1953, 1961a, 1968, 1972); artifactual associations of the child’s skull from Pech de l’Azé 1 (Soressi et al. 2007); facies variability linked to time (Mellars 1965, 1970, 1986, 1996); Le Moustier—dating by TL (Valladas et al. 1986) and ESR (Grün and Stringer 1991); climate correlation of the Combe-Grenal sequence (Laville et al. 1986); continuous facies variation (Freeman 1980); revision of the facies deinitions (Bordes 1981b); Mousterian interassemblage variation in Spain (Freeman 1980) and Ukraine/European Russia (Klein 1969b); activity variation to explain facies (Binford 1973; Binford and Binford 1966, 1969) and lack of support from use-wear analysis (Anderson-Gerfaud 1990; Beyries 1986, 1987, 1988)

Interassemblage Variability in Southwestern Asia

Southwestern Asia (the Near East) rivals western Europe as a focus of research on the Mousterian, thanks to a plethora of productive sites and to the eforts of many dedicated archaeologists for more than seven decades. he extraordinary potential of the region was irst revealed in the 1930s, when Dorothy Garrod excavated rich Mousterian layers at the caves of Tabun, Skhul, and El Wad in the Wadi el Mughara (Valley of the Caves) on the slopes of the biblical Mount Carmel. Today the most informative sites occur in two principal areas. he more fruitful region, with more than ity excavated sites, including the classic Mount Carmel caves, is the Levant, a relatively narrow strip of land between the Mediterranean coast and the Syro-Arabian deserts. In modern political terms it includes Israel, Lebanon, and the adjacent parts of Jordan and Syria, as far north as the southeastern corner of Turkey (ig. 6.4). he second, poorer area is the Zagros foothills of northeastern Iraq and northwestern Iran, with perhaps six excavated sites, including, most notably, Shanidar, Bisitun, Warwasi, and Kunji Caves. Sites with lake tools that could be called Mousterian (or Middle Paleolithic) have been found yet further east, in Afghanistan and particularly on the Indian subcontinent. However, the large majority of reported sites are undated or undatable, and most are surface occurrences that could contain artifacts of varying age. Levallois (prepared-core) lakes characterize many assemblages, and eastern India may mark the eastern limit of the Levallois technique, just as it marks the earlier limit of true Acheulean bifaces.

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he Levantine and Zagros Mousterian assemblages can be described using François Bordes’s typology, and they exhibit the same kind of quantitative interassemblage variability observed in Europe. In the Levant, however, typological change linked to time is more obvious. he sequence that has been established in the deeply stratiied deposits of Tabun Cave seems to characterize the entire region. Although the Tabun deposits lie entirely beyond the range of 14C dating, they are amenable to the TL and ESR methods. Both show that the Mousterian began before 130 ka, but TL implies a particularly long chronology (ig. 6.41). Recall (from chapter 2) that TL and ESR difer in the primary material to which they are usually applied—TL to heated lints and ESR to dental enamel. Both depend on estimates of the annual radiation dose (environmental dose rate) to which buried objects were exposed, and they can provide misleading results if the dose varied signiicantly across a layer or through time. Variation within layers is particularly well-documented at Tabun. As between the two chronologies, the TL version is probably the more trustworthy because the dated lints came from recent excavations, and it was possible to measure the annual radiation dose near their indspots. he dated teeth came mainly from older excavations because the recent excavations provided almost none, and it was necessary to estimate the annual radiation dose from adhering patches of sediment. Such estimates inevitably discount the efects of rocks or other large objects that are likely to afect the dose in a complex cave deposit. Arguably, the TL chronology may be more trustworthy, also because lint, unlike enamel, neither adsorbs nor loses uranium ater burial, and the calculation of TL dates therefore requires no assumptions about the pattern or rate of uranium uptake or loss. Near the base of the Tabun sequence, there is a late Acheulean industry that probably antedates 400 ka, based on TL dating. As discussed in the previous chapter, it grades typologically and technically into the succeeding Acheuleo-Yabrudian (or Mugharan) tradition that likely extended from about 400 to 250–200 ka. Acheuleo-Yabrudian assemblages include some that are relatively rich in small hand axes and others that are much richer in sidescrapers, but in the sidescraper component they are all broadly similar to the Quina-Ferrassie (Charentian) Mousterian of France. As noted in the previous chapter, in the midst of the AcheuleoYabrudian sequence at Tabun, there is an assemblage that is relatively richer in blades (lakes that are at least twice as long as wide) and in burins, endscrapers, and backed knives, alongside typical AcheuleoYabrudian sidescrapers. If the Acheuleo-Yabrudian elements are placed aside, the assemblage might anticipate the Aurignacian and other Upper Paleolithic industries that succeeded the Mousterian in western Asia and Europe ater 45 ka. hus, together with broadly contemporaneous

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3

ka

38/35 Early Ahmarian Emiran 46/47

50

4 5

Cultural Units

Tabun B Mousterian

100

Dederiyeh Kebara Amud Tabun I woman? Qafzeh MP

Dederiyeh? Amud

Kebara Tor Faraj Tor Sabiha Qafzeh

Tabun B

Skhul

Tabun II (jaw) Tabun C Hayonim E

6

ESR chronology

Ksar Akil Qafzeh UP

Skhul

Tabun C Mousterian

TL chronology

Hominin fossils

Tabun C

150 Douara?

200

Tabun D Mousterian

Tabun D

Hayonim F Yabrud I (1-10) Tabun D

7

Rosh Ein Mor

Tabun E

507 FIGURE 6.41. Paleolithic cultural stratigraphy and chronology of the levant (modiied after bar-Yosef and meignen [2001], 271). The thermoluminescence (Tl) and electron spin resonance (esr) methods that lie behind the chronology provide discrepant results, and the igure shows that the discrepancy tends to grow with age. both methods, however, place the near-modern humans from Qafzeh and skhul caves before the neanderthals from amud and Kebara Caves, and both show that the levantine mousterian began before 130 ka. as discussed in the text, the Tl chronology is probably more accurate.

250 Tabun E fragments

8

Zuttieyeh

300

9 10

11

Tabun E

Tabun E AcheuleoYabrudian

350

400

Tabun F Acheulean

Tabun F

blade-rich assemblages from the Zumofen Shelter (Adlun) in Lebanon, Yabrud Shelter I in Syria, and the Haua Fteah in northern Libya, the Tabun sample was once referred to as “Pre-Aurignacian.” However, the blades and other supposed Upper Paleolithic elements of the Pre-Aurignacian tend to be much more casually made than their true Upper Paleolithic counterparts, and detailed analysis suggests that the Pre-Aurignacian at Tabun was just a facies within a continuum of Acheuleo-Yabrudian variation that also includes a facies rich in sidescrapers and another somewhat less rich in small hand axes. he Pre-Aurignacian is thus now more commonly known as the Amudian. Both ESR and TL imply that the Amudian antedates the Upper Paleolithic by more than 100 ky, and

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it clearly also antedates other “precocious” industries, like the Aterian of northern Africa and the Howieson’s Poort of southern Africa, discussed below. At Tabun, the Acheuleo-Yabrudian grades typologically and technically into the true Mousterian that succeeds it. Depending on how the available dates are read, the transition occurred sometime between 250 and 200 ka. Almost everywhere in the Levant, including Tabun, this Mousterian tends to be rich in Levallois lakes, and it is oten called the Levalloiso-Mousterian. Broadly speaking, it resembles the French Typical Mousterian of Levallois Facies, but it can be subdivided into three chronologically successive industries that are designated Types D, C, and B, ater the successive Tabun Cave layers in which they occur. Type D, the earliest industry, is richest in blades and elongated Levallois points. In contrast, Types C and B emphasize squatter, thinner Levallois lakes, and Type B is distinguished from Type C by greater numbers of triangular points and retouched tools. he diference between Types C and B is in keeping with a diference in Levallois core reduction (lake removal), which tended to be bidirectional or radial in C and unidirectional-convergent in B. Assemblages that conform broadly to the D-C-B types occur widely in the Levant, and they can then be used to supplement or conirm dates at sites like Tor Foraj and Tor Sabiha, southern Jordan. Both sites appear to contain variants of the Tabun-B kind of Mousterian. he latest Mousterian assemblages in the Levant sometimes contain Levallois points on which the butt has been bifacially thinned, perhaps to facilitate hating. hese are known as Emireh points, ater Emireh Cave in Israel, and assemblages in which they occur have long been assigned to a separate Emiran phase or industry. he Emiran combines Levallois technology with abundant Upper Paleolithic-like endscrapers and burins, and it has oten been regarded as transitional between the Mousterian and the Upper Paleolithic. As discussed in the conclusion of this chapter, other potentially transitional industries are also known from Lebanon, Syria, and Turkey, and together with the Emiran, they might be better assigned to the “Initial Upper Paleolithic.” he sequence in the Zagros foothills is much less well-documented than the one in the Levant, but ininite 14C dates, together with climate inferences drawn from the sediments suggest that the Zagros Mousterian dates primarily from the earlier part of the Last Glaciation. It is therefore probably contemporaneous with the Type B Levalloiso-Mousterian of the Levant, between roughly 70 and 45 ka. It varies somewhat from site to site, but compared with the Levalloiso-Mousterian, it is poorer in Levallois lakes and richer in sidescrapers, points, and other characteristic Mousterian retouched pieces. he smaller size of the raw nodules available for laking may explain both the more limited use of the Levallois technique and the higher frequency of retouch. A similar kind of Mous-

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terian with relatively few Levallois lakes and numerous retouched pieces may have extended from the Zagros in a broad arc westward through the Taurus Mountains of Turkey, where it is especially well-represented in Karaïn Cave. he topmost Mousterian layers at Karaïn contain small, bifacial, leaf-shaped points like those that have oten been reported from central European and Balkan Mousterian sites, and the change may imply a reorientation in cultural connections from east to north during the Mousterian occupation of Turkey. In sharp contrast to Europe, in western Asia, Mousterian toolmakers included both Neanderthals and near-modern people. Table 6.4 lists the main fossil sites. Neanderthal-Mousterian associations are particularly well-documented in Israel at Tabun, Kebara, and Amud Caves; in Syria at Dederiyeh Cave; in Turkey at Karaïn Cave; and in Iraq at Shanidar Cave. Mousterian artifacts accompany the remains of robust modern or nearmodern people only in Israel, at Skhul and Qafzeh Caves and perhaps also at Tabun Cave. Conceivably, the list of Mousterian human remains from Israel should include a cranial fragment from Zuttiyeh Cave, but its stratigraphic provenience is uncertain, and it is perhaps most likely to have been associated with the hand ax–rich (Acheulean) facies of the Acheuleo-Yabrudian Tradition. If so, it would probably date to between 400 and 250–200 ka. It is too incomplete for absolutely irm diagnosis and as indicated previously, it could represent an early Neanderthal population, a population broadly ancestral to the robust early near-modern people found at Qafzeh, or a separate archaic human population linked to neither the Neanderthals nor Qafzeh. Until the late 1980s, it was commonly assumed that the Skhul and Qafzeh near-modern humans postdated the west Asian Neanderthals, and this appeared to be conirmed by aspects of artifactual change through the Tabun and Qafzeh sequences. However, the associated mammal fauna implied that the Qafzeh near-modern humans actually antedated the Tabun Neanderthals, for Qafzeh provided archaic rodent species that do not occur with Neanderthal fossils at Tabun, and Tabun provided a more modern rodent that is absent at Qafzeh. he faunas also implied a change in climate from warmer, drier interglacial conditions at Qafzeh to cooler, wetter glacial conditions at Tabun. he faunal indications have now been amply conirmed by multiple broadly concordant TL, ESR, and U-series age determinations. hese bracket the Qafzeh and Skhul near-modern people between about 120 and 80 ka, while they place the Kebara and Amud Neanderthals between roughly 65 and 47 ka (ig. 6.41). A reasonable inference is that the Neanderthals actually replaced nearmodern people in Israel at the beginning of the Last Glaciation, but there is one important complication. his concerns the human remains from Tabun Layer C, which comprise Tabun 1, the partial skeleton of a Neanderthal woman, and Tabun 2, an isolated mandible (table 6.4).

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TABLE 6.4. he principal west and central Asian Mousterian sites that have provided Neanderthal or nearmodern human remains.

Site

Human Remains

References

Tabun Cave, Israel

a partial skeleton with skull (Tabun 1) and some isolated postcranial elements and teeth, including seven grouped as “BC7”

(Bar-Yosef 1992; Bar-Yosef 1993; Bar-Yosef 1994b; Bar-Yosef 1995; Bar-Yosef & Callander 1999; Copeland 1975; Coppa et al. 2005; Jelinek 1981; Jelinek 1982a; Jelinek 1982b; Jelinek 1992; Marks 1992; McCown & Keith 1939; Meignen & Bar-Yosef 1989; Meignen & Bar-Yosef 1992).

Kebara Cave, Israel

partial skeletons of an infant and an adult, twenty-seven isolated cranial and postcranial pieces from additional individuals

(Arensburg et al. 1985; Bar-Yosef et al. 1992; Bar-Yosef et al. 1986; Schwarcz et al. 1989; Smith & Arensburg 1977; Valladas et al. 1987)

Amud Cave, Israel

a partial adult skeleton, cranial fragments of two other adults, a partial infant skeleton, and fragmentary remains of perhaps twelve other, mainly very young individuals

(Hovers et al. 1996; Hovers et al. 1995; Rak et al. 1994; Suzuki & Takai 1970; Valladas et al. 1999)

Dederiyeh Cave, Syria

partial skeletons of two infants plus fragments of two other children and at least 6 adults

(Akazawa & Muhesen 2002; Akazawa et al. 1995; Kondo et al. 2000)

Karaïn Cave, Turkey

a fragmentary let mandible, isolated teeth, four hand phalanges, and ive or six other postcranial fragments from an unspeciied number of individuals

(Otte et al. 1998)

Shanidar Cave, Iraq

nine partial skeletons

(Solecki 1963; Solecki 1971; Stewart 1977; Trinkaus 1983b)

Teshik-Tash Cave, Uzbekistan

a child’s skeleton

(Movius 1953b; Weidenreich 1945)

Okladnikov Cave, southern Siberia (Russia)

an adult phalanx and humerus fragment and subadult humerus and femur fragments

(Krause et al. 2007b)

Skhul Cave, Israel

seven partial skeletons and the isolated bones of three other individuals

(Grün et al. 2005; McCown & Keith 1939; Mercier et al. 1993; Stringer et al. 1989)

Qafzeh Cave, Israel

ive partial skeletons and the cranial or postcranial fragments of as many as ten additional individuals

(Schwarcz et al. 1988b; Valladas et al. 1988; Vandermeersch 1981)

Tabun Cave, Israel

an isolated mandible (Tabun 2)

(Quam & Smith 1996; Rak 1998; Stefan & Trinkaus 1998)

Neanderthals

Near-Modern Humans

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In its chin development and in some features of the ascending ramus (ig. 6.12), the Tabun 2 mandible appears modern, but in other features, including especially the large size of its incisors relative to its cheek teeth, it appears more Neanderthal-like. Arguably, it is in fact neither modern nor Neanderthal but “near-modern” in the same sense as the fossils from Qafzeh, Skhul, Klasies River Main, and other African sites. ESR and TL dates show that it dates from broadly the same Last Interglacial period as the Skhul-Qafzeh fossils, which means it antedates the Neanderthals of nearby Kebara and Amud. he Tabun 1 skeleton is more problematic, not for its physical identity, which is unqualiiedly Neanderthal (vs. near-modern), but for its dating. Unlike the Tabun 2 mandible, which came from deep within Layer C, the Tabun 1 skeleton lay at the interface between layers B and C, and the excavator believed it could represent a burial from layer B. It has usually been assigned to Level C, and if this is accepted, its artifact associations and the available dates imply that it would date from the Last Interglacial, like the Skhul/Qafzeh nearmodern humans. However, unpublished excavation records have once again raised the possibility that it originated from Layer B. In this case, it would be broadly coeval with the Neanderthals from Kebara and Amud, and the best-dated west Asian Neanderthals would all postdate the Qafzeh and Skhul near-moderns. Tabun also provided seven isolated maxillary teeth whose stratigraphic provenience is now unclear, but whose uranium content suggests an origin within layer B. hey may represent a single individual, provisionally designated BC7 to allow for the possibility they came from layer C. he teeth appear to represent a Neanderthal, and if their stratigraphic position and age have been correctly assessed, they further suggest that Neanderthals succeeded near-modern humans in Israel. It should be emphasized that the pattern does not depend strictly on numeric dates, which are oten problematic for the period involved, but on the assumed stratigraphic origin of key fossils within sites and on correlations between sites based on artifact assemblages or fauna. Artifact stratigraphy also shows that the Upper Paleolithic succeeded the Mousterian in western Asia, beginning perhaps 50 ka. he last Mousterians appear to have been Neanderthals, and the earliest Upper Paleolithic artifact makers were fully modern humans expanding from Africa. he actual origins of the Upper Paleolithic, however, remain murky. At Tabun Cave, an acceleration in the rate of change in lake thickness in the latest Mousterian layers may anticipate the more radical artifactual change to come. And at Boker Tachit, also Israel, a stratiied alluvial sequence tentatively bracketed between 47 and 38 ka by 14C and U-series dating records progressive change from a Levallois point technology to a true blade technology, with a concomitant increase in Upper Paleolithic tool types, especially endscrapers. As already noted, other sites in the

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Levant have provided artifact assemblages that combine Mousterian-style Levallois technology with typically Upper Paleolithic tool types and in parallel with Boker Tachtit, these could document the birth of the Upper Paleolithic. If so, however, the process was lengthy and gradual and it implies a degree of cultural and probable biological interaction that is diicult to reconcile with the limited genetic and fossil evidence for Neanderthal/modern human interbreeding. he conclusion of this chapter revisits this issue. SOURCES (also table 6.4): Garrod excavations on Mt. Carmel (Garrod and Bate 1937); Middle Paleolithic of India (James and Petraglia 2005); absence of the Levallois technique in China (Gao and Norton 2002); dating of the Levantine Paleolithic—TL (Mercier and Valladas 1994, 2003; Mercier et al. 1995b, 2000), ESR (Rink et al. 2004; Schwarcz 1994; Schwarcz and Rink 1998); Tabun Paleolithic sequence (Bar-Yosef1998a, 2000; Bar-Yosef and Meignen 2001; Copeland 1975; Jelinek 1982b; Ronen 1979; Shea 2007); Pre-Aurignacian (Amudian) (Copeland 2000; Jelinek 1981; McBurney 1975); age of Acheuleo-Yabrudian/Mousterian interface (Porat et al. 2002); Tor Faraj and Tor Sabiha (Henry 1995); Emiran Industry (Garrod 1955; Marks 1983); west Asian Initial Upper Paleolithic (Bar-Yosef 2000; Kuhn et al. 1999); Zagros Mousterian (Baumler and Speth 1993; Dibble and Holdaway 1993; Skinner 1965; Solecki 1963); Taurus Mousterian (Kuhn 2002; Rink et al. 1994); Zuttiyeh artifacts and age (Bar-Yosef 1995; Gisis and Bar-Yosef 1974), human fossil as Neanderthal (Trinkaus 1982, 1983b), as proto-modern (Vandermeersch 1982, 1989), as neither (Simmons et al. 1991); Neanderthals possibly before modern humans in Israel (Jelinek 1981, 1982a, 1982b), later than modern humans (Bar-Yosef and Meadow 1995; Bar-Yosef and Vandermeersch 1981; Tchernov 1988a, 1992, 1994); numeric dating of Neanderthals and modern humans in Israel (Grün et al. 1991; Grün and Stringer 1991; McDermott et al. 1993; Mercier et al. 1995b; Mercier and Valladas 1994; Schwarcz 1994; Schwarcz et al. 1988b, 1989; Stringer et al. 1989; Valladas et al. 1987, 1988); Tabun 2 as modern (Quam and Smith 1996; Rak 1998) or as Neanderthal (Stefan and Trinkaus 1998); stratigraphic position and age of Tabun 1 (Bar-Yosef and Callander 1999; Garrod and Bate 1937, 64); Tabun BC7 (Coppa et al. 2005); Boker Tachit (Marks 1981a, 1983)

Interassemblage Variability in Northeastern Africa

In northeastern Africa, the Mousterian in the narrow sense is best known at the Haua Fteah (Great Cave) in Cyrenaican Libya and at open-air sites in Nubia, straddling the border between Egypt and the Sudan along the Nile. Excavations at the Haua Fteah penetrated to a depth of 13 m without reaching bedrock. he lowermost horizons provided an assemblage with elongated lakes or lake blades that were sometimes modiied into crude angle burins. In its vague resemblance to the much later Upper Paleolithic, the assemblage recalls the Amudian or Pre-Aurignacian of Israel and Lebanon, and it may date to the same interval, sometime between 400 and 250 ka. he overlying layers contained Levallois lakerich Mousterian assemblages that have been likened variously to the Levalloiso-Mousterian of the Levant or to the Nubian Mousterian, described immediately below. Radiocarbon dates show that latest Mousterian layers at the Haua Fteah antedate 40 ka, and associated fauna and sediments suggest the Mousterian sequence minimally spans the Last Interglacial and the earlier part of the Last Glaciation. he succession is divided between earlier and later parts by layers with possible tanged

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or stemmed elements that could represent the Aterian variant of the Mousterian, discussed in the next section. One of the later Mousterian horizons produced two let human ascending mandibular rami that are characteristically modern (as opposed to Neanderthal) in the symmetry of the sigmoid notch and the subequal size of the mandibular condyle and the coronoid process (ig. 6.12), hey thus reconirm the modern or near-modern anatomy of north African Mousterians. he Mousterian was followed at the Haua Fteah by the Dabban Industry, named for its occurrence at nearby ed Dabba Cave. he Dabban is characterized by typical Upper Paleolithic endscrapers, burins, and backed bladelets, but it also contains chamfered (bevel-ended) blades recalling those from the Initial Upper Paleolithic horizons at Ksar Akil and neighboring sites in Lebanon. Future research may show that the Dabban comprises an early phase that might be called “transitional” or Initial Upper Paleolithic in the west Asian sense and a later phase that was fully Upper Paleolithic like the west Asian Ahmarian Industry. Radiocarbon dates indicate that the Dabban extended from 40 ka or before until 20 ka or later, and except for Nazlet Khater 4 in the Nile Valley of Egypt, the Dabban sites in Cyrenaica are the only ones that unquestionably record this interval in northern Africa. Nazlet Khater 4 has provided a classic Upper Paleolithic blade industry radiocarbon-dated to 31.5 ka. In Nubia, the various Mousterian occurrences are scattered along the Nile, and there are no long, deeply stratiied sequences. his makes it dificult to distinguish variation due to time from variation due to activities or location. Several kinds of assemblages have been described, including ones that resemble the French Typical Mousterian, others that are more similar to the Denticulate Mousterian, and yet others that are relatively unique, particularly those with numerous burins that were once assigned to a separate “Khormusan” industry. Levallois lakes tend to be common in all Nubian Mousterian assemblages, but some also contain Levallois points that were produced by a distinctive method. his involved preparation of the surface of a core so that a knapper could then strike of pointed lakes along a central ridge. hese specially produced points are mostly rare, but their presence has been used to deine a “Nubian (Mousterian) Complex” that contains them as opposed to a “Lower Nile Valley (Mousterian) Complex” that does not. In this scheme, the early part of the Nubian Complex is also distinguished by occasional hand axes and leaf-shaped bifaces, and the later part includes the Khormusan, with its abundant burins. he Lower Nile Complex includes all those assemblages that were previously likened to the Denticulate Mousterian and others that occur where Mousterian people quarried chert from alluvial terraces along the lower course of the Nile. he Nubian and Lower Nile Valley Complexes may be artiicial constructs, superimposed on a single, long-lasting, variable Mousterian

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tradition, but if their reality is accepted, their distributions overlapped in the Nile Valley and the adjacent deserts until the beginning of the Last Glaciation. During the Last Interglacial, the Nubian Complex is wellrepresented at Bir Tarfawi and Bir Sahara to the west of the Nile and in Sodmein Cave in the Red Sea Hills to the northeast. At Bir Tarfawi and Bir Sahara, putative Nubian Complex assemblages are oten associated with now-defunct springs or shallow lakes that indicate relatively moist conditions, probably mainly in the Last Interglacial, but perhaps also partly in the early part of the Last Glaciation. At Sodmein Cave, TL analysis of burned lint fragments centers the earliest Nubian Complex occupation on 118 ka, within the Last Interglacial. A later assemblage in the topmost Mousterian layer contains two basally thinned Levallois points like those that mark the Emiran (Initial Upper Paleolithic) Industry of Israel. During the early part of the Last Glaciation, Nubian Complex assemblages disappeared from the Nile Valley and Lower Nile Complex assemblages disappeared from the adjacent deserts. At Taramsa Hill in the Nile Valley, a late manifestation of the Lower Nile Complex is distinguished by a Levallois technique oriented toward blade production, and this may anticipate the Upper Paleolithic in the same sense that the Initial Upper Paleolithic does in western Asia. Radiocarbon dating of charcoal at Taramsa Hill and at the nearby site of Nazlet Safaha shows that the late blade-rich phase antedates 38 ka. At Taramsa Hill, it is associated with a child’s skeleton, bracketed by TL readings on enclosing sands between 80.4 and 49.8 ka. he skeleton was fragmentary and poorly preserved, but in its steeply rising frontal, rounded occipital, large, prognathic face, and slender limb bone shats, it broadly recalls some of the near-modern skeletons from Qafzeh Cave, Israel. It was buried in a trench that late Nile Complex Mousterians dug to extract chert for toolmaking, and until now, it provides the best indication for a Mousterian/MSA grave in Africa. SOURCES: Mousterian at the Haua Fteah (McBurney 1967) and in Nubia (Marks 1968a, 1968b; Wendorf et al. 1979, 1994a; Wendorf and Schild 1976, 1992); similarity of the Haua Fteah and Nubian Mousterian (Crew 1975); Haua Fteah mandible fragments (Rak 1998; Tobias 1967b); Nazlet Khater 4 (Vermeersch et al. 1982); Nubian Levallois points—rarity (Schild 1998) and deining feature of the Nubian (vs. Lower Nile) Mousterian Complex (van Peer 1998; Vermeersch 2001); artiiciality of the Nubian and Lower Nile complexes (Schild and Wendorf 2002–2004); Bir Tarfawi and Bir Sahara (Close et al. 1990; Wendorf et al. 1987, 1993; Wendorf and Schild 1980); Sodmein Cave (Mercier et al. 1999); Taramsa Hill (Vermeersch et al. 1998)

Interassemblage Variability in Northwestern Africa

In northwestern Africa, the Mousterian narrowly deined is represented at no more than ten-to-twelve sites, but the Aterian Industry or facies, with its characteristic stemmed pieces (ig. 6.42), is abundant. Except

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515 FIGURE 6.42. aterian artifacts from algeria (redrawn after vaufrey [1955], ig. 44). The tanged or stemmed pieces are the principal artifacts that distinguish the aterian from the mousterian.

stemmed points

stemmed flake stemmed scrapers 0

5 cm

ordinary (nonstemmed) scrapers

for the stems, the tool types involved are primarily the same kinds (sidescrapers, points, denticulates, etc.) that characterize Mousterian assemblages elsewhere, and most Aterian assemblages contain numerous Levallois lakes. Especially in the Sahara, some are also characterized by well-made bifacial leaf-shaped points. he Aterian is best known in the Mediterranean borderlands of northwestern Africa (the Maghreb) from eastern Morocco to Cap Blanc in Tunisia, but it also occurred through much of the Sahara, southward to Mauritania, Mali, and northern Niger and eastward to Kharga and Dungul Oases in the western desert of Egypt (ig. 6.20). It apparently did not cross the Nile Valley. At Mugharet el ‘Aliya, Taforalt (Pigeon Cave), Rhafas Cave, and the Météo alluvial site in Morocco and in alluvial (wadi) sequences at Kharga Oasis and other Saharan localities, the Aterian postdates the local Mousterian, but in Cyrenaica and the Nile Valley, the Mousterian continued long ater the Aterian had appeared elsewhere.

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Like Mousterian assemblages in northern Africa, Aterian ones often occur in a sedimentologic-paleontologic context indicating relatively moist conditions. his is above all true in the Sahara, where Aterian occupations usually occur in lake sediments in places where standing water is absent today. At stratiied caves such as Taforalt and Dar es Soltan 2 in Morocco, Aterian layers are separated from overlying Upper Paleolithic or Epi-Paleolithic ones by a long occupation gap, from 40 ka or before to 20 ka or later. As indicated previously, northern Africa appears to have been signiicantly wetter only during interglacials, and the occupation hiatus between the Aterian and later Paleolithic industries may thus span the entire early and middle part of the Last Glaciation, from roughly 71 to 20 ka. Except perhaps for Cyrenaica, there is little or no indication for signiicant human populations anywhere in northern Africa during the hiatus, probably because the entire region sufered from extreme aridity in the earlier and middle parts of the Last Glaciation. As indicated above, the relatively sparse human remains associated with Mousterian/Aterian artifacts in northwestern Africa came from people who were clearly distinct from the Neanderthals and whose skull and facial form approached those of modern people. SOURCES: abundance and distribution of Mousterian and Aterian (Camps 1974; Debénath 1994; Ferring 1975; van Peer 1991; Wendorf and Schild 1992; Wengler 1986); long hiatus between the Aterian and the Epi-Paleolithic (Camps 1974, 1975; Close 2002; Cremaschi et al. 1998; Wendorf et al. 1990; Wendorf and Schild 1992); aridity in the earlier and middle parts of the Last Glaciation (Guillien and Laplace 1978)

Interassemblage Variability in Sub-Saharan Africa

MSA artifact assemblages have been recovered in western Africa, but they are much better known in eastern Africa and especially in southern Africa, where the MSA was irst deined. MSA sites are scarce in western Africa partly because few archaeologists have looked for them and partly because they occur mainly in rain forest settings that impede site discovery. In addition, rain forests have probably never supported signiicant hunter-gatherer populations because they provide limited foraging opportunities. Phytoliths (minute silica particles that form in plant tissues) and pollen recovered in or near some west African sites indicate they formed under conditions when the forest was thinned or reduced. he eastern and southern African sites occur in areas that were never heavily forested, and the historic lack of forest facilitated site discovery. he eastern and southern Africa sites are mostly surface occurrences where materials of mixed age may occur and where reliable dating is impossible. However, there are also many sites where artifacts and other debris are stratiied in place, which allows archaeologists to estimate numerical age and to establish cultural successions. Table 6.5 lists the especially well-

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known stratiied MSA sites in southern Africa, deined here as Africa south of 9°S latitude. Figure 6.22 locates the sites. Boldface in the igure marks sites that contain particularly long sequences with well-preserved bones, shells, or both. Some of the southern African sites are particularly signiicant because they are said to anticipate the Later Stone Age. Table 6.5 includes 70 MSA sites, which is far fewer than the number of like-aged, stratiied Mousterian sites in western Europe and signiicantly smaller even than the number in some regions of France. he contrast with France is especially notable because southern Africa is so much larger, but France has a much longer tradition of stone age archaeology, a much larger number of archaeologists, more favorable logistics for site discovery and excavation, and a greater number of locales in which sediments have sealed ancient archaeological objects. France is particularly rich in limestone caves, and in common with other parts of Europe, it also contains spreads of windblown silts (loesses) that rapidly buried sites during glacial intervals. Southern Africa has many fewer limestone caves, and it lacks deposits comparable to the European loess sheets. Compared to Europe and eastern Africa, it also has few lake basins in which archaeological objects could become trapped and buried. Among all southern African regions, South Africa comes closest to France in the density of known MSA sites, partly because it has a relatively long history of archaeological research and partly because it is relatively rich in deeply stratiied caves, particularly along the southern and western coasts. Some of the MSA sites in table 6.5 are relatively uninformative because excavation mixed layers within the MSA or between the MSA and the overlying LSA or because the MSA samples are small or inadequately described. Still, the number of well-excavated sites with large, welldocumented samples is suicient to illuminate the nature of the MSA and to establish what is usual and what is exceptional. his is important because unlike European Mousterians, MSA Africans included the ancestors of both LSA Africans and Upper Paleolithic Europeans, and their archaeological residues might thus be expected to diverge from Mousterian residues in the direction of the LSA/Upper Paleolithic. For example, unlike LSA/Upper Paleolithic artifact assemblages, Mousterian assemblages rarely include bone artifacts or art objects, and where they do, either the artifactual quality or the stratigraphic origin of the objects is usually questionable. he large majority of MSA assemblages are similar—undoubted bone artifacts or art objects are rare, and where they do occur, they are mainly isolated objects whose artifactual nature or stratigraphic associations are debatable. However, as discussed below, sealed MSA layers at two South African sites (Blombos Cave and Diepkloof Rockshelter) have provided relatively abundant bone artifacts, proposed art objects, or both, and this raises the question of whether the layers are

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TABLE 6.5. he principal stratiied Middle Stone Age (MSA) sites in southern Africa. Figure 6.22 shows their approximate locations. Asterisks mark sites with particularly deep sequences, and boldface marks those that preserve bone.

Site

References

ANGOLA Humpata Plateau Cave

(França 1964)

Chambuage Mine

(Clark 1968a)

ZAMBIA Kalambo Falls

(Clark 1964; Clark 1969; Clark 1974a; Clark 2001; Sheppard & Kleindienst 1996)

*Kalemba Cave

(Phillipson 1976)

*Mumbwa Caves

(Avery 2000; Barham 2000a; Barham 1993; Barham 1995; Barham 2000b; Barham & Debenham 2000; Clark 1942; Clark 1950; Dart & Del Grande 1931; Klein & Cruz-Uribe 2000a)

Twin Rivers Fissure

(Avery 2002; Barham et al. 2000; Bishop & Reynolds 2000; Clark 1971; Clark & Brown 2001)

MALAWI Malowa Shelter

(Denbow 1973)

ZIMBABWE *Pomongwe Cave

(Brain 1981; Cooke 1963; Walker 1987; Walker 1990)

Tshangula Cave

(Cooke 1963; Cooke 1971)

Zombepata Cave

(Cooke 1971; Sheppard & Swart 1973)

Nswatugi Cave

(Walker 1978b; Walker 1980; Walker 1987)

*Bambata Cave

(Jones 1940; Jones 1949)

*Redclif Cave

(Brain & Cooke 1967; Cooke 1978a; Cruz-Uribe 1983; Klein 1978b)

Pfupi Shelter

(Robinson 1952)

Mtemwa Rocks Shelter

(Robinson 1952)

BOTSWANA ≠Gi

(Brooks et al. 1980; Brooks & Yellen 1977; Helgren & Brooks 1983)

*White Paintings Shelter

(Donahue et al. 2004; Feathers 1997; Robbins 1990; Robbins et al. 2000)

NAMIBIA

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Zais Shelter

(Cruz-Uribe & Klein 1981/83; Wendt 1972)

Zebrarivier Cave

(Cruz-Uribe & Klein 1981/83; Freundlich et al. 1980; Wendt 1972)

Bremen 1 Shelter

(Cruz-Uribe & Klein 1981/83; Wendt 1972)

Pockenbank 1 Shelter

(Freundlich et al. 1980; hackeray 1979; Wendt 1972)

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TABLE 6.5. (continued )

Site

References

*Apollo 11 Cave

(Freundlich et al. 1980; hackeray 1979; Wendt 1972; Wendt 1976)

LESOTHO Melikane Shelter

(Carter 1976; Vogel et al. 1986)

Ha Soloja Shelter

(Carter 1976)

Moshebi’s Shelter

(Carter & Vogel 1974)

Sehonghong Shelter

(Carter et al. 1988; Carter & Vogel 1974; Mitchell 1993; Mitchell 1994)

SWAZILAND Lion Cavern

(Beaumont 1973b; Boshier & Beaumont 1972; Dart & Beaumont 1968)

Sibebe Shelter

(Price-Williams 1981; Vogel et al. 1986)

SOUTH AFRICA *Cave of Hearths

(Cooke 1962; Latham & Herries 2004; Mason 1962; Mason 1988b; Sampson 1972; Sampson 1974)

Olieboompoort Cave

(Mason 1962)

Kalkbank

(Brown 1988; Cooke 1962; Mason 1988a)

Mwulu’s Cave

(Mason 1962; Sampson 1972; Tobias 1949)

*Bushman Rockshelter

(Brain 1969; Elof 1969; Louw 1969; Plug 1981; Vogel 1969)

Witkrans Shelter

(Butzer et al. 1978b; Clark 1971; Klein 1980)

*Wonderwerk Cave

(Beaumont 1990b; Beaumont & Vogel 2006; Binneman & Beaumont 1992; Malan & Cooke 1941; Malan & Wells 1943; van Zinderen Bakker 1982; Vogel 2001)

Kathu Pan

(Beaumont 1990a; Beaumont et al. 1984; Butzer 1984b; Klein 1988a)

Florisbad Spring Site

(Brink 1987; Brink 1988; Butzer 1988; Douglas 2006; Kuman & Clarke 1986; Kuman et al. 1999; Scott & Rossouw 2005; van Zinderen Bakker 1990)

Orangia 1

(Sampson 1974)

Oakleigh Farm QOB Shelter

(Derricourt 1977)

Grassridge Shelter

(Opperman 1988)

Highlands Shelter

(Deacon 1976)

*Rose Cottage Cave

(Clark 1997a; Clark 1997b; Clark 1999; Harper 1997; Plug & Engels 1992; Wadley 1991; Wadley 1996; Wadley 1997; Wadley & Harper 1989; Wadley & Vogel 1991; Woodborne & Vogel 1997)

Holley Shelter

(Cramb 1952; Cramb 1961; Wadley & Jacobs 2004)

Strathalan Cave B

(Opperman 1992; Opperman 1996; Opperman & Heydenrych 1990)

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TABLE 6.5. (continued )

Site

References

*Border Cave Lincoln Cave, Sterkfontein

(Kuman & Clarke 2000; Reynolds et al. 2007; Reynolds et al. 2003)

*Sibudu Cave

(Cain 2004; Lombard 2003; Lombard 2005; Plug 2004; Schleigl et al. 2004; Villa et al. 2005a; Wadley 2001; Wadley 2004; Wadley 2005a; Wadley 2007; Wadley & Jacobs 2004)

*Umhlatuzana Rock Shelter

(Kaplan 1989; Kaplan 1990)

*Boegoeberg 2

(Klein & Cruz-Uribe 1996; Klein et al. 1999b)

Klipfonteinrand Shelter

(Parkington & Poggenpoel 1971; Volman 1981)

Hollow Rock Shelter

(Evans 1994)

Sibebe Shelter

(Price-Williams 1981)

*Diepkloof Rockshelter

(Parkington & Poggenpoel 1987; Parkington et al. 2005; Rigaud et al. 2006)

Elands Bay Cave

(Butzer 1979; Volman 1981)

Hoedjies Punt Middens

(Berger & Parkington 1995; Matthews et al. 2005; Parkington 2003; Steele & Klein 2005/06)

Sea Harvest Midden

(Steele & Klein 2005/06; Volman 1978)

*Ysterfontein 1 Rockshelter

(Halkett et al. 2003; Klein et al. 2004)

*Montagu Cave

(Butzer 1973a; Goodwin 1929b; Keller 1973; Volman 1981)

*Peers Cave (= Skildergat (Anthony 1967; Jolly 1947; Jolly 1948; Volman 1981) = Fishhoek)

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Tunnel Cave

(Malan 1955)

Skildergatkop

(Malan 1955)

*Die Kelders Cave 1

(Avery et al. 1997; Feathers & Bush 2000; Goldberg 2000; Grine 1998; Grine et al. 1991; Klein & Cruz-Uribe 2000b; Marean et al. 2000b; Schwarcz & Rink 2000; Schweitzer 1979; Tankard & Schweitzer 1976; hackeray 2000)

Herolds Bay Cave

(Brink & Deacon 1982)

*Blombos Cave

(d’Errico et al. 2001; d’Errico et al. 2005; Grine & Henshilwood 2002; Grine et al. 2000; Henshilwood 2004; Henshilwood 2005; Henshilwood et al. 2001a; Henshilwood et al. 2001b; Jacobs et al. 2003a; Jacobs et al. 2003b)

Cape St. Blaize Cave (= Mossel Bay Cave)

(Goodwin & Malan 1935; Keller 1969)

Pinnacle Point Caves

(Marean et al. 2007; Marean et al. 2004)

Bufelskloof Shelter

(Opperman 1978)

*Boomplaas Cave A

(Deacon 1979; Deacon 1989; Deacon 1995; Deacon 2001; Deacon & Deacon 1999; Klein 1978a)

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TABLE 6.5. (continued )

Site

References

Paardeberg Cave

(Deacon & Brooker 1976; Volman 1981)

Kangkara Cave

(Deacon & Brooker 1976; Volman 1981)

Howieson’s Poort Rockshelter

(Stapleton & Hewitt 1927; Stapleton & Hewitt 1928)

*Nelson Bay Cave

(Butzer 1973b; Klein 1972a; Volman 1981)

*Klasies River Main Caves

(Butzer 1978b; Deacon & Deacon 1999; Deacon & Geleijnse 1988; Deacon et al. 1986; Feathers 2002; Grün et al. 1990b; Klein 1976; Singer & Wymer 1982; hackeray 1989; hackeray & Kelly 1988; hackeray 1988; hackeray 1992b; Voigt 1982; Wurz 1997; Wurz 1999; Wurz 2002)

chance exceptions to a general MSA rule or exemplify a widely shared behavioral capacity that only some MSA people expressed. In either case, the contrast with the LSA remains, but in the second case, it could not be attributed to a diference in the behavioral capability of the people. In general, compared to Mousterian assemblages in Europe, southern African MSA assemblages are poor in retouched pieces. However, the same basic retouched types prevail (scrapers, points, and denticulates), and their relative importance varies among MSA assemblages just as it does among Mousterian ones. Denticulates are the most common retouched type in some assemblages, whereas either scrapers or scrapers plus points dominate others. Most specialists in southern Africa have not counted Levallois lakes separately, but MSA people apparently employed the Levallois technique more in the interior than along the southern coast. At least in part, this relects diferences in the rock types available for artifact manufacture. he abundance of blades also varies among southern African MSA assemblages, and both the number and size of blades may increase with time. Like the Mousterian in Europe, the MSA in southern Africa includes variants or facies that replace each other in time and space. In its degree of internal variability, the MSA is intermediate between the preceding Earlier Stone Age (ESA), where regional diferences are hard to detect and temporal diferences are obvious only between assemblages separated by hundreds of thousands of years, and the succeeding Later Stone Age (LSA), where regional diferences are obvious and assemblages separated by only a few thousand years oten difer conspicuously. he deeply stratiied deposits at Diepkloof Cave illustrate temporally successive MSA variants particularly well. hese include (from older to younger) (1) the Still Bay variant, distinguished by well-made, bifacial, leaf-shaped points; (2) a so far unnamed variant marked by relatively numerous denticulated lakes and blades; and (3) the Howieson’s Poort

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variant, distinguished by lakes or blades on which one edge or the ends were steeply retouched (backed or truncated) (ig. 6.43). In plan view, the inished pieces resemble crescents or trapezoids. In three dimensions, they recall the segments of an orange, and they are oten called segments, as previously noted in the section “Mousterian/MSA Tool Function.” OSL and TL dating suggest that the Still Bay variant terminated 80–75 ka, and OSL, TL, and U-series dating ix the beginning of the Howieson’s Poort variant at 70–65 ka. he exact duration of each remains to be established, but the thickness of deposits at key sites suggest spans of 5–10 ky or more. he same is true of the unnamed variant with denticulates, which persists through more than 3 m of deposit at Ysterfontein 1 Rockshelter and 4–5 m at Die Kelders Cave 1. Geographically, with respect to igure 6.22, sites with abundant Still Bay points are best known in the southwestern corner of South Africa, southwest of a line between Hollow Rock Shelter and Blombos Cave, but MSA sites with rarer, oten fragmentary bifacial points occur far to the east and north as far away as Sibudu Cave, and they may imply a much wider distribution. Sites like Blombos Cave and Hollow Rock are parlorsized and they arguably represent specialized Still Bay workshops. he unnamed denticulate variant has so far been identiied only on the South African west coast, between Diepkloof Cave on the north and Die Kelders Cave 1 on the south, but the Howieson’s Poort variant has been found in many sites south of the Zambezi River, or in modern political terms, from the Zambian/Zimbabwean border southward. It is so far unknown in Zambia, but an MSA assemblage with Howieson’s Poort– like segments and trapezoids from Mumba Cave, central Tanzania, may mean that it once extended to eastern Africa. he Howieson’s Poort variant takes its name from the irst reported occurrence, at the Howieson’s Poort Rockshelter in what is now the Eastern Cape Province of South Africa, and it was once thought to represent an intermediate stage between the MSA and the LSA. his is mainly because Howieson’s Poort segments resemble segments that LSA people produced widely throughout southern Africa ater 10–8 ka. At least some LSA segments served as arrow armatures, attached to the ends of arrow shats by vegetal adhesive (mastic). However, in distinction from LSA segments, which are generally between 12 and 16 mm long, Howieson’s Poort segments tend to be about 40 mm long, and as discussed below, if they were mounted, they are more likely to have tipped spears. hey might still be seen to foreshadow LSA segments, but at Apollo 11 Cave, Diepkloof Shelter, Peers Cave, Boomplaas Cave, Klasies River Main, Rose Cottage Cave, Umhlatuzana Cave, Sibudu Cave, and other deeply stratiied sites, MSA variants without segments succeed the Howieson’s Poort, and at least 5–10 ky separated it from the earliest LSA at 50–45 ka. he

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5 cm

segments (“crescents”)

523 FIGURE 6.43. howieson’s Poort artifacts from nelson bay Cave, south africa (drawings by J. deacon). The howieson’s Poort industry is distinguished from other middle stone age industries mainly by the presence of well-made backed tools, including the segments (or “crescents”) pictured here. They resemble later stone age segments that mostly postdate 10 ka, but they are two-to-three times larger on average.

retouched or utilized flakes

retouched or utilized flakes

Howieson’s Poort is thus best seen as a variant of the MSA that was connected no more closely to the succeeding LSA than the French Mousterian of Acheulean Tradition Type B (with numerous backed knives) was connected to the subsequent Upper Paleolithic Gravettian Culture (with broadly similar, but usually more inely made backed elements). Unlike artifacts in other MSA variants, which tend to be made on quartzite or other widespread, local rock types, Howieson’s Poort backed elements are oten made on “exotic” ine-grained types like silcrete and chalcedony. his could mean that Howieson’s Poort groups roamed over much larger territories, perhaps in response to early Last Glaciation climatic deterioration that reduced the abundance of stable, predictable plant and animal resources and increased human dependence on highly migratory, gregarious ungulates. If this interpretation is correct, the Howieson’s Poort variant would provide the oldest demonstrated example of a climatically driven change in human territorial behavior. However, the ine-grained rock types that distinguish the Howieson’s Poort were generally available near Howieson’s Poort and other MSA sites,

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ChaP Ter s iX TABLE 6.6. East African sites that were occupied in the gap between 60 and 30 ka when southern and northern Africa seem to have been largely abandoned.

Site

References

Lukenya Hill, Kenya

(Kusimba 2001; Marean 1992b; Miller 1979)

Enkapune Ya Muto, Kenya

(Ambrose 1998a; Marean 1992a; Marean et al. 1994)

Nasera (Apis) Shelter, Tanzania

(Barut 1994; Mehlman 1977; Mehlman 1989)

Mumba Shelter, Tanzania

(Mehlman 1979; Mehlman 1989)

Matupi Shelter, D. R. Congo

(Cornelissen 2002; Mercader & Brooks 2001; van Noten 1982)

though usually only in nodules that were too small for more ordinary MSA lakes and blades. It may therefore be that Howieson’s Poort people difered from other MSA groups not so much in their larger territories as in their need or ability to utilize relatively small lumps of high-quality stone. In southern Africa, deeply stratiied cave sites like Montagu Cave, Diepkloof Rock Shelter, Nelson Bay Cave, Blombos Cave, Die Kelders Cave 1, and the Klasies River Caves commonly exhibit major occupation gaps between the latest MSA, antedating 50 ka, and succeeding LSA occupations that date from 20 ka or later. As in northern Africa, the occupation hiatus probably relects a reduction in human population density owing to extreme aridity in the middle part of the Last Glaciation. A deep drill core from Lake Malawi suggests that in contrast, eastern Africa was relatively moist between 50 and 35 ka, and table 6.6 lists ive sites that suggest that greater moisture encouraged greater human population density. It is thus to eastern Africa then that we should probably look for LSA origins. Like their north African contemporaries, as detailed above, the people who made MSA artifacts in southern and eastern Africa were morphologically derived in the direction of living people. SOURCES (not including those in tables 6.5 and 6.6): the MSA in western Africa (Allsworth-Jones 1986a; Mercader 2002; Mercader and Marti 1999) and in eastern Africa (Anthony 1972; Clark 1988; Masao 1992; McBrearty 1988; Mehlman 1977, 1979, 1989; Phillipson 1976; Sheppard and Kleindienst 1996; Wendorf and Schild 1974; Yellen et al. 2005); initial deinition of the MSA (Goodwin 1928, 1929a); foraging opportunities in rain forests (Bailey et al. 1989); past luctuations in rain forest distribution (Cornelissen 2002); the frequency of Mousterian sites in regions of France (de Lumley 1976a); frequency of art objects in MSA sites (Cain 2006); overviews of the southern African MSA (Mitchell 2002; Phillipson 2005; hackeray 1992a; Volman 1984); dating of the Still Bay MSA variant (Jacobs et al. 2003a, 2003b); dating of the Howieson’s Poort variant (Deacon 1995; Rigaud et al. 2006; Tribolo et al. 2005a, 2005b; Valladas et al. 2005; Vogel 2001); distribution and basic description of the Howieson’s Poort variant (Deacon and Deacon 1999); Mumba Cave artifacts (Mehlman 1979, 1989; Prendergast et al. 2007); Howieson’s Poort raw material and territory size (Ambrose and Lorenz 1990); aridity during the Last Glaciation in southern Africa (Beaumont and Vogel 2006; Deacon 1995; Deacon and hackeray 1984) and more moist conditions in eastern Africa (Scholz et al. 2007)

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Artifacts in Wood and Bone As indicated previously, use-wear studies show that Mousterian stone artifacts were oten applied to wood. Wooden artifacts are rare, due to adverse preservation conditions at the vast majority of Mousterian/MSA sites, but table 6.7 lists four signiicant exceptions. Mousterian woodworking could have been anticipated from the wooden spears or spear points recovered at the earlier Paleolithic sites of Clacton (England) and Schöningen (Germany), discussed in the previous chapter. Unlike wood, bone is oten plentiful and well-preserved at Mousterian/MSA sites, and butchery marks repeatedly conirm Mousterian/ MSA interest in meat and marrow. Damage marks also indicate that some bones were used as retouchers, hammers, scrapers, cutting boards, or anvils, though in some cases, the marks could have been produced by carnivore chewing or crunching. Few Mousterian/MSA sites have produced deliberately shaped, formal bone tools, and at some sites where shaping has been reported, the actual cause could be a natural factor like abrasion in a stream or trampling on a sandy substrate. Natural forces can polish bones, sometimes to a smooth point, and they might explain twenty-eight polished mammoth ribs and ibulas that have been interpreted as artifacts at the Salzgitter-Lebenstedt Mousterian site, Germany. At other sites where human shaping is not in doubt, the worked bone objects are mainly isolated and their stratigraphic origin is questionable. he MSA deposits at Sibudu Cave and Klasies River Main, South Africa, present cases in point. Sibudu has provided three bone artifacts, described as a hide-working tool, a pin or needle-like implement, and a projectile point. All are said to have been in place, but the accompanying MSA animal remains included bones of a burrowing rodent, and its activity could account for some other bones, including human specimens, whose preservation or concenstration suggests they postdate the MSA. he extensive MSA deposits at Klasies River have provided a single polished (ground) LSA-like bone point among tens of thousands of unworked bone fragments. Like the Sibudu pieces, the point has been taken to imply LSA-like bone-working in the MSA, although LSA bone assemblages that are far smaller almost always contain multiple points and other bone artifacts. he contrast suggests that the Klasies point may be intrusive, and it came from a part of the deposit that the excavators thought was disturbed. Disturbance isn’t always obvious, however, and even the most careful excavators cannot preclude it, particularly when small, isolated or unique objects are involved. he demonstration of Mousterian/MSA formal bone-artifact manufacture thus depends on sites with multiple, fundamentally similar specimens. At the moment, only two sites or site clusters qualify. hese are Blombos Cave, South Africa, and the riverside sites of Katanda 2, 9, and 16, Zaire.

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ChaP Ter s iX TABLE 6.7. Mousterian/MSA sites with wooden artifacts. Sedimentary circumstances have not favored wood preservation at the vast majority of other sites.

Site

Artifacts

References

Lehringen, Germany

2.4 m long wooden spear found among the ribs of a straighttusked elephant; tip possibly ire-hardened

(Movius 1950; hieme & Veil 1985)

Abric Romani, northeastern Spain

two possible wooden shovels or (Carbonell & Castro-Curel trays; also cavities or hollows 1992; Castro-Curel & that may have originated from Carbonell 1992) additional, rotted wooden artifacts or irewood. Stone artifacts with typical wood-working microwear.

Chambuage Mine, Angola

possible throwing stick

(Clark 1968a)

Florisbad, South Africa

fragments of three possible wooden artifacts, including one tentatively identiied as a throwing stick

(Bamford & Henderson 2003; Clark 1955)

Blombos Cave

At Blombos Cave, the bone artifacts occur in and immediately below layers with numerous bifacial, leaf-shaped stone points that deine the Still Bay variant of the MSA. he same deposits have provided an ocher fragment with an incised pattern and forty-one putative shell beads, discussed below. OSL dating of sand grains and TL dating of burned silcrete artifacts place the ensemble near 78–75 ka. he bone artifacts include twenty-six “awls”—bone splinters of various sizes and shapes on which the broader end or “butt” is generally unworked, while the opposite, pointed end or “tip” is conspicuously polished from use—and three “points” that may be simply larger versions of the “awls,” although they vary less in cross-section from butt to tip and they appear polished along their entire length. he MSA awls and points originated from below LSA layers that provided six awls and a single point, but the MSA awls are thought to have been shaped mainly by whittling, laking, and use, while the LSA awls were fashioned mainly by abrasion against coarse rock. he MSA points more closely resemble their LSA counterparts at Blombos and other sites, and the irst MSA point was stratigraphically associated with three charcoal samples that have been radiocarbon-dated between 2,100 and 1,950 years ago. he charcoal was said to have come from three stratigraphically successive MSA horizons, but it must all have originated from the LSA occupation, and it argues for substantial mixture or intrusion. he remaining two points came from parts of the site that

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lack comparable evidence for mixing, and they are more likely to have been in place. he MSA points are important not only because they resemble LSA counterparts in workmanship and form but also because, by analogy to LSA points, they could have been projectile armatures. hey might then imply an enhanced ability to hunt, but if so, this is not apparent in the Blombos animal remains, which closely resemble those from nearby MSA sites that lack bone points. he sections on MSA ecology and population numbers below show that the supposed LSA-like artifacts at Blombos Cave were not associated with an LSA-like ability to hunt and gather or with LSA-like population density. his means they did not confer an LSA-like itness (survival-and-reproduction) advantage on their makers. Katanda

he three Katanda sites have provided eight whole or partial barbed bone points (ig. 6.44) and four other indisputable bone artifacts. In the degree and quality of shaping, the Katanda objects far exceed any other bone artifacts that have been reported from Mousterian/MSA (or older) sites. he accompanying stone artifacts are not classically MSA, but ESR dates on associated hippopotamus teeth and luminescence dates on enclosing sands bracket the sites between roughly 90 and 60–70 ka, within the MSA time range. he dates are particularly striking because the next oldest barbed points in Africa date from no more than 25 ka and most specimens are younger than 10 ka. he implication is for a tradition of point manufacture (and associated point use in ishing) that spanned up to 80 ky. An alternative possibility is that the Katanda luminescence and ESR dates do not bear on the bone artifacts. he individual luminescence determinations scatter widely for reasons that remain unclear, and unlike the bone artifacts, the Katanda mammal bones and teeth tend to be conspicuously abraded and rounded. his suggests they were transported by moving water and that they may rest in a river bar on which much younger bone artifacts were dropped. he “modern” quality of the Katanda points is undeniable, and their discovery initiated the argument that MSA people were capable of fully modern behavior and that the MSA/LSA transition marks neither the advent of novel advanced behaviors nor a change in the capacity for novel behavior. he “modern” quality of the Blombos Cave inds is less compelling, but those who seek modern behavioral origins within the MSA now emphasize Blombos Cave, and they oten ignore Katanda. he reason is probably concern about the antiquity of the Katanda points. he issue might be resolved by direct 14C dating or by comparing mineral crystallinity and elemental content between the points and associated mammal bones.

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FIGURE 6.44. Two of eight barbed bone points excavated at Katanda, democratic republic of the Congo (redrawn after Yellen [1998], 189). luminescence readings on enclosing sands and esr determinations on associated animal teeth suggest that the points date from between 90 and 70–60 ka, and they would then be the most elaborate bone artifacts from any site older than 50–40 ka. arguably, however, their age has been overestimated, and like most similar points from eastern and northern africa, they are actually no older than 12 ka.

0

5 cm

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In sum, even if Blombos and Katanda bone artifacts are accepted at face value, they are exceptional. Systematically shaped, standardized points, awls, and so on are rare or absent in other Mousterian/MSA sites, and in this respect, the Mousterian and MSA resemble the preceding Acheulean much more than they do the succeeding Upper Paleolithic and LSA. In addition, the Blombos bone artifacts appear casual and expedient compared to many Upper Paleolithic/LSA examples, and by themselves, they might be regarded as an intriguing but not particularly important exception to the general rule. he Katanda bone artifacts, in contrast, are as elaborate and sophisticated as any known Upper Paleolithic/LSA examples, and they unquestionably imply an LSA-like level of technical ingenuity. hey also indicate an LSA-like ability to hunt or ish that would probably have promoted human population growth, and if they genuinely date from 60 ka or before, it is diicult to understand how and why they remained geographically localized for so long. SOURCES (also tables 6.5 and 6.7): Clacton (Warren 1911); Schöningen (hieme 1997); carnivoremodiied bones (Binford 1982); rarity of formal bone artifacts in Mousterian/MSA sites (d’Errico

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and Henshilwood 2007; Kozlowski 1990a; Mellars 1996; Patou-Mathis 2000; hackeray 1992a; Volman 1984); bones pointed by trampling (Brain 1967); Salzgitter-Lebenstedt bone artifacts (Gaudzinski 1999a); Sibudu Cave bone artifacts (Backwell et al. 2008) and animal remains (Plug 2004); Klasies River ground bone point (Singer and Wymer 1982); LSA-like bone-working in the MSA (McBrearty and Brooks 2000); Blombos Cave bone artifacts (Henshilwood et al. 2001a); Katanda bone artifacts (Brooks et al. 1995; Yellen 1996, 1998; Yellen et al. 1995); antiquity of barbed points in Africa (Brooks et al. 1995; Phillipson 2005; Robbins et al. 1994, 2000); Katanda TL dating (Feathers and Migliorini 2001); Katanda central to advanced behavior in the MSA (McBrearty and Brooks 2000) and ignored (d’Errico et al. 2005; Mellars 2006c)

Art and Ornamentation Further, like earlier peoples and unlike later ones, Mousterian/MSA peoples let remarkably little evidence for art or ornamentation. he sample of proposed art objects from Mousterian/MSA sites depends on the author, but table 6.8 lists the items that are most commonly cited. In support of Mousterian/MSA art objects, enthusiasts also oten note even older bones with incised lines from the Lower Paleolithic site of Bilzingsleben, Germany, and a volcanic pebble recalling a female igurine from the Acheulean site of Berekhat Ram on the Israeli/Syrian border. Both specimens antedate 250 ka, and as discussed in the precious chapter, as between the two, the Berekhat Ram igurine is the more diicult to dismiss. Some of the proposed MSA/Mousterian art objects are problematic because they could be intrusive from overlying horizons or because their stratigraphic origin is not well-documented. With respect to items in table 6.8, intrusion seems likely to explain the perforated fox canine and perforated wolf incisor from La Quina and Repolusthöhle, respectively, since both items closely resemble Upper Paleolithic counterparts and Upper Paleolithic people occupied both sites. For the same reason, the excavator believes that intrusion could explain the two ostrich eggshell beads from the MSA layers of Boomplaas Cave. he MSA associations of the painted slab and incised ostrich eggshell fragments from Apollo 11 Cave are questionable, although the slab almost certainly antedates 19 ka and is thus the oldest painted object so far found in Africa. he stratigraphic origin of the naturally perforated shells from Oued Djebbana and Skhul Cave is uncertain, and interpretation of the Skhul specimens as beads depends as much on from their possible association with nearmodern human remains as on their form. he thirteen putative beads from Pigeon Cave (Taforalt) are comparably problematic. he blanks are from the same genus (Nassarius) that provided blanks for proposed beads at Blombos Cave, discussed below. However, unlike the Blombos shells, the Pigeon Cave specimens appear to have been picked up empty ater some had been abraded and choked with sediment on a beach. he excavators attribute the Pigeon Cave objects to Aterian horizons dated by luminescence to roughly 82 ka, but

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Mousterian/MSA sites with proposed art or personal ornaments. Some specialists. such Bednarik (1992), recognize a few additional Mousterian sites with crudely incised or shaped objects resembling those in the list below.

TABLE 6.8.

Site

Objects

La Quina, France

a reindeer phalanx and a fox canine, each (Marshack 1976a; Marshack 1976b) punctured or perforated by a hole to form a “pendant”

La Ferrassie, France

a bone fragment with incised parallel lines

La Roche-Cotard, France

a lint nodule that may have been laked to (Marquet & Lorblanchet 2003) enhance its resemblance to a face or mask

Bocksteinschmeide, Germany

a perforated wolf metapodial and a perforated swan vertebra

(Marshack 1991a)

Repolusthöhle, Austria

a perforated bone fragment and a perforated wolf incisor

(Bednarik 1992)

Divje Babe Cave 1, Slovenia

a juvenile bear femur shat with four evenly spaced perforations suggesting a lutelike musical instrument

(d’Errico et al. 1998a; Lau et al. 1997; Turk et al. 1997a; Turk et al. 1997b)

Tata, Hungary

a fragment of mammoth molar plate that may have been deliberately carved and polished to an oval shape and an invertebrate fossil (nummulite) on which a supposedly engraved line intersects a natural one to form crosses on both surfaces

(Marshack 1976a; Marshack 1976b; Vértes 1964)

Bacho Kiro, Bulgaria

bone fragment with incised lines forming a crude zigzag pattern

(Marshack 1976b)

Molodova 1, Ukraine

a mammoth scapula and other bones with possible incised marks

(Chernysh 1982; Hofecker 2002)

Prolom II, Crimea (Ukraine)

3 bone fragments and horse canine with possibly incised lines, and 111 perforated saiga antelope phalanges

(Enloe et al. 2000; Stepanchuk 1993)

Quneitra, Golan Heights, Israeli/Syrian border

a plate of lint cortex with incised semicircles and other lines

(Goren-Inbar 1990)

Skhul Cave, Israel

3 perforated marine snail shells (Nassarius) that could represent beads, though the perforations are probably natural

(Bar-Yosef Mayer 2005; Garrod & Bate 1937; Vanhaeren et al. 2006)

Qafzeh Cave, Israel

4 naturally perforated cockle shells (Glycymeris sp.)

(Bar-Yosef Mayer 2005; Hovers et al. 1997; Taborin 1996; Vanhaeren et al. 2006)

Pigeon Cave (Grotte des Pigeons), Taforalt, Morocco

13 perforated Nassarius shells, broadly (Bouzouggar et al. 2007) similar to ones from Blombos Cave, listed below.

Oued Djebbana, Bir-el-Ater, Algeria

a perforated Nassarius shell similar to the ones from Skhul Cave listed above

(Morel 1974; Vanhaeren et al. 2006)

Apollo 11 Cave, Namibia

7 fragments of painted rocks slabs (two conjoining) and incised ostrich egg shell fragments

(Wendt 1972; Wendt 1975; Wendt 1976)

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TABLE 6.8. (continued )

Site

Objects

References

Boomplaas Cave A, South Africa (OLP Member)

one complete and one uninished ostrich egg shell bead

(Deacon 1995)

Diepkloof Rock Shelter, South more than 80 fragments of intentionally (Parkington et al. 2005; Rigaud et al. Africa (uppermost Howiemarked ostrich egg shell, mostly < 25 mm 2006) son’s Poort horizon) across Blombos Cave, South (Still Bay an incised bone fragment, nine incised red and/or immediate pre-Still ocher fragments, and 41 perforated Bay horizons) Nassarius shell shells

(d’Errico et al. 2001; Henshilwood et al. 2004; Henshilwood et al. 2002)

they don’t mention that previous, far more extensive excavations recovered abraded, perforated and nonperforated coastal shells in overlying Epipaleolithic (Iberomaurusian) layers dated by radiocarbon to between 20 and 10 ka. he new excavation was in the trench wall of the earlier one, but it has been only weakly described, and the excavators do not address the possibility that the supposed beads originated by slumping or other disturbance from the Epipaleolithic layers. heir identitiication as beads depends mainly on irregular perforations that are not demonstrably artiicial and on their resemblance to the proposed beads from Blombos Cave. However, they are less compelling than the Blombos specimens, and they might have been ignored altogether, if the Blombos inds had not already been publicized. he same fundamental issues afect the other proposed art objects— their modiication may not be artiicial, and on most, it is not persuasively artistic. he phalanx “pendant” from La Quina, for example, could have been punctured by a carnivore’s bite, while a bone with parallel engraved lines from La Ferrassie could simply represent a cutting board on which skin or some other sot material was repeatedly sliced. Similar nonartistic, repetitive cutting could explain apparently patterned incisions on many ancient bones, including those from Bacho Kiro or Molodova, and large animal trampling on a sandy or gritty substrate could account for others. Elephant trampling of bones near African waterholes occasionally produces clusters of subparallel marks that could be confused with incised patterns. Chewing or partial digestion by spotted hyenas or other carnivores probably perforated the phalanges at Prolom II and other sites, and carnivore biting probably produced the holes on the proposed lute from Divje Babe 1 (ig. 6.45). In retrospect, given the known human and natural actions that can simulate crude human attempts at art and the huge sample of available Mousterian/MSA (and earlier) objects, it would be truly remarkable if occasional pieces did not appear deliberately incised, perforated, or otherwise artistically shaped. he contrast with even the earliest Upper

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FIGURE 6.45. The putative mousterian lute from divje babe Cave 1, slovenia (drawn by Kathryn Cruzuribe from a photo in Turk et al. [1995]). The object is the shaft of a juvenile cave bear femur with four round, evenly spaced perforations, two of which are only partially preserved at the broken ends. divje babe 1 is far richer in cave bear bones than in mousterian artifacts, and it appears to have been mainly a cave bear den. in cave bear dens that totally lack artifacts, 4%–5% of the cave bear bones show similar perforations or punctures, and they were probably produced by cave bear or other large carnivore biting (d’errico et al. 1998a, 2003). The supposed divje babe 1 lute exhibits no stone-tool marks, and it could have been created by biting that accidentally produced four evenly spaced perforations. The oldest known, unequivocal musical instruments come from upper Paleolithic sites, after 35 ka.

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perforations

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juvenile bear femur shaft (Divje Babe I) Paleolithic/LSA sites is striking since these oten contain shaped objects whose artistic (as opposed to natural or utilitarian) origin is incontestable. he diference from the Upper Paleolithic/LSA is all the more notable since the Mousterian and the MSA span a far longer interval. However, accidents or natural forces may not explain all Mousterian/ MSA art objects, and as of this writing, the ones that merit special attention are possible shell beads from Blombos Cave, an ocher fragment with an engraved pattern, also from Blombos Cave, and incised fragments of ostrich eggshell from Diepkloof Rockshelter (references in table 6.8). Blombos Cave

he beads and ocher fragment from Blombos Cave come from layers that have been ascribed to the Still Bay variant of the MSA, an immediately preceding variant, or both. It will be recalled that well-made bifacial leaf-shaped points separate the Still Bay from other MSA variants, and Blombos has provided more than 400 bifacial points of various sizes. OSL analysis of sand grains and TL dating of burned silcrete artifacts indicate that the beads and the engraved ocher fragment accumulated about 78–75 ka, and their association with MSA artifacts is irm. LSA deposits that formed ater about 2 ka unconformably overlie the MSA layers at Blombos, but over most of the site, 5–50 cm of archaeologically sterile eolian sand separates the MSA and LSA deposits, and the ocher fragment is too large (about 6 × 2 × 2 cm) to have easily slipped through. he beads (ig. 6.46, top) are more likely prospects since they were made on tiny (7–10 mm long) shells of the snail-like, estuarine scavenger, Nassarius kraussianus, known in the vernacular as the tick shell,

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and indisputable tick shell beads occur in the LSA layers. However, bead identiication depends irst and foremost on perforations through the shell walls, and the MSA and LSA people employed diferent perforation methods. he LSA inhabitants of Blombos and other sites commonly perforated the walls from the outside, and their eforts let telltale surface traces. MSA shell walls show no such traces, and replication experiments on modern specimens indicate that MSA people could have perforated the Blombos tick shells by inserting a lat, sharp bone point through a natural aperture and applying pressure from the inside. Decalciication ater burial afected all Blombos MSA shells and bones, and compared to the perforations and surfaces of LSA beads, the perforations and surfaces of the MSA beads tend to be less obviously worn or polished from friction with thread or other objects. he sum raises the question of whether postdepositional pressure from trampling or sediment compaction could have produced the MSA perforations. Such pressure probably explains why the shells at Blombos (and at most other South African coastal sites) tend to be highly fragmented, regardless of species. It could also account for perforations in snail-like LSA shells that have not been interpreted as beads because the surfaces lack conspicuous polish or signs of human working. At the same time, the occurrence of tick shells in the MSA layers of Blombos is diicult to explain if they were not collected for beads since they contain little lesh. hey could have been inadvertently introduced on bundles of estuarine eelgrass (Zostera capensis) that oten shelter live tick shells and that LSA people are known to have used as bedding. However, the Blombos analysts argue that such introduction is unlikely in the MSA because the closest eelgrass was probably at least 20 km away. his is further than hunter-gatherers were likely to seek it. In addition, eelgrass bundles would have contained tick shells of all ages (sizes), and the Blombos MSA shells come exclusively from adults. In sum, the Blombos perforated tick shells could have been beads, but even if they were, they still difer from many well-known LSA and Upper Paleolithic beads in one crucial respect—even if they were humanly perforated, they were not humanly shaped. he oldest known LSA beads, discussed below, were carefully cut, carved, and ground from ostrich eggshell, and their intended use or meaning is unquestioned. Many early Upper Paleolithic beads were carved and polished with equal or greater care from bone, antler, or ivory, and their shaping alone identiies them as decorative items. Unlike the proposed Blombos perforated shells that natural forces might explain, the incision on the ocher fragment is unequivocally human in origin. As discussed in the next section, MSA people throughout southern Africa routinely collected lumps of naturally occurring ocher, and the Blombos MSA people accumulated more than 8,000 pieces. As

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FIGURE 6.46. Top: Proposed tick shell beads from the middle stone age layers of blombos Cave, south africa. Bottom: a red ocher fragment with an incised, cross-hatched abstract pattern from the same deposits (drawings by K. Cruz-uribe from photos in henshilwood et al. [2002], 1279; [2004], 404). The deposits have been dated to 78–75 ka, and if the describers’ interpretation of the objects is accepted, they might be the oldest examples of jewelry and art in the world, antedating the next oldest by at least 25 ka.

post-mortem perforations

post-mortem perforations

0

5 mm

post-mortem perforations

0

incised cross-hatched pattern

10 mm

Blombos Cave

at other MSA sites, many of the Blombos lumps were scraped or rubbed to produce powder or smears, and at least seven were scored with stone tools. On six specimens, the scoring may have been mainly intended to test the quality of the ocher or to roughen the surface before grinding, but on one the scored lines form a cross-hatched pattern within a partial frame (ig. 6.46, bottom). An ocher lump from Klein Kliphuis Rockshelter, roughly 400 km northwest of Blombos, exhibits eight intersecting lines that might represent a simpler version of the Blombos pattern. he Klein Kliphuis fragment is associated with Howieson’s Poort or post–Howieson’s Poort MSA artifacts that probably postdate the Blombos occupation, and the implication might be for an enduring tradition of engraving. As discussed in the next section, European Mousterians also collected pigment fragments, which they also ground or rubbed, and two (out of 450) black (manganiferous) fragments from the French site of of Pech de l’Azé I are reported to exhibit engraved abstract patterns. he patterns at Blombos, Klein Kliphuis, and Pech de l’Azé I suggest intent,

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and they present the most compelling case yet for art before 50 ka. he Blombos piece has received special attention because it might contradict the idea that advanced behavior evolved in Africa only ater fully modern anatomy, but geography aside, it would obviously be more compelling if the scoring outlined an animal or human form. So far, such representational or igurative art remains unknown anywhere before 50–40 ka. Diepkloof Rockshelter

he incised fragments of ostrich eggshell at Diepkloof Shelter are associated with artifacts of the Howieson’s Poort variant of the MSA. As noted previously, the characteristic stone-artifact type of the Howieson’s Poort variant is a backed or truncated lake or blade that resembles a small, thin segment of an orange. OSL and TL dating at Diepkloof and other sites suggest that the Howieson’s Poort variant spans 5–10 ky beginning roughly 70–65 ka. Diepkloof also contains layers with Still Bay points, stratiied below the Howieson’s Poort layers, and both the Still Bay and Howieson’s Poort layers contain bones and ocher lumps. So far, however, it is only the incised eggshell fragments from the Howieson’s Poort layers that suggest precocious LSA-like behavior. he interiors of the Diepkloof eggshell fragments retain the light color of fresh ostrich eggshell, but the surfaces have been mostly stained brown to black by the containing deposits. he deeper grooves on the incised pieces tend to be light-colored, which could mean that incision postdated the staining and that it resulted from abrasion against coarse particles in the ground. he fragments are small, and on most the grooves exhibit no obvious pattern. However, on some they appear more organized, such as those sometimes observed on decorated LSA ostrich eggshell canteens. he incised fragments do not occur throughout the Howieson’s Poort layers, but in a relatively narrow band near the top, and they might represent crushed canteens that LSA people buried at Diepkloof long ater the Howieson’s Poort occupation. LSA caching of water-illed eggshell canteens is well-documented, and two Diepkloof incised fragments show rounded indentations that might mark the canteen mouths. Direct 14C dating could preclude the possibility that these fragments or others are LSA intrusions. For the moment, the most persuasive argument against intrusion (or LSA burial) is that the incised fragments are distributed over an area of many square meters near the top of the Howieson’s Poort sequence. Conclusion

In sum, the tick shell beads, incised ocher fragment, and bone artifacts from Blombos Cave and the incised ostrich eggshell fragments from Diepkloof Shelter may show that LSA behavior originated within the MSA, 10–25 ky earlier than the LSA itself. However, if we accept that

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such behavior, or perhaps more precisely, its cognitive underpinnings, enhanced human survival and reproduction, it’s puzzling that similar objects are not equally abundant in other layers at the same sites or in well-excavated, broadly contemporaneous MSA sites like Ysterfontein 1 Rockshelter, Die Kelders Cave 1, Boomplaas Cave, or the Klasies River Main Caves. he contrast with the LSA is sharp since bone artifacts, art objects, ornaments, and other advanced behavioral markers abound in all LSA sites where preservation is suitable, even when the samples are small. Some items, above all ostrich eggshell beads, occur in virtually every LSA assemblage. In addition, the summary of MSA ecology below points out that animal remains fail to distinguish Blombos Cave and Diepkloof Shelter from other regional MSA sites, and like the remains at other sites, they suggest that MSA people hunted and gathered less effectively than their LSA predecessors. he bottom line is that even when the precocious items from Blombos Cave and Diepkloof Shelter are considered, the MSA and the LSA still contrast sharply. If there’s a problem with this conclusion, it’s that the early LSA, dating between 50–45 and 20 ka, remains poorly known, particularly in southern Africa. he characterization of the LSA is thus based mainly on sites that postdate 20 ka, and the case that the early LSA was equally derived depends mainly on the rapidity with which early LSA people spread to Eurasia and on ensuing, abrupt change in the Eurasian archaeological record. SOURCES (not including those in table 6.8): Bilzingsleben incised bones (Bednarik 1995; Mania and Mania 1988); Berekhat Ram igurine (Bahn 1996; Goren-Inbar 1986; Goren-Inbar and Peltz 1995): possible intrusion of perforated teeth at La Quina and Repolusthöhle (Mellars 1996; White 1989); questionable associations at Apollo 11 (Butzer et al. 1979); Epipaleolithic horizons at Pigeon Cave (Lubell 2001); patterned marks on bone possibly produced by nonartistic repetitive cutting (Chase and Dibble 1987; Davidson 1990) or trampling (d’Errico 1995; Haynes 1988, 1991); bone perforations produced by carnivore chewing or partial digestion (Chase 1990; d’Errico et al. 2003; d’Errico and Villa 1997; White 1995a); uncontestable early LSA and Upper Paleolithic art objects (Ambrose 1998a; Conard 2003; Hahn 1993; White 1989, 1990, 1995a, 2003a); Pech de l’Azé pigment (d’Errico 2003; d’Errico et al. 2003; d’Errico and Soressi 2002)

Pigment Collection and Utilization he last section pointed out that the MSA inhabitants of southern Africa routinely collected naturally occurring red ocher (iron oxide), and ocher lumps have been reported in virtually every well-excavated MSA site that postdates 130 ka (site references in table 6.5). In MSA sites, unlike LSA ones, ocher smudges only occasionally occur on possible receptacles and grindstones, but MSA people frequently scratched on ground lumps to produce powder or smears, and the grinding transformed some lumps into crude crayons with faceted tips. MSA people mostly collected individual ocher fragments from scattered outcrops, but at Lion Cav-

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ern, Swaziland, they literally mined dense ochreous deposits with large stones. Tailings overlying the mined surface contain MSA laked artifacts that identify the miners. he most ancient example of MSA ocher collection and grinding is perhaps at Sai Island, northern Sudan, where both ocher lumps and ocher-smudged grindstones accompany MSA artifacts in a deposit that is likely to approach 200 ka in age. MSA people probably inherited their widespread penchant for pigment from even more distant ancestors since Acheulean people living before 250 ka brought naturally occurring iron-based pigments to Kapthurin (site GnJh-15), Kenya, and to Wonderwerk Cave, Kathu Pan, and Duinefontein 2, South Africa. Like-aged non-Acheulean people also introduced pigment to Twin Rivers Cave, Zambia, and the Twin Rivers fragments are particularly notable because they exhibit traces of rubbing or grinding like those on subsequent MSA lumps. Besides red ocher, the later MSA inhabitants of Hollow Rock Shelter, Ysterfontein 1 Shelter, and Border Cave (all South Africa) collected and ground naturally occurring black pigment (manganese dioxide). Pigment lumps have been reported from more than forty Eurasian Mousterian sites, and the same Mousterian layers at Qafzeh Cave, Israel, that furnished particularly informative near-modern human fossils have produced more than eighty-four ocher pieces. More than a dozen French Mousterian sites have also yielded ocher fragments, and a larger number have provided fragments of black colorant (manganese dioxide). he Mousterian layers of Pech de l’Azé Cave I have yielded more than 8 kg of black colorant. As in Africa, the pigment lumps at Eurasian sites were oten ground or rubbed, and they sometimes resemble crude crayons. As noted above, two fragments from Pech de l’Azé I exhibit incised abstract patterns, and they thus recall the incised ocher fragment from the MSA layers, Blombos Cave, South Africa. At Pech de l’Azé I and other European sites, the collectors were undoubtedly Neanderthals, who may have collected pigments for the same purpose(s) as their near-modern African and Israeli contemporaries. Conceivably, MSA/Mousterian interest in pigment implies a widespread inclination to draw or paint on skins, bark, or other perishable materials, including human bodies. In southern Africa, it could even imply wall art, since the walls in most MSA caves and rockshelters were too exposed to retain paintings or drawings that date from 40 ka or before. However, it is at least equally likely that both Mousterian and MSA people employed ocher and other pigments as antibiotics, as tanning agents, and especially as emulsiiers and hardening agents in the mastic (adhesive) used to attach stone tools to wooden handles or shats. At Blombos Cave, ocher was incorporated in the polish on eight MSA bone “awls,” as would happen if the tools were repeatedly used to pierce ocher-stained

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hides. Modern experiments conirm that powdered ocher improves the quality of resinous mastic, and ocher incorporation in mastic is so far the only function that MSA artifacts directly suggest. he section above titled “Mousterian/MSA Tool Function” stressed that ocher concentrations on points and segments at Sibudu Cave coincide with microscopic plant residues that probably derive from hating glue or ibers. he value of ocher as an ingredient in adhesive could explain its extraordinary abundance at Blombos Cave, where the associated stone artifacts include more than 400 well-made bifacial, leaf-shaped (Still Bay) points, many of which were probably mounted on wooden shats. Upper Paleolithic/LSA people employed ocher in hating, and if Mousterian/MSA people did the same, the implication would be for a shared degree of technological sophistication. Still, in contrast to their successors, Mousterian/MSA people let no direct evidence that they painted with ocher or other pigments, they did not concentrate ocher in graves, and they did not intentionally burn (oxidize or reduce) ocher fragments to obtain a wider range of colors. hus, in their pigment use, as in so much else of their behavior, they appear to have been both similar to later humans and fundamentally diferent from them. SOURCES (not including those in table 6.5): ocher at Sai Island (van Peer et al. 2003), at Kapthurin (Tryon and McBrearty 2002), at Wonderwerk and Kathu Pan (Beaumont and Vogel 2006), at Duinefontein 2 (Cruz-Uribe et al. 2003), and at Twin Rivers (Barham 2002); manganese dioxide in MSA sites (Klein et al. 2004; Watts 2002); ocher at Qafzeh Cave (Hovers et al. 2003) and in French Mousterian sites (Mellars 1996); manganese dioxide at Pech de l’Azé (d’Errico 2003; d’Errico et al. 2003; d’Errico and Soressi 2002); possible Mousterian painting on perishable material (Bordes 1952, 1968; Combier 1988); possible utilitarian uses of pigment (Keeley 1980; Wadley et al. 2004); experimental veriication of the utility of ocher as an ingredient in vegetal mastic (Wadley 2005b); potential utilitarian signiicance of ocher at Blombos Cave (Henshilwood et al. 2001a) and Sibudu Cave (Lombard 2005, 2007; Wadley et al. 2004); rarity of ocher in Mousterian graves (Harrold 1980); Mousterian failure to alter ocher with ire (Perlés 1976)

An Overview of Mousterian/MSA Artifacts he Mousterian/MSA exhibited more variability through time and space than did earlier (Lower Paleolithic/Earlier Stone Age) industries, but it was still remarkably uniform over vast areas and timespans compared with the succeeding Upper Paleolithic/LSA. he reason is partly that Mousterian/MSA peoples made a much smaller variety of readily discernible artifact types. Together, the relative homogeneity of the Mousterian/MSA through time and space and the relatively small number of distinguishable MSA/Mousterian artifact types suggest that the behavior of Mousterian/MSA people difered fundamentally from that of their Upper Paleolithic/LSA successors. As detailed in the section titled “Settlement Systems” below, it is further noteworthy that Mousterian people rarely transported raw materials more than 25 km, whereas Upper Paleolithic people sometimes

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transported or imported them over scores of kilometers. Some of the items that Upper Paleolithic people moved were nonutilitarian goods, such as marine shells and amber, that may not have interested Mousterian people, but some Upper Paleolithic sites contain large quantities of exotic (nonlocal) stone that was used to manufacture utilitarian artifacts. On the relatively rare occasions when exotic stone occurs in Mousterian/MSA sites, it usually involves isolated, inished artifacts, whereas exotic stone in Upper Paleolithic sites commonly consists of raw nodules and intermediate laking products. he sum implies that, relative to the Mousterians, at least some Upper Paleolithic people roamed over far larger territories or participated in much broader social networks. he rarity or absence of formal bone artifacts and art objects in the Mousterian/MSA only adds to the contrast. he most general implication is that Mousterian/MSA people were behaviorally more conservative than their successors, with more limited inclination or ability to innovate even in the face of signiicant environmental variability through time and space. At least tentatively, it seems reasonable to propose that the behavioral limitations implied by Mousterian/MSA artifacts relect biological (genetic) diferences from later humans. SOURCES: Mousterian vs. Upper Paleolithic transport of raw materials (Gamble 1986; Klein 1969a; Mellars 1973, 1989a, 1996; Schild 1984; Sofer 1989b)

Site Distribution he overall distribution of Mousterian/MSA sites closely resembles the distribution of earlier (Lower Paleolithic/Early Stone Age) sites, except that Mousterian sites occur in easternmost Europe (Ukraine and European Russia) and in southwestern Siberia, where traces of earlier people are rare and mostly equivocal. Since the commercial and archaeological activities that promote site discovery have been as intense in eastern Europe as in central and western Europe, where many earlier sites are securely documented, it seems increasingly likely that the rarity or lack of pre-Mousterian sites on the east relects the rarity or absence of people. he most probable reason is that pre-Mousterians could not cope with the especially cold winters that have always characterized easternmost Europe and southern Siberia, even during interglacial periods. At the same time, it is important to stress that Mousterian sites occur only on the southern and western margins of easternmost Europe, and no sites have been irmly documented in the more continental—central and eastern—portions. his is despite substantial commercial and archaeological activity, which has revealed numerous, rich Upper Paleolithic sites discussed in the next chapter. Based on the distribution of known sites, Mousterians occupied only those parts of Europe where the mean January temperature presently exceeds −15°C. Upper Paleolithic

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people were the irst to colonize much colder regions, and they did it in glacial times when mean January temperatures were commonly 50°C or more below present-day ones. Upper Paleolithic people were also apparently the irst to inhabit the especially harsh subarctic and arctic latitudes of central and northern Siberia (above 50°N latitude), where, in spite of dedicated eforts, no convincing Mousterian (or earlier) sites have so far been found. SOURCES: (Hofecker 1987, 2002; Hofecker and Elias 2003; Klein 1973b)

Site Types Mousterian/MSA debris occur both in open-air sites and in rock shelters or cave mouths. With rare exceptions, open-air sites occur near active or once active springs, lakes, or streams, partly because these were favorable places for camping or obtaining food but also because these were places where sediments accumulated and sites were most likely to be preserved. Most ancient open-air sites are no longer visible at the surface, and archaeologists depend on mining, construction, or other commercial activity to expose them. Table 6.9 lists prominent Mousterian/MSA open-air sites in Europe, western Asia, and Africa. At some of the sites in table 6.9, bones of a single large ungulate species dominate heavily, and it seems likely that these were kill/butchery sites, where the people concentrated on a single species that was abundant nearby. he most frequently noted sites with their dominant species are La Borde with aurochs (Bos primigenius); Mauran, Wallertheim, Sukhaya Mechetka, and Il’skaya, all with bison (Bison priscus); Zwoleń with horse (Equus caballus); and Salzgitter-Lebenstedt with reindeer (Rangifer tarandus). Lehringen may also qualify since it preserved a 2.4m-long yew wood “lance” among the ribs of a single straight-tusked elephant (Palaeoloxodon antiquus). Other open-air sites, notably at Rosh Ein Mor and Nahal Aqev in the central Negev Desert of Israel, are unusually rich in laking debris, and they appear to have been specialized stone-tool workshops, located near sources of high-quality lint. At the majority of open-air sites, however, no specialized function is apparent, and the cultural debris demonstrates only human presence. As discussed in the next section, unequivocal remnants of structures that might imply camps are uniformly absent. his stands in contrast to later openair sites, particularly Upper Paleolithic sites in Europe, where traces of structures occur regularly. Mousterian/MSA caves and rockshelters are better known than open-air sites because Mousterian/MSA peoples lived so recently that most of their caves are still conspicuous and the caves constitute obvious targets for archaeologists. In general, they retain artifacts, bones,

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Prominent open-air Mousterian and MSA sites in Europe, western Asia, and Africa. Figures 6.2, 6.4, 6.20, 6.21, and 6.23 locate the sites. TABLE 6.9.

Site

Source

FRANCE Mauran

(Farizy & David 1992; Farizy et al. 1994)

La Borde

(Jaubert et al. 1990)

Riencourt-les-Bapaume

(Tufreau 1993)

Hénin-sur-Cojeul

(Marcy et al. 1993)

Biache-Saint-Vaast

(Auguste 1995; Tufreau 1988; Tufreau 1989)

Seclin

(Révillon & Tufreau 1994; Tufreau et al. 1985)

NETHERLANDS Maastricht-Belvédère

(Roebroeks 1986; Roebroeks 1988; Roebroeks et al. 1993)

GERMANY Weimar-Ehringsdorf

(Behm-Blancke 1960; Blackwell & Schwarcz 1986)

Salzgitter-Lebenstedt

(Butzer 1971; Gaudzinski 1999a; Gaudzinski & Roebroeks 2000; Gaudzinski & Roebroeks 2003; Munson & Marean 2003; Tode et al. 1953)

Lehringen

(Gaudzinski 2004a; Movius 1950; hieme & Veil 1985)

Wallertheim

(Conard 1999; Conard et al. 1995a; Conard et al. 1995b; Conard & Prindiville 2000; Gaudzinski 1995)

Wannen

(Justus & Urmersbach 1987)

Ariendorf

(Bosinski et al. 1983, pp. 163, 164)

Tönchesberg

(Conard 1992; Conard & Prindiville 2000)

Königsaue

(Grünberg 2002; Mania & Toepfer 1973)

Neumark-Nord

(Brühl & Mania 2003; Gaudzinski 2004a)

Rheindahlen

(hieme 1983; hieme 1990; hissen 1986)

POLAND Zwoleń

(Gautier 1989; Schild et al. 1988)

HUNGARY Tata

(Schwarcz & Skolik 1982; Vértes 1964)

Erd

(Gábori-Czánk 1968)

MOLDAVIA Ripiceni-Izvor

(Păunescu 1965; Păunescu 1989)

UKRAINE Molodova I and V

(Chernysh 1961; Chernysh 1982; Haesaerts et al. 2003; Hofecker 2002; Klein 1973b; Sofer 1989b)

Korman’ IV

(Chernysh 1977; Hofecker 2002; Klein 1973b)

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TABLE 6.9. (continued )

Site

Source

RUSSIA Sukhaya Mechetka (Volgograd, ex-Stalingrad)

(Hofecker 2002; Klein 1969b; Vereshchagin & Kolbutov 1957; Zamyatnin 1961)

Il’skaya

(Baryshnikov & Hofecker 1994; Hofecker et al. 1991; Hofecker et al. 1989; Klein 1967; Zamyatnin 1929)

Rozhok I

(Hofecker 2002; Klein 1969b; Praslov 1968)

SYRIA Uum el Tlel (andother sites in the El Kowm Basin)

(Le Tensorer & Hours 1989; Le Tensorer et al. 2001)

ISRAEL Quneitra

(Davis et al. 1988; Goren-Inbar 1990)

Rosh Ein Mor

(Marks 1977; Marks 1981a; Marks 1989; Rink et al. 2003)

Nahal Aqev

(Marks 1977; Marks 1981a; Marks 1989)

Boker Tachtit

(Marks 1981a; Marks 1983)

TUNISIA El Guettar

(Gruet 1954; Gruet 1958)

LIBYA Hajj el Sidi Creiem (Wadi Derna)

(McBurney & Hey 1955)

EGYPT Bir Tarfawi

(Close et al. 1990; Wendorf et al. 1993)

NIGER Seggedim

(Tillet 1984; Tillet 1985)

KENYA Prospect Farm

(Anthony 1972; Michels et al. 1983)

ZAMBIA Kalambo Falls

(Clark 2001)

BOTSWANA ≠Gi

(Brooks et al. 1980; Brooks & Yellen 1977; Helgren & Brooks 1983)

SOUTH AFRICA

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(Brown 1988; Cooke 1962; Mason 1988a)

Kathu Pan

(Beaumont 1990a; Beaumont et al. 1984; Butzer 1984b; Klein 1988a)

Florisbad

(Douglas 2006; Kuman & Clarke 1986; Kuman et al. 1999; Scott & Rossouw 2005; van Zinderen Bakker 1990)

Orangia 1

(Sampson 1974)

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and other human refuse, oten in deeply stratiied deposits, and they are abundant in areas such as southwestern France, where investigation of the Mousterian has been underway longest. Most Mousterian/ MSA caves appear to have been living places as opposed, for example, to kill/butchery sites. However, many Mousterian caves, including for example, Krapina Cave, Croatia, famous for its large assemblage of Neanderthal bones, contain abundant remains of bears, hyenas, or wolves in addition to artifacts. Artifacts and other evidence of human presence are oten sparse, and the impression is that carnivores, particularly bears and hyenas, occupied the caves at least as oten as Mousterian people. In contrast, Upper Paleolithic cave sites tend to be much richer in cultural debris, and they rarely suggest that carnivores denned inside. As discussed below, the cave bear disappeared 5–10 ky ater Upper Paleolithic people appeared in Europe, and Upper Paleolithic monopolization of suitable den sites could explain why. SOURCES (not including those in table 6.9): Mousterian kill/butchery sites (Gaudzinski 1996, 1998); Krapina Cave contents (Simek and Smith 1997); the Upper Paleolithic and cave bear extinction (Grayson and Delpech 2001; Kurtén 1958)

Structural Traces Fossil ireplaces or hearths occur in virtually every well-excavated Mousterian/MSA cave where preservation conditions were suitable, and they provide the most persuasive evidence for site modiication. In some sites, such as Gorham’s Cave (Gibraltar), Kebara Cave (Israel), and the Klasies River Main Caves (South Africa), stacked hearths comprise much of the deposit, and they relect recurrent, intensive ire building over many millennia. he ubiquity of hearths implies that Mousterian/MSA people could make ire at will, and the hearths conirm that the caves served as camps. Hearths presumably functioned to provide warmth, light, and protection from predators and, of course, to prepare food. At lower latitude sites like Kebara and Klasies River Main, plant-food preparation may be especially indicated. However, even where hearths are especially abundant in Mousterian/MSA contexts, they tend to be relatively simple lenses of ash and charcoal 50 cm to 1 m across and a few centimeters thick. Some are surrounded or underlain by rocks, but none are as elaborate as some Upper Paleolithic hearths, which were itted with stone liners, air-intake ditches, or other features to control airlow and heat dissipation. At some Mousterian/MSA sites, hearths may mark the position of a hut or a lean-to, and the conines of a hut may explain why artifacts and other cultural debris tend to be clustered, oten near hearths. However, more direct evidence for housing is all but absent. Table 6.10 lists ot-cited examples, and igures 6.47 and 6.48 illustrate two. None

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TABLE 6.10. Mousterian/MSA sites that may preserve remains of structures. Structural traces from the MSA site of Orangia 1 in South Africa (Sampson 1974) have been omitted, because the excavator subsequently concluded that they postdate the MSA artifacts at the site. As explained in the text and the captions to Figures 6.47 and 6.48, no Mousterian/MSA ruins are especially compelling, and it appears that Mousterian/MSA people rarely, if ever built structures that were durable enough to leave an archaeological trace.

Site

Proposed Structural Remnants

References

Cueva Morín, Spain

A line of stone piles that may represent the remnant of a low wall

(Freeman 1983; Freeman 1989)

La Ferrassie Cave, France

A 5 m by 3 m spread of limestone fragments

(de Lumley & Boone 1976)

Pech de l’Azé Cave 1, France

A small segment of what might have been a low, dry stone wall

(de Lumley & Boone 1976)

Combe Grenal Cave, France

A probable posthole

(Bordes 1972)

Ariendorf (ind-level II), Germany

Artifacts and other debris concentrated in a depression

(Bosinski et al. 1983; Turner 1986)

Rheindahlen, Germany

Artifacts and other debris concentrated in a depression

(hieme 1983; hieme 1990; hissen 1986)

Molodova I and V, Ukraine

Large rings or partial rings of large (mainly mammoth) bones surrounding dense concentrations of artifacts, fragmentary bones, and ash spreads (hearths).

(Chernysh 1982; Hofecker 2002; Klein 1969b; Sofer 1989b)

Mumbwa Cave, Zambia

3 possible postholes on the margin of a dense semicircular concentration of ash, sediment, quartz artifactual debris, and fragmentary animal bones

(Barham 1995)

are compelling, and a reasonable conclusion is that if Mousterian/MSA people built structures, most were not substantial enough to leave unambiguous traces. he persistence of hearths, which are relatively fragile, shows that postdepositional blurring or destruction is an inadequate explanation. he absence of unequivocal ruins is particularly striking in Europe, where many Mousterian sites have now been carefully excavated and where the contrast with subsequent Upper Paleolithic sites is particularly sharp. As detailed in the next chapter, well-excavated Upper Paleolithic sites almost always contain obvious and oten spectacular evidence of buildings, in the form of artiicially excavated depressions and pits; patterned arrangements of large bones or stones; postholes; or some combination of these. Even Upper Paleolithic caves commonly preserve structural traces such as stone walls, artiicial pavements, and pits, and to the annoyance of archaeologists, Upper Paleolithic cave dwellers sometimes dug out preexisting deposits, perhaps to enlarge the living area or to level out a loor. As in many other aspects of culture, in terms of site modiication, Mousterian/MSA people seem to have been quantitatively, if not qualitatively, diferent from their successors. At least in part, their

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FIGURE 6.47. floor plan (top) and section (bottom) through “ind level ii” at the ariendorf open-air site, Germany (redrawn after bosinski et al. [1983], 163, 164). Quarrying through wind blown silt (loess) exposted the occurrence, and it unfortunately destroyed part of the site. however, careful excavation showed that elephant ribs, other large bones, and stone artifacts were concentrated in and near an ancient depression 2–3 m across. The depression may have been dug by early middle Paleolithic (mousterian) people to level the loor of a hut whose frame was made partly of elephant ribs. alternatively, it could be a natural feature, perhaps created when a large tree was uprooted. less equivocal structural traces are remarkably rare in middle Paleolithic and older sites, suggesting that the people rarely built substantial houses.

Ariendorf edge of excavation

conjoinable stone artifacts

0

2m

quarry face

bones

545

N

antler artifacts bones

small stones yellow, homogeneous loess

brownish gray loess

limited architectural ability may explain why they failed to occupy the most continental parts of Eurasia. SOURCES (not including those in table 6.10): stacked hearths at Gorham’s Cave (Barton 2000), Kebara (Meignen et al. 1989), and Klasies River Main (Deacon 1989, 1995); possible plant food preparation at Klasies River Main (Deacon 1989); structure of Mousterian hearths (Perlés 1976; Villa 2006, 15); abundance of structural remnants in Upper Paleolithic vs. Mousterian sites (Conard 2005; Klein 1973b; Kuhn 1995; Mellars 1973, 1996)

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rings of large bones

N

m

mammoth tusks mammoth molars other animal bones bones with stone tool cut marks

hearth natural depression with bones cores other flint artifacts

Molodova I, level 4 FIGURE 6.48. Plan of the excavations in molodova i, level 4, ukraine (redrawn after Chernysh [1982], ig. 8). The excavations uncovered two rings of large (mainly mammoth) bones enclosing dense concentrations of artifacts, fragmentary bones, and ash spreads (hearths). The bones may represent weights that held down skins that were stretched over a wooden framework that disintegrated long ago. however, each ring was about 8 m long and 5 m wide, and it would have required considerable architectural skill to cover. The full meaning of the molodova features remains unclear, and it is possible that slope wash or some other natural process created the patterning.

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Settlement Systems Most Mousterian/MSA peoples were certainly nomadic, occupying different sites for diferent purposes or at diferent seasons to take advantage of seasonal changes in resource availability. However, in most regions the distribution of known sites probably does not closely mirror the distribution of utilized sites. More likely it relects the distribution of caves and other places that favor preservation and the distribution of modern settlements and roads used by archaeologists. Since it is impossible to determine whether diferent sites were occupied by essentially the same people, in most instances it is also virtually impossible to reconstruct Mousterian/MSA settlement systems. However, the stone raw materials that Mousterian/MSA peoples used may provide some insight into their settlement patterns or at least into the extent of their movements since wherever Mousterian/MSA stone sources have been established, they turn out to be heavily local. In southwestern France, where the most detailed studies have been conducted, 70%–98% of the stone in various Mousterian sites came from less than 5 km away. Only 2%–20% originated between 5 and 20 km away, and less than 5% traveled more than 30 km. A broadly similar pattern has been found in eastern Morocco, and it also appears to characterize eastern Africa (Kenya and northern Tanzania) and central and eastern Europe. here is the diference, however, that in eastern Africa and in central Europe (Poland, Czech Republic, Slovak Republic, and Hungary), desirable stone sometimes traveled greater distances, even exceeding 100–200 km. he contrast may mean that compared to their French contemporaries, east African and central European Mousterian/MSA people occupied larger territories because their environmental conditions were less favorable, or perhaps more likely, they occupied regions where highly desirable raw material sources were more dispersed. he interplay between environment and raw material dispersion is clear in the eastern Sahara, where the Mousterians of Bir Tarfawi and Bir Sahara East quarried raw chunks of suitable stone at outcrops, transported the chunks to lake edges for core manufacture, and then took the cores elsewhere (possibly to nighttime sleeping sites) to produce tools. Like Mousterians elsewhere, however, those in the Sahara rarely moved stone raw material more than 30–40 km. Everywhere, Mousterians showed their appreciation for high-quality exotic stone by conspicuously transforming it into tools, but they still acquired exotic material infrequently. In contrast, Upper Paleolithic people oten acquired 20%–25% of their stone more than 30 km away, and they frequently transported not just desirable lakes but also cores of nonlocal stone. he sum suggests that Upper Paleolithic groups routinely roamed over larger territories or that they interacted (traded) more intensively with other, widely

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dispersed groups. Whichever possibility is accepted, it implies a signiicant shit in settlement patterns, social organization, or both. So far, stone raw materials provide no evidence for a diference between Mousterian and Upper Paleolithic settlement patterns in western Asia, but analyses of growth increments (or annuli) in animal teeth suggest a possible diference between the settlement patterns of west Asian near-modern people and their Neanderthal successors. he relevant growth increments occur in acellular cementum that anchors tooth roots to the surrounding socket and that forms in successive bands, each comprising a thicker subband formed during the season of maximum growth and a thinner subband formed during the season of reduced growth. Under transmitted light, the thicker bands tend to be translucent and the thinner bands opaque. When the cementum is appropriately sectioned and examined under a microscope, the total number of bands can reveal the age of an animal at time of death, and the translucency or thickness of the last (outermost) subband can reveal the season of death. Analysis of increments on the roots of Qafzeh and Kebara Cave gazelle teeth imply that Qafzeh was occupied seasonally (the gazelles died only in the local dry season), while Kebara was occupied more or less throughout the year (the gazelles died in both the dry and the wet seasons). his might mean that the Qafzeh (near-modern) people circulated from one base camp to another seasonally, whereas the Kebara Neanderthals tended to occupy the same base camp year-round. he Neanderthals may have been equally mobile, however, if they frequently radiated to smaller, more ephemeral camps that have not yet been identiied. he Qafzeh/ Kebara contrast is echoed by a broadly similar one between Skhul and Tabun Cave level C, on the one hand (seasonal), and Tabun level B, on the other (nonseasonal or multiseasonal). Recall that Skhul and Tabun C broadly parallel Qafzeh in time and in occupation by near-modern people and that Tabun B broadly parallels Kebara in time and in probable occupation by Neanderthals. If the seasonality contrast is accepted, it is the only documented behavioral diference between near-modern people and Neanderthals in western Asia. As discussed above and below, both groups made broadly similar Levalloiso-Mousterian artifacts, both built simple ireplaces, both buried their dead in uncomplicated graves, and both hunted mediumsized mammals, at least on occasion. However, a diference in seasonality may say more about climate than about culture since near-moderns occupied Qafzeh, Skhul, and Tabun C during the Last Interglacial, and Neanderthals inhabited Kebara and Tabun B during the early and middle phases of the Last Glaciation. he way climate could force such a change in seasonality of settlement is perhaps illustrated by a later, similar change that has been inferred between Mousterian and Upper Paleolithic patterns in the Avdat/Aqev area of the central Negev Desert, Israel.

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Here Mousterian sites appear to comprise semipermanent base camps near permanent water and outlying, more temporary sites near probable sources of food and stone raw material. Upper Paleolithic sites include no semipermanent base camps but only “ephemerally occupied multipurpose camps and even more ephemeral loci of unknown function.” he Mousterian pattern suggests a radiating system in which the people moved back and forth between the central base camp and the peripheral resource sites, whereas the Upper Paleolithic pattern suggests a circulating system in which the people simply moved from multipurpose camp to multipurpose camp. he shit from a radiating to a circulating pattern may have been an adaptive response to contemporaneous environmental deterioration (increasing aridity) that has been documented from geomorphic and palynologic evidence in the central Negev. he adoption of a circulating system could explain why Upper Paleolithic people abandoned the lake technology of the Mousterians since Upper Paleolithic blade technology reduced the need to deviate from a circulating pattern to renew stone raw material reserves at quarry sites. his is because, by comparison with the Mousterian lake technology, Upper Paleolithic blade technology produced more usable blanks per core, that is, more usable blanks per quarry visit. SOURCES: Mousterian/MSA stone sources in southwestern France (Féblot-Augustins 1993; Geneste 1988a, 1988b, 1990; Mellars 1996; Turq 1990), eastern Morocco (Wengler 1990a, 1990b, 1991), eastern Africa (Barut 1994; Merrick and Brown 1984; Merrick et al. 1994), and eastern and central Europe (Féblot-Augustins 1993; Roebroeks et al. 1988); Mousterian raw material transport at Bir Tarfawi and Bir Sahara East (Wendorf and Schild 1992); Upper Paleolithic transport of stone raw material (Kozlowski 1990a; Mellars 1996; Sofer 1985, 1989b); analysis of gazelle cementum annuli at Qafzeh and Kebara (Lieberman 1993a, 1998; Lieberman and Shea 1994); Mousterian and Upper Paleolithic settlement patterns in the central Negev (Marks 1988; Marks and Freidel 1977) and climatic change (Marks 1977)

Mousterian/MSA Ecology Mousterian/MSA peoples were restricted to mid- and low-latitude environments where they lived entirely by hunting and gathering. In their ecology they probably resembled recently observed low-latitude huntergatherers such as the Hadza people of northern Tanzania. he Hadza hunt and scavenge medium-sized and large mammals including impala antelope, hartebeest, wildebeest, warthog, and zebra, but for a steady nutrient low they depend heavily on berries, roots, fruits, honey, tortoises, and other gatherable resources. Except perhaps when peak cold reduced plant availability in midlatitude Europe, MSA/Mousterian people probably relied at least as much on gathered foods, but most gathered items unfortunately leave little archaeological trace. his is particularly true of plants, and apart from nondietary charcoal, pollen, and phytoliths (minute silica particles formed within plant tissues), Mousterian/MSA

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sites rarely preserve plant fossils. Table 6.11 lists the most prominent exceptions. hey have mostly furnished seeds, particularly ones whose survival was enhanced by carbonization before burial. Interpretation at each site is limited by the possibility that birds, rodents, or another nonhuman agent could have introduced many of the seeds. Arguably, table 6.11 should be extended to include the MSA deposits of Klasies River Main, South Africa, where careful excavation revealed extensive spreads of carbonized organic debris around hearths. In historic and late prehistoric times, local people relied extensively on the underground storage organs (corms) of iris-like geophytes, and they discarded the inedible parts at living sites, where, over time, they might carbonize to form spreads exactly like those at Klasies River Main. If the Klasies evidence is accepted, table 6.11 should perhaps also include those occasional sites that have provided “grindstones,” artiicially smoothed or polished pieces of coarse rock like those that historic people used to crush nuts or seeds. he most noteworthy MSA examples come from Sai Island, northern Sudan, where physicochemical analysis of the surfaces revealed phytoliths and starch granulates that could originate from ground grass seeds. Some Mousterian/MSA grindstones were surely used to pulverize nonedible materials like pigment (ocher), but in South Africa, MSA grindstones occur mainly in the northeast where nuts or seeds have probably always been particularly abundant compared to soter vegetal foods (like corms) that do not require crushing. In contrast to plant fossils, bones that relect the lesh component of Mousterian/MSA diet occur at many sites, and they commonly exhibit stone-tool marks from dismemberment, deleshing, and marrow extraction. he principal animals in Mousterian/MSA sites are mediumsized and large ungulates that were doubtless common near the sites. In Eurasia, depending on the time and place, the species include red deer, fallow deer, reindeer, bison, wild oxen (aurochs), wild sheep and goats, gazelles, and horses (including asses and onagers). In Africa the ungulates include various antelopes, zebras, and wild pigs. he largest available species—elephants and rhinoceroses, present in both Eurasia and Africa—tend to be rare. In addition, at Molodova (Ukraine) and other sites where they are common, their bones rarely exhibit stone-tool marks, and they may have been scavenged for construction. Succeeding Upper Paleolithic people in central and eastern Europe unambiguously amassed large numbers of mammoth bones to build houses (discussed in the next chapter), but even they let few stone-tool marks to suggest the bones came from still meaty carcasses. To acquire animals, Mousterian/MSA people probably relied heavily on stone-tipped thrusting spears, and they may have run down large animals cooperatively much as wolves do. To make a kill, they probably had to approach prey closely, and this may explain why (as noted again be-

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Site

Plants Represented

References

Gorham’s Cave, Gibraltar

Carbonized seeds of wild olive (Oliva sp.) and nut shells of stone pine (Pinus pinea)

(Barton et al. 1999)

Douara Cave, Syria

Hackberry (Celtis) seeds

(McLaren 1998)

Kebara Cave, Israel

Carbonized legume seeds and fragments of acorn and pistachio shells

(Lev et al. 2005)

Bushman Rockshelter, South Africa

Seeds of bufalo thorn (Ziziphus macronatus) and fragments of marula nuts (Sclerocarya birrea)

(Plug 1981; Wadley 1987)

Border Cave, South Africa

Charred seeds and nuts

(Beaumont 1973a)

Sibudu Cave, South Africa

Seeds mainly from evergreen fruit trees

(Wadley 2004)

Strathalan Cave B, South Africa

Corm scales or bases from the iris genera Watsonia and Tritonia-Freesia

(Opperman & Heydenrych 1990)

low), Neanderthals resemble modern rodeo riders in the frequency and anatomical location of skeletal trauma. Still, the stable isotopes in thirteen adult Neanderthal bones—from Jonzac, Saint Césaire, Les Pradelles (Marillac), and Rochers-de-Villaneuve in France, Sclayn (Scladina), Spy, and Engis Caves in Belgium, and Vindija Cave in Croatia—imply that meat dominated Neanderthal diets, both under relatively mild interglacial conditions (Sclayn) and cold glacial ones (the remaining sites). At each site, the bones retain protein (collagen) residues with the key isotopes, carbon-13 (13C) and nitrogen-15 (15N). In midlatitude, terrestrial settings, protein that is relatively rich in 13C implies an origin in grassy (as opposed to shadier, more wooded) conditions, while protein that is rich in 15N implies a highly carnivorous (as opposed to herbivorous) diet. Elevated 13C levels indicate that the ive analyzed Neanderthals hunted and gathered mainly in open, grassy or steppe situations, while high 15 N levels imply that they fed largely on animals. In their 13C levels, the sampled Neanderthals resembled many of their herbivore and carnivore contemporaries, but in their 15N levels, they recalled only wolves, lions, and spotted hyenas. he implication is that they were top carnivores, at the peak of the food chain. Artifactual similarities may imply that their near-modern African MSA contemporaries also fed largely on animals, but they lived in places where plant foods were generally far more abundant, and like historic African foragers, they probably relied much less on large mammals. Unfortunately, stable-isotope analysis will be slow to reveal their diet since climatic conditions over most of Africa do not favor protein survival in bones of such antiquity. In many Mousterian cave sites, bones of large carnivores—mainly cave bears but also hyenas and wolves—outnumber Mousterian artifacts and other traces of human presence, and the sites appear to have been

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mainly carnivore dens that Mousterians occupied only sporadically. In caves that Mousterians occupied intensively, large carnivore bones are rare, and in general, Mousterian/MSA people and large carnivores probably avoided each other. In apparent contradiction, a Neanderthal bear cult has been inferred from apparently patterned arrangements of cave bear bones in European caves such as Le Regourdou (France), Drachenloch (Switzerland), and Petershöhle (southern Germany). In these caves and others, complete bear bones, especially skulls, tend to occur near the walls or near large limestone slabs that have been imaginatively interpreted as stone lockers built by the Neanderthals. However, there are no stone artifacts or cutmarked bones to demonstrate human activity, and bear trampling could have destroyed many complete bones and displaced others to cave walls or to large fallen slabs. Almost certainly, the bones accumulated naturally from hibernating bears that occasionally died in the caves. Cave bear bones abound in Chauvet Cave, southern France, famous for its early Upper Paleolithic wall art, and the prominence of cave bears in the wall art and the superposition of their claw marks on some paintings conirm chronological overlap. Radiocarbon dating of the paintings, of ireplaces on the cave loor, and of cave bear bones bracket the overlap between 32 and 28 ka. However, by 25 ka, the cave bear had apparently vanished from Europe, and the most likely reason is that expanding Upper Paleolithic human populations progressively limited the number of suitable hibernation caves. Larger, denser Upper Paleolithic populations probably also explain why cave sites dominated by hyenas or wolves also declined sharply ater 40–35 ka. In general, it is diicult to estimate how successful the occupants of any particular Mousterian/MSA site were at obtaining animals. At sites where artifacts, humanly damaged bones, or other clear evidence of human presence are sparse, it is even diicult to tell what proportion of the animals were obtained by people and what proportion may have been killed by carnivores or died naturally. he problem tends to be particularly acute at open-air sites, where cultural debris and fossilized feces (coprolites), chewed bones, or other evidence of carnivore activity may be equally abundant. Even at a site where context and associations indicate that people were the principal bone accumulators, it is generally impossible to establish whether the relative abundance of various species relects human practices or preferences, the natural abundance of the species near the site, or some combination of these factors. Separating the relative roles of human behavior and environment is possible only when species abundance can be compared between two or more sites. In this event it may be possible to control for diferences in environment by reference to pollen, sediments, or geochemistry and to control for diferences in behavior by reference to artifacts or other

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cultural debris. hus if pollen and sediment analyses indicate that two sites shared a similar environment, human behavior becomes a likely cause for diferences between them in species abundance. Conversely, environment is implicated if artifacts or other cultural remains indicate that the occupants of two sites were similar in their behavior. In most instances where controlled comparisons are possible, differences in species abundance among Mousterian/MSA sites or between Mousterian/MSA sites and yet older or younger ones appear to relect diferences in site environment rather than in occupant behavior. hus, in France, luctuations in the relative abundance of red deer versus reindeer, traced from the Acheulean before 200 ka through the Mousterian aterward, correlate with climatic luctuations indicated by sediments and pollen. Red deer peaks are closely associated with warm periods and reindeer peaks coincide with cold ones. In Spain, Italy, and other parts of Mediterranean Europe, where climatic luctuations were less extreme, reindeer were always rare or absent and red deer prevailed in both interglacial and glacial periods. Similarly, in Ukraine and European Russia, where climate was always continental and climatic variation through time was perhaps less important than variation through space, bison are more common in Paleolithic sites on the south versus deer and horses in sites on the west, regardless of time or Paleolithic culture. In Africa too, most faunal diferences or changes among Mousterian/MSA sites relate more clearly to climate than to human behavior. he most dramatic example comes from the Sahara, where episodes of nonoccupation alternated with periods when Mousterian or Aterian people coexisted with typical African grassland or savanna ungulates. Associated deposits from now defunct springs or lakes show clearly that climatic change (not human behavior) accounts for the alternation. More subtle climatic change controlled the abundance of grazing versus browsing species within MSA and LSA archaeological sequences at the extreme southern tip of Africa. Here, independent of associated artifacts, deposits that suggest relatively cool, dry (glacial) conditions are dominated by grazers, whereas deposits suggesting relatively warm, wet (interglacial) climate are dominated by browsers. he same pattern marks deposits with bones that were collected by carnivores, conirming a climatic—versus a human-behavioral (cultural)—explanation for the species diferences. Among the comparatively rare instances of species diferences that are probably due to Mousterian/MSA behavior (as opposed to environment), perhaps the best examples are the shit from gazelle to fallow deer abundance within the Mousterian sequence at Tabun Cave in Israel and contrasts in ish, bird, and mammal frequencies between coastal MSA and LSA sites in South Africa, outlined in the next section. Fallow deer prefer moister conditions than gazelle, and the sharp increase in deer

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late in the Tabun Mousterian sequence was originally thought to relect a regional increase in moisture. However, local and regional sediment and pollen observations do not indicate a climatic change at the same time, though they do suggest an earlier change toward drier (not wetter) conditions. In fact, analysis of the Tabun sedimentary sequence now suggests that fallow deer increased because of a change in the way people used the cave. In the earlier Mousterian, it was probably a camp to which gazelles were brought from the nearby coastal plain. Later, ater a chimney had opened through the roof, it was unsuitable for habitation and became a trap into which deer were driven from the wooded slopes above. Arguably, the list of culturally (as opposed to environmentally) induced faunal changes should also include a shit in small animals between Mousterian and Upper Paleolithic sites in western Italy and northern Israel. he analysis in each region depends mainly on the diference between a single Mousterian site—Moscerini Cave in Italy and Hayonim Cave in Israel—and one or two neighboring Upper Paleolithic sites. he small animal remains divide between sluggish or sessile prey (mainly tortoises, shellish, or both) and more agile or elusive species (primarily hares and birds). In both regions, Upper Paleolithic sites are richer in the more elusive species, and the diference might mean that population pressure had forced Upper Paleolithic people to expend more efort on harder-to-catch prey items. he population increase might have begun in the late Mousterian, but it would be easier to explain if it dated to the early Upper Paleolithic, when it could relect advances in hunting-gathering technology. he more fundamental problem, however, is that the Italian and Israeli samples that document the supposed faunal shit are small, and most other regions in Europe and western Asia do not show it. Some, such as northern Spain, even suggest a reverse trend in which Upper Paleolithic people concentrated much more heavily on shellish. In South Africa, large MSA faunal samples document statistically secure shits in the ratio of (sluggish) tortoises plus shellish to (more agile) hares plus birds, but the shits occur as oscillations within the MSA, they continue within the LSA, and they are more clearly tied to climate than to culture. As outlined immediately below, South African faunal samples could support the idea that LSA people focused more heavily on elusive small prey, if shellish and tortoises were removed from the sluggish category and ish were added to the elusive one. SOURCES (also table 6.11): Hadza foraging (O’Connell et al. 1990, 1992, 1988); carbonized plant remains at Klasies River (Deacon 1993; Deacon and Deacon 1999); Sai Island grindstones (van Peer et al. 2003); distribution of MSA grindstones in South Africa (Volman 1984); use of mammoth bones in construction—Mousterian (Klein 1973b; Sofer 1985, 1989b) and Upper Paleolithic (Gaudzinski et al. 2005); Neanderthal skeletal trauma (Berger and Trinkaus 1995); stable-isotope analysis of Neanderthal bones (Beauval et al. 2006; Bocherens et al. 1999, 2001, 2005; Fizet et al. 1995; Richards et al. 2000, 2008); Neanderthal bear cult—pro (Bergounioux 1958) and con (Kurtén 1976); Chauvet cave bears and Upper Paleolithic people (Bocherens et al. 2006); disappearance of the cave bear (Estévez

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2004); red deer/reindeer abundance in France (Bordes and Prat 1965) and in Mediterranean Europe (Altuna 1972, 1992; Stiner 1994; Straus 1992); large ungulate abundance in Ukraine and European Russia (Hofecker 1986; Hofecker et al. 1991; Klein 1969b, 1973b; Sofer 1989b); savanna ungulates in the Sahara (Close et al. 1990; Wendorf et al. 1987, 1993; Wendorf and members of the Combined Prehistoric Expedition 1977; Wendorf and Schild 1980, 1992); grazer/browser abundance at southern African sites (Klein 1980, 1983); gazelle/fallow deer abundance at Tabun Cave (Jelinek 1982b); small animal abundance in Italy and Israel (Stiner et al. 1999, 2000); Upper Paleolithic shellish in northern Spain (Straus 1992)

MSA Coastal Ecology in South Africa

MSA and LSA sites on the south and west coasts of South Africa have provided one of the richest and most detailed records of Stone Age ecology anywhere in the world. Table 6.12 (top) lists the principal MSA sites and their probable ages. Supplemented by a handful of Mousterian/ MSA sites in northern Africa, southwestern Asia, and Europe listed in table 6.13, the South African MSA sites provide the oldest known evidence for intensive human exploitation of coastal resources, dating from the Last Interglacial and the early part of the Last Glaciation, between 130 and 57 ka. Table 6.12 (bottom) lists the LSA coastal sites with which the South African MSA sites are most fruitfully compared. As a group, the LSA sites date partly from the end of the Last Glaciation, between 24 and 11 ka, but mainly from the Present Interglacial (or Holocene), ater 11 ka. MSA and LSA sites from the middle part of the Last Glaciation, between 57 and 24 ka are rare and usually poor in southern Africa, probably because conditions were mainly too dry to support substantial human populations. he long occupational hiatus is vexing because it includes the time, about 50 ka, when anatomically modern Africans expanded from Africa to swamp or replace the Neanderthals and other nonmodern humans in Eurasia. he expansion is discussed in detail near the end of this chapter, and its timing implies it was rooted in a behavioral shit from the MSA to the LSA. he expansion alone indicates that the shit enhanced human itness—the ability to survive and reproduce—and the diferential should be visible in an enhanced human ability to obtain natural resources ater 50 ka. he rich Last and Present Interglacial occupations on South African coasts ofer a unique opportunity to investigate human foraging capacity before and ater the Out-of-Africa expansion, even if they cannot answer the question of whether any signiicant difference arose abruptly about 50 ka or evolved more gradually between 50 ka and perhaps 20 ka. he South African sites occur on both the south (Indian Ocean) coast and the west (Atlantic) coast (ig. 6.49), and the distinction is important because historically, the two coasts contrasted sharply. Terrestrial foods were much sparser on the west coast, owing to much lower rainfall, but marine foods were more abundant, due to more persistent

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TABLE 6.12. South African sites that illuminate Middle and Later Stone Age coastal foraging. he listings proceed

from southeast to northwest along the South African coastline. Site

Geologic Age (basis)

References

Klasies River Main

Between 127 and 57 ka (ESR, geologic context; faunal associations)

(Deacon 1989; Deacon 1995; Deacon & Geleijnse 1988; Deacon & Shuurman 1992; Klein 1976; Singer & Wymer 1982; hackeray 1988; Voigt 1982)

Herolds Bay Cave

Between 120 and 80 ka (U-series dating, geologic context, and faunal associations)

(Brink & Deacon 1982)

Pinnacle Point

Between 164 and 40 ka (artefacts and OSL)

(Marean et al. 2007; Marean et al. 2004)

Blombos Cave

Between 140 and 75 ka (OSL and TL)

(d’Errico & Henshilwood 2007; d’Errico et al. 2001; d’Errico et al. 2005; Grine & Henshilwood 2002; Grine et al. 2000; Henshilwood 1996; Henshilwood 1997; Henshilwood 1998; Henshilwood 2004; Henshilwood 2005; Henshilwood et al. 2001a; Henshilwood & Sealy 1997; Henshilwood & Sealy 1998; Henshilwood et al. 2001b; Jacobs et al. 2003a; Jacobs et al. 2006; Jacobs et al. 2003b; Tribolo et al. 2006)

Die Kelders Cave 1

Between 71 and 45 ka (ESR, geologic context; faunal associations)

(Avery et al. 1997; Grine et al. 1991; Klein & CruzUribe 1996; Marean et al. 2000b; Tankard & Schweitzer 1976)

Ysterfontein 1 Rockshelter

Between 119 and >= 46 ka (14C and geologic inference)

(Halkett et al. 2003; Klein et al. 2004)

Sea Harvest

Between 130 and > 40 ka (14C and geologic inference)

(Grine & Klein 1993; Volman 1978)

Hoedjies Punt 1 & 3

Between 130 and > 40 ka (14C, (Berger & Parkington 1995; Butzer 2004; Parkington Infrared Stimulated Lumines2006, 99–100; Stynder et al. 2001) cence and geologic inference)

Boegoeberg 2

Between 130 and > 40 ka (14C and geologic inference)

Halkett, Hart, and Parkington (personal communication, 1996) (Klein & Cruz-Uribe 1996)

Nelson Bay Cave

12 - 0.5 ka (for layers with shells, ish, and other coastal food debris) (14C)

(Deacon 1984; Inskeep 1987; Klein 1972a; Klein 1972b)

Noetzie (Knysna)

5 - 1.5 ka (artefact and faunal associations)

(Orton & Halkett 2007)

Die Kelders Cave 1

2 - 1.6 ka (14C)

(Schweitzer 1974; Schweitzer 1979; Schweitzer & Scott 1973)

Byneskranskop Cave 1

13 - 0.8 ka (14C)

(Schweitzer & Wilson 1982)

Hawston Midden

1.9 ka ( C)

Avery unpublished and (Avery 1976)

Pearly Beach Middens

2.9 - 0.8 ka ( C)

MIDDLE STONE AGE

LATER STONE AGE

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TABLE 6.12. (continued )

Site

Geologic Age (basis)

References

Bakoond

4.3 - ?0.7 ka (14C and artefact and faunal associations)

(Orton 2007)

Paternoster 1062

2.8 ka (14C)

(Yates 1998)

Kasteelberg A and B

1.8 - 0.4 ka ( C)

Duiker Eiland

1.9 - 1.7 ka (14C)

Tortoise Cave

7.7 ka and 4.4 - < 0.75 ka ( C)

(Jerardino 1995; Robey 1987)

Pancho’s Kitchen Midden

3.7 - 0.6 ka ( C)

(Jerardino 1997; Jerardino 1998)

Elands Bay Cave

13 - 8 ka and 4 - 0.3 ka (for layers with shells, ish, and other coastal food debris) (14C)

(Buchanan 1988; Orton 2006; Parkington 1981; Parkington 1987; Parkington 1990; Parkington 2006; Parkington in press)

Hail Stone Midden

1 ka (14C)

(Parkington 1979; Parkington et al. 1988)

Connies Limpet Bar

0.4 ka ( C)

Duneield Midden

0.9 - 0.6 ka ( C)

(Orton 2002; Orton 2002/2003; Parkington 2006; Parkington et al. 1992; Tonner 2005)

Steenbokfontein

6.1 - 2.1 ka (14C)

(Jerardino & Swanepoel 1999; Jerardino & Yates 1996)

(Klein & Cruz-Uribe 1989; Smith 1987; Smith 1992a; Smith 2006a; Smith 2006b)

14

(Robertshaw 1979) 14

14

(Parkington 1979; Parkington et al. 1988)

14

14

North African, European, and west Asian coastal or near-coastal sites to which Mousterian/MSA people brought shells of edible species. he Aterian layers of Smugglers’ Cave are said to have provided particularly large numbers of limpets and mussels. he Abdur Reef site on the Red Sea Coast of Eritrea may one day be added to this list, since it has provided artifacts, shells, and large mammal bones dated to about 125 ka by Uranium-series analysis of associated corals (Bruggemann et al. 2003; Walter et al. 2000). However, the artifacts comprise a mix of Acheulean hand axes and MSA lakes and blades scattered irregularly through the deposits, most of the artifacts probably do not lie where they were dropped, and none are directly stratiied with shells or bones. As presently known, Abdur Reef indicates that people occupied the Red Sea coast 125 ka or before, but it says nothing about how they made a living.

TABLE 6.13.

Site

References

Smugglers’ Cave (Grotte des Contrebandiers) (Témara), Morocco

(Bouzouggar et al. 2002; Roche & Texier 1976)

Zouhrah Cave (El Harhoura 1), Morocco

(Aouraghe 2004; Debénath & Sbihi-Alaoui 1979)

Mugharet el ‘Aliya (Tangiers), Morocco

(Bouzouggar et al. 2002; Howe 1967; Howe & Movius 1947)

Bérard open air-site, Algeria

(Roubet 1969)

Haua Fteah, Libya

(Klein & Scott 1986; McBurney 1967)

Ras el-Kelb Cave, Lebanon

(Copeland & Moloney 1998; Reece 1998)

Moscerini Cave and possibly other coastal caves, Italy

(Stiner 1994; Stiner 1999; Stiner 2001)

Ramandils Cave, France

(Cleyet-Merle & Madelaine 1995)

Devil’s Tower Rockshelter, Gibraltar

(Garrod et al. 1928; Lalueza-Fox & Pérez-Pérez 1993)

Gorham’s Cave, Gibraltar

(Barton 2000; Barton et al. 1999; Eastham 1989; Pettitt & Bailey 2000; Rink et al. 2000; Waechter 1964)

Vanguard Cave, Gibraltar

(Barton 2000; Barton et al. 1999; Fernández-Jalvo & Andrews 2000; Pettitt & Bailey 2000)

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FIGURE 6.49. The approximate locations of the south african sites that illuminate msa and lsa coastal ecology. boldface marks sites with msa occupations.

20o

25o

0

30o

300 km

25o

25o

SOUTH AFRICA

Boegoeberg 30o

Blombos Pinnacle Point Herolds Bay Noetzie

Atlantic Ocean

Nelson Klasies Bay 25o River

35o 20o

Sibudu 30o

Indian Ocean 35o 30o

18o 32o

Dune Field Midden & Connies Steenbokfontein Limpet Bar Tortoise Cave Elands Bay Cave & Hail Stone Diepkloof Shelter Midden Panchoʼs Kitchen Midden Duiker Eiland PaterKasteelberg noster Sea Harvest 33o Hoedjiespunt

Ysterfontein

N

Bakoond Duinefontein

CAPE TOWN Swartklip

o

34

0

Hawston Die Kelders 1 Byneskranskop 1 Pearly Beach 30 km

18o

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ofshore upwelling of nutrient-rich waters. West coast waters also tended to be considerably cooler, and they promoted a distinctive intertidal community that shared few invertebrate species with the south coast. Terrestrial vertebrates were much more similar between the coasts, but higher humidity on the south coast tended to promote denser large mammal populations. he fundamental environmental contrast apparently endured global climatic luctuations, for regardless of age, shellish from MSA and LSA sites that span the past 125–115 ky anticipate their local historic counterparts on the same coast, and at any particular time, large mammals tend to be signiicantly rarer at west coast sites. he persistent environmental diference is important to keep in mind because it means that culture (rather than environment) is likely to explain the MSA/LSA faunal contrasts that the two coasts share. Overall, independent of coast, MSA faunas difer from LSA faunas in four respects: 1. 2. 3. 4.

MSA assemblages are signiicantly poorer in ish bones; MSA assemblages tend to contain fewer bones of airborne birds; MSA assemblages include fewer bones of the most dangerous ungulates; and on average, MSA shellish and tortoises tend to be signiicantly larger than their LSA counterparts.

Larger average shellish and tortoise size implies lighter, more selective MSA predation, probably because MSA people were less numerous. he section titled “Population Numbers” below provides supporting detail. Smaller MSA populations and more selective predation could further explain why large MSA shellish samples on the west coast contain a smaller range of species than LSA samples and, also, why they are poorer in a common and readily accessible but relatively small limpet that less numerous collectors are more likely to have ignored. MSA and LSA shellish assemblages from south coast sites have not been studied in suicient detail to determine if they exhibit a similar contrast in species diversity and richness. On both coasts, MSA and LSA people exploited Cape fur seals (Arctocephalus pusillus), and the four main MSA/LSA contrasts could be supplemented by a diference in individual seal ages between MSA and LSA sites. Fur seals breed exclusively on of-shore rocks, and they give birth mainly during a three-week period in late November and early December. Roughly nine months later, the adults force the young from the rocks, and many nine-to-eleven-month-old pups wash up on the adjacent mainland exhausted or dead. he section “Seasonality of Site Occupation” below shows that LSA people clearly recognized the potential boon, and nine-to-eleven-month-old seals dominate their sites. However, the same section shows that older seals, including numerous adults,

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rule the large MSA sample from Klasies River Main (KRM), and the diference suggests that the KRM people visited the coast more evenly through the year. Both LSA and MSA sites commonly provide ostrich eggshell fragments, but only LSA sites unquestionably contain fragments from shells that were modiied into canteens, and this may mean that MSA people were more closely tied to permanent water like the Klasies River. If so, the disadvantage could have been considerable since it would have limited their ability to move among locales that varied seasonally in desirable resources. However, a signiicant mobility diference from the LSA must remain speculative until the KRM MSA seal-age result is observed in another large sample, preferably from the west coast. Among all the known or proposed MSA/LSA contrasts, the rarity or lack of ish bones in MSA sites is the most conspicuous and consistent. Fish bones occur at MSA sites, particularly at Klasies River Main and Blombos Cave, but in each case, mammal bones outnumber them by an order of magnitude, and the ish were large individuals that could have been wash-ups. In coastal LSA sites, the reverse is true: ish bones commonly outnumber mammal bones by an order of magnitude, and the ish were primarily smaller individuals that were probably caught mainly with lines or nets. At both MSA and LSA sites, abundant intertidal shells and numerous bones of fur seals and shore birds show that the coast was nearby, and the rarity of ish bones in MSA sites must thus relect a cultural (behavioral) diference. It might mean that MSA people simply didn’t like ish, but a more likely alternative is that they lacked efective ishing technology. Only LSA sites have provided probable ishing gear, including grooved stones that could have sunk lines or nets and carefully shaped, double-pointed toothpick-sized bone “gorges” that could have been tied to a line and baited. Local indigenous people used similar gorges to catch both ish and shore birds. he remaining contrasts between MSA and LSA faunas are subtler, and they are less clearly universal. he diference in the frequency of airborne birds is conspicuous mainly on the south coast where penguins outnumber cormorants in the MSA occupations at Klasies River Main and Die Kelders Cave 1, while cormorants outnumber penguins in local LSA sites. As discussed below, only LSA people unquestionably had projectile weapons, and the utility of these in fowling could explain the diference. he contrast is not so clear on the west coast, however, where penguins outnumber cormorants in a small sample from the MSA layers at Ysterfontein 1 but not in a larger sample from Boegoeberg 2. he contrast in ungulate frequencies is also clearest on the south coast, particularly between the Last Interglacial MSA layers at Klasies River Main and the Present Interglacial LSA layers at Nelson Bay Cave. he environs of both sites are similar today, and sediments and geochemistry indicate that the environs were similar to the present and to each other during

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Nelson Bay Cave LSA (Present Interglacial deposits) Klasies River Mouth Cave 1 MSA (Last Interglacial deposits)

Byneskranskop 1 LSA (end Last Glaciation and Present Interglacial deposits) Die Kelders Cave 1 MSA (early Last Glaciation deposits)

eland

Cape buffalo

wild pig

6

39

38

80

29

3

5

28

20

16

3

0

the Last and Present Interglacials. In the historic environment, bufalo and bush pigs signiicantly outnumbered eland, and this situation is mirrored in the LSA layers at Nelson Bay Cave (ig. 6.50). However, in the MSA layers at Klasies River Main, eland strongly outnumber both buffalo and pigs, and if eland were as rare nearby as they were historically, MSA people must have deliberately concentrated on them. Eland similarly dominate bufalo in the medium-sized MSA samples from Blombos Cave and Die Kelders Cave 1, both on the same south coast as Klasies River, and in a much smaller sample from Ysterfontein 1 on the west coast. Blombos and Ysterfontein 1 lack nearby LSA counterparts with equally large faunal samples, but Die Kelders 1 can be fruitfully compared to Byneskranskop Cave 1. As at Nelson Bay Cave, the Byneskranskop 1 deposits span the Present Interglacial and they suggest the same contrast with the MSA (ig. 6.50): compared to the Die Kelders 1 MSA layers, the Byneskranskop 1 LSA deposits are much poorer in eland and richer in bufalo and wild pigs. In this instance, an environmental explanation for the contrast is possible because MSA people occupied Die Kelders 1 in the early part of the Last Glaciation (oxygen-isotope stage 4), when local climate was relatively cooler and moister. Still, temperature variation had no obvious impact on historic bufalo and eland numbers in Africa, and wherever both species occurred together, bufalo tended to be much more numerous. In addition, the eland-to-bufalo ratio remained constant when “glacial” conditions replaced “interglacial” ones at Klasies River Main. At least circumstantially then, Byneskranskop 1 and Die Kelders support an apparent MSA preference for eland over bufalo and wild pigs.

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561 FIGURE 6.50. Top: The minimum numbers of eland, Cape buffalo, and bushpig in samples from the last interglacial deposits of Klasies river main Cave 1 and in the Present interglacial deposits of nearby nelson bay Cave, south africa. Bottom: The minimum number of individuals from the same species in the early last Glaciation deposits of die Kelders Cave 1 and in the end last Glacial/ Present interglacial deposits of nearby byneskranskop Cave 1. The relatively greater abundance of buffalo and bushpig at nelson bay and byneskranskop 1 probably relects the technological superiority of local lsa hunters over their msa predecessors.

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As among the various ungulates available to MSA and LSA people, eland were probably the least dangerous, while bufalo and wild pigs were among the most. Unlike bufalo and wild pigs, which tend to counterattack potential predators, eland are more inclined to lee, and they can even be driven in a desired direction. If as seems likely, MSA hunting weapons consisted entirely of thrusting spears, eland would have presented far less personal risk to hunters, and if people found an eland herd in the right position, they could have driven it over clifs like those that exist at Klasies River and the other three MSA sites where eland dominate. LSA people could also have driven eland, but from at least 20 ka, they probably had projectile weapons since their artifact assemblages include bone rods that historic people incorporated in arrow shats and tiny backed and nonbacked stone bits that historic people used to tip arrows. Armed with arrows or other projectiles, LSA people could have attacked more dangerous prey from a distance, and even if their success rate was relatively low, the larger number of trials would have increased the number of dangerous ungulates in their sites. he LSA advantage would have been substantial since they surely encountered the more dangerous species far more oten than they encountered eland. Driving over clifs would result in the death of all members of a herd regardless of age, and it would produce a mortality proile in which prime-age (reproductively active) adults were as abundant as they are in live groups. Hunting individual animals, whether with thrusting spears or projectiles, would net primarily the very young and the old, which tend to be especially vulnerable because of small size, limited experience, or weakened special senses. he next section shows that the ages of the Klasies eland conform to the pattern expected from driving, while the ages of the bufalo conform to the pattern that would result from stalking individual animals. Eland, bufalo, and wild pigs are subequally represented in the MSA deposits at Sibudu Cave near the east coast of South Africa, and this could mean that an MSA emphasis on less dangerous prey was geographically localized. However, the Sibudu sample is small, and no local LSA sample is available for a controlled comparison. Suitable MSA and LSA samples are also lacking elsewhere in Africa, and even if they existed, past geographic variation in the composition of large mammal communities means that the details of an MSA/LSA contrast would vary from region to region. his may always complicate eforts to conirm a fundamental contrast in MSA/LSA ability to obtain large ungulates. A diference in the ability to acquire ish should be much easier to corroborate since it would take the same form everywhere. he rarity or absence of ish in South African costal MSA sites is striking, and MSA sites elsewhere all support the pattern, with three possible exceptions: the Katanda MSA sites on the Semliki River (Zaire); Mousterian Site 440

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on the Nile River (northern Sudan), and the Aduma MSA sites on the Awash River (Ethiopia). At the Katanda sites, the ish bones are associated with barbed points that are likely to have tipped ish spears, and the Katanda ish come mainly from two species that MSA people might have speared in seasonally shallow loodplain waters. As previously discussed, however, the Katanda stone artifacts are not classically MSA, and the barbed points closely resemble ones that mostly postdate 12 ka in eastern and northern Africa. For this reason and others, it is thus possible, even probable that the Katanda points and by extension the ish bones have mistakenly been assigned to the MSA. he ish bones at site 440 are more securely associated with Mousterian (or MSA) artifacts, but the bones and artifacts were difused (“mixed”) through two sandy layers 30–50 cm thick, and the bone/artifact association might not have resulted from human activity. he Aduma sites have provided few identiiable bones, and they could represent natural background on a river loodplain more than human food debris. Fish bones dominate the samples, but they are mainly from catish (Clarias sp.) that could have died naturally in shallow loodplain pools. In sum, neither Katanda, site 440, nor the Aduma sites show that MSA/ Mousterian people ished routinely, and the most plausible explanation is that they lacked the necessary technology. he utilization of aquatic foods is never as apparent in European Mousterian sites as it is in South African MSA caves, perhaps because relevant coastal sites are rare. Still, in the few sites that are known, primarily in western Italy and on Gibraltar, ish bones are rare or absent in the Mousterian levels, even though intertidal mollusks show that the sea was nearby. Fish bones are also mostly absent in Mousterian sites that were located near rivers, though they are suiciently abundant in some similarly situated Upper Paleolithic sites to suggest active ishing, particularly ater 22 ka in southwestern France and northern Spain. Stable-isotope analysis of human bone protein (collagen) conirms that at least some Upper Paleolithic people relied on ish or other freshwater resources by 28 ka, but it has so far detected no ish component in Mousterian diets. he addition of ish is suicient in itself to explain why LSA/Upper Paleolithic people appear to have signiicantly outnumbered earlier people living under similar conditions. If Katanda and Sudanese Mousterian site 440 are discounted, LSA Africans were probably the irst people to ish actively, but it may be diicult to determine exactly when they started because the pertinent sites probably date from between 60 and 20 ka when much of southern and northern Africa appear to have been depopulated. Perhaps even more important, this was an interval of lower sea levels, and coastal sites that could provide the most persuasive evidence for early LSA ishing are now mostly submerged on the continental shelf.

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ChaP Ter s iX SOURCES (also tables 6.12 and 6.13): human population and Last Glacial aridity in southern African (Beaumont and Vogel 2006; Deacon and hackeray 1984); MSA/LSA transition in eastern Africa (Ambrose 1998a); contrasts in shellish between MSA and LSA coastal sites in South Africa (Klein et al. 2004; Parkington 2003); seal ages in MSA and LSA sites (Klein et al. 1999c); marine bird frequencies in MSA and LSA sites (Avery 1990); historic abundance of eland and bufalo in Africa (Estes 1992; Skinner and Chimimba 2005); antiquity of the bow and arrow in southern Africa (Deacon 1984); MSA fauna from Sibudu Cave (Plug 2004); ish bones at Katanda (Brooks et al. 1995; Yellen et al. 1995), Mousterian site 440 (Greenwood 1968; Shiner 1968), and Aduma (Yellen et al. 2005); rarity of ish bones in Mousterian sites on European coasts (Barton et al. 1999; Garrod et al. 1928; Stiner 1994) and rivers (Mellars 1973; Straus 1983, 1985); freshwater component in Upper Paleolithic diets revealed by stable-isotope analysis (Richards et al. 2001)

Hunting versus Scavenging

For the sake of argument, it has been assumed so far that Mousterian/ MSA people actively hunted large animals, albeit less successfully than their Upper Paleolithic/LSA successors. But we must consider the possibility that Mousterian/MSA groups acquired most large carcasses by scavenging. Most specialists believe that at least some Mousterian/MSA people routinely hunted large animals, but some argue that Mousterian/ MSA populations mainly scavenged. he issue is complicated partly because hunting and scavenging are not as distinct as they might appear. For instance, recently observed Hadza hunter-gatherers in northern Tanzania obtain about 80% of their large mammal carcasses by bowand-arrow hunting and about 20% by scavenging. heir scavenging often involves driving leopards, spotted hyenas, or lions of kills, however, and if the predators refuse to leave, they may themselves be killed. In both practice and result, this kind of “confrontational” scavenging may not be usefully separated from active hunting, and a more meaningful distinction is probably between early and late access to carcasses. It is this diference that archaeologists must address if they are to evaluate the hunting prowess of MSA/Mousterian people. Among the available clues are the pattern of skeletal part representation, the ages of prey animals at death, and the frequency and positioning of bone damage marks. he MSA animal bones from Klasies River Main (KRM) provide a useful example. At KRM, the large animals are mainly bufalo and eland, and these are represented primarily by bones of the feet (metapodials, carpals, tarsals, and phalanges) and of the skull (mainly teeth). Proximal or “upper” limb bones (humeri, radii, femurs, and tibiae) are relatively scarce, although they tend to be very rich in meat, marrow, and other edible tissue and are thus the bones that both people and other large predators tend to carry away irst. heir scarcity at KRM might therefore mean that the people were mainly picking over carcasses on which other predators had already fed. he implication would be for scavenging in the narrowest sense of the term.

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here is the problem, however, that foot and skull (mainly dental) elements from large ungulates tend to outnumber limb bones not only at KRM but at most prehistoric sites throughout the world, including, for example, African Iron Age sites, where the large ungulates were mainly domestic cattle that are unlikely to have been scavenged. he pervasiveness of the pattern suggests it is due to factors that all large ungulates share, and the most conspicuous factors are relative bone durability and identiiability. he skull and foot bones that tend to dominate large ungulate assemblages are among the densest parts of the skeleton, and they are also among the easiest to identify even ater they have been fragmented. heir density and relative lack of nutrients probably combined to ensure that unlike less dense, more nutritious limb bones, they were not extensively butchered and broken during food preparation. hey were thus more likely to be buried intact, and this, together with their initial density, helped maintain their identiiability even ater relatively intense postdepositional leaching and proile compaction. At KRM and many other sites, in contrast to large ungulates, small species (like steenbok and bushbuck) are relatively well represented by limb bones (vs. skull and foot elements). his could mean that the KRM people hunted small ungulates more oten. However, wherever small ungulate limb bones are abundant, it is also notable that they tend to be much more complete than their large ungulate counterparts. his might mean that people processed smaller bones less intensively, that small size enhanced postdepositional survival, or both. Another possibility is that people preferentially smashed large ungulate limb bone shats into fragments that analysts usually consider useless for estimating limb bone numbers and that hyenas then selectively removed the large ungulate limb bone epiphyses (ends). he supposed diference between small and large ungulate limb bone numbers is based strictly on the epiphyses, and a program to reit previously neglected large ungulate shat fragments might show that it is analytic artifact. Reitting is impractical at KRM because the excavators thought that shat fragments retained little information, and they kept relatively few. However, computer-aided reitting has been applied to the large ungulate fragments from the MSA layers of Die Kelders 1, and it suggests that the number of large ungulate limb bones would have been seriously underestimated otherwise. he result is problematic because the method requires an analyst to determine not only the kind of animal and the skeletal part from which a fragment came but also the exact position of the fragment on a shat, and most analysts ignore most shat fragments because they believe that they cannot be identiied and placed precisely enough. here is the additional problem that even if selective human destruction explains the apparent underrepresentation of large ungulate limb bone shats, selective hyena removal must be invoked to explain the underrepresentation of the associated

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epiphyses, and there is no evidence that hyenas prefer larger epiphyses over smaller ones. here is also little or no evidence for hyena activity at KRM, Die Kelders, and most other sites where large ungulate limb bones seem underrepresented. Still, a hyena role cannot be excluded, and the bottom line remains the same with or without it: the apparent underrepresentation of large limb bones at KRM and other sites need not mean that prehistoric people didn’t obtain them. In contrast to eland and bufalo skeletal part representation at KRM, whose implications are at best equivocal, bone damage marks and prey death ages point more directly to active hunting. Carnivore tooth marks are extremely rare, and the few that have been observed could relect occasional carnivore visits when the people temporarily abandoned KRM. Stone-tool cut marks are far more abundant, and none overlie tooth marks. heir placement shows that the people were both disarticulating intact bufalo and eland carcasses and deleshing individual bones. he tip of a stone point still embedded in a bufalo vertebra, mentioned previously, also demonstrates hunting. he ages of individual bufalo, eland, and other ungulates can be estimated from the heights of their teeth above the roots. When the ages are summed to make an age proile for each species, two patterns emerge. he bufalo exemplify the irst pattern, in which most individuals are either very young (less than 10% of maximum lifespan) or relatively old (more than 50% of lifespan) (ig. 6.51). Prime-age adults (between 10% and 50% of lifespan) are rare, particularly compared with their abundance in live herds. Paleobiologists call such an age proile “attritional” because it could result from normal everyday mortality factors—like endemic disease, accidents, and carnivore predation—that tend to bypass prime-age individuals and to afect mainly the very young and the old. Since very young and old bufalo are precisely those that die most oten of natural causes, their abundance at KRM might imply human scavenging. However, it is unlikely that the people could have located so many very young bufalo before hyenas or other scavengers did, and modern observations indicate that these scavengers consume young carcasses quickly and completely. he abundance of very young bufalo is thus more likely to relect active hunting. he second age proile is exempliied by the eland (ig. 6.51), in which relative to the very young and the old, prime-age adults are wellrepresented, roughly in proportion to their abundance in live herds. Paleobiologists call this kind of age proile “catastrophic” because it could be ixed in the ground only by a great lood, volcanic eruption, or other catastrophe that kills individuals in rough proportion to their live abundance, regardless of their age. Bufalo are too poorly represented at other MSA sites to determine the nature of mortality, but eland are abundant enough at Die Kelders Cave 1 to show that they also died “catastrophi-

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number of individuals

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attritional mortality profile corresponding to the idealized catastrophic profile to the left

idealized catastrophic mortality profile for a population of large mammals

older

older

crown height

crown height

number of individuals

? individuals lost postdepositionally

? individuals lost postdepositionally

40

40

30

30 20

20 KRM 1 eland (n = 93)

10 0

20 40 60 80 percentage of life span (22 yrs)

10

100% 0

KRM 1 Cape buffalo (n = 39) 20 40 60 80 100% percentage of life span (24 yrs)

cally.” At both KRM and Die Kelders 1, the most likely catastrophe was human driving of whole groups into traps or over clifs such as those that occur near both sites. Modern observations show that among all the species represented at KRM and Die Kelders, eland are the most amenable to driving, and it appears that the KRM and Die Kelders people had discovered this special vulnerability. he KRM and Die Kelders people could not have driven eland very oten, however, or the repeated removal of numerous prime adults would have sapped the reproductive vitality of the species, and there is no evidence that eland numbers declined during or ater the occupation of either site. Historically, eland were extremely widespread in Africa but they were nowhere very numerous, and eland herds tended to be widely dispersed and diicult to locate. his means that the KRM and Die Kelders people probably rarely found eland in a position suitable for driving— and, thus, that they probably killed only a small fraction of the available animals. If eland were rare in the environment and if the KRM and Die Kelders people did not obtain them very oten, it follows that they must have been even less successful at obtaining other ungulates, such as buffalo, which were probably more common in the environment but which are rarer in the sites. In sum, the KRM and Die Kelders prey mortality proiles imply that MSA people actively hunted. Combined with the dominance of eland

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567 FIGURE 6.51. Top left: schematic catastrophic age (mortality) proile for a population of large mammals that is basically stable in size and age structure. The open bars represent the number of individuals that survive in each successive age cohort, the hatched bars the number that die between successive cohorts. Top right: separate plot of the hatched bars, showing the corresponding schematic attritional age proile. The basic form of corresponding catastrophic and attritional proiles is the same for all large mammals, but the precise form will differ from population to population depending on species biology and speciic mortality factors. Middle: eland and buffalo lower third molars (m3s) showing the crown height dimension from which individual age at death may be estimated. Bottom: mortality proiles based on crown heights for eland and Cape buffalo from the msa layers of Klasies river main (Krm) Cave 1. it is probable that postdepositional leaching, proile compaction, and other destructive factors have selectively destroyed teeth of the youngest eland and buffalo. if this is accepted, the eland proile is clearly catastrophic and the buffalo proile is attritional. as discussed in the text, this suggests that the Krm people obtained eland by driving entire social groups over cliffs or into other traps where differences in individual vulnerability due to age would be meaningless. in contrast, the people must have obtained buffalo by using methods to which prime-age adult buffalo were relatively immune.

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over bufalo, however, they also suggest that the people obtained very few ungulates overall.

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SOURCES: Mousterian/MSA hunting (Chase 1986, 1988, 1989; Hofecker and Baryshnikov 1998; Klein and Cruz-Uribe 1996; Kozlowski 1990a; Otte 1990a; Stiner 1994) vs. scavenging (Binford 1984, 1985, 1989); Hadza hunting and scavenging (O’Connell et al. 1992, 1988); early and late carcass access (Bunn 1991; Bunn and Ezzo 1993; Potts 1984); skeletal part representation and scavenging at KRM (Binford 1984); foot and skull bone dominance in African Iron Age sites (Voigt 1983); hyena selection of large ungulate limb bone epiphyses (Marean and Kim 1998); reitting of shat fragments at Die Kelders 1 (Marean et al. 2000a); stone-tool marks on KRM bones (Milo 1994, 1998); estimation of ungulate age from crown height (Klein and Cruz-Uribe 1984)

Seasonality of Site Occupation

he analysis of gazelle teeth summarized in the section titled “Settlement Systems,” above, shows that west Asian near-modern Mousterians moved camp with the seasons. hey were presumably driven by seasonal variability in local resource availability, which oten prompted movements among historically observed hunter-gatherers. African fossil ungulate teeth have not yet been analyzed for indications of seasonal bone accumulation, but South African coastal MSA and LSA sites invariably contain bones of the Cape fur seal (Arctocephalus pusillus), and as mentioned above, these aford an independent opportunity to determine the season(s) of site occupation. he reason is that until 1941, when fur seals were irst legally protected from human predation, they bred almost exclusively on ofshore rocks. he vast majority of births occurred within a few weeks in late November and early December, and adults forced the young from the rocks about nine months later. In recent times, large numbers of nine-to-eleven-month-old seals then washed up ashore, exhausted or dead. It is the short fur seal birth season and the consequent seasonal peak in onshore availability that allow estimates of when fur seal bones accumulated at fossil sites. In cases where fossil seal ages cluster tightly around nine-to-eleven months, bone accumulation probably centered tightly on the August–October period of superabundance in nine-to-eleven-month-old seals. Where ages cluster more loosely around nine-to-eleven months, bone accumulation probably included not only the August–October period but also other times of year. And where ages fail to cluster near the nine-to-eleven-month average, then accumulation probably fell largely outside August–October. Fossil seal bones may be “aged” by comparison with bones from known-age animals. Various skeletal elements are useful for age determination, but the distal humerus is especially apposite because it is relatively durable and thus tends to dominate fossil samples. he most consistently available dimension is the mediolateral diameter or “breadth” of the distal end. Figure 6.52 uses a boxplot format to summarize breadths of fur seal distal humeri in samples from key MSA and LSA sites, in two sam-

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75

Cape Fur Seal (Arctocephalus pusillus)

Newborns ca. 9 months (49)

LSA SAMPLES (ca. 11 - 0.4 ka) Dune Field Midden (67) Elands Bay Cave 1-5 (14) Elands Bay Cave 10-19 (30)

0

10 20 30 mm

Kasteelberg B (204) Kasteelberg A (34) Die Kelders 1 LSA (189) Nelson Bay Cave (87)

MSA SAMPLES (ca. 127 - 57 ka) Boegoeberg 2 (4) Die Kelders 1 MSA (7) Klasies River Mouth (71)

BROWN HYENA SAMPLES Boegoeberg 1 (22) Central Namib Dens (all) (134) Central Namib Dens (> 9 mos.) (80) 15 30 45 60 75 mediolateral diameter of the distal humerus (mm) FIGURE 6.52. boxplots summarizing distal humerus breadth in known-age fur seals and in fossil or subfossil samples from lsa, msa, and brown hyena sites on the coast of southern africa. The key elements of each plot are the vertical line near the middle, which represents the median measurement or iftieth percentile in each sample, and the open rectangles, which enclose the middle half of the measurements, between the twenty-ifth and seventy-ifth percentiles (velleman 1997). The shaded rectangles mark the 95% conidence limits for the medians. When two shaded rectangles fail to overlap, chance (sampling error) is unlikely to explain the difference between the associated medians. The msa samples from boegoeberg 2 and die Kelders 1 are probably too small for meaningful comparisons, but the sample from Klasies river main differs from the various lsa samples in two important ways: it contains signiicantly larger humeri on average (as indicated by the median and its conidence limits), and it is characterized by greater dispersion around the median (as indicated by the open rectangle). in median size and especially in dispersion, the Klasies river main sample recalls the fossil brown hyena sample from boegoeberg 1. as discussed in the text, the boxplots suggest that lsa people timed their coastal visits to coincide with the august– october period when nine-to-eleven-month-old seals are especially abundant, whereas the msa inhabitants of Klasies river main followed a less focused seasonal round. it is even possible that msa people occupied the coast year round, like the boegoeberg 1 hyenas.

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ples accumulated by brown hyenas (Hyaena brunnea), and in samples of modern newborns and nine-month-olds. In each plot, the vertical line near the center is the median, the open rectangle encloses the middle half of the data (between the twenty-ith and seventy-ith percentiles), the shaded rectangle is the 95% conidence period for the median, and the vertical lines at the ends mark the range of more or less continuous data. Circles or asterisks indicate extreme values (points that are far removed from the main body of data). he number of specimens in each sample is given in parentheses. Samples for which the 95% conidence limits do not overlap difer signiicantly in the conventional statistical sense. To aid visual assessment, a vertical gray bar extends the range for nine-month-olds through the igure. Figure 6.52 shows that, in most samples, median fur-seal-distalhumerus breadth lies within or just outside the range for known ninemonth-olds. In addition, breadths in the LSA samples tend to be tightly clustered around the median, and the sum suggests that LSA people focused their coastal visits on the August–October period when they could harvest nine-to-eleven-month-old seals on nearby beaches. he most glaring departure from the LSA pattern involves the MSA sample from Klasies River Main Cave 1, where distal humerus breadths are much more broadly dispersed and the median is substantially and signiicantly greater than that for nine-month-olds. his implies that MSA people occupied Klasies River Main mainly outside the peak period of availability of nine-to-eleven-month-old seals and, more generally, that MSA people followed a signiicantly diferent seasonal round than their LSA successors. From the available data, it could even mean that they did not follow a seasonal round at all. Arguably, the MSA sample from Boegoeberg 2 supports the same inference, while the one from Die Kelders Cave 1 suggests an LSA-like pattern. Unfortunately, however, both samples are too small for any irm conclusion. As an aid to further interpretation, it is instructive to compare the LSA and MSA boxplots with those for the two brown hyena samples at the bottom of igure 6.52. he sample labeled “Central Namib Dens” was accumulated between 1990 and 1996 in dens adjacent to a fur seal breeding colony about 20 km south of Luderitz, Namibia. he other sample was accumulated at the Boegoeberg 1 fossil hyena den near the Boegoeberg 2 MSA site mentioned above. In keeping with hyena access to a breeding colony, the Central Namib sample includes ity-four humeri from newborn individuals. he abundance of newborn specimens explains why Central Namib median distal humerus breadth is so small, and the rarity or absence of newborn specimens in the various fossil sites shows that none were near breeding colonies. For heuristic purposes, the Central Namib sample is plotted twice, with and without the newborn specimens. When the newborn specimens are removed, median

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humerus breadth more closely approximates that in the various LSA samples, but the degree of breadth dispersion becomes much greater. his is because the Central Namib brown hyenas remain at the coast throughout the year, including times when the only available seals are older than nine-to-eleven months. he comparable or greater dispersion in the Boegoeberg 1 fossil hyena sample also illustrates nonseasonal bone accumulation. In sum, the boxplots in igure 6.52 imply that LSA people generally timed their coastal visits to include the seasonal peak availability of young fur seals, whereas the MSA occupants of Klasies River Main obtained many, if not most, of their seals during other seasons. If larger fur seal samples from other MSA sites replicate the Klasies River Main pattern, the implication would be that MSA people occupied the coast nonseasonally, perhaps like brown hyenas. MSA people may have difered from LSA people because they failed to perceive seasonal variability in fur seals and other resources or because they were technologically more limited. In this regard, as mentioned previously, it may be pertinent that no MSA site has provided unambiguous evidence for water containers, whereas LSA sites have repeatedly provided fragmentary or even whole ostrich eggshell canteens. he contrast is particularly striking because ostrich eggshell abounds at Boegoeberg 2 and other west coast MSA sites. SOURCES: seasonal movements by historic hunter-gatherers (Kelly 1983, 1992); fur seal breeding habits (David 1989; Skinner and Chimimba 2005); Central Namib hyena dens (Skinner et al. 1995, 1998; Skinner and van Aarde 1991); LSA ostrich eggshell canteens (Deacon 1984)

Disposal of the Dead One of the reasons the Neanderthals are so well-known is that they buried their dead, at least on occasion. Few excavators have identiied burial pits, perhaps because these were usually shallow and easily obscured by proile compaction long ago. Some pits may also have been overlooked in the relatively unsystematic excavations that produced important Neanderthal skeletons in the late nineteenth and early twentieth centuries. In addition, there are no instances where Neanderthal skeletons are accompanied by special artifacts or other indisputable grave goods. However, burial pits have been reported in France from La Chapelle-aux-Saints, La Ferrassie, and Le Roc de Marsal and in Israel at Kebara Cave. Even if a pit were not obvious, burial could be inferred for Kebara since the deposits accumulated far too slowly to protect a body exposed on the surface and coprolites show that hyenas intermittently visited the site. At other Neanderthal sites, including the original type locality (Feldhofer Cave) in Germany, Spy Cave in Belgium, and Le Moustier, La Quina, Le Regourdou, and La Roche à Pierrot (Saint-Césaire) caves in

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France, deliberate burial is the most plausible explanation for the presence of articulated Neanderthal skeletons. Intentional burial probably also accounts for the Neanderthal remains found at Kiik-Koba in Crimea (Ukraine), at Mezmaiskaya Cave (southern Russia), at Tabun and Amud Caves (Israel), at Dederiyeh Cave (Syria), at Shanidar Cave (Iraq), and at Teshik-Tash Cave (Uzbekistan). La Ferrassie was a veritable cemetery of eight graves, and Shanidar may have contained ive, although some of the Shanidar skeletons may have lain under natural rockfalls. Specialists employing diferent criteria estimate diferent numbers of Neanderthal burials, but there is consensus on about thirty-ive, split more or less evenly between Europe and western Asia. he near-modern Israeli contemporaries of the Neanderthals at Skhul and Qafzeh Caves also buried their dead, and the graves tend to resemble those of the Neanderthals in their essential simplicity. A boar mandible found partly under the let radius of Skhul individual 5 and the antlers of a fallow deer associated with Qafzeh individual 11 may represent grave goods, but the case depends mainly on specimen completeness. his may relect only the fortuitous inclusion of each specimen in grave ill, which protected it from the various pre- and postdepositional forces that fractured bones of the same species elsewhere in the deposits. Certainly neither Skhul nor Qafzeh provides the striking evidence for burial ritual or ceremony that characterizes some Upper Paleolithic/Later Stone Age graves, as discussed in the next chapter. Except perhaps at Taramsa Hill, Egypt, and by inference from Skhul and Qafzeh, Israel, there is no evidence so far that the (strictly) African contemporaries of the Neanderthals buried their dead, but the site sample is relatively small, and, as discussed above, the conditions for grave preservation were generally poorer than in Europe and western Asia. he three modern or near-modern human skulls found isolated on or near the surface at Herto, Ethiopia, may present the best case for ritual in the Neanderthal time range, if it is accepted that they bear cut or scrape marks from postmortem deleshing that was not for consumption and if it can be excluded that the marks were produced by abrasion ater burial. Neanderthal graves present the best case for Neanderthal spirituality or religion, but more prosaically, they may have been dug simply to remove corpses from habitation areas. In sixteen of twenty welldocumented Mousterian graves in Europe and western Asia, the bodies were tightly lexed (in near fetal position), which could imply a burial ritual or simply a desire to dig the smallest possible burial trench. Ritual has been inferred from well-made artifacts or once meaty animal bones found in at least fourteen of thirty-three Mousterian graves for which information is available, but there are no Mousterian burials in which the grave goods difer signiicantly from the artifacts and bones in the

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surrounding deposit, and accidental incorporation in the grave ill is thus possible in virtually all cases. In the lack of truly special or distinctive items, Mousterian graves contrast sharply with many later, Upper Paleolithic burials. Perhaps the best case for ceremonial treatment of Neanderthal bones comes from Grotta Guattari (Monte Circeo), near Rome, Italy, where construction work in 1939 exposed a cave that had been sealed by rock debris for perhaps 50 ky. Inside, on the cave loor, an irregular ring of rocks reportedly surrounded a Neanderthal skull (known as Circeo 1), whose base had been broken away as if to extract the brains. Unfortunately the original position of the skull inside the ring is uncertain since it was moved and replaced before it was irst seen by scientists. In addition, the skull exhibits no evidence of human manipulation, and the only diagnostic antemortem damage is from carnivore teeth. Among other possible evidence for ritual by the Neanderthals are items associated with Neanderthal burials at Teshik-Tash and Shanidar Cave. At Teshik-Tash, the skull of an eight-to-nine-year-old boy in a shallow grave was surrounded by ive or six pairs of mountain goat (ibex) horns. Goat horns were common throughout the Teshik-Tash deposit, however, and no plot of their overall distribution has ever been published to demonstrate that the horn circle stood out. At Shanidar, the grave ill associated with skeleton 4 contained numerous clumps of lower pollen. However, the ill was heavily disturbed by rodent burrows, and the pollen may have been intrusive. he Shanidar lower burial will probably remain problematic so long as it is unique. In sum, there is little or no evidence that the Neanderthals or their near-modern contemporaries practiced ritual or ceremony when they buried their dead. Both kinds of people clearly dug graves, at least sometimes, but the motivation need not have been religious, and the graves tended to be much simpler than those of their fully modern successors. SOURCES: Neanderthal burial pits—absent or undetected at most sites (Gargett 1989, 1999), present in France (Bordes and Laille 1962; Bouysonnie et al. 1913; Bricker 1989; Capitan and Peyrony 1911; Villa 1989) and Israel (Bar-Yosef and Vandermeersch 1991; Bar-Yosef et al. 1992); hyenas at Kebara (BarYosef and Pilbeam 2000); burial to explain nearly complete or articulated Neanderthal skeletons in Europe (Bouyssonie 1954; Deleur 1993; Frayer and Montet-White 1989; Straus 1989b; Vandermeersch 1976), in western Asia (Akazawa et al. 1995; Akazawa and Muhesen 2002; Binford 1968; Deleur 1993; Harrold 1980; Rak et al. 1994; Tillier et al. 1991); rockfalls to explain Neanderthal skeletons at Shanidar (Solecki 1989); accepted number of Neanderthal graves in Europe and western Asia (Pettitt 2002); Skhul and Qafzeh graves (Bar-Yosef and Vandermeersch 1993; Belfer-Cohen and Hovers 1992); boar mandible with Skhul 5 (McCown 1937); fallow deer antlers with Qafzeh 11 (Vandermeersch 1970); Taramsa Hill burial (Vermeersch et al. 1998); cut or scrape marks on the Herto skulls (Clark et al. 2003); possible religious signiicance of Neanderthal graves (Bergounioux 1958; Deleur 1993); lexing of the body and possible Mousterian grave goods (Harrold 1980); contrast between Mousterian and Upper Paleolithic graves (Chase and Dibble 1987; Harrold 1980); context of the Grotta Guattari skull (Blanc 1958; Stringer 1986b) and antemortem damage (Giacobini 1991; White and Toth 1991); TeshikTash grave (Movius 1953a; Weidenreich 1945); lower pollen in place at Shanidar (Solecki 1975, 1989) or intrusive (Chase and Dibble 1987; Sommer 1999)

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Cannibalism he Neanderthals and their contemporaries may have been the irst people to bury the dead, but older burials may be missing because there are so few relevant sites, particularly caves, where graves are most likely to preserve. Even in most of the caves in which the Neanderthals and their contemporaries camped, isolated, fragmentary human remains, scattered among artifacts and animal bones, far outnumber intentional burials. his abundance of fragmentary elements raises the possibility of cannibalism, and if it occurred, it would not be the oldest on record. he previous chapter noted that the oldest known inhabitants of Europe—the people who occupied the Gran Dolina cave about 800 ka—accumulated numerous butchered human bones alongside those of other mammals. he Gran Dolina people may have been reacting to acute dietary stress, and the Neanderthals and their contemporaries surely faced this as well, at least on occasion. Two forms of evidence may imply cannibalism—the sheer abundance of human fragments in a site and, more persuasively, the occurrence of cut-marked or burned fragments. Krapina Shelter, Croatia, is the most famous site for sheer abundance (almost 900 pieces from at least fourteen individual Neanderthals), followed by El Sidrón Cave, northern Spain (about 1,323 highly fragmentary specimens from at least eight individuals), and Hortus Cave, southern France (approximately 100 fragments from perhaps twenty individuals). Moula-Guercy Shelter, southeastern France, is the most notable site for cut-marked human bones, but some have also been reported from Combe-Grenal Cave, southwestern, France, and from Klasies River Main, South Africa. At Moula-Guercy and Combe-Grenal, the humanly damaged bones represent Neanderthals, while at Klasies River Main, they represent modern or near-modern people. he list could be expanded to include three modern or near-modern human skulls from Herto, Ethiopia, if it can be deinitively shown that stone tools account for the marks they bear. If so, the disposition of the marks may suggest postmortem deleshing for purposes other than consumption. Detail is available mainly for Krapina, El Sidrón, and Moula-Guercy. he nearly 900 Neanderthal bones from Krapina represent nearly all parts of the skeleton, and they are mainly broken. Among the fourteen or more individuals represented, the majority were either teenagers or young adults. Nineteen of the bones appear to have been scored by stone tools and some exhibit carnivore tooth marks, but preservatives now coat most bone surfaces, and this impedes any efort to estimate the actual extent of damage from either stone tools or carnivore teeth. Substantial carnivore damage might be expected because the accompanying animal bones come in part from bears, hyenas, or wolves, any of which could have occupied the cave on occasion. Human occupation does not

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appear to have been intense since Mousterian artifacts only slightly outnumber the Neanderthal bones. Still, cannibalism remains a plausible explanation for the numerous human bones and for their high degree of fragmentation. he age bias toward individuals who are least likely to die of natural causes may imply the intentional destruction of one Neanderthal group by another. El Sidrón recalls Krapina in the abundance of Neanderthal bones, and it has likewise produced relatively few artifacts. However, in contrast to Krapina, it has also provided few bones of other mammals. In this respect, its closest analogue might be the much older Atapuerca SH site, discussed in the previous chapter, except that the abundant Atapuerca SH human fossils exhibit no stone-tool cut or percussion marks, while the El Sidrón bones oten do. Cannibalism at El Sidrón might be linked to an unusual degree of developmental stress since the El Sidrón teeth display an extraordinary number of hypoplastic enamel defects, even for Neanderthals. Such defects relect periods of arrested growth, as discussed in the section titled “Pathology” below. he Moula-Guercy evidence for cannibalism is the most direct and compelling. It comes from layer XV, which has provided seventy-eight Neanderthal bones, most of which were conspicuously cut or fractured by people. he same layer has also produced about 300 bones of red deer (Cervus elaphus), which dominates the mammalian sample. he human bones represent at least six individuals ranging in age from six or seven years to mature adult at time of death, while the deer bones represent at least ive individuals from newborn or even fetal to adult. Humanly produced damage is equally apparent on the majority of deer bones, and it tends to occur in the same anatomical positions as on the human bones. he basic pattern suggests that, regardless of species, the butchers used their tools irst to disarticulate bodies and cut away lesh and then to open the skull and long bones for brains and marrow. When the butchers were done, they scattered the human and deer bones equally across the surface of the site. Like the Gran Dolina people about 700 ky earlier, the Moula-Guercy Neanderthals thus consumed people the same way that they consumed other animals. However, from the perspective of a species competing with others, cannibalism is obviously a zero-sum game, and despite the evidence of Moula-Guercy XV, Neanderthals probably rarely ate each other. At most sites where broken and dispersed Neanderthal bones occur, they could alternatively relect carnivore consumption of bodies that were buried in shallow graves or that were let on the surface. Partly empty graves such as those at Teshik-Tash and Kiik-Koba provide circumstantial evidence for carnivore disturbance. he spotted hyena is known to exhume human bodies in Africa today, and spotted hyenas were widespread throughout both Africa and Eurasia in the Neanderthal

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time range. As already noted, hyenas or other carnivores could account for much of the bone fragmentation at Krapina Shelter, and hyena or other carnivore damage has been reported on other human fossils, including on the partial skull from the Florisbad MSA site, South Africa, tentatively on the most complete skull from the Aduma MSA site, Ethiopia, on a let irst metatarsal from Klasies River Main, South Africa, on the limb bones of the Neanderthal child from the Teshik-Tash, Uzbekistan, on the Circeo 1 Neanderthal skull from Grotta Guattari, Italy, and on a Neanderthal femoral shat from Les Rochers-de-Villeneuve Cave, France. SOURCES: cannibalism at the Gran Dolina (Fernández-Jalvo et al. 1999); abundance of Neanderthal bones at Krapina (Malez 1970; Radovcic et al. 1988; Trinkaus 1995a; Wolpof 1979), El Sidrón (LaluezaFox et al. 2005; Rosas et al. 2006), and Hortus (de Lumley 1972; Villa 1992); cut-marked human bones at Moula-Guercy (Deleur et al. 1993, 1999), Combe Grenal (Garralda and Vandermeersch 2000; Le Mort 1989), Klasies River Main (Deacon and Shuurman 1992; White 1987a), and Herto (Clark et al. 2003); Krapina bones with tool marks (Cook 1991) and carnivore tooth marks (Russell 1987a, 1987b); preservative obscuring Krapina marks (White 2001b); abundance of carnivore bones at Krapina (Simek and Smith 1997); carnivore damage on human bones from Florisbad (Clarke 1985; Tappen 1987), Aduma (Haile-Selassie et al. 2004a), Klasies River Main (personal observation), Teshik-Tash (Movius 1953a; Weidenreich 1945), Grotta Guattari (Giacobini 1991; White and Toth 1991), and Les Rochers-de-Villeneuve (Beauval et al. 2005)

Population Numbers here are no practical or theoretical grounds for estimating the absolute population densities of the Neanderthals or their contemporaries, though there is strong evidence that climatic change forced population luctuations. hus the rarity or absence of Mousterian sites in deposits relecting peak cold in northern France and Poland almost certainly relects population shrinkage under adverse conditions, and comparable site paucity during the later Mousterian (Aterian)/MSA in northern and southern Africa suggests a similar population crash, probably owing to extreme aridity during the corresponding (middle) part of the Last Glaciation. Continuing midglacial aridity probably accounts for the rarity of early LSA/Upper Paleolithic sites in both northern and southern Africa. Together with likely preservation bias against older sites, the possible efects of climatic change must be considered before site numbers or densities can be used to suggest that human ability to survive and reproduce—to sustain larger populations—varied through time. Still, including only sites that are reasonably rich in occupational debris and that were probably occupied about the same length of time under broadly similar climatic conditions, it appears that per unit time, Mousterian sites were much less abundant in Europe than succeeding Upper Paleolithic sites were. In southwestern France and neighboring northeastern Spain, there is probably less than one Mousterian cave for every ive Upper Paleolithic ones. In addition, Mousterian caves tend

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Cape turban shell (Turbo sarmaticus) operculum length (mm)

LSA (12 - 0.4 ka)

12.5

25.0

37.5

50.0

12.5

25.0

37.5

50.0

Die Kelders 1 (2308) Blombos Cave (198) Klasies River (37) Soetfontein 1 (305) Byneskranskop 1 1-4 (703) Noetzie (967) Byneskranskop 5-12 (1381) Nelson Bay Cave J-RA (49) Byneskranskop 11-19 (7) Nelson Bay Cave GBSL (80)

MSA (ca. 127 - ca. 71 ka) Klasies River MSA III/IV (11) Klasies River HP (346) Klasies River MSA II (69) Klasies River MSA I (15) Blombos Cave MSA 1 (187) Blombos Cave MSA 2 (272) Blombos Cave MSA 3 (463)

to be much richer in bones of bears, hyenas, or other large carnivores that probably denned in the sites when people were absent, and they imply that Mousterian occupation was relatively light. he bottom line is that the Mousterians appear to have been less numerous than their successors, probably because they exploited the available animal and plant resources less eiciently. In addition, the broad, thick trunks and short legs of the Neanderthals, which were important for heat retention, also increased individual energy requirements, irst to maintain the large body mass centered in the trunk and second to forage for food, based on the observation that living humans with shorter legs use more energy to walk. In brief, Neanderthals almost certainly needed more calories per individual than their successors and a set number of calories would have supported fewer individuals. hus even if Neanderthals extracted as much energy from nature as their successors, their populations would have been smaller.

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577 FIGURE 6.53. box plots summarizing the maximum length of Cape turban shell opercula from successive msa units at Klasies river main and blombos Cave and from lsa units at nelson bay Cave, byneskranskop Cave 1, soetfontein 1, noetzie, Klasies river main, blombos Cave, and die Kelders Cave 1. The sites are all on the south coast of south africa. The number in parentheses after each sample name is the number of specimens in the sample. The legend to igure 6.52 explains the box plot format. The most essential elements are the median, indicated by the vertical line near the center of each plot; the 95% conidence limits for the median, indicated by the shaded rectangle around each median; and the number of measured opercula, shown in parentheses. medians whose 95% conidence limits do not overlap are distinct in the conventional statistical sense. at blombos Cave, the msa 1 (still bay), msa 2, or both have provided widely publicized bone artifacts, putative shell beads, and incised ocher. The associated turban shells are large, implying that msa 1 and msa 2 people pressed relatively lightly on the species. This suggests that, despite the precocious artifacts, msa 1 and msa 2 populations were relatively small and that the blombos artifacts provided no special itness (survival and reproduction) advantage.

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maximum shell length (mm)

granite limpet Patella (Cymbula) granatina

10 min & fossil beach

45

60

75

90

45

60

75

90

Bekbaai (27) Cape St. Martin (38) Noordwesbaai (35) SBF “Eemian” beach (435)

LSA (11 - 0.7 ka) Bakoond 1/1 (13) Bakoond 1/2 (209) Bakoond 1/3 (156) Bakoond 1/4 (102) Bakoond 1/5(197) Bakoond 1/6 (279) Bakoond 1/7 (191) Bakoond 1/8 (48) Steenbokfontein 4a (46) Steenbokfontein 4b (297) Elands Bay Cave D (255)

MSA (ca. 127 - ca. 71 ka) Boegoeberg 2 (199) Hoedjies Punt 3 (29) Sea Harvest (114) Ysterfontein 1/1 (21) Ysterfontein 1/2 (22) Ysterfontein 1/3 (53) Ysterfontein 1/4 (46) Ysterfontein 1/5 (27) Ysterfontein 1/6 (40) Ysterfontein 1/7 (82) Ysterfontein 1/8 (5) Ysterfontein 1/9 (33) Ysterfontein 1/10 (43) Ysterfontein 1/11 (20) Ysterfontein 1/12 (24) Ysterfontein 1/13 (42) FIGURE 6.54. box plots summarizing the maximum length of granite limpets from the msa deposits at boegoeberg 2, hoedjiespunt 3, sea harvest, and Ysterfontein 1; from the lsa deposits at elands bay Cave, steenbokfontein, and bakoond; from a probable last interglacial (“eemian”) fossil beach at steenbokfontein; and from three recently collected “ten-minute samples” at bekbaai, Cape st. martin, and noordwesbaai. all the localities are on the west coast of south africa. The number in parentheses after each sample name is the number of specimens in the sample. The legend to igure 6.52 explains the boxplot format. The most essential elements are the median, indicated by the vertical line near the center of each plot; the 95% conidence limits for the median, indicated by the shaded rectangle around each median; and the number of measured humeri, shown in parentheses. medians whose 95% conidence limits do not overlap are distinct in the conventional statistical sense. The modern samples were collected by students at ten-minute intervals, from intertidal rocks that are not being exploited today (buchanan et al. 1978), and they show how large granite limpets can grow. The fossil beach sample documents average (as opposed to maximum) granite limpet size long before the lsa. The limpets in the modern and fossil beach samples are remarkably large, but they are particularly large relative to lsa specimens, and they imply that lsa limpet collection was especially intense. The msa/lsa contrast in average size suggests that msa people collected much less intensively, probably because their populations were smaller.

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angulate tortoise (Chersina angulata)

LSA (13 - 0.6 ka)

mediolateral diameter of the distal humerus (mm) 5.0

7.5

10.0

Die Kelders 1 LSA (932) Blombos Cave LSA (32) Byneskranskop Cave 1 1–4 (693) Byneskranskop Cave 1 5–12 (1366) Byneskranskop Cave 1 13–17 (655) Byneskranskop Cave 1 18–19 (115)

MSA (ca. 115 - ca. 65 ka) Die Kelders 1 MSA 4/5 (1728) Die Kelders 1 MSA 6 (2433) Die Kelders 1 MSA 7–8 (65) Die Kelders 1 MSA 9–15 (412) Blombos Cave MSA 1 (254) Blombos Cave MSA 2 (299) Blombos Cave MSA 3 (296)

Smaller site numbers could also be used to infer that African Mousterian/MSA populations were smaller than those of succeeding Upper Paleolithic/LSA people, but the data are more problematic because the total site sample is small and because the impact of culture on site occupation is diicult to separate from the impact of recurrent aridity. However, in southern Africa smaller MSA populations are probably implied by the tendency for slow-growing shellish species and tortoises to be signiicantly larger in MSA sites than in LSA sites occupied under broadly similar climatic conditions. Both shellish and tortoises grow more or less continuously through life, and larger average size in MSA sites implies more limited human predation pressure, probably because MSA populations were generally smaller. Figures 6.53–6.56 show that the MSA/LSA contrast in shellish and tortoise size characterizes both the southern and western coasts of South Africa, and the southern coast contrast is perhaps particularly signiicant. his is because the MSA shellish and tortoises

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579 FIGURE 6.55. box plots summarizing the mediolateral diameter (“breadth”) of angulate tortoise distal humeri from the msa and lsa layers of die Kelders Cave 1 and blombos Cave and from the lsa layers of byneksranskop Cave 1. all three sites are on the south coast of south africa. The legend to igure 6.52 explains the box plot format. The most essential elements are the median, indicated by the vertical line near the center of each plot; the 95% conidence limits for the median, indicated by the shaded rectangle around each median; and the number of measured humeri, shown in parentheses. medians whose 95% conidence limits do not overlap are distinct in the conventional statistical sense. The site list is different from the one for south coast turban shells in igure 6.53 because Klasies river main, nelson bay Cave, noetzie, and soetfontein 1 (included in that igure) have provided few tortoise bones and because the msa deposits of die Kelders 1 (only in this igure) preserves shells poorly. The plots in the igure show that msa tortoises tend to be signiicantly larger on average than lsa specimens. The difference suggests that msa people collected tortoises less intensively, probably because msa human populations were generally smaller.

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i

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

angulate tortoise (Chersina angulata)

mediolateral diameter of the distal humerus (mm)

5.0

7.5

10.0

12.5

LSA (12.5 - 0.7 ka) Bakoond (17) Kasteelberg B (2175) Elands Bay and Tortoise Caves 1 (147) Elands Bay and Tortoise Caves 2 (53) Elands Bay and Tortoise Caves 3 (303) Steenbokfontein 4–5 (231) Elands Bay Cave 4 (1683) Elands Bay Cave 5 (3453)

MSA (ca. 127 - ca. 65 ka) Diepkloof Rockshelter 1 (295) Diepkloof Rockshelter 2 (224) Diepkloof Rockshelter 3 (258) Diepkloof Rockshelter 4 (132) Diepkloof Rockshelter 5 (89) Diepkloof Rockshelter 6 (54) Hoedjies Punt 1 (26) Ysterfontein 1 1+2 (16) Ysterfontein 1 3+4 (60) Ysterfontein 1 5-13 (42) FIGURE 6.56. box plots summarizing the mediolateral diameter (“breadth”) of angulate tortoise distal humeri from the middle stone age (msa) layers at diepkloof rock shelter, hoedjies Punt 1, and Ysterfontein 1 and from the later stone age (lsa) layers of elands bay Cave, Tortoise Cave, steenbokfontein Cave, Kasteelberg b, and bakoond. all six sites are on or near the west coast of south africa (references in table 6.12). The lsa layers are presented in approximate chronological order. The chronological relationships among the three msa sites are uncertain. They could overlap in time, and the order in the igure is for presentation purposes only. The legend to igure 6.52 explains the box plot format. The most essential elements are the median, indicated by the vertical line near the center of each plot; the 95% conidence limits for the median, indicated by the shaded rectangle around each median; and the number of measured humeri, shown in parentheses. medians whose 95% conidence limits do not overlap are distinct in the conventional statistical sense. The msa site list is different from the ones for west coast granite limpets in igure 6.54 because sea harvest (presented there) has provided few tortoise bones, the tortoise bones from boegoeberg 2 (also presented there) come largely from other species, and diepkloof (presented here) has provided no limpets. The plots in the igure show that msa tortoises tend to be signiicantly larger on average than lsa specimens. The difference suggests that msa people collected tortoises less intensively, probably because msa human populations were generally smaller.

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include those from Blombos Cave, where, as noted above, the artifacts are said to imply LSA-like behavior. he large Blombos MSA mollusks and tortoises suggest that, whatever the cognitive or cultural implications of the artifacts, they do not appear to have conferred a itness (reproductive and survival) advantage. his point is crucial, for evolution is about itness, and if the Blombos artifacts did not enhance itness, they relect only history not evolution. heir failure to enhance itness may explain why they do not appear to have spread from Blombos Cave and why they did not prompt an Out-of-Africa expansion. SOURCES: rarity or absence of Mousterian sites during peak cold in northern France (Tufreau 1992) and Poland (Féblot-Augustins 1993); Mousterian and Upper Paleolithic site numbers in southwestern France and northeastern Spain (Clark and Straus 1983; Mellars 1973, 1982; Straus 1977); Neanderthal energy requirements (Sorensen and Leonard 2001; Weaver and Steudel-Numbers 2005); MSA and LSA shellish and tortoises (Steele and Klein 2005–2006)

Life History Living humans are distinguished from the apes by a signiicantly slower life history, in which individuals reach sexual maturity at a much later age and their life expectancy at maturity is much longer. Observations outlined in the previous chapters suggest that the australopiths and early Homo, including H. ergaster, had faster, more apelike life histories. Among the questions that arise from this, two are pertinent here: irst, when in later human evolution did fully modern life history emerge and, second, was there a diference in life history between the Neanderthals and their early (or near-) modern human contemporaries? Dentitions are the most useful skeletal elements for establishing life history since the timing and pace of dental development are closely linked to the rate of overall maturation. Across the primates, for example, the brain reaches 90% of growth at about the time the irst molar (M1) erupts and the skeleton reaches maturity at about the same time that the third molar (M3) erupts. A mandible of early (or near-) modern Homo sapiens from Jebel Irhoud, Morocco, is so far the oldest hominin fossil to imply the modern, slow rate of maturation. he mandible is known as Irhoud 3, and composite U-series/ESR analysis suggests it dates from about 160 ka. he individual possessed fully erupted irst molars, lateral incisors that were nearly fully erupted, and fully developed, but unerupted permanent premolar and second molar crowns on which the roots were just beginning to form. he same eruption schedule characterizes living humans, for whom the eruption state of Irhoud 3 would imply an age between seven and eight years. he use of noninvasive x-ray synchrotron microtomography to produce high-resolution images of Irhoud 3 dental enamel revealed daily growth increments and showed that longer period

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increments (perikymata) formed every ten days. Enamel formation time can be estimated by multiplying the number of perikymata by their periodicity, and the direct observation of periodicity in this instance allowed unusually precise estimates of crown formation times. he estimates indicate that the Irhoud 3 teeth developed slowly, at roughly the modern rate. Since the schedule of dental eruption and the time of crown formation both resembled those in living humans, it seems reasonable to conclude that the timing of dental eruption also resembled that in living humans and by extension, that the rate of overall maturation was similar. So far, analyses of dental development in Neanderthals do not include direct observation of perikymata periodicity, and studies designed to estimate the rate of dental crown development conlict. Some analyses suggest that crown formation was faster in Neanderthals than in modern humans, while others suggest that the rate was similar between the two or, perhaps more precisely, that they overlapped in rate variability. If Neanderthals did develop their dental crowns more rapidly, this would further underscore their evolutionary divergence from modern humans, and it would imply a signiicant diference in life history that might help explain why only modern humans survived the contact experience. However, in advance, similarities in body and brain mass suggest that Neanderthal and modern human maturation rates were probably similar, and pending irm evidence to the contrary, this is the tentative assumption here. Since the Neanderthal sample is relatively large and includes both sexes and children of various ages, it might seem feasible to establish the fundamental pattern of Neanderthal mortality. here is the complication, however, that the sample is a composite of multiple subsamples. In the terminology used to describe large ungulate mortality in the section on Mousterian/MSA ecology above, the various subsamples could relect “attritional” mortality, “catastrophic” mortality, or some mix of the two. here is also the likelihood of pre- and postdepositional bias, particularly against younger individuals, whose bones are much less durable than those of adults and are therefore far more likely to have been fragmented or destroyed by carnivore feeding, proile compaction, and leaching. In short, the available sample probably cannot reveal the basic, time-averaged pattern of everyday (attritional) mortality among Neanderthals. It might, however, show the extent to which mortality was similar between the Neanderthals and their Upper Paleolithic (CroMagnon) successors, assuming the Upper Paleolithic sample was biased in roughly the same way. he Neanderthals closely resembled living humans in their relatively small birth canals (pelvic inlets) and large adult brains, which

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implies a similar amount of postnatal brain growth. In the absence of contrary evidence, this further supports the prediction that Neanderthals and modern humans shared equally prolonged childhoods. If so, as noted previously, they probably also shared equally prolonged lifespans since age at sexual maturity and longevity are closely correlated across the mammals. However, skeletal age determinations have so far failed to reveal any Neanderthals who lived beyond their mid-forties, and wear seriation of dentitions suggests that compared to their early Upper Paleolithic successors, Neanderthals died much more oten in young adulthood. As discussed in the last chapter, adults in the wear seriation study were deined as individuals who had erupted their third molars (M3s) since M3 eruption coincides roughly with age at sexual maturity in living humans. Younger adults were deined as ones whose M3 wear indicated they were no more than twice as old as the age at which M3 erupted. Assuming an M3 eruption age around iteen or sixteen, they could thus have been parents but probably not grandparents. Arguably, tooth wear and the other skeletal indicators that have been used to age individual Neanderthals have underestimated the ages of the oldest, just as the same indicators oten do for recent people of known age. Problems in scoring dental wear, the exact composition of the Neanderthal and early Upper Paleolithic samples, and a possible shit in the mix of factors that caused death could explain why young adult mortality seems to have been much higher among the Neanderthals than among their Upper Paleolithic successors. he proposed elevated rate of young adult Neanderthal mortality is puzzling since it would have signiicantly exceeded the rate in historic hunter-gatherers, in Japanese macaque monkeys, and in chimpanzees, as summarized in the last chapter. In sum then, pending the accumulation of contrary observations, it seems reasonable to hypothesize that childhood and lifespan were extended in Neanderthals more or less as they are in living humans. Future research may show that Neanderthals and living humans inherited their shared life history from Homo heidelbergensis, their last shared ancestor at 600–500 ka. As discussed in the previous chapter, the natural selective factors that favored slow maturation may have included a longer interval over which children could internalize culture, the value of older adults as cultural repositories, or the potential for older women to enhance their reproductive itness more by provisioning their daughters’ or nieces’ ofspring than by bearing additional young of their own. SOURCES: dental eruption and enamel microstructure as indications of overall maturation rate (Dean 2006); Irhoud 3 mandible (Dean 2007; Smith et al. 2007a); rate of dental crown formation in Neanderthals—similar to that in modern humans (Guatelli-Steinberg et al. 2005, 2007; Macchiarelli et al. 2006) or more rapid (Ramirez Rozzi and Bermúdez de Castro 2004; Ramirez Rozzi and Sardi 2007; Smith et al. 2007b); age determination of individual Neanderthals (Trinkaus 1986,1995a; Trinkaus and hompson 1987); Neanderthal vs. Upper Paleolithic mortality (Caspari and Lee 2004); tendency for

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skeletal indicators to underestimate known ages (Miles 2001); possible natural selective advantages of modern human life history (Hawkes et al. 1998; Kennedy 2003)

Pathology If for the sake of argument, we were to accept that mortality was remarkably high among Neanderthal young adults, Neanderthal skeletal pathology and antemortem trauma could explain why. Both imply imply exceptional physical stress. hus, in a sample of 669 carefully examined teeth from roughly 165 Neanderthals, hypoplastic enamel defects (pitting or grooving indicating periods of arrested enamel growth) appeared on 36%, representing 57% of the individuals. he frequency of hypoplastic defects can reach this level in recent human foragers, but it was much lower in Upper Paleolithic people. he implication is that young Neanderthals were much more commonly subjected to food shortage, trauma, or disease. In addition, the skeletons of older Neanderthals oten exhibit healed fractures of the skull or limbs, degeneration of the joints, advanced osteoarthritis of the vertebral column, and periodontal disease. he most famous case is certainly the “old man” of La Chapelle-aux-Saints, who at an estimated age of forty (or less) sufered from severe periodontal disease that caused substantial antemortem tooth loss and from joint degeneration or arthritis that are manifest in the articulation of the mandible to the skull and in the spinal column, hip, and foot. Other Neanderthals with obvious pathologies or injuries include Kebara 2, Tabun 1, Shanidar individuals 1 and 3–5, La Ferrassie 1 and 2, La Quina 5, the original (Feldhofer) Neanderthal from Germany, as many as ive of the Krapina Cave Neanderthals, and the Sala 1 individual (Czech Republic). Kebara 2 had healed fractures of the ith thoracic vertebra and the let second metacarpal. Shanidar 1 was alicted by numerous partly interrelated traumatic and degenerative lesions, including especially a crushing fracture of the let orbit and cheekbone (ig. 6.57), a withered right upper arm from which the forearm had been lost before death, and degenerative or posttraumatic deformities of both legs that probably caused a limp. Shanidar 3 sufered from debilitating arthritis of both the right ankle and adjacent foot joints and probably died of a stab wound that pierced the lung, leaving an ugly scar on the let ninth rib. Shanidar 4 had a healed rib fracture (also seen in the individual from La Chapelleaux-Saints), and Shanidar 5 had a large scar, caused by a sharp blow to the head, on the let frontal. One of the Krapina Neanderthals had a broadly similar scar on one parietal. Like other aged Neanderthals, La Ferrassie 1 endured periodontal disease. He also had a healed fracture on the right femur and bony lesions on both femurs, tibiae, and ibulae that relect either systemic infection or carcinoma. Tabun 1 and La Ferrassie 2 had damaged ibulae. La Quina 5 sufered a withered let arm. he origi-

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0

5 cm

585 FIGURE 6.57. face of shanidar 1, showing the crushed outer margin of the left orbit (drawn by Kathryn Cruz-uribe from a photograph). The crushing was not fatal and the orbit healed, but associated brain damage may have partially paralyzed the man’s right side, accounting for a withered right arm. like other injured or aged neanderthals, he survived only with the help of other members of his group.

Shanidar 1

nal Neanderthal individual and one of the Krapina Neanderthals had injured ulnae that deformed their forearms. One or more of the Krapina people had a broken clavicle (collarbone) and a concussed parietal. he Sala 1 person had a healed lesion on the right supraorbital torus. Antemortem trauma has also been observed in adult skeletons of the earliest fully modern people, but much less frequently. In fact, in the frequency of trauma and in its anatomical patterning, the Neanderthals closely recall modern rodeo riders, although the activities that produced the Neanderthal injuries were surely more mundane. he most fundamental implication is that Neanderthal technology (or culture in general) was relatively inefective at reducing wear and tear on Neanderthal bodies. However, the same skeletal pathologies and injuries that show the Neanderthals lived risky lives and aged early also reveal a strikingly human feature of their social life. he La Chapelle-aux-Saints and Shanidar 1 individuals, for example, must have been severely incapacitated and might have died even earlier without substantial help and care from their comrades. If this is agreed, group concern for the old and sick may have permitted Neanderthals to live longer than any of their predecessors, and it is the most recognizably human, nonmaterial aspect of their behavior that can be reasonably inferred from the archaeological record. SOURCES: evidence of physical stress in Neanderthal skeletons (Berger and Trinkaus 1995; Trinkaus 1995a); hypoplastic enamel defects in Neanderthals (Ogilvie et al. 1989; Skinner 1996), recent human foragers (Guatelli-Steinberg et al. 2004), and Upper Paleolithic people (Brennan 1991); hypoplastic enamel defects as evidence for stress on young Neanderthals (Neiburger et al. 1990); skeletal pathologies in La Chapelle-aux-Saints (Dawson and Trinkaus 1997; Trinkaus 1985) and other Neanderthals (Berger and Trinkaus 1995; Trinkaus 1978, 1983a); skeletal trauma in Shanidar 1 (Crubézy and Trinkaus 1992) and Shanidar 3 (Franciscus and Churchill 2002; Stewart 1977); systemic infection or carcinoma in La Ferrassie 1 (Fennell and Trinkaus 1997); lower frequency of trauma in earliest fully modern humans (Trinkaus 1989b); pattern of trauma in rodeo riders (Berger and Trinkaus 1995)

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Conclusion: The Fate of the Neanderthals It is not diicult to understand why the Neanderthals failed to survive. he archaeological record shows that in virtually every detectable aspect—artifacts, site modiication, ability to adapt to extreme environments, subsistence, and so forth—the Neanderthals lagged their modern successors, and their more primitive behavior limited their ability to compete for game and other shared resources. To judge from their distinctive morphology and their genetic diference from living humans, their more primitive behavior may have been rooted in their biological makeup. It is more diicult to address how the Neanderthals succumbed. One particularly intriguing question is whether they exchanged genes and culture with their successors. Another is whether they vanished rapidly ater contact or whether they held on for many millennia, perhaps especially in southern Europe. he next chapter summarizes the genetic evidence for modern human origins in Africa, but in advance, it can be said that there is little to suggest a signiicant Neanderthal genetic contribution to subsequent, modern Europeans. he genetics of living humans and the extraction of DNA from Neanderthal bones ever more strongly argue against signiicant gene exchange. he genetic observations do not preclude fossil evidence for hybridization between Neanderthals and their immediate successors. hey also do not rule out cultural exchange, and they bear on the timing of Neanderthal extinction only insofar as prolonged overlap with modern humans is more likely to have let bony or genetic traces of hybridization. he fossil and cultural (archaeological) evidence for Neanderthal contact with their successors and the timing of Neanderthal disappearance are best discussed region by region. Figure 6.58 encapsulates some of the key time-space relationships underlying the following discussion.

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Western Europe Antiquity of the Latest Mousterian and Earliest Upper Paleolithic. Assum-

ing that only Neanderthals produced Mousterian artifacts in western Europe, Neanderthal persistence to at least 40 ka is conirmed by TL dates from Le Moustier in France and U-series dates from Abric Romaní in Spain. At each site, the dates occur in expected stratigraphic order within a deep sequence, which enhances the likelihood that they are valid. he geologic antiquity of the oldest anatomically modern human fossils in western Europe is not well-ixed, but early Aurignacian (early Upper Paleolithic) artifact associations in France suggest ages between 40 and 30 ka for a fully modern frontal bone and fragmentary juvenile maxilla from La Crouzade, two fully modern juvenile mandibles from La Grotte des Rois, and nine diagnostic isolated adult teeth from Brassempouy. Late Aurignacian artifact associations imply a date near 30 ka for ive-to-eight much more complete early-modern skeletons from the

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1

EASTERN ASIA

AFRICA

3

50

20 Aurignacian & modern H. sapiens

Ahmarian & modern H. sapiens

core/scraper industry & modern H. sapiens

Châtelperronian & H. neanderthalensis

50

4

90

5b

100

5c 5d

70 Mousterian & H. neaderthalensis

Last Interglacial

5a

6

flake/chopper industry & evolved H. erectus

Howieson’s Poort, Aterian & nearmodern H. sapiens

80

100 ??? Mousterian & H. neaderthalensis

5e

130

Mousterian & H. neaderthalensis

90

Penultimatate Glaciation

120

40

60

80

110

FIGURE 6.58. approximate chronological arrangement of various late Pleistocene cultural units and fossil humans types in africa and eurasia (modiied after Klein [1992], 6).

30 LSA/Upper Paleolithic & early modern H. sapiens

60 70

ka 10

2

30 40

EUROPE

WESTERN ASIA

Recent

10

Last Glaciation

ka

O-isotope stages & climate stratigraphy

587

MSA/Mousterian & near-modern H. sapiens

Mousterian & near-modern H. sapiens

110 120 130

???

190

190

famous Cro-Magnon Rockshelter, also France, and radiocarbon dating of a possibly associated marine shell shows the skeletons are at least 28 ky old. If early Aurignacian artifacts signal modern human presence in Europe even when human fossils are absent, then radiocarbon dates on early Aurignacian layers at Willendorf II in Austria, at Geissenklösterle Cave in southern Germany, at El Castillo, L’Arbreda, Romaní, and Reclau Viver in northern Spain, and possibly at Trou Magrite in Belgium, imply that modern humans reached Europe as much as 40 ka. At Geissenklösterle Cave, thermoluminescence dates on burned lints broadly support the radiocarbon ages. However, the time when the Aurignacian penetrated central and western Europe depends both on the validity of stratigraphic associations between 14C dates and Aurignacian artifacts and on the accuracy of the dates, and experts can disagree on both matters. A conservative estimate now is that the Aurignacian had penetrated central and western Europe by 37–36 ka (uncalibrated radiocarbon years ago). here may have been two separate early Aurignacian entry routes—one through

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the Danube Valley marked by classic early Aurignacian antler and bone split-base points, and the other along the Mediterranean coast marked mainly by small, abruptly retouched blades (bladelets). his possibility is addressed again below and in the next chapter. No sites with early Aurignacian artifacts have so far been found south of the Ebro and Tagus Rivers in southern Iberia (Gibraltar, Spain and Portugal), and a small number of radiocarbon determinations from ive-to-six sites suggest that Mousterian artifact makers survived locally until perhaps 30 ka. he most notable site is Zafarraya Cave, near Malaga, southern Spain, where the Mousterian artifacts accompany a Neanderthal mandible and other fragmentary Neanderthal bones. Radiocarbon dating of the organic residue in associated Zafarraya animal bones and uranium (U)-series disequilibrium dating of associated animal teeth suggest the Neanderthal fossils accumulated between 35 and 30 ka. Together with the other sites, Zafarraya could place Neanderthals south of the Ebro and Tagus for 4–7 ky ater they had disappeared to the north. Radiocarbon dates at Gorham’s Cave, Gibraltar, might imply even later Neanderthal persistence, to 28 or possibly 24 ka. At 28–24 ka, Neanderthals would have overlapped with unquestionably modern later Aurignacian people, who penetrated southern Spain by 32 ka. he Gorham’s Cave dates are problematic, however, partly because they oten depart from stratigraphic order (they don’t get progressively older with depth) and, perhaps more important, because they bear on sparse artifacts, not on Neanderthal bones. he artifacts include no diagnostic Upper Paleolithic types, but they comprise mostly nondescript laked pieces that either Mousterians or early Upper Paleolithic people could have produced, and the artifact sample that postdates 30 ka may lack diagnostic Upper Paleolithic types by chance alone. Excavations elsewhere in the cave suggest that the Upper Paleolithic was probably present by 30 ka, but that the Mousterian may have persisted locally until then. However, it is also possible that all Mousterian radiocarbon dates near 30 ka in southern Iberia are minima and that the Mousterian ended everywhere by 37 ka. his is because only a minute amount of undetectable recent carbon contamination is suicient to make an object that is actually 37 ky old appear 5–7 ky younger ater radiocarbon assay. Such contamination is commonplace and it most plausibly explains, for example, the inconsistencies or stratigraphic inversions that characterize even the most carefully obtained 14C dates at French early Upper Paleolithic sites. Contamination is especially likely to afect dates based on bone, and the late Mousterian dates from southern Iberia are mainly on bone or on even more problematic substances like carbonaceous sediment. he Zafarraya U-series dates do not truly corroborate the 14C readings since they are on teeth, which are among the least suitable materials for the method, and the dates are somewhat younger than the correspond-

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ing 14C determinations. Both theory and paired 14C and U-series dates on more suitable materials elsewhere suggest that U-series dates should be 3–5 ky older in the 30-ky time range. he reason (discussed in chapter 2) is that uncontaminated U-series readings provide true calendar (solar) ages, whereas 14C ages systematically tend to underestimate them. If in fact the Mousterian actually ended in southern and western Iberia about 37 ka, the local absence of the basal Aurignacian could mainly relect adverse climatic conditions that restricted human occupation from 40 ka or before until 30 ka or later. In nearby northwestern Africa, Mousterian/Aterian (or Middle Stone Age) occupations that antedate 40 ka are commonly separated from Upper Paleolithic (Iberomaurusian, or Oranian) occupations dated to 22–21 ka, and the hiatus may relect extreme aridity in lower midlatitudes during the middle of the Last Glaciation. SOURCES: dates conirming Neanderthal persistence to 40 ka at Le Moustier (Valladas et al. 1986) and Abric Romani (Bischof et al. 1988, 1994); early Aurignacian human fossils from France (Bailey and Hublin 2005; Gambier 1989; Hublin 2000); dating of Cro-Magnon skeletons based on artifact associations (Gambier 1997; Movius 1969a; Petit-Maire et al. 1971) and radiocarbon on associated marine shell (Henry-Gambier 2002); dating of early Aurignacian at Willendorf II (Damblon and Haesaerts 1997), Geissenklösterle (Conard and Bollig 2003; Richter et al. 2000), El Castillo, L’Arbreda, Romaní, and Reclau Viver (Bischof et al. 1989, 1994; Straus 1997), and Trou Magrite (Otte and Straus 1995; Straus 1993–1994); dating of earliest Aurignacian in central and western Europe (Mellars 2006a; Zilhão 2006; Zilhão and d’Errico 1999); early Aurignacian entry routes to Europe (Mellars 2004, 2005, 2006b); survival of Mousterian artifact makers to 30 ka in southern Iberia (d’Errico et al. 1998b; Hublin et al. 1995; Straus 1997); Mousterian at Gorham’s Cave dated to 28–24 ka (Delson and Harvati 2006; Finlayson et al. 2006) or only 31–30 ka (Barton et al. 1999; Pettitt and Bailey 2000); inconsistent dates for the French early Upper Paleolithic (Gowlett and Hedges 1986; Mellars et al. 1987); contamination and 14C dates on bone (Damblon and Haesaerts 1997; Kuzmin and Tankersley 1996; Staford et al. 1987); dating of the Mousterian and Upper Paleolithic in northwest Africa (Camps 1975; Close 2002; Close and Wendorf 1990; Cremaschi et al. 1998; Wendorf 1992)

Skeletal Evidence for Neanderthal/Modern Contact. he absolute age of

the latest Mousterian and early Upper Paleolithic aside, west European cave sites oten contain both Neanderthal/Mousterian and modern human/Upper Paleolithic occupations, and the Upper Paleolithic layers usually overlie Mousterian ones with no evidence either for population contact or for a substantial gap in time. his argues for a rapid replacement with little or no interaction. However, an Upper Paleolithic layer at the Lagar Velho rockshelter, Portugal, provided a child’s skeleton (“LV1”) that is said to demonstrate Neanderthal/modern human hybridization, and occasional archaeological sites contain layers where a blend of Mousterian and Upper Paleolithic types could imply Mousterian/Upper Paleolithic cultural contact. Bulldozing badly damaged the Lagar Velho rockshelter before its signiicance was fully appreciated, but rescue excavations showed that the child’s skeleton lay in a grave and that it was associated with artifacts that typify the Gravettian Culture, which succeeded the Aurignacian Culture about 28 ka. Direct radiocarbon dating of charcoal from

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immediately below and of animal bones from immediately above bracket the skeleton between 24.9 and 23 ka. Except for the cranium, which the bulldozing had shattered, the skeleton was nearly complete. he state of dental eruption showed that the child died at age four or ive, and in virtually all morphological respects, its bones closely resembled those of a modern four-to-ive-year-old. However, they are also said to reveal some possible Neanderthal traits, including the backward slope of the mandible below the lower incisors and especially the shortness of the tibia relative to the femur. It will be recalled that short tibias typify the Neanderthals, and they suggest that the Neanderthals were morphologically adapted to cold climate. he short tibia and other less compelling Neanderthal-like features may imply partial Neanderthal ancestry for the Lagar Velho child. However, the skeleton is overwhelmingly modern with no derived, classic Neanderthal traits, and there is the problem that only a irst- or secondgeneration hybrid would be expected to show a clear mix of modern and Neanderthal characteristics. he Lagar Velho child was conceived at least 200 generations ater the last Neanderthals occupied Portugal, and there is a strong likelihood that it is “simply a chunky Gravettian child, a descendant of the modern invaders who had evicted the Neanderthals from Iberia several millennia earlier” (Tattersall and Schwartz 1999, 7119). he case for hybridization would be resuscitated, however, if the 28–24 ka Mousterian dates at Gorham’s Cave are corroborated, if skeletons from freshly excavated Gravettian sites in Portugal or southern Spain exhibit the same or other supposed Neanderthal features, or most directly if one or more such skeletons provides Neanderthal mitochondrial or nuclear DNA. As discussed in the next chapter, this is a reasonable expectation if hybridization occurred. Unfortunately, the Lagar Velho child’s bones lack residual protein (collagen), and they are thus unlikely to retain endogenous DNA that might close the issue now. SOURCES: Lagar Velho—dating and skeleton as a hybrid (Duarte et al. 1999; Zilhão 2001b; Zilhão and Trinkaus 2002), skeleton as fully modern (Hublin 2000; Tattersall and Schwartz 1999), similarity of the grave and associated artifacts to Gravettian graves dated to roughly the same time (Pettitt et al. 2000)

Artifactual Evidence for Mousterian/Upper Paleolithic Contact. he most

compelling artifact assemblages that suggest Mousterian/Upper Paleolithic contact are from a restricted area of northern Spain and western and central France (west of the Rhône River) where they have been assigned to the Châtelperronian Industry or Culture (ig. 6.59). In deeply stratiied, carefully excavated sites, Châtelperronian layers rest conformably on Middle Paleolithic layers narrowly understood and below early Upper Paleolithic (basal Aurignacian) horizons. Dates on the Châtelperronian and early Upper Paleolithic overlap signiicantly, and the time

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diference between the two may have been too short to measure with current dating technology. From the small sample of available dates and from site stratigraphies, a reasonable inference is that the Châtelperronian began in Spain and France about 45 ka and that it persisted until perhaps 36 ka, when the Aurignacian had already appeared nearby. A juvenile temporal fragment and twenty-nine isolated teeth from Arcysur-Cure (Grotte du Renne) and a partial skeleton from Saint-Césaire (la Roche á Pierrot Rockshelter) indicate that Châtelperronian people were Neanderthals. 14C dates on the organic extract from animal bones suggest that the Châtelperronian at Arcy could have extended to 34 ka, while TL dates on burned lint artifacts at Saint-Césaire place the skeleton between 39 and 33 ka. he Neanderthals at both sites were probably among the last in France. If only stone artifacts were involved, the Châtelperronian might be considered simply a inal Middle Paleolithic stemming from the local Mousterian of Acheulean Tradition Type B (with numerous backed knives), and the earlier part of the Châtelperronian, before 38–37 ka, may have been just that. In support, the Châtelperronian and the preceding Mousterian of Acheulean Tradition B have almost identical geographic distributions. At Arcy, however, Châtelperronian people not only produced a mix of typical Middle and Upper Paleolithic stone-artifact types, they also manufactured quintessential Upper Paleolithic bone tools and

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FIGURE 6.59. The distribution of the aurignacian, Châtelperronian, uluzzian, and szeletian/Jerzmanowician industries in europe and western asia roughly 37 ka (redrawn after mellars [1996], ig. 13.10). The aurignacian represents the earliest undisputed upper Paleolithic industry in central and western europe. it may have coexisted briely with the Châtelperronian, uluzzian, and szeletian/ Jerzmanowician industries before replacing them by about 36 ka. it also appeared in southwestern asia 37–36 ka where it replaced an earlier kind of upper Paleolithic. its place of origin remains uncertain, but southeastern europe or anatolia are possibilities.

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personal ornaments (ig. 6.60). he Châtelperronian layers provided 142 bone implements, including some that appear decorated, and thirty-six animal teeth and pieces of ivory, bone, or shell that are pierced or grooved, presumably for hanging as beads or pendants. he Arcy Châtelperronians also modiied their living space to an extent that is common only in the Upper Paleolithic. he Châtelperronian layers preserve traces of several “hut emplacements,” of which the most conspicuous is a rough circle of eleven postholes enclosing an area 3–4 m across that was partially paved with limestone plaques (ig. 6.61). Pollen analysis indicates that wood was scarce nearby, and the postholes could have supported mammoth tusks, which are more abundant in the Arcy site than in any other known Paleolithic cave. Conceivably, the Châtelperronian represents a totally independent Upper Paleolithic development from the preceding Mousterian, but the most compelling Upper Paleolithic elements appear only near its end, and their difusion from an Upper Paleolithic source is more plausible. he most likely source would be the Upper Paleolithic Aurignacian Industry, which clearly intruded into western Europe (ig. 6.62). he artifact type that has long deined the basal Aurignacian (or Aurignacian I) across Europe is the split-base bone or antler point, although some sites, primarily near the Mediterranean coast, may record an even earlier manifestation (the Aurignacian 0 or Proto-Aurignacian) characterized primarily by numerous small blades (bladelets), oten with semiabrupt marginal retouch. he retouched bladelets divide between the Dufour and Font Yves types, which also characterize the early Upper Paleolithic of southwestern Asia, as discussed below. Even the earliest Aurignacian assemblages typically contain numerous bone or antler artifacts, together with indisputable art objects and personal ornaments (beads or pendants). Alternation (interstratiication) of Châtelperronian and early Aurignacian layers that would demonstrate chronological overlap has been proposed at El Pendo Cave in northern Spain and at three French caves—the Roc du Combe, Le Piage, and the Grotte des Fées de Châtelperron. he Grotte des Fées is particularly notable since it is the name site for the Châtelperronian, but the interstratiication there and elsewhere is controversial. he case for Châtelperronian/Aurignacian contact thus rests on the numeric dates, summarized above, that imply contemporaneity 37–34 ka. Unfortunately, besides independent Neanderthal invention or borrowing from early Aurignacian neighbors, there is a third, less interesting explanation for the remarkable Châtelperronian at Arcy-sur-Cure. he excavations were conducted between 1948 and 1966, and the methods were excellent for their time. However, hindsight suggests that they would not meet present standards, particularly in a deposit that was stratigraphically complex, and it is possible that the excavators inadvertently failed to separate a typical Châtelperronian occupation without

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burins

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Grotte du Renne, Arcy-sur-Cure

593 FIGURE 6.60. stone and bone artifacts from the Châtelperronian layers of the Grotte du renne, arcy-sur-Cure, northcentral france (redrawn after movius [1969b], igs. 3–8). numerous wellmade burins, bone artifacts, and items of personal adornment justify placement of the Châtelperronian within the upper Paleolithic, but the morphology of the backed knives suggests links to the preceding mousterian of acheulean Tradition Type a, and the possibility exists that the Châtelperronian resulted from the diffusion of upper Paleolithic traits into a basically mousterian context. however, the Grotte du renne Chatelperronian is unique for its abundance of upper Paleolithic artifacts, and the reason may be an excavation procedure that mixed the contents of mousterian and aurignacian layers. so far, Châtelperronian human remains are known from only two sites, but they indicate the people were neanderthals.

bone artifacts

ornaments and well-made bone tools from an early Aurignacian occupation that had both. It has been argued that the Châtelperronian people used distinctive methods to manufacture ornaments and other special pieces, but a comprehensive comparative analysis shows that both the manufacturing methods and the inal products it comfortably within the early Aurignacian range. In sum, excavation, not ancient behavior, may have created the singular characteristics of the Arcy Châtelperronian. he case for stratigraphic mixing or misreading recalls the arguments that have been ofered to dismiss Châtelperronian/Aurignacian interstratiication elsewhere. he Arcy instance underscores the extent to which archaeological interpretation depends on excavation quality. Outsiders must usually

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FIGURE 6.61. Plan and reconstruction of the Châtelperronian hut emplacement in level Xi of the Grotte du renne, arcysur-Cure, north-central france (redrawn after farizy [1990a], ig. 5). The arrangement of postholes, plaques, hearths, and large bones suggests that the hut was immediately in front of the cave. The apparent patterning on the living loor is typical only for the upper Paleolithic.

Grotte du Renne posthole ashy spot pebble limestone plaque

hearth

hearth

tusk

N

0

1m

take quality for granted, but even the most careful, experienced excavators may fail to detect mixture between occupations in a complex cave like the Arcy Grotte du Renne. his reinforces what should be an archaeological maxim: the irst discovery of a unique or unexpected assemblage should be regarded as a possible accident, and even the second could be a coincidence. In general, only repeated, independent discoveries establish a pattern that merits serious behavioral interpretation. Nearly ity years ater the Arcy Châtelperronian was irst reported, it remains nearly unique. Only the Châtelperronian layers at Quinçay Cave, southwestern France, provide meager corroboration, in the form of four pierced animal teeth. Rich, well-excavated Châtelperronian layers elsewhere, for example, at Grotte XVI, also southwestern France, lack Upper Paleolithic-like ornaments and bone tools, and among the eighteen or so commonly accepted Châtelperronian sites, Grotte XVI illustrates the

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41 Danu b

e

41/42

41 41

42/44 Proto-Aurignacian

Aurignacian split-base points 0

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rule. he four sites that suggest Châtelperronian/Aurignacian interstratiication actually outnumber sites where Châtelperronian stone tools are unquestionably associated with Upper Paleolithic bone, ivory, or antler artifacts. If we continue to accept that Upper Paleolithic features of the Arcy Châtelperronians could relect difusion from early Aurignacian neighbors, then a southward spread of the Aurignacian ater 37–36 ka could also explain the Uluzzian Industry of central and southern Italy, which broadly recalls the Châtelperronian in its mix of Middle and Upper Paleolithic retouched stone tools. he most common Middle Paleolithic types are sidescrapers and denticulates. Upper Paleolithic types include numerous endscrapers and occasional burins and large curved backed geometric elements (lunates and crescents), some of which resemble Châtelperronian backed knives. Two Uluzzian sites have also provided bone “awls” (stout fragments of bone with artiicially pointed tips) that could relect Upper Paleolithic contact. A human upper deciduous premolar from the Grotta del Cavallo suggests that Uluzzian people were Neanderthals. 14C dates indicate that the Aurignacian and Uluzzian may have overlapped briely, although the Aurignacian horizons are consistently on top at the three reported sites with both Aurignacian and Uluzzian layers. his could mean that like the Châtelperronian, the Uluzzian represents an indigenous Mousterian development that Aurignacian invaders irst inluenced and then truncated.

595 FIGURE 6.62. Two proposed routes for basal aurignacian dispersal across europe (redrawn after mellars [2006b], ig. 2). The numbers attached to the routes are calendar ages calibrated from 14C ages according to a highly provisional scheme. The more northerly route—along the danube valley to western france—is suggested for the classic aurignacian or aurignacian 1, marked by carinate (keeled) endscrapers, by blades with scalelike lateral retouch, and above all by bone or antler split-base bones. The more southerly route—paralleling the mediterranean Coast to southern france and then the Pyrenees to northern spain—is proposed for a separate Proto-aurignacian or aurignacian 0,characterized primarily by retouched bladelets. if two distinct dispersals are accepted, the Proto-aurignacian may have originated in the ahmarian industry of southwestern asia, and it would have spread slightly earlier. The classic aurignacian could have originated in the “levantine aurignacian,” which followed immediately on the ahmarian.

SOURCES: Châtelperronian Culture (Harrold 1983, 1988, 1989; Lévêque and Miskovsky 1983; Zilhão and d’Errico 1999); Arcy-sur-Cure Châtelperronian human remains (Bailey and Hublin 2006; Hublin et al. 1996) and dating (Girard et al. 1990; Hedges et al. 1994); Saint Césaire Châtelperronian human remains (Lévêque et al. 1993) and dating (Mercier et al. 1991); Arcy Châtelperronian artifacts (Farizy 1990a, 1990b, 1994; Leroi-Gourhan 1965a; Movius 1969b), especially bone implements and beads or

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pendants (Bahn 1998; d’Errico et al. 1998b); the Châtelperronian as an independent Upper Paleolithic development from the Mousterian (Zilhão and d’Errico 1999) or as a product of difusion from the Upper Paleolithic (Demars and Hublin 1989; Harrold 1983, 1989, 2000; Klein 1973b); the Aurignacian 0 (Carbonell et al. 2000; d’Errico et al. 1998b; Mellars 2004); lack of evidence for Châtelperronian/Aurignacian interstratiication (Bordes 2002; Rigaud 2000; Zilhão 2001b; Zilhão et al. 2006) and evidence from the Grotte des Fées (Gravina et al. 2005); excavation methods at Arcy-sur-Cure (Schmider 2002; White 2001a); Châtelperronian ornaments—outside the Aurignacian range (d’Errico et al. 1998b) or inside (Taborin 2002; White 2001a); Quinçay Cave (Granger and Lévêque 1997; White 1998); Uluzzian Industry (Bietti 1997; Gioia 1988, 1990; Kuhn and Biette 2000; Mussi 1990b, 1998; Palma di Cesnola 1980); Grotta del Cavallo tooth (Palma di Cesnola and Messeri 1967)

Central Europe

Like western Europe, central Europe provides indications that Neanderthals and modern humans perhaps overlapped in time and that they may have exchanged genes and culture. Possible Chronological Overlap. Most important with respect to possible

overlap, direct radiocarbon dating suggests that a Neanderthal mandible and a parietal fragment from Vindija Cave layer G1, Croatia, date from about 32 ka, while direct dating places a fully modern mandible from Peştera cu Oase (Cave with Bones), Romania, near 35 ka. he Peştera cu Oase mandible was found on the surface, without archaeological associations, but it is the right age to represent the Aurignacian population that had invaded central Europe by 36–37 ka. A parietal fragment, two very incomplete mandibles, and ive isolated teeth from Bacho Kiro Cave, Bulgaria, may come from even earlier modern humans, antedating 40 ka, but the Peştera cu Oase mandible is the oldest irmly dated modern human fossil in Europe. Other surface inds from Peştera cu Oase include a largely complete human cranium representing a second, younger individual that is equally modern and that may be equally old. Direct radiocarbon dating shows only that it probably antedates 29 ka. Direct radiocarbon dating places teeth of four fully modern individuals from the Mladeč (Lautsch) Caves, Czech Republic, at 31 ka or before, while radiocarbon analysis of calcite that probably overlay the fossils indicates they could approximate the 35-ka age of the Peştera cu Oase mandible. he Mladeč sample was recovered in weakly controlled excavations between 1881 and 1922, but it includes up to seven variably complete, rugged early-modern skulls, other cranial fragments, and numerous postcranial bones, most or all of which were reportedly associated with early Aurignacian bone points and pierced animal teeth. Elsewhere in central Europe, early Aurignacian artifacts lack irm human fossil associations. A modern human skull, a mandible, and a humerus that were long thought to come from rich early Aurignacian deposits at Vogelherd Cave (Stetten), southwestern Germany, must be placed aside since direct dating, more than seventy years ater their discovery, shows

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that they probably date from only 4–5 ka. Together with other south German sites, Vogelherd still conirms that the early Aurignacian Culture with spectacular ivory igurines was in place between 36 and 32 ka, but it also illustrates how seemingly careful, well-reported excavations can overlook intrusions from much younger layers. he Vogelherd modern human fossils are the most famous of those that direct dating has removed from the early Upper Paleolithic human sample, but there are other examples, including especially those from Velika Pečina in Croatia, from Zlatý kůň (Koněprusy), Svitávka, Svatý Propkop in the Czech Republic, and Kostenki XIV (Markina Gora) in Russia. Radiocarbon analysis places these variously between 12.9 ka (Zlatý kůň) and 1.2 ka (Svitávka). At Kostenki XIV, a skeleton associated with a cultural horizon dated to 30–31 ka, was directly dated to 4–5 ka, but the bones were permeated with a preservative that could not be removed, and the redating is thus only tentative. he juxtaposition of the 32-ka Vindija date with the 35-ka ages from Peştera cu Oase and Mladeč and the 36–32-ka dates on early Aurignacian layers like those at Vogelherd might mean that some central European Neanderthals survived the modern human invasion by up to 3–5 ky. However, the Vindija radiocarbon dates face the same objection as the late Neanderthal/Mousterian dates from Zafarraya Cave, Spain: only a miniscule amount of undetectable recent carbon contamination could make specimens that are actually more than 37 ky old appear many thousands of years younger. It was noted in the last section that such contamination is particularly likely to afect bone, and it appears to be commonplace. Vindija itself illustrates the point since the same two Neanderthal bones that have been directly dated to 32 ka were previously directly dated to 29–28 ka, and the previous dating was thought to be reliable because it was based on a reined technique for removing contaminant. he redating is based on an even more reined technique, but the analysts acknowledge that the new 32-ka age might be only a minimum. Dating aside, it has been suggested that the Vindija Neanderthals might have produced probable early Aurignacian objects, including a split-base bone point, that were found in the the same G1 layer as dated fossils. he stratigraphic association of Neanderthal bones and artifacts in Vindija G1 is questionable, however, partly because cryoturbation (frost heaving) and cave-bear occupation disturbed the surrounding deposits and partly because published reports on the stratigraphy are inconsistent. SOURCES: direct dating of human fossils at Vindija (Higham et al. 2006; Smith et al. 1999) and Peştera cu Oase (Trinkaus et al. 2003c); Bacho Kiro human remains (Kozlowski 1982); Peştera cu Oase cranium (Rougier et al. 2007; Trinkaus et al. 2003b); dating of Mladeč human fossils (Wild et al. 2005) and overlying calcite (Svoboda 2000, 2001, 2004; Svoboda et al. 2002); Mladeč artifact associations (Vlček 1971); Vogelherd human remains (Churchill and Smith 2000a), dating (Conard et al.

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2004), and ivory igurines (Conard 2003; Conard et al. 2003); dating of human remains at Velika Pečina, Zlatý kůň, Svitávka, and Svatý Propkop (Smith et al. 1999; Svoboda 2005; Svoboda et al. 2002; Trinkaus 2005); dating of human remains at Kostenki XIV (Haesaerts et al. 2004); Vindija G1 artifacts (Ahern et al. 2004; Karavanić 1995; Karavanić and Smith 2000; Smith and Ahern 1994) and stratigraphic uncertainty (Allsworth-Jones 1986b; d’Errico et al. 1998b)

Skeletal Evidence for Neanderthal/Modern Human Continuity or Contact.

Based mainly on geological context and artifact or faunal associations, central European Neanderthal fossils have been divided between two successive groups, and the diferences between them have been said to illuminate the relation of Neanderthals and modern humans. he earlier group comes from Krapina in Croatia, Ochoz (Švédov stůl) Cave in the Czech Republic, the Gánovce spring limestones (travertines) in Slovakia, and Subalyuk Cave in Hungary, and it probably dates from the Last Interglacial and the earliest part of the Last Glaciation. he later group comes from Vindija Cave unit G3, Croatia, from Kůlna and Šipka Caves in the Czech Republic, and perhaps from the Sala alluvial site in Slovakia, and it probably dates from the middle of the Last Glaciation. ESR readings on animal teeth from Krapina and Kůlna support this sequence, as do radiocarbon dates on bones from Vindija G, even if we accept that the two dated Neanderthal bones from Vindija are actually several thousand years older than the dates suggest. Most of the Neanderthal fossils come from Krapina (about 900 pieces) and Vindija (about ity pieces). Compared with earlier central European Neanderthals, the later ones had smaller, thinner, and less projecting browridges, frontal bones (“foreheads”) that were perhaps less receding, smaller and somewhat less prognathous faces, mandibles with a greater tendency for chin development, and perhaps smaller anterior teeth. With these diferences in mind, some specialists have suggested that central European Neanderthals evolved into local early-modern humans, such as those from the Mladeč Caves and the somewhat later Upper Paleolithic site of Předmostí, also in the Czech Republic. Both the Mladeč and Předmostí people arguably approached the late Neanderthals in browridge size and general cranial ruggedness. However, the argument for phylogenetic continuity between central European Neanderthals and their early-modern successors has always been problematic because the central European Neanderthal fossils tend to be fragmentary and the sample of truly diagnostic specimens is relatively small, especially with regard to the proposed later Neanderthals who would represent the transitional population. In addition, the postulated diferences between the earlier and the later Neanderthals involve reductions in craniofacial size or ruggedness more than they do changes in morphology. he early-modern (Aurignacian and Pavlovian) inhabitants of central Europe, especially the males, certainly had rugged skulls, oten with large browridges, prominent muscle markings, and occipitals

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midfacial projection juxtamastoid crest

mastoid

Shanidar 1 process (reconstruction) retromolar space Neanderthal 0

receding mandibular symphysis (no chin)

5 cm

mastoid process

Předmostí 3 (reconstruction) modern human

(no retromolar space)

vertical mandibular symphysis (with chin)

599 FIGURE 6.63. reconstructed skulls of individual 1 from the mousterian levels of shanidar Cave, iraq, and of individual 3 from the early upper Paleolithic site of Prˇedmostí, Czech Republic (drawn by Kathryn Cruz-Uribe from photographs and casts). The Shanidar skull is from a typical Neanderthal, whereas the Prˇedmostí one comes from a robust, early anatomically modern European. Like other Neanderthal skulls, the one from Shanidar differs from modern skulls in its relatively longer, lower braincase and long, forwardly projecting face, as well as in more detailed features such as its large juxtamastoid crest. The striking protrusion of the Shanidar face is perhaps most obvious in the forward placement of the nose and front teeth relative to the orbits. Like other Neanderthal mandibles, the Shanidar specimen has a large retromolar space, relecting the combination of a long mandible, a short postcanine (premolar and molar) row, and a relatively narrow ascending ramus.

that exhibit some posterior projection and extensive nuchal planes. In overall morphological pattern, however, both cranially and postcranially they are unmistakably modern, with no derived Neanderthal features (ig. 6.63). In light of the burgeoning evidence for modern human origins in Africa, specialists who previously advocated evolutionary continuity between central European Neanderthals and early-modern humans now commonly argue for interbreeding, in which more numerous modern human invaders assimilated less numerous local Neanderthals. In support of assimilation, advocates continue to cite the Mladeč and Predmosti specimens, supplemented now by the Peştera cu Oase fossils and somewhat younger ones from the Peştera Muierii (Cave of the Old Woman) and Peştera Cioclovina Uscată (Cioclovina Dry Cave), both also in Romania. Like the Peştera cu Oase fossils, those from Peştera Muierri and

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Peştera Cioclovina Uscată lack stratigraphic provenience and artifact associations, but they have been directly dated by the accelerator radiocarbon method. he radiocarbon ages place both near 30–29 ka, and the people might thus have made late Aurignacian artifacts like those found separately from the fossils at Peştera Muierri. he Peştera cu Oase, Peştera Muierri, and Peştera Cioclovina Uscată fossils are overwhelmingly modern in appearance, but they exhibit features that might indicate a small or remote degree of Neanderthal ancestry. Interbreeding between modern humans and Neanderthals might particularly explain the coniguration of the opening or foramen for the mandibular nerve on the inner (lingual) surface of the ascending ramus of the 35-ky-old Peştera cu Oase mandible. On the mandibles of living humans, this foramen is V-shaped more than 98% of the time, while it is horizontal-oval in about 60% of the Neanderthals on which it has been observed (ig. 6.13). he horizontal-oval coniguration also occurs on about 20% of early Upper Paleolithic mandibles, and advocates of genetic continuity between Neanderthals and their modern successors have oten cited this similarity. Both Neanderthal and modern mandibles sometimes exhibit a V-shaped foramen on one side and a horizontaloval coniguration on the other. he Peştera cu Oase specimen falls in this category, with a V-shaped opening on the right side and a horizontal-oval coniguration on the let. he Peştera Muierri fossils comprise a skull, a mandible, a scapula, and a tibia, and as emphasized above, their overall appearance is unambiguously modern. his is especially true of the skull and mandible, which are more or less complete. However, the skull has a relatively lat (vs. arched) frontal bone and a prominent occipital bun (a marked bulge or projection at the rear), and the ascending branch of the mandible exhibits a moderately asymmetric sigmoid notch (ig. 6.12 illustrates the tendency for the notch to be more asymmetric in Neanderthals than in modern or near-modern humans). All three features occur in varying frequency on the skulls of modern, including living people, but they are much more common in Neanderthals. he scapula has a narrow glenoid fossa (the hollow or socket that articulates with the head of the humerus) that again recalls the Neanderthal or the more general nonmodern human condition and that may not be replicable in living humans. he Cioclovina Uscată skull is also unquestionably modern in overall form, but it exhibits an elongated patch of roughened (resorptive) bone above a moderately developed transverse occipital torus. he torus itself is non-Neanderthal like in its relatively weak lateral development, but the roughened patch vaguely recalls the suprainiac fossa of the Neanderthals (illustrated in ig. 6.5.) he morphological case for assimilation is easier to make than the case for evolutionary continuity, but it is not compelling. he horizontal-

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oval mandibular foramen on one side of the Peştera cu Oase mandible, for example, need not imply interbreeding, partly because foramen shape may speak more to chewing during development than to genes and partly because the relevant anatomy is observable on only three mandibles from the early-modern (or near-modern) African contemporaries of the Neanderthals. hey all have V-shaped foramina, but a larger sample could show that early-modern Africans sometimes also had horizontal-oval foramina, and the frequency may have been as high as in early Upper Paleolithic populations. he issue has been clouded further by the redating of the Vogelherd human fossils to 4–5 ka since the Vogelherd sample includes a mandible that has oten been used to demonstrate the presence of the horizontal-oval coniguration in early Upper Paleolithic people. In the same vein, the proposed suprainiac fossa on the Cioclovina Uscată occipital would be more compelling if it matched the Neanderthal condition much more closely and if it were not combined with otherwise completely modern cranial morphology. Late prehistoric and historic skulls sometimes exhibits patches of roughened bone in the same position on the occipital, and their developmental basis is unclear. he proposed Neanderthal-like features of the Peştera cu Oase, Peştera Muierii, and Cioclovina Uscată fossils cannot be summarily dismissed, but they would support hybridization more strongly if they were more abundant and if their genetic basis were known. In the absence of this information, there is no way to determine whether to expect them in Neanderthal/modern human hybrids, particularly ones that were probably 100–200 generations removed from the proposed hybridization event. Observations on interbreeding between modern baboon species suggest that only an early generation hybrid might be readily detectable in the fossil record and that the principal clues might be developmental (ontogenetic) anomalies like excess (supernumerary) teeth or extra cranial sutures that relect divergent developmental schedules. In the absence of possible early generation hybrids that match this description, a decision on the extent to which modern humans and Neanderthals interbred in Romania or anywhere else in Europe will probably depend on the extraction of endogenous DNA from the largest possible number of relevant fossils. As discussed in the next chapter, both ancient and modern DNA so far suggest that if interbreeding took place, it was on a scale that may be too small to detect. Dental characters reinforce this conclusion since they are highly heritable, and so far, no early-modern and living Europeans exhibit dental traits that are otherwise uniquely Neanderthal. SOURCES: earlier and later Neanderthals in central Europe (Smith 1982, 1984; Svoboda 2005); ESR dates from Krapina (Rink et al. 1995) and Kůlna (Rink et al. 1996); radiocarbon dates from Vindija G (Ahern et al. 2004); Neanderthal fossils from Krapina (Radovcic et al. 1988; Simek and Smith 1997)

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and Vindija (Ahern et al. 2004; Smith et al. 1985; Smith and Ahern 1994; Wolpof et al. 1981); trends in central European Neanderthals (Smith 1982, 1984; Smith et al. 1985; Wolpof 1996); similarities between later Neanderthals and early-modern humans (Frayer 1992; Smith 1991); possible modern human assimilation of Neanderthals (Ahern et al. 2004; Smith et al. 1999, 2005; Trinkaus 2007); Peştera Muierii (Soicaru et al. 2006); Peştera Cioclovina Uscată—dating (Olariu et al. 2004), suprainiac fossa present (Soicaru et al. 2007; Trinkaus 2007) or absent (Harvati et al. 2007); mandibular foramen shape (Frayer 1992; Trinkaus et al. 2003c); continuity in size and robusticity between earlier and later central European Neanderthals (Howells 1982; Stringer 1982); Neanderthal traits in the Mladeč skulls—present (Frayer 1992), absent (Bräuer and Broeg 1998); mandibular foramen shape in earlymodern Africans (Stringer and Bräuer 1994); depressed areas of rugose bone on recent human occipitals (Bräuer 2006); baboon hybrids (Ackermann et al. 2006); distinctive Neanderthal dental traits (Bailey 2002; Bailey and Lynch 2005)

Archaeological Evidence for Neanderthal/Modern Human Interaction.

Parallel to the skeletal remains, central European artifact assemblages may imply continuity between Mousterian and Upper Paleolithic populations, but the case is at best equivocal. To begin with, there is the questionable assumption that stone artifacts can reveal population continuity in the same way that human fossils might. he mechanisms underlying artifactual (cultural) change are much more poorly understood, and it is even more diicult than with fossils to separate similarities that are due to shared descent (homologies) from ones that relect parallel development or simply chance (analogies). Nonetheless, central European archaeologists have identiied an array of broadly similar artifact industries that are said to be transitional between the Mousterian and the Upper Paleolithic. he most commonly cited examples are the Altmühlian, Micoquian (or Micoquian-Prondnikian), Blattspitzengruppe (“leaf-point group”), Bohunician, Bryndzenian (or Moldavan Szeletian), Jankovichian, Jerzmanowician, and Szeletian Industries, which difer mostly in where they occur. he artifactual distinctions between the Bohunician, Bryndzenian, Jankovichian, Jerzmanowician, and Szeletian are particularly vague, and for present purposes they can all be lumped into the Szeletian, which was described irst. Even ater lumping, however, there are still few Szeletian assemblages that comprise more than a few dozen retouched pieces, and the implication is that Szeletian populations were generally very sparse. Cave bear bones far outnumber infrequent Szeletian artifacts at key sites, including the type site of Szeleta Cave (Hungary). he Szeletian is uniied by the presence of bifacially worked pieces, mainly (but not always) leaf-shaped (foliate) points (ig. 6.64). In other respects Szeletian assemblages are remarkably diverse. Most are dominated by Mousterian sidescrapers, denticulates, and so forth, but some (particularly those assigned to the Bohunician) are relatively rich in Levallois blades, and some contain numerous endscrapers and burins. It is the latter assemblages that especially suggest continuity from the Mousterian to the Upper Paleolithic. So far, Szeletian human remains are

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603 FIGURE 6.64. bifacial leaf-shaped points from the upper deposits of szeleta Cave, hungary (redrawn after allsworth-Jones [1986b], ig. 18). such points typify the szeletian variant of the middle Paleolithic in central europe.

cm

Bifacial leaf-shaped points from Szeleta Cave (“Szeleta Upper”)

restricted to three isolated teeth (two mandibular incisors and a mandibular canine probably from the same individual) associated with Szeletian (or Jankovichian) artifacts at Upper Remete Cave (Máriaremete-Felsö) in Hungary and a lower second molar that may have been associated with Szeletian artifacts at Dzeravá skala (cave) in Slovakia. Based on large dimensions, all four teeth have been tentatively attributed to Neanderthals, which could mean that Neanderthal Szeletians were directly ancestral to early-modern central Europeans. here is the problem, however, that some important Szeletian assemblages come from surface sites, and others were excavated long ago with very weak stratigraphic control. In both circumstances, an apparent mix of Mousterian and Upper Paleolithic stone-artifact types could result from the inadvertent amalgamation of initially separate Mousterian and Upper Paleolithic components. Moreover, even if an Upper Paleolithic Szeletian truly existed, it need not imply a local origin for the Upper

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Paleolithic. Instead, like the late Châtelperronian and the Uluzzian, it could signal the difusion of Aurignacian traits into a Mousterian context. Radiocarbon dating implies that the Szeletian may have persisted until 35 ka when the Aurignacian had appeared locally. A few Szeletian sites, including Szeleta Cave itself, have provided well-made bone points but in each case, there is the possibility that they come from mixed Szeletian and Aurignacian deposits, and no site has produced unequivocal ornaments like those that may relect Aurignacian inluence on the Châtelperronian at Arcy, France. In addition, there is no site where Szeletian and Aurignacian layers alternate to demonstrate Szeletian/Aurignacian contemporaneity. At the rare sites where Szeletian and Aurignacian layers occur together, the Aurignacian layers always overlie the Szeletian ones, and it remains possible that the Szeletian was a strictly indigenous Mousterian development that was ultimately truncated by the Aurignacian. As in western Europe, so in central Europe, the Aurignacian represents the earliest unequivocal Upper Paleolithic, and it was unquestionably intrusive. TL dates from Temnata in Bulgaria and 14C dates from Istállóskö in Hungary, Bacho Kiro in Bulgaria, and Willendorf II in Austria suggest that the Aurignacian may have encroached on central Europe as much as 40 ka. If so, it may have appeared a few millennia earlier than in western Europe, but additional dates from indisputable early Aurignacian layers in both central and western Europe may eventually show that it appeared everywhere roughly 37–36 ka and that existing methods cannot establish geographic diferences in time of appearance. SOURCES: distinctive Neanderthal dental traits (Bailey 2002; Bailey and Lynch 2005); transitional central European artifact industries (Allsworth-Jones 1986b, 1990; Bolus and Conard 2001; Conard and Fischer 2000; Kozlowski 2000, 2004; Svoboda 1993, 2004; Svoboda and Simán 1989; Svoboda and Svoboda 1985; Valoch 1969, 1972) and their relation to the Szeletian (Allsworth-Jones 1986b, 1990); cave bear bones in Szeletian sites (Adams and Ringer 2004); human teeth at Upper Remete Cave (Gábori-Czánk 1983) and Dzeravá skala (Svoboda 2001, 2005); possible difusion into the Szeletian (Allsworth-Jones 1986b, 1990; Valoch 1969, 1972, 1982a); bone points in Szeletian sites (Svoboda 2001, 2004); intrusion of the Aurignacian into central Europe (Allsworth-Jones 1990; Kozlowski 1990a; Mellars 1996; Svoboda 2004; Valoch 1969, 1972), perhaps by 40 ka (Allsworth-Jones 1990; Damblon and Haesaerts 1997; Kozlowski 1982, 1992b; Mellars 1993, 1996; Straus 1993–1994; Svoboda and Simán 1989) or only 37–36 ka (Zilhão and d’Errico 1999)

Eastern Europe

Until recently it could be claimed that both Neanderthals and modern humans produced Mousterian artifacts in eastern Europe and, thus, that Neanderthals evolved locally into modern humans. he evidence came mainly from Crimea, where Mousterian artifacts accompanied Neanderthal bones at Kiik-Koba and Zaskal’naya VI and fully modern remains at Starosel’e Cave. However, the Starosel’e association was problematic from the time it was irst reported since the excavator could not distinguish any stratigraphy in a 4-m thick Mousterian proile, and chemical analy-

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sis suggested that the modern human bones postdated accompanying animal bones. A careful reexcavation of Starosel’e subsequently uncovered two additional modern skeletons at roughly the same depth in the Mousterian deposit as the original skeleton. However, the two new skeletons plainly lay in intrusive seventeenth-century or later Tatar graves. One of them was laid out exactly like the original one, and since the positioning accords with known Tatar custom, the previously proposed Mousterian/early-modern human association at Starosel’e should be discarded. As in western and central Europe, so in eastern Europe, it still appears that only Neanderthals made Mousterian artifacts. here is no compelling evidence for continuity between the Mousterian and the Upper Paleolithic in eastern Europe, but the Upper Paleolithic was established locally by 40 ka, as early or earlier than in central and western Europe. he early date is particularly well-established at Kostenki (roughly 52°N, 39°E) on the Don River, where assemblages at seven separate open-air Upper Paleolithic sites occur below a volcanic ash that originated in Italy and that has been dated by 40Ar/39Ar to 38.5–41 ka. OSL dates on the containing deposits suggest the assemblages accumulated between roughly 45 and 40 ka. he oldest—at Kostenki XIV and XVII—variously comprise typical Upper Paleolithic prismatic blades, endscrapers, and burins, together well-made bone points and awls, shaped antler digging tools (mattocks), drilled stone beads, perforated shells, and worked ivory objects, including the head of an ivory igurine. he shells originated on the Black Sea coast, more than 500 km away, and most of the stone for artifact manufacture originated 150–200 km away. At Kostenki and other sites in eastern Europe, the earliest Upper Paleolithic industries are remarkably diverse, and only three sites—Syuren’ 1 Cave in Crimea, Kostenki I layer III, and Kostenki XIV (“cultural layer in the volcanic ash horizon”)—have provided assemblages that meet the central and west European deinition of the early Aurignacian, primarily because they include numerous bladelets or microblades with ine or semi-steep retouch. By dating and typology, only the Kostenki XIV assemblage unquestionably approximates the age of the basal Aurignacian on the west. he Syuren’ I and Kostenki III assemblages could represent later variants, but even at Kostenki XIV the Aurignacian assemblage is not the oldest Upper Paleolithic at the site. In both Crimea and at Kostenki, the oldest Upper Paleolithic assemblages have no west or central European counterparts, and Buran-Kaya-III shelter, Crimea, has provided a probable initial Upper Paleolithic variant that is so far known only there. It comprises thin, bilaterally asymmetric bifacial points, trapezoidal microliths shaped by ine bifacial retouch on two or three edges, and standardized bone artifacts. Use wear indicates that the trapezoidal microliths were hated and used for cutting or chopping.

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If the Upper Paleolithic signals the arrival of modern humans everywhere in Europe, its early course was clearly diferent on the east than on the west and center. he reason may be the distribution of Neanderthals, who lived throughout western and central Europe but occurred only on the western and southern margins of eastern Europe when Upper Paleolithic people arrived roughly 40 ka. Over most of eastern Europe then, Upper Paleolithic people colonized regions where they had no human competitors and they quickly established themselves in places where no one had lived before. SOURCES: human fossils in Crimea (Kolossov et al. 1975; Ullrich 1958; Vlček 1975); antiquity of Starosel’e human fossils (Klein 1969b; Marks et al. 1997); earliest Upper Paleolithic in eastern Europe (Anikovich 1992; Hofecker 1986, 1997, 2002) and especially Kostenki (Anikovich et al. 2007; Sinitsyn and Hofecker 2006); Aurignacian in eastern Europe (Sinitsyn 2003); Buran-Kaya III artifacts (Chabai 2001; Marks and Monigal 2000, 2004) and use-wear on microliths (Hardy et al. 2001)

Western Asia

Western Asia arguably provides the strongest case for a local evolution of Neanderthals into modern humans—from Neanderthals like those represented at Tabun, Amud, Kebara, Dederiyeh, and Shanidar Caves into near-modern people like those at Skhul and Qafzeh Caves. Such a succession is contradicted, however, by associated faunas and artifacts, and above all by TL and ESR dates showing that the Neanderthals mainly or entirely postdate the Skhul-Qafzeh people. With this in mind, it is notable that the near-modern people are known only from Israel, on the extreme southwest Asian periphery of Africa, and that they were associated with typical African or Afro-Arabian mammals. he Neanderthals in contrast have been found throughout western Asia, and they were associated with exclusively Eurasian faunal elements. he sum suggests that the SkhulQafzeh people were simply near-modern Africans who extended their range slightly to the northeast during the relatively mild and moist conditions of the Last Interglacial, between 130 and 71 ka. he Neanderthals may have been migrants from the north or east who displaced the Israeli near-moderns when climatic climatic conditions deteriorated at the beginning of the Last Glaciation, roughly 71 ka. In contrast to the Neanderthals, who had broad trunks and short limbs that probably evolved as an adaptation to cold, the Skhul-Qafzeh people had slender trunks and long limbs that ix their ancestry in equatorial latitudes (ig. 6.65). he (Tabun C-type) artifact assemblages that accompany the nearmodern Skhul-Qafzeh people have no compelling northeast African antecedents, but the (Tabun B-type) assemblages that accompany the Neanderthals at Tabun, Kebara, Amud, and Dederiyeh Caves could derive from earlier assemblages farther north and east in Europe or western Asia, and they could thus support a Neanderthal incursion. he

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FIGURE 6.65. The contrast in body proportions between the early-modern or near-modern last interglacial occupants of israel and the last Glacial neanderthals of europe (redrawn after Pearson [2000], 241). The narrow trunks and long extremities of the nearmodern israelis suggest an equatorial origin, while the broader trunks and shorter extremities of the neanderthals suggest adaptation to recurrent glacial conditions in europe.

broad hips and trunk

long forearm

short forearm

long lower leg

607

short lower leg

Israeli near-modern human

French Neanderthal

(Skhul IV)

(La Ferrassie I)

Neanderthals may have accomplished the replacement because they had relatively thick trunks and short limb segments that specially adapted them to cooler climatic conditions. here is nothing in the cultural debris of the Israeli near-moderns (or of their putative near-modern African ancestors and contemporaries) to suggest that they had a cultural (behavioral) advantage over the Neanderthals. In fact, from a strictly archaeological (behavioral) perspective, there is nothing to diferentiate the Qafzeh-Skhul people (or their near-modern African contemporaries) from the Neanderthals in either western Asia or Europe. In western Asia as in Europe, the disappearance of the Neanderthals is closely tied to the appearance of the Upper Paleolithic Industrial Complex. his is distinguished from the preceding Mousterian by essentially the same features as in Europe, including the manufacture of formal bone artifacts and art objects or items of personal adornment ( jewelry). Upper Paleolithic human remains are scarce in western Asia, but where they occur they are uniformly modern. his is true not only of later Upper Paleolithic people, postdating 20 ka, but also of earlier ones, especially those accompanying early Upper Paleolithic (Ahmarian) artifacts at Qafzeh Cave, Israel, and Ksar Akil Cave, Lebanon. At Qafzeh, associated radiocarbon dates indicate that two partial adult skulls date from at least 28 ka. At Ksar Akil, extrapolation from overlying radiocarbondated layers implies that the remains of a seven-to-nine-year-old child (“Egbert”) date from 35 ka or before.

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he best available dates place the Upper Paleolithic in western Asia by 40 ka and perhaps by 45 ka. he earliest currently recognized west Asian Upper Paleolithic assemblages combine Levallois lakes, blades, and points of Mousterian aspect with typically Upper Paleolithic endscrapers, burins, and retouched blades. In the past, such assemblages have been considered transitional from the Mousterian to the Upper Paleolithic, but the consistent appearance of Upper Paleolithic types may mean that they are better referred to as “Initial” (or “Early”) Upper Paleolithic. hey are known from at least eight sites, including especially Emireh Cave and Boker Tachtit in Israel; Ksar Akil Cave, Antelias Shelter, and Abu Halka in Lebanon; Umm el Tlel in Syria; and Üçağizli Cave in Turkey. Regional variants of the Initial Upper Paleolithic share distinctive, inished tool types, but there are no special types common to all. hus, in Israel a deining type is the previously mentioned Emireh point, a Levallois point on which the base was bifacially thinned, presumably to facilitate hating; in Lebanon, it is the chamfered piece or chanfrein, a blade or elongated lake with a lat, beveled end produced when a blow from the side removed the previous tip; and in northern Syria, it is the Umm el Tlel point, an elongated, pointed blade on which the base was thinned before removal from the core (ig. 6.66). At Üçağizli, where the Initial Upper Paleolithic is irmly dated by radiocarbon to 40 ka or before, the artifacts include numerous shaped bone artifacts and more than 500 perforated marine shells that underscore the Upper Paleolithic assignment. Similar perforated shells occur with Initial Upper Paleolithic stone artifacts at Ksar Akil, and if it is assumed that the perforations were humanly produced, the shells are among the oldest beads or pendants in the world. Only the putative shell beads from Blombos Cave, South Africa, would be unequivocally older. At Ksar Akil and Üçağizli, the Initial Upper Paleolithic was followed by the Ahmarian Industry, named for Erq el Ahmar rockshelter in the West Bank south of Jerusalem. he Ahmarian is now known from southern Israel through southeastern Turkey, and it was far more homogeneous than the Initial Upper Paleolithic. Typical prismatic Upper Paleolithic blades and bladelets predominate everywhere, and the bladelets were oten modiied by ine, abrupt, ventral retouch to produce pieces known in Europe as Dufour bladelets (with nonconvergent edges) and Font-Yves points (with edges that converge to a sharp tip). As discussed above, Font Yves points and Dufour bladelets mark the earliest Upper Paleolithic (Proto-Aurignacian or “Aurignacian 0”) in southwestern Europe, where they appeared at the same time or slightly later than the Ahmarian. Font Yves points, long known in western Asia as el Wad points, particularly characterize the Ahmarian. At Kebara Cave, Israel, radiocarbon dating places the Ahmarian before 40 ka, which means either that it overlapped with the Initial Upper Paleolithic in time or that

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Emireh Points

609 FIGURE 6.66. artifacts that characterize different variants of the early upper Paleolithic in southwestern asia: emireh points from boker Tachtit, israel; chamfered pieces from Ksar akil, lebanon; and umm el Tlel points from umm el Tlel, syria (redrawn after bar-Yosef [2000], igs. 8, 9, and 10).

tip removed from side to produce a flat, bevelled end

Chamfered Pieces (Chanfreins)

basally thinned before removal from the core

Uum el Tlel Points

the Initial Upper Paleolithic is even older than radiocarbon currently suggests. Ahmarian layers at Üçağizli and Kebara have provided bone artifacts and perforated shell beads that broadly resemble those from the Initial Upper Paleolithic at Üçağizli and Ksar Akil. he Ahmarian or a derivative persisted until about 18 ka, but in some places it was interrupted between 36 and 28 ka by the Aurignacian (or “Levantine Aurignacian”) Industry, distinguished by the same artifact types that characterize the classic Aurignacian in Europe. As in Europe, Aurignacians in the Levant emphasized lakes as much as blades, and the lakes were commonly transformed into nosed and keeled (carinate) endscrapers like those that mark the French Aurignacian 1, as described in the next chapter. Levantine Aurignacians also produced split-base antler points and pendants on carnivore, red deer, and horse incisors or

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canines that closely resemble their European counterparts. In both Europe and western Asia, unlike later Upper Paleolithic people who oten perforated canine or incisor roots by drilling, Aurignacians gouged out depressions to thin the roots and applied pressure to perforate the depression. Conceivably, the Aurignacian 1 originated in western Asia, but it is known in central and western Europe at least as early, and its actual origin may have been in southeastern Europe, from which it expanded both west and south. he more fundamental point is that the west Asian Initial Upper Paleolithic and the early Ahmarian antedate the earliest Upper Paleolithic in Europe, including the Aurignacian. his was to be expected if modern humans were the sole makers of the Upper Paleolithic and they dispersed from Africa. Nonetheless, the actual origins of the Upper Paleolithic remain obscure. Conceivably, its development is recorded in the Initial Upper Paleolithic–to-Ahmarian sequence or in the already noted progressive change from Mousterian to Upper Paleolithic technology and tool typology dated between 47 and 38 ka at Boker Tachtit, Israel. However, in either case, the transition from the Mousterian to the Upper Paleolithic would have been lengthy and gradual, and it is diicult to reconcile such gradualism with the contemporaneous replacement of resident Neanderthals by modern human invaders. If the gradual artifactual change implies prolonged cultural contact and exchange, its occurrence let no traces in the genes of modern humans. Perhaps future research will show that the Initial Upper Paleolithic relects long-distance difusion of ideas or techniques from modern human populations on the verge of expansion and that the Ahmarian relects their actual spread. he diference might explain why the Ahmarian is conspicuously more homogeneous. SOURCES: dating of near-modern humans and Neanderthals in western Asia (Bar-Yosef 1987; Howell 1957; Tchernov 1992, 1994; Vandermeersch 1981, 1982); origins of Tabun C-type and Tabun B-type artifact assemblages (Marks 1992); Mousterian/Upper Paleolithic diferences in western Asia (Gilead 1981, 1991); bone artifacts and ornaments in the west Asian Upper Paleolithic (Bar-Yosef 2000; Bar-Yosef and Belfer-Cohen 1988; Hole and Flannery 1967); west Asian Upper Paleolithic human fossils (BarYosef and Belfer-Cohen 1988; Hershkovitz et al. 1995; Simmons et al. 1991); dating of Upper Paleolithic human fossils at Qafzeh (Bar-Yosef and Pilbeam 2000) and Ksar Akil (Bergman and Stringer 1989); dating of earliest Upper Paleolithic in western Asia (Bar-Yosef et al. 1996; Bar-Yosef and Belfer-Cohen 1988; Hole and Flannery 1967; Marks 1990; Mellars and Tixier 1989; Phillips 1994; Solecki 1963); earliest west Asian Upper Paleolithic as “transitional” (Copeland 1975) or “Initial” (Kuhn 2002; Kuhn et al. 1999); Üçağizli (Kuhn 2002; Kuhn et al. 2001); Ahmarian (Gilead 1981; Marks 1981b); Dufour bladelets and Font-Yves points (Mellars 2005); dating of Ahmarian (Bar-Yosef et al. 1996); Levantine Aurignacian (Bar-Yosef 2000); early Aurignacian perforated teeth (White 1989); possible west Asian origin of the Aurignacian (Mellars 2006b; Mellars and Tixier 1989); Boker Tachtit (Marks 1990)

Eastern Asia (Including Australia and Vicinity)

Eastern Asia—deined as the eastern portion of the Indian subcontinent and the area yet farther east that Europeans call the Far East—has provided less irm evidence for modern human origins than any other

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major region, and the situation must be improved before the issue of modern human origins can be conclusively resolved. At the moment, however, for the time period between 200 and 50 ka that is of principal concern, both the fossil and archaeological records suggest that eastern Asia continued on a unique trajectory that was established hundreds of thousands of years earlier. hus between roughly 200 and 50–40 ka, when western Asia was variably occupied by near-modern people of probably African origin or by Neanderthals of possible European origin, eastern Asia seems to have been occupied by two or more distinctive human types that were neither Neanderthal nor modern and that probably evolved locally. In Java, there were the Ngandong (or Solo) people whose cranial morphology suggests direct descent from classic Javan Homo erectus. In China there were the populations represented by the Dali, Xujiayao, Maba, and Jinniushan skulls, which might be assigned to “archaic” Homo sapiens (or Homo heidelbergensis) if they had been found in Europe or Africa, but which are more likely to represent an evolved end product of Chinese Homo erectus. he east Asian archaeological record is equally distinctive. It ofers no evidence that a typical Upper Paleolithic blade technology ever intruded southern China and adjacent southeastern Asia. Instead, a relatively crude lake-and-chopper technology appears to have continued there, largely unchanged, from the time of classic Homo erectus, before 250 ka, until 12–10 ka or even later. Even more important, there is no sign of the relatively abrupt appearance of formal bone artifacts, art, sophisticated graves, or other innovations that signal a behavioral metamorphosis in the west between 50 and 40 ka. However, the impression of remarkable continuity and conservatism in the Far East is based on a tiny number of excavated sites, oten poorly dated and unevenly described, and it may not be sustained as research expands. One hint of what future research may show comes from Sri Lanka to the southwest, where a layer dated to 31 ka at Fa Hien Cave and a second dated to 28.5 ka at Batadomba Cave have provided sophisticated stone tools (“geometric microliths”) associated with bones of modern humans. he Batadomba Cave assemblage also includes well-made bone tools and ostrich eggshell beads. he pattern that may eventually emerge is also suggested by the relatively well-known archaeological records of northern Asia (Siberia) to the north and, even more strongly, by the record of Sahul, the glacial age continent that included Australia, New Guinea, and Tasmania, to the southeast. he next chapter presents the details, but the harsh climatic conditions of northern Asia seem to have largely precluded human occupation until 50–40 ka, when people making Upper Paleolithic–like stone tools, formal bone artifacts, and art objects spread across the southern, more temperate margin of Siberia. Sometime between 35 and 20 ka, they extended their range northward into arctic and subarctic latitudes, and

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by 14–13 ka they were at the Bering gateway to North America. Substantial houses and ireplaces like those of their western contemporaries help explain how they managed to live where no one had before. hey also buried their dead in graves that resembled western ones in complexity. Stone-artifact assemblages imply that the same broadly “Upper Paleolithic” complex penetrated southward to Mongolia, northern China, Korea, and Japan. It could in fact be represented at the famous Upper Cave at Zhoukoudian, with its elaborate burials of fully modern people. Radiocarbon dates on animal bones that probably accompanied some of these burials range between 30 and 10 ka. he situation in Sahul is even more directly relevant, for the irst occupants surely migrated from southeastern Asia. he timing of their entry to Sahul is debatable, and some specialists advocate a date of 60 ka or before. he bulk of available evidence, however, favors a more recent entry between 45 and 40 ka. In this instance, a fully modern skull from a layer radiocarbon dated to about 40 ka at Niah Cave, Borneo could represent the parent population. To the extent the irst Australians are known, they were fully modern in both anatomy and behavior. hey made nondescript stone artifacts, but they let behind formal bone artifacts, art objects, and relatively elaborate graves that broadly recall those of their Upper Paleolithic/LSA contemporaries in western Asia, Europe, and Africa. he initial colonization of Sahul might itself be regarded as an index of behavioral modernity since it required the invention of watercrat that could cross 90 km of open ocean. In this light, the timing of the irst crossing becomes critical. If it occurred as much as 60 ka, the colonists would probably be the oldest known behaviorally modern people in the world. hey would antedate comparably modern Europeans by roughly 20 ky, and the Out-of-Africa theory of modern human origins, detailed in the next chapter, would have to be modiied to allow for at least two modern human dispersals—an earlier one eastward to southern Asia and Sahul and a later one northward and westward to western Asia and Europe. Depending on future research on the Asian mainland, the theory might even have to be abandoned in favor of one that accords a more central role to the Far East. he bottom line is that speculation surrounding a possible 60-ky-old colonization of Australia underscores the need to clarify the contemporaneous east Asian archaeological and fossil records.

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SOURCES: continuity in east Asian lake-and-chopper technology (Pope 1988); quantity and quality of the east Asian site sample (Bellwood 1990; Chen and Olsen 1990; Higham 1989; Jia and Huang 1985; Jones 1989; Olsen and Miller-Antonio 1992); Fa Hien Cave (James and Petraglia 2005); Batadomba Iena (Abeyratne et al. 1997; Kennedy and Deraniyagala 1989); Paleolithic in southern China (Gao and Norton 2002); Upper Paleolithic in Siberia (Vasil’ev 1993), in Mongolia and northern China (Brantingham et al. 2001; Chen and Olsen 1990; Qiu 1992), and in Korea and Japan (Norton 2000; Ono et al.

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2002; Reynolds and Kaner 1990); Zhoukoudian Upper Cave (Kamminga 1992); initial colonization of Sahul (O’Connell and Allen 2004)

Africa

Among all the places from which skeletal remains are available, Africa presents by far the best case for a local or regional evolution of modern anatomy. his follows not only from the fossil evidence summarized in this chapter but also from genetic evidence summarized in the next. An African origin for modern humans could explain not only what happened to the Neanderthals but also why, unlike them, their fully modern European successors had relatively long distal limb segments (forearms and lower legs), suggesting an ancestral adaptation to warmer climes. If, as genes and fossils imply, anatomically modern humans originated in Africa as much as 200 ka, a puzzling question is why they failed to expand from Africa before 50 ka. A partial answer, supported by archaeological indings summarized in this chapter and reiterated in the next, is that it was only about 50 ka that they acquired an obvious competitive advantage over their nonmodern Eurasian contemporaries. his advantage is signaled by the appearance of the Later Stone Age (Africa) and Upper Paleolithic (western Asia and Europe), and it may have followed on a mutation that promoted the development of the fully modern brain. A neurological explanation for the Upper Paleolithic/LSA and nonbiological alternatives to it are explored further in the next chapter. In the present chapter, it is more pertinent that a behaviorally based spread from Africa requires that Africa contain the earliest evidence for the behavioral advance. Much of Africa seems to have been depopulated during the period between 60 and 40 ka when the advance occurred, but Enkapune ya Muto, Kenya, implies that the LSA had begun in eastern Africa by at least 46 ka. his is 6 ky before the Upper Paleolithic had appeared in Europe and perhaps 3 ky before it appeared in western Asia. Enkapune Ya Muto and other early LSA sites in the same region imply that eastern Africa is the place to seek not only the earliest evidence of fully modern behavior but also additional human fossils to substantiate the African origin of modern humans. Only eastern Asia stands in potential contradiction, if Sahul was colonized as much as 60 ka and if the colonization required the inventive ability relected in the LSA and Upper Paleolithic. SOURCES: Enkapune ya Muto (Ambrose 1998a)

Overview

In sum, two long-standing questions concerning the fate of the Neanderthals and the origin of modern humans are largely settled. Anatomically

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modern humans evolved in Africa between roughly 300 and 100 ka when the Neanderthals and other nonmodern humans were the sole occupants of Eurasia. he vast majority, if not all, of early-modern or near-modern Africans were behaviorally indistinguishable from the Neanderthals, and it was only about 50 ka that they developed the fully modern capacity for culture. his conferred an adaptive (survival-and-reproductive) advantage that allowed them to spread rapidly to Eurasia, where they swamped or replaced the Neanderthals and other nonmodern people. Specialists difer on whether the shit to fully modern behavior was gradual or abrupt and, if it was abrupt, on whether it might have been rooted in a biological (genetic) transformation or in a social or demographic change. hey also dispute the extent to which expanding Africans interbred with resident Eurasians. he next chapter, which is devoted to the earliest fully modern humans, revisits these issues.

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anaTOmICaLLY mODern humanS

Living humans exhibit remarkably little genetic diversity compared to nearly all other mammals, and the implication is that all living people share a remarkably recent common ancestor. Based on the estimated mutation rates that underlie present diversity, the ancestral population existed about 200 thousand years ago (ka), give or take a few tens of thou­ sands of years. Fossil and genetic observations together locate the popula­ tion in Africa, and archaeological discoveries imply an initial dispersal out of Africa about 50 ka. A more conclusive, more precise statement will require a much denser fossil record and either new dating methods or signiicant reinements to existing ones that permit dates between 200 and 50 ka. Much new research is also needed to determine whether there were multiple dispersals from Africa, perhaps at slightly diferent times and in diferent directions, and to establish whether African migrants exchanged genes and behavioral traits with some of the (nonmodern) Eurasians they encountered. But even if important details remain to be ixed, the signiicance of modern human origins cannot be overstated. Before the emergence of modern people, the human form and human behavior evolved together slowly, hand in hand. Aterward, fundamental evolutionary change in body form ceased, while behavioral (cultural) evolution accelerated dra­ matically. A reasonable explanation is that the modern human form—or more precisely the modern human brain—permitted the full develop­ ment of culture in the modern sense and that culture then became the primary means by which people responded to natural selective pres­ sures. As an adaptive mechanism, culture is not only far more malleable than the body but cultural innovations can accumulate far more rapidly than genetic ones, and this explains how, in a remarkably short time, the human species has transformed itself from a relatively rare, even insig­ niicant life form to the dominant mammal on the planet. his chapter

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summarizes what is known about modern human origins, emphasizing especially the highly signiicant behavioral diferences between even the earliest fully modern people and their nonmodern predecessors.

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History of Discovery Until the mid­nineteenth century, human origins were a subject mainly for theologians, not scientists. Medieval clerics summed the ages of post­ Adamite generations in the Bible and concluded that the present world was created about 6,000 years ago. his estimate itself became gospel, and most eighteenth­ and early nineteenth­century geologists and pale­ ontologists accepted it. To explain extinct species they postulated a se­ ries of earlier, imperfect creations, each terminated by a great lood or some other global catastrophe. his interpretation, commonly termed “creationism” or “catastrophism,” was widely questioned only ater 1859, when Darwin’s Origin of Species showed how natural selection could ex­ plain the evolution of living species from earlier ones. One of creationism’s central tenets was that people had not coex­ isted with long extinct species. During the irst half of the nineteenth century, however, fossil hunters searching in European caves repeatedly found bones of anatomically modern humans alongside those of extinct animals. Excavation methods were crude, and in some cases the associa­ tions were probably erroneous, created, for example, when excavators failed to recognize a relatively recent grave dug into much older depos­ its. In other cases, however, the associations were undoubtedly valid and they implied that modern people had lived in Europe long before popular theology allowed. Since the theology was largely unquestioned, the associations were generally discounted, though some have been vin­ dicated in the present century. Perhaps the most famous example was from Goat’s Hole (Cave) at Paviland, South Wales, where the Reverend William Buckland, professor of geology at Oxford, excavated a skeleton in 1822–1823. It was covered by a layer of red ocher and was accompanied by numerous mammoth ivory artifacts, but Buckland concluded it be­ longed to a Welsh woman of the Roman era whose kinfolk worked tusks they found in the cave. In fact, direct radiocarbon dating now shows the skeleton dates from more than 26 ka, and it thus represents a person (ac­ tually a young male) who was a contemporary of the mammoth. Probably the best known discovery ater Paviland occurred in 1852, when a road repairman pulled a human bone from a rabbit hole in a hill­ side near Aurignac in the lower Pyrenees, southwestern France. He dug a trench into the hillside and found a cave whose mouth was blocked by a collapsed limestone slab. Behind the slab lay the skeletons of seventeen people, while backdirt from the trench contained bones from extinct animals, some of them engraved. he skeletons were reburied in a local

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Christian cemetery, but the site attracted the attention of the pioneer French prehistorian Edouard Lartet, who excavated the cave loor systematically in 1860. He found some isolated human bones apparently associated with those of extinct mammals, but the association was never conclusively established. It is now thought the skeletons were of Neo­ lithic (Holocene) age, postdating the extinct animals. he irst widely accepted discovery of unquestionably very ancient but anatomically modern human remains occurred only in 1868, when railway workers exposed deposits with human bones at the Cro­Magnon rockshelter in the town of Les Eyzies in the Dordogne region of southwest­ ern France. An excavation by Edouard Lartet’s son Louis showed that the bones came from a layer that also contained bones of mammoths, lions, and reindeer, along with numerous artifacts. Stone tools (of the kind now called Late or Evolved Aurignacian) were especially common, and there were also artiicially perforated seashells and animal teeth. he human bones represented ive­to­eight people, including minimally a middle­ aged male (subsequently called the “Old Man of Cro­Magnon”) (ig. 7.1), two younger adult males, a young adult female, and a very young infant. he associated artifacts and animal bones showed that the human remains were very ancient, and radiocarbon dates on late Aurignacian horizons uncovered at the nearby Abri Pataud site in the 1950s suggest the skeletons may date from near 30 ka. A radiocarbon assay of one of the perforated seashells provided an age of 28 ka, and if this estimate is not simply a mini­ mum, it might mean that the people were not late Aurignacians, but their immediate (Gravettian) successors. Ironically, for a brief period the Cro­ Magnon skeletons could be cited against the concept of human evolution, for it could be argued that the Cro­Magnon people were in Europe as early as or earlier than nonmodern kinds of people, particularly the Neander­ thals, who were discovered at about the same time. However, even before the turn of the twentieth century it became apparent that, though the Cro­ Magnons were indeed very ancient, they still postdated the Neanderthals. Not long ater the discovery at the Cro­Magnon shelter, numerous other European sites provided modern human remains clearly associated with animal bones or artifacts indicating great antiquity. Some of the most important early discoveries were made at Mladeč (Lautsch) (1881– 1904), Brno (Cerveny Kopec) (1885), Brno (Francouzská Street) (1891), and Předmostí (1894), all in Moravia, Czech Republic; at the Chancelade (Raymonden) Shelter (1888) and Combe­Capelle Cave (1909) in France; and at the Grimaldi Caves (Grotte des Enfants, Grotta del Caviglione, Barma Grande, and Bausso da Torre) in Italy (1874–1901) (locations in ig. 7.2). Together these inds and those from many other more recently excavated sites show that anatomically modern people have occupied Europe since at least the 35 ka. he exact period may vary from place to place, as discussed below.

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FIGURE 7.1. skulls (reconstructed) associated with early upper Paleolithic artifacts at the Cro-magnon rockshelter and with mousterian artifacts at la Chapelle-aux-saints Cave, both in france. The Cromagnon skull is modern in every important respect, including its short, flat face tucked in beneath the fore part of the braincase and the great height of the braincase compared to its length.

Cro-Magnon 1 (early modern European)

La Chapelle-aux-Saints (classic Neanderthal) 0

5 cm

Remains of ancient anatomically modern people have also been found in other parts of the world, though in smaller numbers than in Europe. he Wajak skeletons, found in central Java in 1888 and 1890 are oten re­ garded as the irst, although it now appears they may date from only 6.5 ka. Table 7.1 lists subsequent discoveries that are especially well­known or informative. he most signiicant might be a skull found in 1952 in a dry river valley near the town of Hofmyer in the Eastern Cape Province of South Africa, he stratigraphic origin and associations of the skull are unknown, but optically stimulated luminescence (OSL) dating suggests that quartz grains inside were last exposed to sunlight about 36 ka, and the skull resembles skulls of early­modern (Upper Paleolithic) Europe­ ans more than it does those of later prehistoric or indigenous Africans. It may thus supplement already abundant evidence for the African origin of the earliest modern Europeans, but the reliability of OSL dating is oten questionable even where sediment samples can be extracted from stand­ ing sections and the radiation dose rate can be assessed in place. In ad­

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Oberkassel, Gonnersdorf & Andernach Magrite, Maisières Paviland Creswell & Spy Stellmoor Ahrensburg & Meiendorf Spadzista, Vogelherd, Předmostí & Nietoperzowa Hohlenstein-Stadel Petrkovice & Mamutowa Geissenklosterle, Mladeč Pincevent Hohle Fels Brno & Verberie Le Flageolet & Peterfels Dolní Vĕstonice, Pavlov & Milovice Aurignac, Enlène & Mas d’Azil Willendorf, Rochereil & Galgenberg, Isturitz, Duruthy, Chancelade Grubgraben Brassempouy & Dufaure & Krems Istállóskö Solutré & Szeleta Grimaldi & V indija & Fumane Arene Candide Velika Pecina Combe-Capelle Tito Bustillo, Niaux La Riera & Aitzbitarte Los Azules & Ekain Crvena Stijena Altamira, El Juyo, La Crouzade Palidoro & El Pendo, Cueva Morín, Polesini El Castillo & El Rascaño Paglicci L’Arbreda & Bacho Kiro Parpallo Reclau Viver Casa da Moura & Temnata Ambrosio Romanelli Kastritsa & Nerja & Asprochaliko Romito La Ferrassie, La Madeleine, Laugerie-Haute, Cro-Magnon, Pataud, Font-de-Gaume, Lascaux, Blanchard, Castanet, Souquette

Gough’s Cave

Franchthi Afalou-Bou-Rhummel Taforalt

dition, even if the 36­ky OSL result is accepted, it bears on the deposits into which the skull was buried, not necessarily on the skull itself, which could be younger, even much younger. Unfortunately, the skull did not retain residual protein (collagen) from which radiocarbon dating could provide a far more reliable estimate of its antiquity, and in the absence of information on its context and associations, its relevance to modern human origins may always be debatable. Colloquially, all early­modern Europeans are oten called Cro­ Magnons, ater the early and historically signiicant French discovery, and this chapter continues the custom. Sometimes, the term Cro-Magnon is

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FIGURE 7.2. approximate locations of the western and central european upper Paleolithic sites mentioned in the text.

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Non-European sites, in order of discovery, that have provided especially informative fossils of early, fully modern humans. Except where otherwise speciied radiocarbon provided the age estimates. he table excludes fossils of supposed Pleistocene age whose actual antiquity is highly uncertain. his includes some from Korea (Norton 2000) and especially 23 isolated skeletal elements recovered in the early 1920’s and late 1970’ s in the Xarusgol (= Sjara-osso-gol = Salawusu) River valley, on the Ordos Plateau, Inner Mongolia. None of the Xarusgol fossils were sealed in place, and direct radiocarbon dating shows that one (a partial femur found in 1923) is less than 300 years old (Keates et al. 2007).

TABLE 7.1.

Site and Discovery Date

Fossils (and geologic antiquity)

References

Wajak (Wadjak) Cave, Java, 1888 and 1890

2 skulls [long thought to date from the late Pleistocene, now dated between 10 and 6.5 ka]

(Dubois 1922; Storm 1995; Storm 2001)

Zhoukoudian “Upper Cave” 3 nearly complete skulls (known as 101, 102, (Shandingdong), China, 1933–34 and 103), 3 additional cranial fragments, 4 mandibles, 4 additional mandibular fragments, numerous isolated teeth, vertebrae, 2 sacrums, 3 scapula fragments, 1 radius fragment, 6 fragmentary pelves, numerous femoral fragments, 3 patellas, 2 tibias, 6 metatarsals, and 1 tarsal [variously estimated between 29 and 10 ka by 14C on possibly associated animal bones)

(Cunningham & Jantz 2003; Cunningham & Wescott 2002; Hedges et al. 1992; Jia 1980; Kamminga 1992; Neves & Pucciarelli 1998; Pei 1939b; Weidenreich 1939; Wu & Poirier 1995)

Qafzeh Cave, Israel, 1933–35

two fragmentary human skulls [30–28 ka]

(Bar-Yosef 2000; Bar-Yosef & Belfer-Cohen 2005; Smith 1995)

Ksar Akil (Ksar ‘Aqil) Cave, Lebanon, 1938

skull (“Egbert”) [35 ka]

(Bergman & Stringer 1989; Coon 1962)

Keilor, New South Wales, Australia, 1940

a skull and femur fragments [12 ka]

(Brown 1987; Jones 1944)

Niah Great Cave (West Mouth), Sarawak, Borneo, 1958

a partial skull (= the “Deep Skull”), let femur, and proximal right tibia, probably from a single individual [>= 40 ka]

(Barker 2002; Barker et al. 2003; Barker et al. 2007; Brothwell 1960; Harrisson 1978; Kennedy 1979)

Nacurrie, Victoria, Australia, 1949

several skeletons, one nearly complete [11.4 ka]

(Brown 1993)

Coobool Creek, New South Wales, Australia, 1950

skeletal remains of 126 individuals, includ­ ing 33 more or less complete skulls [14–9 ka (morphological similarity to Kow Swamp)]

(Brown 1987; Brown 1993)

Ziyang (construction site) (Huang­ cranium without face [late Pleistocene shanxi), Sichuan, China, 1951 (mammalian associations)]

(Wu & Poirier 1995)

Hofmyer, Eastern Cape Province, South Africa, 1952

cranium and mandible [36.2 + 3.3 ka, OSL on sediment inside the cranium]

(Grine et al. 2007)

Laibin (Gaitou Cave), Guangxi, southern China, 1956

partial skull (maxilla, right zygomatic arch, and an occipital fragment) [between 44 and 38 ka (U­series on carbonate deposits below and above)]

(Shen et al. 2007; Wu & Poirier 1995)

Liujiang (Tongtianyan Cave), Guangxi, southern China, 1958

skull and partial skeleton [possibly intrusive burial into deposits dated >= 68 ka (U­series)]

(Shen et al. 2002; Wu & Poirier 1995)

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TABLE 7.1. (continued )

Site and Discovery Date

Fossils (and geologic antiquity)

References

Tabon Cave, Palawan Island, Philippines, 1962 and 2000

fragments of frontal, temporal, and occipital bones; two fragmentary mandibles; and 9 mostly fragmentary postcranial bones [age uncertain, possibly 24–22 ka assuming association with 14C-dated charcoal]

(Détroit et al. 2004; Dizon et al. 2002; Fox 1978)

Minatogawa Quarry, Okinawa, Japan, 1967–74

partial skeletons of between 5 and 9 individuals [18–16 ka, assuming association with 14 C-dated charcoal]

(Suzuki & Hanihara 1982)

Yamashita-cho Cave, Okinawa, Japan, 1968

juvenile femur and tibia [> 32 ka]

(Ikawa-Smith 1986; Trinkaus & Ruf 1996)

Kow Swamp, Victoria, Australia,1968–72

partial skeletons of 30+ individuals [14–9 ka (14C) or possibly 22–19 ka (OSL)]

(Brown 1987; Stone & Cupper 2003; horne 1977; horne & Macumber 1972)

Lukenya Hill, Kenya, 1971

partial skull [18 ka]

(Gramly 1976; Rightmire 1975)

Lake Mungo, New South Wales, Australia, 1969 (Mungo 1 and 2), 1974 (Mungo 3)

3 partial skeletons, including the especially complete one known as Mungo 3 [40+2 ka (OSL)]

(Bowler et al. 2003; Bowler et al. 1970; Bowler & horne 1976; Brown 1987)

Nahal Ein Gev 1, Israel, 1976

a fragmented skull and skeleton [ca. 19 ka (artifact associations)]

(Arensburg 1977; Nadel & Hershkovitz 1991; Smith 1995)

Nazlet Khater, 1980

poorly preserved skeletons of two adults and a foetus or neonate [37.5 ka on possibly associ­ ated charcoal]

(Bräuer & Rimbach 1990; Pinhasi & Semal 2000; homa 1984; Vermeersch 2002; Vermeersch et al. 1984a)

Wadi Kubbaniya, Egypt, 1982

skull and partial skeleton [between 25 and 20 ka (stratigraphic position and artifact associa­ tions)]

(Wendorf et al. 1986)

Batadomba Cave, Sri Lanka, 1981–82

two partial skeletons and various cranial and postcranial bones from at least 14 additional individuals [16 ka]; a mandible, 4 parietal fragments, 2 isolated teeth, and 8 postcranial bones [28 ka]

(Abeyratne et al. 1997; Kennedy & Deraniyagala 1989; Kennedy et al. 1987)

Willandra Lakes Hominid (WLH) 50, New South Wales, Australia, about 1982

a fragmentary skull and postcranial fragments [undated, but presumed late Pleistocene]

(Brown 1993; Simpson & Grün 1998; Stringer 1998; horne 1984)

Qianyang Cave, Liaoning, north­ eastern China, 1982

Cranial vault, mandible and limbbones from one individual; six isolated teeth from a second [between 22 and 15 ka (14C and U­series)]

(Fu et al. 2007)

Moh Khiew Cave, hailand, 1990–91

fragmentary skull [26 ka]

(Matsumura & Pookajorn 2005)

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TABLE 7.1. (continued )

Site and Discovery Date

Fossils (and geologic antiquity)

References

Ohalo II, Israel, 1991

a nearly complete skeleton [19 ka]

(Hershkovitz et al. 1995; Nadel et al. 1995; Nadel & Hershkovitz 1991; Smith 1995)

Tianyuan Cave, Zhoukoudian, China, 2001–2003

34 mostly fragmentary fossils, including a (Shang et al. 2007; Tong et partial mandible and isolated postcranial al. 2004; Trinkaus & Shang elements, believed to represent a single indi2008) vidual [35.5–33.5 ka, based primarily on AMS dates of associated animal bones]

also applied to the African and Asian contemporaries of early-modern Europeans, although they are probably better known simply as early moderns, meaning fossil people whose skeletons are not meaningfully distinguished from those of living people. Many specialists lump the Cro-Magnons, their African and Asian contemporaries, and all living people in the subspecies Homo sapiens sapiens. However, if it is accepted that all modern humans, living and fossil, share a recent African ancestor, they should probably be separated from other humans at the species level, as Homo sapiens, and this is the practice adopted here. SOURCES: history of thought on human origins (Eiseley 1961; Gillespie 1951; Grayson 1983, 1990); early nineteenth­century discovery of associations between modern humans and extinct mammals (Grayson 1983; Oakley 1964); Paviland Cave skeleton—discovery (Molleson 1976) and dating (Oakley 1980; Pettitt 2000); Aurignac human skeletons (Oakley 1964); Cro­Magnon excavations (Lartet 1868), human remains (Gambier 1989; Petit­Maire et al. 1971), associated artifacts and geologic age (Movius 1969a), and 14C dating of an associated shell (Henry­Gambier 2002); early discoveries of Paleolithic modern human remains in the Czech Republic (Smith 1982, 1984; Svoboda 2008; Vlček 1971) and in France and Italy (Formicola 2004; Petit­Maire et al. 1971; Stringer et al. 1984); antiquity of the Wajak skeletons (Storm 1995)

Morphology Any osteological diagnosis of modern humans must encompass a far greater range of variation than is known for earlier nonmodern groups, each of which is represented by far fewer specimens. A succinct diag­ nosis is thus almost bound to fail since marginal exceptions will always exist. Nonetheless, the following is a reasonable working description of modern humans as they are distinguished osteologically from other hominins. Cranium

Average endocranial capacity variable from population to population but generally greater than 1,350 cc; frontal bone (forehead) relatively ver­ tical; vault (braincase) relatively high, more or less parallel­sided, usu­

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short, flat face tucked in beneath the forepart of the brain

smoothly rounded occipital

0

10 cm

parietal bossing

623 FIGURE 7.3. skull of a six-to-seven-year-old child from an elaborate grave (fig. 7.27) dated to roughly 30 ka at Kostenki XV (Gorodtsovskaya) on the Don River in European Russia (redrawn after Yakimov [1957], fig. 5). Stippling indicates parts of the skull that were poorly preserved or missing. The wellpreserved parts show that the braincase was completely modern in shape and that the face occupied the modern position, below the forepart of the brain. X rays of the temporal bones reveal tympanic antra (cavities) that were smaller than usual and that were bounded by hardened (sclerosed) bone, suggesting chronic inflammation of the mastoid processes and middle ear.

Kostenki XV (Gorodtsovskaya) ally with some outward bulging (bossing) in the parietal region (ig. 7.3); occipital contour relatively rounded and lacking a prominent transverse torus; browridge development generally greater in males than in females and variable among populations, rarely forming a continuous bar (su­ praorbital torus) across the top of the orbits but instead usually consist­ ing of two parts—a variably swollen supraorbital trigone at the upper outer corner of each orbit separated by a (supraorbital) notch or groove from an elongated, variably bulging superciliary arch along the upper inner margin (ig. 7.4) (the superciliary arches merge at glabella above the bridge of the nose); face relatively lat and tucked in beneath the an­ terior portion of the braincase; a distinct hollowing of the bone (canine fossa) below each orbit, between the nasal cavity and the cheek bone (zy­ gomatic arch); mandible variably robust, partly in keeping with signii­ cant interpopulational variation in tooth size, but mostly with a distinct chin; usually no gap (retromolar space) between the third molar and the

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FIGURE 7.4. skull of an anatomically modern, fifteen-to-eighteenyear-old woman buried roughly 22 ka in the Pataud Rockshelter, southwestern France (drawn by Kathryn Cruz-Uribe from a slide). The drawing has been deliberately oriented and shaded to emphasize the characteristically modern, two-part structure of the browridge. The two distinct parts are the supraorbital trigone and the superciliary arch, separated by the supraorbital notch or groove. Superciliary arches tend to bulge further forward in males than in females.

glabellar swellling

superciliary arch superorbital notch superorbital trigone

Abri Pataud 0

5 cm

ascending ramus when viewed from the side, relecting the retraction of the face under the skull. he listed features oten vary in expression among modern human populations, and individually, they sometimes fail to distinguish mod­ ern human skulls from the skulls of other kinds of people. However, measures of individual features can be combined into indices that pro­ vide 100% discrimination; and indices that relect braincase globularity (three­dimensional roundedness of the cranial vault) and facial retrac­ tion below the braincase particularly distinguish modern humans from others. he separation conirms that modern humans are uniquely de­ rived (specialized) in both features, and it implies that they are reason­ ably separated at the species level from nonmodern humans, including recent ones such as the Neanderthals. Diferences in both features among human species probably depend mostly on diferences in when and how rapidly diferent parts of the skull grow shortly ater birth or even before. In modern humans, a distinctive pattern of early skull growth, perhaps linked to early expansion of the temporal and frontal lobes, produces increased lexion of the cranial base, lengthening of the anterior part of the cranial base, and reduction in the size of the face (particularly from front to back), all of which help explain why braincase globularity and facial retraction are so conspicuous in adults.

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Postcranium

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

Limb bones sometimes robust, with pronounced muscle markings, particularly in the earliest modern people, but still signiicantly less robust than in earlier kinds of people. Distinctions from the Neanderthals are especially clear, thanks to many available Neanderthal postcranial bones. Besides decreased overall robusticity, the diferences from Ne­ anderthals include: axillary margin of the scapula sometimes bisulcate, but most commonly unisulcate, with the sulcus or groove on the ventral surface, rather than on the dorsal surface as in Neanderthals; distal pha­ lanx of the thumb usually about two­thirds the length of the proximal one, rather than being nearly equal in length as in the Neanderthals; api­ cal tuts or tuberosities on the terminal phalanges of the hand relatively small and elongated, versus large and round in Neanderthals; cortical bone of the femur and tibia signiicantly thinner than in the Neander­ thals; limbs long relative to the trunk, and distal limb segments (lower arm and lower leg) usually longer relative to the entire limb than in the Neanderthals; and pubis signiicantly shorter and thicker than in the Neanderthals. he probable signiicance of these features was considered in chap­ ter 6. In brief, the relatively lat, retracted face, together with some as­ pects of skull shape, may relect decreased use of the anterior teeth as tools, at least by comparison to the Neanderthals. he distinctive shape of the skull may also partly mirror underlying structural changes in the brain that permit fully modern human behavior, including the unique innovative capability that Neanderthals and other nonmodern people apparently lacked. Alternatively, craniofacial diferences from the Ne­ anderthals may partly or largely relect stochastic processes, in which Neanderthals drited one way and ancestral moderns drited another. he decline in overall postcranial robusticity and muscularity, coupled with the cited morphological changes in the scapula, hand, femur, and tibia, probably signal a shit toward reduced reliance on bodily strength and greater reliance on culture (especially technology) in performing everyday tasks. he selective advantage of reduced robusticity was that it reduced individual food requirements and permitted a given quantity of food to sustain more people. At least in Europe, where skeletal data are most abundant, in the 20 ky ater modern people appeared, bodily dimensions and robusticity continued to decline, probably in relation to ongoing cultural advances. Relatively long limbs and short trunks and long distal limb segments probably relect an ancestral adaptation to relatively warm climates and thus an African origin for all modern people, as discussed below. In keeping with an especially recent African origin, the earliest fully mod­ ern (early Upper Paleolithic) Europeans had even longer limbs relative

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to trunks than their later prehistoric and historic successors. he shorter, thicker pubis might relect a smaller average birth canal (pelvic inlet), perhaps because cultural advances enabled a shorter gestation period. As a result, infants would have been exposed to critical environmental stimuli when they were neurologically especially responsive. A shorter gestation period would also allow a reduction in birth spacing, increas­ ing the potential for population growth. As discussed in the previous chapter, however, it is more likely that the diferences between Neander­ thal and modern human pelvises are due to slight, if consistent, difer­ ences in overall body structure as opposed to diferences in reproductive physiology. Much has been written on the racial ainities of early­modern peo­ ple, and it has sometimes been said that “racial types” occurred in areas where they were absent historically—as, for example, black Africans in Europe (Grimaldi in northern Italy and Kostenki XIV in Russia), Eski­ mos in Europe (Chancelade in France), and Ainu (or possibly Europe­ ans), Melanesians and Eskimos in northern China (Zhoukoudian Upper Cave). However, most early­modern skulls do not exhibit unequivocal characteristics of any present­day race, and the modern pattern of cra­ nial variability is mainly apparent only ater 10 ka. It might then relect the expansion of “Neolithic” (agricultural) peoples from geographically separate centers. he expansion of rice agriculturalists in eastern Asia could, for example, explain the origin of the “Mongoloids” ater 7–6 ka, while the spread of mixed farmers with cereals and livestock from south­ western Asia ater 8 ka could mainly explain the origins of European “Caucasoids.” Early­modern human remains are more abundant in Eu­ rope than anywhere else, and in a broad morphological sense the earliest modern Europeans did anticipate living ones. But as a group they tended to have larger, higher, and broader skulls with broader faces, more robust browridges, and larger mastoid processes, and they should probably not be lumped with living Europeans in a Caucasoid race. In the present context, the more important point is that the skeletal diferences that separate early­modern Europeans from living humans are trivial com­ pared with the diferences that distinguish them and all living humans from the Neanderthals.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: modern human craniofacial distinctions (Lieberman 2008); variability among historic populations in “modern” craniofacial features (Howells 1973a, 1973b); craniofacial indices that sepa­ rate modern and nonmodern humans (Lieberman et al. 2002); postcranial diferences between Neanderthals and modern humans (Trinkaus 1983a) and their implications for behavior (Trinkaus 1987a); reduction in skeletal robusticity in Europe ater 20 ka (Frayer 1984); African limb proportions of early­modern Europeans (Holliday 1997); failure of early­modern human skulls to it in modern “races” (Howells 1973b, 1989, 1995; Sarich 1997); agriculture and population expansions, including the case of the Mongoloids (Bellwood 1996, 2001; Brown 2001); diferences between early­modern and later European skulls (van Vark et al. 1992)

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Modern Human Origins In the previous chapter, it was shown that the human form came to differ markedly between Europe and Africa ater 500 ka and that by 100 ka Europe was occupied exclusively by the highly distinctive Neanderthals, whereas Africa was inhabited by people who resembled living people far more closely. European fossils from sites such as Atapuerca Sima de Los Huesos in Spain, Swanscombe in England, Biache­Saint­Vaast in France, and Reilingen and Ehringsdorf in Germany, all probably dating to be­ tween 600 and 150 ka, already anticipate the Neanderthals and imply that they were an autochthonous European development. Table 7.2 lists the principal sites that have provided fossils of the near­modern or modern African contemporaries of the Neanderthals. he table should perhaps also include the famous and especially com­ plete modern or near­modern human remains from es Skhul and Jebel Qafzeh Caves in Israel, since associated “Ethiopian” mammalian species imply that the human fossils date to a time when Africa had expanded ecologically to incorporate Israel. hermoluminescence dates on asso­ ciated lints and electron spin resonance dates on animal teeth ix this time between roughly 110 and 80 ka. As a group, the key African fos­ sils reveal people with relatively short, high braincases overhanging the face in front, in stark contrast to the long, low braincases and forwardly mounted faces of the Neanderthals. his diference provides the main fossil support for the now famous Out­of­Africa theory, according to which modern humans originated in Africa and subsequently spread to replace the Neanderthals and other equally archaic humans in Eurasia.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Neanderthal origins in Europe (Arsuaga et al. 1993, 1994, 1997b; Hublin 1996; Stringer and Gamble 1993); Skhul/Qafzeh human fossils—associated mammals (Tchernov 1992, 1994) and radio­ metric dates (Mercier and Valladas 1994; Schwarcz 1994)

The Out-of-Africa Hypothesis and Its Multiregional Alternative

he Out­of­Africa hypothesis for modern human origins might be better called Out of Africa 2 to distinguish it from Out of Africa 1, the widely accepted original human dispersal from Africa between 2 and 1 Ma. It might even be called Out of Africa 3, given fossil and archaeological evi­ dence (summarized in chapter 4) that African hand ax makers expanded to Europe about 600 ka. Following common practice, however, only the last expansion, involving modern humans, will be referred to here as “Out of Africa.” In efect, Out of Africa posits that earlier expansions from Africa were followed by evolutionary divergence, culminating by 100 ka in the emergence of at least three continentally distinct human populations. In Africa there were early­modern or near­modern people, in Europe there were the Neanderthals, and in eastern Asia, there were equally nonmodern people who could represent evolved end­products

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he principal African sites that have provided modern or near-modern human fossils older than 50 ka. For more precise age estimates, see Table 6.2 For locations see igs. 6.20–6.22.

TABLE 7.2.

Site

Fossils

References

Klasies River Main Caves

Five partial mandibles, two partial maxillas, fronto-nasal, zygomatic, temporal, and other cranial fragments, isolated teeth, an atlas, a lumbar vertebra, three metatarsals, a phalange, and portions of a clavicle, radius, and ulna

(Rightmire & Deacon 1991; Rightmire & Deacon 2001)

Blombos Cave

nine isolated teeth or tooth fragments

(Grine & Henshilwood 2002; Grine et al. 2000; Henshilwood 2005)

South Africa

Die Kelders Cave 1 Twenty-three isolated teeth and two phalanges

(Grine 1998; Grine 2000; Grine et al. 1991)

Sea Harvest

An upper premolar and a phalanx

(Grine & Klein 1993)

Hoedjies Punt

cranial fragments, isolated teeth, and postcranial bones

(Berger & Parkington 1995; Matthews et al. 2005; Stynder et al. 2001)

hree isolated molars

(Bräuer & Mehlman 1988; Bräuer & Rimbach 1990)

Omo­Kibish

A partial skull and associated postcranial bones (Omo 1); a second partial skull (Omo 2) and fragments of a third (Omo 3)

(Bräuer 1984a; Day & Stringer 1982; Day & Stringer 1991; McDougall et al. 2005)

Herto

A nearly complete adult skull; a partial juvenile skull; and 24 fragments from a second adult skull

(White et al. 2003)

Aduma

fragments of four skulls

(Haile­Selassie et al. 2004a)

he partial, poorly preserved skeleton of an 8–10­year old child

(Vermeersch et al. 1998)

Ascending rami of two let mandibles

(Hublin 2000; McBurney 1967; McBurney 1975; Rak 1998; Tobias 1967b)

Tanzania Mumba Shelter Ethiopia

Egypt Taramsa Hill Libya Haua Fteah Morocco Dar es Soltan 2

A partial adult skull, with the upper face, and (Bräuer & Rimbach 1990; Ferembach associated let mandible, a fragmentary child’s 1976b; Hublin 1993) skull, and an adolescent mandible and associated maxillary fragment

Mugharet el ‘Aliya

A juvenile let maxillary fragment and three isolated teeth

(Debénath 1982; Howe 1967; Hublin 1993; Hublin 2000)

Zourah Cave

A mandible and an isolated canine tooth

(Debénath 1980; Debénath 1994; Debénath 1982)

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of the H. erectus lineage. It its most extreme form, Out of Africa posits that modern people expanded from Africa beginning 60–50 ka and then replaced the Neanderthals and equally archaic east Asians with­ out gene exchange (or interbreeding). In its less extreme form, some­ times dubbed the Weak Garden of Eden hypothesis, Out of Africa allows for some gene low between expanding moderns and resident archaic populations. he only scientiic alternative to Out of Africa is the theory of mul­ tiregional evolution, which postulates that modern humans originated essentially everywhere that nonmodern humans had lived previously— in Africa, but also in Europe and Asia. Proponents of the multiregional model agree that widely dispersed human populations tended to diverge morphologically immediately following the initial Out­of­Africa event, but they argue that continuous gene low ensured the rapid spread of highly adaptive novelties (like larger brains) and thereby kept all human populations on the same fundamental evolutionary track toward mod­ ern people. An obvious objection to multiregionalism is that it postulates sub­ stantial gene low among small populations that were thinly scattered across three continents. In this light the multiregional model is not so much a theory as it is an ater­the­fact explanation for proposed mor­ phological resemblances between nonmodern and modern populations in Asia and Europe. Multiregionalists argue, for example, (1) that the skulls of the living Chinese share relatively nonprotruding (nonprogna­ thous) jaws, upper facial latness, a tendency for the development of a (sagittal) keel or torus along the top of the skull, extrasutural (“Inca”) bones between the main bones of the skull, upper incisors shaped like tiny coal shovels, and other features with fossils that have been tradi­ tionally assigned to Chinese Homo erectus; (2) that historic Australian Aborigines share large, sometimes shellike browridges, long, lat, reced­ ing frontal bones (“foreheads”), a ridge of bone (an occipital or nuchal torus) around the back of the skull for attachment of the neck muscles, forwardly protruding (prognathous) jaws, large teeth, and other char­ acters with fossils assigned to Indonesian Homo erectus; and (3) that early­modern Europeans share large, prominent noses, a tendency for backward projection (“bunning”) of the occiput (rear of the skull), and a propensity for a “horizontal­oval” mandibular foramen (a perforation for the mandibular nerve on the lingual [inner] surface of the ascending ramus) with the Neanderthals. Ironically, multiregionalists do not cite comparable indications of continuity between archaic and modern Af­ ricans, perhaps because the most conspicuous similarities are ones that modern Africans share more broadly with other modern people. hese include “a high, convex frontal positioned directly above a vertical face, a chin, a rounded occiput, and a short, lexed basicranium” (Lieberman

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1995, 177), whose early appearance in Africa in fact comprises crucial support for the Out­of­Africa model of modern human origins. he multiregional theory has been questioned because most key fea­ tures are not simply present or absent but vary in frequency among far­ lung human populations, both fossil and living. Many are actually most common in recent populations where the multiregional theory supposes them to be rare. In addition, some features that do prevail where mul­ tiregionalists specify, such as large browridges and occipital tori, are primitive characters that may simply have been conserved more in some populations than in others, while other apparent regional characters, such as large noses or especially lat faces, may have evolved repeatedly (convergently) in successive archaic and modern populations. Recur­ rent, independent evolution would be particularly likely for traits that represented adaptive responses to regional conditions, and any trait that truly indicated long­term continuity would almost have to be regionally adaptive. Otherwise its regional character would have been blurred by the interregional gene low that multiregionalism requires. Yet other supposed indicators of regional continuity, such as occipi­ tal bunning or sagittal keeling, may not be developmentally homologous between archaic and modern humans, or they may be mechanically forced by partially shared cranial dimensions that themselves are not homologous (that is, that do not relect close shared descent). Finally, apparent evidence for multiregional continuity may be inevitable so long as regional fossil samples remain small compared with the number of anatomical features among which multiregionalists can search for similarities. In sum, the multiregional model cannot be summarily dismissed, but the fossil record far more strongly favors its Out­of­Africa alterna­ tive. his is particularly true with respect to Europe and western Asia, where both fossils and archaeology indicate that modern humans quickly replaced the Neanderthals. he trunk and limb proportions of the earli­ est modern Europeans imply a tropical or subtropical origin, and the Af­ rican tropics or subtropics are the obvious choice. From a strictly fossil perspective, the biggest obstacle to Out of Africa is the murkiness—mul­ tiregionalists would say contrariness—of the Far Eastern fossil record. Most relevant Chinese fossils, including those from Maba (Tianshuigou), Dali, Yunxian, Jinniushan (Yinkou), Xujiayao, Yanshan (Chaoxian), and Changyang (Walongdong), remain poorly described or weakly dated, and when only fossils are considered, it is still conceivable that modern humans appeared as early in China as in Africa or that the living Chinese originated from gene exchange between invading Africans and resident archaics. Regional continuity or interbreeding between invaders and residents might similarly explain the origins of the modern Australian Aborigines, but this possibility is complicated by the remarkable mor­

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phological variability of Australian fossils, only some of which recall archaic Javan Homo in any meaningful sense. his variability is discussed briely below and is a puzzle that no existing theory of modern human origins parsimoniously accommodates.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Out of Africa 2 (Stringer and Gamble 1993); Weak Garden of Eden hypothesis (Harpending et al. 1993; Harpending and Rogers 2000) and interbreeding between expanding modern humans and resident archaics (Bräuer 1992; Eswaran 2002; Eswaran et al. 2005; Smith 1994b); multiregionalism (Frayer et al. 1993, 1994; Wolpof 1996; Wolpof et al. 1994b, 2000); regional features common where mul­ tiregionalism posits they should be rare (Lahr 1994, 1996); possible convergent evolution in successive populations (Stringer and Bräuer 1994); gene low and the masking of regional characters (Stoneking 1993); indicators of regional continuity may not be homologous (Lieberman 1995); regional sample size and the number of anatomical features indicating continuity (Harpending 1994; Rogers 1995)

Genetics and Modern Human Origins

he fossil record must be the inal arbiter between the Out of Africa and multiregional models, but the pattern of genetic diversity in living humans afords a useful, independent means of assessment, and the re­ markable recovery of Neanderthal DNA now provides another. Occa­ sional genetic assessments support the Multiregional model, at least to a degree, but the large majority, involving modern and Neanderthal DNA far more strongly back the Out­of­Africa alternative, and they suggest that early­modern Europeans exchanged few, if any, genes with their Ne­ anderthal predecessors.

Evolutionary Implications of Modern Human DNA Advances in technology mean that the human genome is now far easier to sample than the fossil record, and evolutionarily signiicant genetic analyses appear far more oten than fresh fossil discoveries. To recon­ struct prehistoric population relationships, geneticists have focused par­ ticularly on mitochondrial (mt) DNA and the Y chromosome because of their relatively simple mode of inheritance. mtDNA resides in organelles in the cytoplasm of cells, and unlike the DNA in cell nuclei, which is passed on in both eggs and sperm, mtDNA is transmitted only in eggs. his means that individuals inherit their mtDNA exclusively from their mothers, and barring the occasional mutation, a particular woman will pass on her mtDNA unchanged to all her ofspring. Again barring muta­ tion, her daughters’ ofspring, their ofspring, and so on down the line will inherit the same mtDNA, which means that mtDNA can be used to trace long lines of female descent. Mutation is rare, but when it occurs, it produces new mtDNA vari­ ants (haplotypes). hese usually have no implications for the survival and reproduction of the owner, and natural selection will thus not afect their frequency. However, random genetic drit (chance) can ix a new variant,

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and diferent mutational histories mean that the ixed variants will tend to difer among populations. Since older variants will tend to exhibit more mutational changes, geneticists can infer the order of mtDNA di­ versiication, and they can use the degree of similarity among variants to construct branching trees (or phylograms) that broadly relect popu­ lation relationships (ig. 7.5). mtDNA mutates more rapidly than most other DNA, and the rate is rapid enough to detect whether all extant human populations share a very distant mtDNA ancestor, compatible with the multiregional theory, or a much more recent one, compatible only with Out of Africa. For purposes of tracing ancestry, the Y chromosome may be regarded as the male counterpart of mtDNA. Unlike other chromosomes in the cell nucleus, the bulk of the Y chromosome is not subject to gene swap­ ping (crossing over and recombination) during meiosis, and again bar­ ring the occasional mutation, this means that every male inherits a Y chromosome that is identical to his father’s. he male ofspring of his sons, their male ofspring, and so forth continue to inherit the same Y chromosome, and the Y chromosome can thus be used for tracing long lines of male descent. It is important to distinguish the relatively simple inheritance of the Y chromosome from the more complicated mode that characterizes other chromosomes, which can be altered by gene swapping during meiosis. As a result, excepting the Y chromosome in males, an in­ dividual’s nuclear DNA generally difers in signiicant and unpredictable ways from the nuclear DNA of either parent. his complicates the use of most nuclear DNA in tracing descent, and the problem is roughly akin to the diiculty that genealogists would face if children were allowed to choose the surname of their father, their mother, or a mix of the two. Genealogical construction is obviously far more straightforward when there is a rule—analogous to the strictly maternal inheritance of mtDNA or the strictly paternal inheritance of the Y chromosome—that forces children to assume the surname of a previously speciied parent, male or female. he potential of genetics to illuminate modern human origins has not always been obvious, and its prominence today can be traced to a landmark study of mtDNA by Cann, Stoneking, and Wilson (1987). Cann and her colleagues analyzed variability in a portion of the mtDNA genome among 147 individuals characterized as sub­Saharan Africans (20), Asians (34), Caucasians (Europeans, north Africans, and Near Easterners) (46), Aboriginal Australians (21), and New Guineans (26). he 147 subjects possessed 133 diferent variants of mtDNA, all presumed to originate ultimately from a single variant by mutation—or, more pre­ cisely, by a series of mutations. Assuming that the smallest number of possible mutations is closest to the actual number that created the ob­ served DNA diversity, Cann and her colleagues computed a genealogical

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1 Chukchi 2 Australian Aboriginal 3 Australian Aboriginal 4 Piman 5 Italian 6 Highland Papua New Guinea 7 Coastal Papua New Guinea 8 Highland Papua New Guinea 9 Georgian 10 German 11 Uzbek 12 Saam 13 Crimean Tatar 14 Dutch 15 French 16 English 17 Samoan 18 Korean 19 Chinese 20 Asian Indian 21 Chinese 22 Coastal Papua New Guinea 23 Australian Aboriginal 24 Evenki 25 Buriat 26 Khirgiz 27 Warao 28 Warao 29 Siberian Inuit 30 Guarani 31 Japanese 32 Japanese

Non-African African

33 Mkamba 34 Ewondo 35 Bamileke 36 Lisongo 37 Yoruba 38 Yoruba 39 Mandenka 40 Effik 41 Effik 42 Ibo 43 Ibo

44 Mbenzele 45 Biaka 46 Biaka 47 Mbenzele 48 Kikuyu

48 Hausa

50 Mbuti 52 Mbuti

52 San 53 San

633 FIGURE 7.5. a phylogram based on complete mtdna sequences from fifty-three geographically dispersed individuals (redrawn after Ingman et al. [2000], 709). The branching pattern reflects the probable order of sequence differentiation, and differences in branch length reflect differing degrees of genetic distance. The phylogram shows that the three deepest (and presumed oldest) branches include only Africans, while the next deepest branch includes some Africans and everyone else. The implication is that the last shared mtDNA ancestor of all living humans lived in Africa. The mean genetic distance between the various human sequences and the chimpanzee sequence allows an estimate of the mutation rate that underlies the human branching pattern. This estimate suggests that the branch that includes all the human sequences diverged from other branches (now extinct) 171.5 ± 50 ka, and the deepest branch that includes non-Africans diverged from the exclusively African branches 52+27.5 ka.

chimpanzee

diagram or tree linking the observed DNA variants by their degree of similarity. he tree turned out to have two main branches—one includ­ ing only Africans and the other some Africans and everyone else. Since the individual branches that required the most mutations to explain (and that had thus probably existed the longest) were African, Cann and her colleagues concluded that the most plausible root for the tree was in Africa. By itself the mtDNA tree did not indicate when the common (fe­ male) mitochondrial ancestor existed, but as noted above, mutations in mtDNA are mostly not subject to natural selection, and they tend to ac­ cumulate at a constant rate. From an estimate of this rate, Cann and her

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colleagues calculated that all modern human mtDNA derived from a single woman who lived in Africa about 200 ka. his was a time when archaic Eurasian lineages, including most notably the Neanderthals, were already distinct, and it followed that archaic Eurasian women had contributed little, if any, mtDNA to living people. he sum suggested an exclusively African origin for living humans, and the single female whose mtDNA underlay all later variants inevitably became known as “African Eve.” It may seem counterintuitive that a single woman could be the source of the mtDNA in all living people, but to understand how and why, consider what would happen to surnames, if we started with a inite number, each unique to one male and each passed on only through the male line. By chance, some families in each generation would have no male ofspring, and their surnames would disappear. he winnowing process would end in just one surname at a rate that would depend on how many surnames there were originally. he reduction to one would obviously not mean that everyone in the inal generation had only one male ancestor in the irst generation, only that the surnames of the other males were lost by chance through time. By reverse analogy, the fact that every living person’s mtDNA derives from the mtDNA of a single woman does not mean that she was the only female ancestor of all living people, only that the mtDNA variants of her female contemporaries were lost by chance over time. In fact, variants of all genes, not just those of mtDNA, must coalesce back to an original version in a single individual, and mtDNA occupies a special place in evolutionary studies mainly because of its strictly maternal inheritance and its relatively rapid, broadly clocklike rate of diversiication. A subsequent, more comprehensive and methodologically more so­ phisticated analysis of variation in part of the mtDNA genome recon­ irmed the structure of the tree that Cann and her colleagues inferred, and it continued to suggest that the last shared mitochondrial ancestor of all living humans had lived in Africa roughly 200 ka. An even more thorough analysis based on the entire mtDNA genome of ity­three geo­ graphically dispersed individuals produced fundamentally the same tree (ig. 7.5) and demonstrated conclusively that it was rooted in Africa. he age of the last shared mtDNA ancestor is debatable because it depends on varying estimates of the mtDNA mutation rate and of past population size and structure, but all studies accept an age of 200 ka or less and some project that non­African mtDNA variants arose as recently as 60–50 ka. his is roughly the time when fossil and archaeological observations im­ ply that modern humans dispersed from Africa to the rest of the world. Global analysis of mtDNA diversity identiies three sub­Saharan macroclusters of similar mtDNA haplotypes or haplogroups designated L1, L2, and L3. L3 can be divided between two subgroups, M and N, and it was early members of the M and N subgroups that irst let Africa.

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Outside of Africa, the M and N lineages progressively diversiied into new haplogroups, whose branching pattern of origin and geographic distribution track the expansion of modern humans across the globe. hus, late­arising haplogroups designated A, B, C, D, and X mark Na­ tive American populations, and in theory, they can be used to determine when and where Native Americans originated. he Y chromosome closely echoes mtDNA in its implications for human evolution. he deepest (oldest) Y­chromosome lineages are in Africa, and the Y­chromosome tree is rooted there. Based on estimated mutation rates, “African Adam,” the last shared Y­chromosome ancestor of all living males, certainly lived ater 200 ka, and the oldest non­African Y­chromosome variants arose ater 100 ka and perhaps closer to 50 ka. Most age estimates place African Adam somewhat later than African Eve, but there is no inherent reason why they had to be contemporaries. hey need not even have been anatomically modern, although their es­ timated ages suggest that they probably were. he key point is that both lived in Africa ater the Neanderthals and other archaic humans had al­ ready emerged in Eurasia. Separately and together then, mtDNA and the Y chromosome strongly support the recent Out­of­Africa model of modern human origins, as opposed to its multiregional alternative. Other forms of genetic evidence bear less conclusively on Out of Af­ rica, but none contradict it. Perhaps most signiicant is the overall degree and geographic patterning of genetic variation among extant human populations. Two facts stand out. First, living humans exhibit far less mtDNA and nuclear DNA variation than do chimpanzees, indicating that extant human populations diversiied from a shared ancestor much more recently than extant chimpanzee populations did. If we assume that the rate of genetic divergence has been more or less constant and that the last common ancestor of chimpanzees and people lived about 7 Ma, it would follow that the last shared ancestor of all living humans existed between 175 and 125 ka. Similar computations based on a single nuclear genetic locus (alpha­globin 2 Alu 1) that is especially invariable in living people place their last shared ancestor at 45 or at 30 ka, depend­ ing on whether the date of chimpanzee/human divergence is taken as 7 or 4.7 Ma. Estimates of this order obviously support Out of Africa far more strongly than its multiregional alternative. Second, genetic overlap is far greater among geographically dis­ persed human populations than it is among populations of great apes for which data are available. hus, no geographically conined common chimpanzee population is known to exhibit more than 41% of the total genetic variability in chimpanzees, and most populations exhibit less. In contrast, individual human populations typically exhibit 85% or more of the total known genetic variation in humans. he diference suggests that geographically dispersed human populations have had far less time

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to diverge genetically, in keeping with the idea that they shared a relatively recent common ancestor. In addition, analyses of mtDNA and of nuclear DNA microsatellites show that human genetic diversity is greater in Africa than on any other continent. (Microsatellites, also known as short tandem repeats or STRs, are blocks of DNA that contain multiple repeats of the same simple DNA sequence, such as CACACACA. Most microsatellites are believed to be selectively neutral, and they mutate at an especially rapid rate.) Africans are also the most diverse people in dental morphology and in cranial form. Since dental form is known to be highly heritable and trees of craniometric similarity broadly resemble gene trees, greater dental and cranial heterogeneity provide support for greater genetic diversity in Africa. By itself, greater genetic diversity might not mean that African populations have been evolving longer, only that they have generally been larger than those on other continents. However, the degree of diversity declines in close step with distance from eastern Africa, and the steplike decline implies a rolling succession of founder populations, each smaller and less diverse than the population closer to Africa from which it derived. It simultaneously implies that the further a population is from Africa, the less time it has had to develop comparable genetic diversity, and the sum is consistent only with a recent Out-of-Africa expansion. he decline simultaneously implies that the further a human popula­ tion is from Africa, the less time it has had to develop an African level of genetic diversity, and the sum is consistent only with a recent Out­ of­Africa expansion. Helicobacter pylori, a bacterium that infects the stomachs of most humans, exhibits basically the same pattern, in which genetic diversity peaks in eastern Africa and declines away from there in parallel to the decline in human diversity. Simulations imply that H. pylori spread from eastern Africa about 58 ka, and the implication is that H. pylori and its host share their geographic pattern of genetic diversity because they expanded from Africa together. In the face of growing genetic and fossil evidence for Out of Africa, the original multiregional theory has all but disappeared, to be replaced by what can be called “Mostly Out of Africa.” his is the idea that mod­ ern humans expanding from Africa interbred to some extent with the Neanderthals and other archaic Eurasians. Such interbreeding could explain the anatomical traits that arguably indicate continuity between regional archaic Eurasian populations and their modern successors. For those who advocate Mostly Out of Africa, the absence of archaic Eur­ asian mtDNA and Y chromosomes in historic human populations may mean only that archaic variants were swamped and then lost by chance in the 50–40 ky interval since modern and archaic humans irst met. Mostly Out of Africa may be supported by an apparent contradiction

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between mtDNA, which exhibits a remarkable number of rare variants (haplotypes), and some selectively neutral nuclear genes, which exhibit far fewer. he excess of rare mtDNA types suggests rapid population growth following Out of Africa 60–50 ka, while the smaller number in some nuclear genes may argue for little or no growth, perhaps because the efective (breeding) population for these genes included not only ex­ panding Africans but also resident Eurasians. he discrepancy, however, could also relect ascertainment bias—the failure of sampling so far to locate rare variants of some nuclear genes—or it could mean that the genes are not truly neutral and that natural selection has pruned many rare variants. Interbreeding between expanding modern Africans and nonmod­ ern Eurasians would also be implied by nuclear genes whose evolution­ ary roots lie outside Africa before 50 ka. Among such genes, the most widely publicized is probably microcephalin (MCPH1). his has two primary haplogroups (clusters of closely related variants) that appear to have separated about 1.7 Ma and to have been conined to reproductively isolated populations from about 1.1 Ma. One of the haplogroups—known as haplogroup D or the D allele—commonly reaches frequencies of 70% or more in Eurasia, while its frequency in sub­Saharan Africa is gener­ ally less than 30%. Mathematical analysis of D allele variability suggests that it rose to high frequency abruptly in early­modern Eurasians about 37 ka. his could mean that it was conined to archaic Eurasians before 37 ka, that expanding modern humans acquired it from Neanderthals about this time, and that it increased rapidly among early moderns be­ cause it conferred a strong natural selective advantage. he argument is intriguing, but there is nothing in the pattern of mi­ crocephalin variation to suggest strong recent selection, and no evidence that the D allele confers an advantage. An efect on brain size is unlikely since the earliest modern or near­modern Africans had brains as larger as or larger than the Neanderthals, and an efect on intelligence seems equally improbable since the archaeology of the Neanderthals does not suggest they were especially intelligent. here is also nothing in the ar­ chaeology of Africans ater 37 ka to suggest they were less intelligent than their Eurasian contemporaries. Among the possible explanations for the inconsistency is that the D allele actually confers no special selec­ tive advantage and that it rose to high frequency among early­modern Eurasians by chance (random genetic drit) as the founding population rapidly expanded ater 37 ka. here is also the possibility that future sam­ pling will show that it is actually much more common in Africa. his leads to the more general point, suggested by modeling, that Out of Af­ rica can accommodate even those neutral nuclear genes that appear to have longer histories in Eurasia than in Africa. he proportion of such genes is well below 20% of the total, and it would have to exceed this

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threshold to preclude the possibility that they relect either inadequate sampling or the chance loss of key variants in Africa.

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SOURCES: a genetic assessment rejecting out Out-of-Africa replacement (Templeton 2007); char­ acteristics of mtDNA (Wallace et al. 1999) and of the Y chromosome (Kaessmann and Pääbo 2002); irst mtDNA trees to imply a recent African origin for all living humans (Cann et al. 1987; Vigilant et al. 1991); tree based on the entire mtDNA genome (Ingman et al. 2000); age of the last shared mtDNA ancestor (Quintana­Murci et al. 1999); global mtDNA haplogroups (Forster 2004; Watson et al. 1997); Y­chromosome tree (Underhill et al. 2000); age of last shared Y­chromosome ancestor (Pritchard et al. 1999; Underhill et al. 2001); oldest non­African Y­chromosome variants (homson et al. 2000); genetic diversity in humans vs. chimpanzees (Cann et al. 1994; Kaessmann and Pääbo 2002; Rogers and Jorde 1995; Ruvolo et al. 1993); last shared alpha­globin 2 Alu 1 ancestor of living humans (Knight et al. 1996); proportion of total genetic variability in individual human and ape populations (Kaessmann and Pääbo 2002; Ruvolo 1997); genetic diversity in Africa (Bowcock et al. 1994; Cann et al. 1994; Relethford 1995; Relethford and Jorde 1999; Tishkof et al. 1996; Zhivotovsky et al. 2003); diver­ sity in African dentitions (Irish 1998; Irish and Guatelli­Steinberg 2003) and cranial form (Relethford 1995; Relethford and Harpending 1994); genetic diversity in Africa and population size (Harpending 1994; Relethford 1995; Relethford and Harpending 1995; Relethford and Jorde 1999); genetic diversity and distance from Africa in human populations (Liu et al. 2006; Prugnolle et al. 2005; Ramachandran et al. 2005; Weaver and Roseman 2008) and in Helicobacter pylori (Linz et al. 2007); Mostly Out of Africa (Relethford 2001a, 2001c; Relethford and Jorde 1999); swamping and chance loss of ancient mtDNA and Y­chromosome haplotypes (Relethford 2001b); failure of some nuclear genes to indicate population expansion following 60–50 ka (Eswaran et al. 2005); trees for neutral nuclear genes that root outside of Africa (Garrigan and Hammer 2006); microcephalin (Currat et al. 2006; Evans et al. 2005, 2006; Mekel­Bobrov et al. 2006); modeling that supports Out of africa even genes with long apparent histories in Eurasia (Fagundes et al. 2007); chance loss or inadequate sampling in Africa to explain nuclear genes that seem to have originated in Eurasia before 50 ka (Satta and Takahata 2002); ascertainment bias to explain the failure of some nuclear genes to relect population expansion ater 50 ka (Weaver and Roseman 2005)

Evolutionary Implications of Ancient DNA So far, eforts to recover DNA from ancient bones have focused on mtDNA, mainly because it is about 1,000 times more abundant than nu­ clear DNA and thus easier to detect. DNA degrades rapidly, and bones of great antiquity rarely retain any. However, in the proper circumstances, probably mostly involving rapid burial under cool, dry conditions, frag­ ments of mtDNA have been shown to survive for up to 100 ky, and be­ ginning with the original Feldhofer Cave Neanderthal specimen in 1997, sixteen Neanderthal bones from eleven sites have now yielded mtDNA (table 7.3). he sites are scattered across southern Eurasia, from Spain on the west to southern Siberia on the east. Some of the fossil mtDNA sequences are extremely short, but they cluster together and apart from modern human sequences. Based on the degree of mtDNA diference and the assumed mutation rate, the last shared mtDNA ancestor of Ne­ anderthals and living humans existed around 500 ka. his estimate is associated with a statistical error of roughly 200 ky either way, but both the midpoint and the error range are consistent with the 600–500 ky estimated age for the last shared fossil ancestor, labeled Homo heidelbergensis in previous chapters.

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Sites with Neanderthal bones that have provided mitochondrial DNA. Figure 6.1 in the previous chapter locates the sites.

TABLE 7.3.

Site

References

El Sidrón Cave, Spain (2 bones, possibly from one individual)

(Lalueza­Fox et al. 2006; Lalueza­Fox et al. 2005; Rosas et al. 2006)

La Chapelle­aux­Saints Cave, France

(Serre et al. 2004)

Les Rochers­de­Villeneuve Cave, France

(Beauval et al. 2005)

Engis Cave, Belgium

(Serre et al. 2004)

Scladina Cave (Sclayn), Belgium

(Orlando et al. 2006)

Feldhofer Cave, Germany (2 bones from 2 individuals)

(Krings et al. 1999; Krings et al. 1997; Schmitz et al. 2002)

Riparo Mezzena, Lessini Mountains, Italy

(Caramelli et al. 2006)

Vindija Cave, Croatia (3 bones, possibly from 3 individuals)

(Krings et al. 2000; Serre et al. 2004)

Mezmaiskaya Cave, Russia

(Ovchinnikov et al. 2000)

Teshik­Tash Cave, Uzbekistan

(Krause et al. 2007b)

Okladnikov Cave, Russia (2 bones from a single individual)

(Krause et al. 2007b)

Like modern human mtDNA, Neanderthal mtDNA exhibits a remarkably low degree of variability, and this is in striking contrast to the much greater degree in chimpanzees and gorillas. In living humans, limited mtDNA (and other genetic) diversity is usually thought to imply both a recent shared ancestor and an origin from a breeding population that numbered 10,000 individuals or fewer. he latter inference in­ directly favors Out of Africa since such a small founding population is unlikely to have been spread across three continents. he low diversity of Neanderthal mtDNA may mean that the sampled Neanderthals stem from an even smaller, geographically restricted population that existed perhaps 300–250 ka. On its own, the uniqueness of Neanderthal mtDNA does not mean that Neanderthals failed to interbreed with the earliest modern Europe­ ans, and if it is fair to assume that Neanderthal females (not just males) participated, the mtDNA of the earliest modern Europeans could pro­ vide the acid test. he main obstacle is the impossibility of diferentiating mtDNA that originated in early­modern human bones from contami­ nating mtDNA that was introduced by recent handling, and the problem is not abstract since handling must explain why ancient animal bones commonly provide modern human mtDNA sequences. he likelihood of modern human contamination is especially great where laboratory controls were weak, as in the case of the mtDNA extracted from a mod­

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ern human skeleton dated to roughly 40 ka at Lake Mungo, Australia. However, it persists even where controls were tighter, as in the case of mtDNA extracted from two Upper Paleolithic (Cro­Magnon) skeletons dated to about 24 ka at Paglicci Cave, Italy. he Lake Mungo skeleton yielded an aberrant mtDNA variant that might have been introduced by interbreeding with nonmodern Eur­ asians and then lost in the ensuing 40 ky. However, it resembles mtDNA that was long ago inserted in the nuclear genome and that has been pre­ viously shown to represent contamination in ancient DNA studies. he Paglicci skeletons provided mtDNA that is comfortably within the range of modern human variability, and the result is surely more reliable, given the stricter protocols employed in extraction. Nonetheless, specialists re­ main skeptical that the Paglicci mtDNA originated in the fossil bones. To circumvent the contamination issue, the most compelling analy­ sis so far focused on bones of ive Cro­Magnons (early­modern Europe­ ans), not to see if they would yield modern human mtDNA, which they surely would, if only as contaminant, but whether they would also pro­ vide distinctly Neanderthal mtDNA. hree of the bones came from the sites of Cro­Magnon, Abri Pataud, and La Madeleine in France and two from the Mladeč Caves in the Czech Republic. he Mladeč specimens are particularly pertinent because they have been dated to between 35 and 30 ka, and they are thus among the oldest known Cro­Magnons. Also, as discussed in the previous chapter, some specialists believe they exhibit Neanderthal features. Amino­acid content and degree of racemization (a time­ and temperature­dependent shit from the original amino­acid form to its mirror image described in chapter 2) have been shown to be particularly good predictors of whether ancient bones retain DNA, and in their amino­acid state and overall quality of preservation, the targeted Cro­Magnon bones equaled or exceeded those of the Neanderthals from which authentic (endogenous) mtDNA has been successfully extracted. Assuming Neanderthal/Cro­Magnon interbreeding, some of the bones could thus be reasonably expected to provide Neanderthal mtDNA. None did, and a similar dedicated efort to detect Neanderthal mtDNA in the well­preserved Paglicci Cave Cro­Magnon bones also failed. his does not mean that Neanderthals contributed no mtDNA to early­ modern Europeans, but if we accept that the early­modern population started small and then grew substantially over time, simulation shows that even a tiny Neanderthal contribution at the beginning should re­ main evident in living humans. A model that assumes Cro­Magnons and Neanderthals interbred over six­to­thirty­seven generations on a front that moved from east to west across Europe between roughly 42 and 30 ka produces essentially the same result. It may never be possible to rule out some assimilation between mod­ ern humans and Neanderthals or other archaic Eurasians, but the analy­

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sis of nuclear DNA from Neanderthal bones promises further insight. Recovery has only just begun, and it is complicated by the need to focus on those rare bones that are relatively uncontaminated by modern human DNA. Specimens that are proportionately rich in Neanderthal (vs. modern human) mtDNA sequences present good candidates, and the most favorable bones so far include two from El Sidrón Cave, Spain, one from Riparo Mezzina Cave (Monti Lessini), Italy, and one from Vindija Cave, Croatia. he two bones from El Sidrón yielded Neanderthal Fork­ head Box 2 or FOXP2, which as discussed below, was particularly desir­ able because FOXP2 is the only gene so far that is known to inluence speech and language. he El Sidrón FOXP2 difered from the chimpan­ zee version in the same respects as modern human FOXP2, and if it is authentically Neanderthal (as opposed to modern human contaminant), it implies that FOXP2 did not restrict Neanderthal language ability. However, even if the modern version of FOXP2 is essential for language, other genes are surely required too, and future research may show that Neanderthals and modern humans difered in one or more of those. he Neanderthals and modern humans probably inherited their shared FOXP2 from their last common ancestor, and the existence of this ancestor only 600–500 ka implies that Neanderthals and modern humans will share most nuclear genes. he alternative explanation for shared genes—interbreeding—is unlikely for FOXP2 because the El Sidron Neanderthals died before modern humans had appeared nearby. In addition to FOXP2, one of the El Sidrón bones and a specimen from Riparo Mezzina have provided a previously unknown variant of the Melanocortin 1 Receptor or MC1R gene. he discovery of the same vari­ ant in Neanderthal bones from two sites supports its likely Neanderthal origin. MC1R is one of many genes that are known to inluence pigmen­ tation in humans, and it includes so­called full­function variants (alleles) that maximize the production of dark pigment and loss­of­function variants that reduce it. People with loss­of­function variants have light skins and red hair, and they concentrate in Europe where natural se­ lection has probably favored light skins because they facilitate ultravio­ let penetration and consequent Vitamin D synthesis. Although the El Sidrón/Riparo Mezzina version of MC1R is unknown in living humans, it shows a loss of function when it is inserted into cultured cells. he im­ plication is that the Neanderthals had light skins and perhaps red hair. he result was predictable since light skin would have conferred the same selective advantage on the Neanderthals that it did on their European successors. he El Sidrón bones have also provided Y­chromosome DNA se­ quences that are unambiguously Neanderthal because they difer from those of modern males in the nucleotides that occur at ive key positions. Nucleotides (or bases) are the basic structural elements of DNA, and at

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any given position, there are four possibilities, abbreviated A, T, C, or G. he recovery of Neanderthal Y chromosomes makes it even more likely that the accompanying FOXP2 and MC1R are authentic, and the failure of Neanderthal MC1R and Y chromosomes to persist in living humans supplements mtDNA indications that Neanderthals and early­modern humans rarely, if ever, interbred. he Vindija Neanderthal bone has yielded a sequence of 65,250 nu­ cleotides to one lab and a sequence of more than 1 million to a second. he diference mainly relects a diference in recovery techniques. Separate analyses of the sequences produced somewhat diferent results regard­ ing the time when Neanderthal and modern human nuclear sequences diverged and the likelihood of subsequent interbreeding. Evaluation of the longer sequence suggested a divergence date centered on about 560 ka, and it opened the possibility of subsequent interbreeding since only interbreeding or a much younger divergence date could explain the rela­ tively high number of derived single nucleotide polymorphisms that Ne­ anderthals and modern humans appeared to share. A single nucleotide polymorphism or SNP is one where the structural DNA element (A, T, C, or G) at a single position difers between two genetic sequences, and derived in this instance refers to an SNP that humans do not share with chimpanzees. Evaluation of the shorter sequence suggested a divergence date centered on 706 ka, and it found a much smaller proportion of de­ rived SNPs shared with modern humans. he shorter sequence actually suggested little or no interbreeding since it included none of the rare, derived SNPs that occur only in living Europeans and that might thus be expected to have originated from Neanderthals. An independent third­party analysis of both Vindija sequences has shown that they are inconsistent, and the most likely reason is that the longer one was contaminated by modern human DNA. hus, the shorter sequence probably allows more reliable inferences, and this pertains par­ ticularly to the issue of interbreeding. he conidence limits around the two calculated divergence dates overlap substantially, which means they could difer simply by chance, and both are consistent with a last shared ancestor for Neanderthals and living humans around 600 ka. he recovery and analysis of Neanderthal nuclear DNA is in its in­ fancy, and the challenge of avoiding contamination with modern human DNA cannot be exaggerated. he Neanderthal bones that have provided characteristically Neanderthal mtDNA sequences have usually also pro­ vided a much larger number of characteristically modern sequences, and the explanation must be contamination. he problem is more severe with respect to nuclear DNA, partly because it will be rare relative to mtDNA in Neanderthal bones and partly because, even when it is authentic, it will mostly be identical to modern human nuclear DNA. However, as discussed below, when additional examples of authentic Neanderthal

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nuclear DNA are available, they will bear not only on the dating of the Neanderthal/modern human split and the likelihood of interbreeding but also on the question of whether the Neanderthals difered from mod­ ern humans in a gene or genes that afect communication or cognition or in a gene or genes that regulate genes that afect communication or cognition. A genetically based diference in communicative or cognitive capacity could explain why, unlike modern human populations that are thrust together, Neanderthals and early­modern Europeans exchanged few if any genes. Genetic incompatibility is unlikely since they had been geographically isolated for only 500–600 ky, but a genetically founded diference in cognition or communication could have kept them apart.

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SOURCES (not including those in table 7.3): circumstances that promote ancient DNA preservation (Hofreiter et al. 2001; Pääbo et al. 2004); consistent diference between Neanderthal and modern human mtDNA (Knight 2003); age of the last shared Neanderthal and modern human mtDNA an­ cestor (Krings et al. 1999); low degree of variability in Neanderthal vs. ape mtDNA (Kaessmann et al. 2001; Kaessmann and Pääbo 2002; Schmitz et al. 2002); mtDNA diversity and inferred size of the founding modern human breeding population (Harpending et al. 1998; Zhivotovsky et al. 2003); origin of Neanderthals from a small founder population 300–250 ka (Lalueza­Fox et al. 2005); con­ tamination of fossils by modern human mtDNA (Cooper et al. 2004; Pääbo et al. 2004); ancient animal bones with modern human mtDNA (Hofreiter et al. 2001; Serre et al. 2004); L. Mungo mtDNA (Adcock et al. 2001); Paglicci Cave mtDNA (Caramelli et al. 2003); L. Mungo mtDNA as contami­ nant (Hofreiter et al. 2001); skepticism on Paglicci Cave mtDNA (Abbott 2003; Gilbert et al. 2005); lack of Neanderthal mtDNA in Cro­Magnon bones (Caramelli et al. 2003; Serre et al. 2004); an­ tiquity of Mladeč fossils (Svoboda 2004; Wild et al. 2005); Neanderthal features in Mladeč skulls (Frayer 1992); amino­acid racemization and DNA preservation (Hofreiter et al. 2001; Pääbo et al. 2004); simulation to determine the Neanderthal contribution to extant mtDNA variability (Weaver and Roseman 2005); modeling to test ancient Neanderthal and modern human interbreeding (Currat and Excoier 2004); Neanderthal nuclear DNA—FOXP2 and the Y chromosome (Krause et al. 2007a), MC1R (Lalueza­Fox et al. 2007), sequences of 65,250 bases (Noonan et al. 2006), more than 1 million bases (Green et al. 2006), and possible contamination (Wall and Kim 2007); skin color varia­ tion—natural selection (Jablonski and Chaplin 2000) and genetics (Barsh 2003; Norton et al. 2006b)

Archaeology and Modern Human Origins

he remaining sections of this chapter focus mainly on the archaeology of the earliest modern humans. he present subsection draws on mate­ rial in the previous chapter to show how archaeology illuminates modern human origins and it anticipates regional detail presented further on. A potentially fatal objection to Out of Africa concerns the failure of near­modern people to expand to Eurasia immediately ater they ap­ peared there between 200 and 100 ka. Instead, they seem to have been conined to Africa until roughly 50 ka, and it is even possible that they were replaced by Neanderthals on the southwest Asian margin of Af­ rica—in what is now Israel—about 80 ka. he lag between the appearance of modern anatomy and its spread from Africa may be more apparent than real since, in general, African and Israeli fossils older than 50 ka are modern only in a cladistic sense. hey exhibit derived anatomical features of modern humans, but the speciic features vary from sample

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to sample or even from fossil to fossil within samples, and it’s only people who lived ater 50 ka that were as conspicuously and uniformly mod­ ern as living people. Conceivably then, signiicant additional anatomical change preceded the expansion of modern Africans to Eurasia. Putting aside the issue of whether the modern human expansion substantially lagged modern anatomy, archaeology helps to explain why the expansion occurred only about 50 ka. As summarized in chapter 6, Africans who lived between 100 and 50 ka diverged sharply from the Ne­ anderthals in their physical form but not in their behavior. heir cultures have been lumped in the Middle Stone Age or MSA, in distinction to the Mousterian cultures of the Neanderthals in Europe. However, the MSA and Mousterian were closely similar, and the degree of interassemblage variability is about the same whether they are kept separate or combined. As discussed in the previous chapter, stone laking in both depended oten but not always on Levallois (prepared­core) technology, and the dominant artifact types were variously sidescrapers, denticulates, uni­ facial and bifacial points, and backed pieces. he historical reason for the distinction is perhaps best illustrated in northern Africa, where ar­ chaeologists have traditionally related artifacts of MSA age to the Mous­ terian Tradition not because the artifacts resembled European artifacts more closely than sub­Saharan ones but because the archaeologists knew Europe better. In contrast, those who irst investigated the MSA in sub­ Saharan Africa, beginning in southern Africa, were more commonly lo­ cals, with little or no European experience. In addition, until 1970, site stratigraphies and 14C dates suggested that the MSA was mostly younger than the Mousterian. he turning point came in 1972, when a suite of 14C determinations showed that the MSA at Border Cave, South Africa, lay mainly beyond the reach of the radiocarbon method. Both site stratigra­ phies and repeated datings by OSL, TL, ESR, U­series, and 40Ar/39Ar now underscore the fundamental contemporaneity of the MSA and Mouste­ rian between 250–200 ka and 50–40 ka. MSA and Mousterian people not only shared sophisticated stone laking technology and similar stone­tool types, they collected naturally occurring iron or manganese compounds that they could have used as pigments; they apparently built ires at will; they buried their dead, at least on occasion; and they routinely acquired large animals as food. In all these respects and perhaps others, they may have been relatively advanced over earlier, archaic people. However, in common with ear­ lier people, they manufactured a relatively small range of recognizable stone­artifact types; their artifact assemblages tended to vary little over thousands of years or thousands of square kilometers (in spite of no­ table environmental variation); they obtained stone raw materials over­ whelmingly from local (vs. far distant) sources (suggesting relatively

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small home ranges or simple social networks); they rarely utilized bone, ivory, or shell to produce formal artifacts; they let little or no evidence for structures or for any other formal modiication of their campsites; they were relatively inefectual hunter­gatherers who lacked, for exam­ ple, the ability to ish; their populations were apparently sparse, even by historic hunter­gatherer standards; and they rarely, if ever produced unequivocal art or jewelry. In addition, their graves contain no unam­ biguous evidence for burial ritual or ceremony. In all these respects, they difered conspicuously from many later prehistoric and historically ob­ served hunter­gatherers. he sum implies that MSA Africans lacked a competitive behavioral (cultural) advantage over their Eurasian contemporaries, and it was only when a conspicuous advantage appeared, about 50 ka, that MSA people or their immediate Later Stone Age (LSA) descendants dispersed to Eurasia. Most specialists accept this point, but they disagree sharply on whether the advantage appeared abruptly about 50 ka or evolved more gradually within the MSA, between 100 and 50 ka. he diference is important for explaining how it evolved since an abrupt appearance could relect a genetically based neurological change that transformed behavioral po­ tential. A gradual development would imply a strictly environmental or demographic driver that afected only performance, not capacity. he last chapter emphasized that the case for abrupt development depends on the strong contrast between LSA sites and the overwhelming majority of MSA sites. Animal remains imply that LSA people hunted and gathered more efectively and that their populations were larger and denser. LSA sites also routinely contain formal bone artifacts and art ob­ jects, even when the samples are small. MSA sites rarely do, and the re­ corded instances involve mainly isolated objects that could be intrusive from overlying LSA layers. he case for gradual development depends mostly on two South African MSA sites where numerous precocious, LSA­like artifacts occur: Blombos Cave, which has provided bone arti­ facts, possible tick shell beads, and an engraved ocher lump from the Still Bay and/or pre­Still Bay MSA layers, and Diepkloof Rockshelter, which has produced incised ostrich eggshell fragments from near the top of the Howieson’s Poort MSA occupation. he Blombos and Diepkloof inds cannot be simply dismissed, but even if they are accepted, it remains true that on average, the MSA contrasts with the LSA in basically the same way that the Mousterian contrasts with the Upper Paleolithic in Europe. his still implies that behavior changed signiicantly at the MSA/LSA interface, and as discussed in the next section, the question then is what drove the change. he choice is mainly between new social or demo­ graphic factors on the one hand or biologically based cognitive change on the other.

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TABLE 7.4. African sites with ostrich egg shell beads that antedate 30 ka.

Site

Age (and dating method) 14

References

Enkapune Ya Muto, Kenya

39.9 ka ( C); 40–50 ka (hydration of associated obsidian artifacts)

(Ambrose 1998a)

Mumba Cave, Tanzania

33–29 ka (direct 14C dating of beads); >37–30 ka (14C dates on associated bones); 40–45 ka (amino­acid racemization of bead eggshell)

(Brooks 1996; Conard 2005; Hare et al. 1993)

Kisese II Rockshelter, Tanzania

ca. 32 ka (14C on ostrich eggshell)

(Deacon 1966b; Inskeep 1962)

Border Cave, South Africa

40–33 ka (concordant 14C and amino­acid racemization dates)

(Beaumont et al. 1978; Grün & Beaumont 2001; Miller et al. 1993)

Out of Africa predicts that Africa will contain the oldest secure evidence for art and other indicators of advanced (“modern”) behavior, and the Blombos and Diepkloof inds could be taken as substantiation. How­ ever, even if Blombos and Diepkloof are placed aside, Africa may still provide the oldest reliable dates for indisputable decorative items. hese are ostrich eggshell beads from the four sites listed in table 7.4. Unlike the Blombos shell beads, the artifactual quality of the eggshell beads is unequivocal since they were deliberately shaped, and at each site, radio­ carbon indicates they are older than 30 ka or even 40 ka. At Enkapune Ya Muto, Kenya, their actual age may approach 50 ka, based on the esti­ mated rate at which the surfaces of accompanying obsidian artifacts were altered in the ground. he ancient ostrich eggshell beads are notable not only because they document ancient personal ornamentation but also because they may signal the beginnings of an elaborate exchange system like the one recently observed among !Kung­San hunter­gatherers in the Kalahari Desert, Botswana. !Kung groups in diferent places regularly ex­ change beadwork, and the result is a network of relationships that en­ hances group survival in times of environmental stress. As discussed in the previous chapter, Üçağizli Cave, Turkey, has pro­ vided the oldest widely accepted decorative items outside Africa. hey comprise more than 500 perforated marine shells interpreted as beads, and they are accompanied by numerous shaped bone artifacts that might also be the oldest of their kind outside Africa. Radiocarbon dates the ensemble to at least 40 ka. Similar shell beads have also been recovered from what are probably like­aged layers at Ksar Akil Cave, Lebanon. he available dates from Europe tentatively suggest that ornaments, formal bone artifacts, and other markers of advanced behavior appeared on the east between 45 and 40 ka and only between 40 and 36 ka on the west. Many more observations are required to conirm a pattern, but the west Eurasian dates broadly suggest what Out of Africa would predict: popu­

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Arcy-sur-Cure

40/36 ka 40/36 ka 45/40 ka

FIGURE 7.6. hypothetical routes of modern human dispersal from africa beginning roughly 50 ka (partly after bar-Yosef [2002], fig. 1). The dispersal dates must be imprecise because the event occurred beyond the effective range of the radiocarbon dating method and currently alternative methods are less reliable.

45/40 ka

40/30 ka 45/40 ka

Taforalt

647

?45ka

45/40 ka

?45ka ?60 ka

Katanda

Enkapune Ya Muto

putative modern human epicenter 50-45 ka

45/40 ka

45/40 ka 45 ka Diepkloof Blombos

lation expansion from eastern Africa through southwestern Asia to eastern Europe and inally to western Europe (ig. 7.6). SOURCES: dating of Neanderthal appearance in Israel (Tchernov 1992, 1994); Mousterian of northern Africa (Alimen 1955; Balout 1955; Camps 1974; Marks 1968b; Schild 1998); formulation of the MSA in southern Africa (Goodwin 1928, 1929a; Goodwin and van Riet Lowe 1929); MSA dated to ater the Mousterian (Clark 1959b; Klein 1970); MSA dating at Border Cave (Beaumont and Vogel 1972; Vogel and Beaumont 1972); abrupt behavioral change 50 ka (Klein and Edgar 2002); gradual change between 100 and 50 ka (McBrearty and Brooks 2000); evolution of modern behavior among Neanderthals (d’Errico 2003; Zilhão 2001a, 2006); oldest LSA beads (table 7.4); historic !Kung­San exchange sys­ tems involving beadwork (Ambrose 1998a; Wiessner 1982); Üçağizli Cave shell beads and bone arti­ facts (Kuhn 2002; Kuhn et al. 2001); Ksar Akil shell beads (Mellars and Tixier 1989); Upper Paleolithic earlier in eastern Europe than in western (Hofecker 2005a; Mellars 1993, 1996, 2004)

The Relation between Biological and Cultural Change he archaeological evidence bearing on modern human origins is uneven and incomplete, but wherever it is full enough for judgment, it implies that human behavior changed dramatically about 50 ka. Before this time, morphology and behavior appear to have evolved slowly, more or less in tandem, but ater this time, morphology remained relatively stable while behavioral (cultural) change accelerated rapidly. Arguably, barring only the development of those typically human traits that produced the oldest known archaeological sites between 2.5 and 2 Ma, the behavioral trans­ formation that occurred 50 ka represents the most dramatic behavioral event that archaeologists will ever detect. It is oten referred to as the

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shit to fully modern behavior, for it is only ater 50 ka that the material residues of prehistoric foragers commonly anticipate those of historic foragers in every detectable respect. he most frequently cited “mod­ ern” behavioral markers are art and ornamentation, but new technolo­ gies that facilitated hunting and gathering were even more fundamental, for they enhanced human reproduction and survival and they promoted population growth. Most specialists would probably agree that the sub­ sequent growth propelled modern humans out of Africa and that their augmented ability to reproduce and survive explains the rapidity of their expansion across Eurasia. Some might also add that climate in the middle of the Last Glaciation could have facilitated the modern African expansion and the roughly simultaneous demise of nonmodern Eurasians, particularly the Nean­ derthals. It will be recalled from chapter 2 that the Greenland ice­cores record twelve­to­iteen closely successive, abrupt temperature oscilla­ tions between 60 and 25 ka. he warmer ones are known as Greenland Interstadials (or Dansgaard­Oeschger events ater their discoverers) and the colder ones as Greenland Stadials. he stadials grew colder and more frequent with time, and in some instances, the shit from stadial to in­ terstadial conditions occurred within a few decades. However, both ice cores and deep­sea cores show that broadly similar stadial/interstadial luctuations occurred before 60 ka within the earlier part of the Last Gla­ ciation and closely packed climatic oscillations also characterized prior glaciations, including the Glaciation­before­Last, between 186 and 130 ka. he earlier instances precipitated neither an expansion of modern hu­ mans from Africa nor the extinction of the Neanderthals. he difer­ ence ater 60 ka ago was the development of more sophisticated human behavior in Africa, and it is this, therefore, more than climate change, that is likely to have stimulated the modern human expansion. However, even if the major role is granted to new behavior, there remains the need to explain what prompted the behavior. Two major alternatives exist: irst, that people had been neurologically capable of such behavior long before 50 ka, but expressed their capacity only ater some momentous social or demographic change, or second, that they acquired the neural capability for the behavior only about 50 ka. Social change independent of biological (neural) change could have involved, for example, the initial development of the nuclear family as the fundamental productive unit and together with this, the division of labor by sex and age. Such a division was universal in historic hunter­gatherer societies, in which women and children tended to focus on plants and other gatherable foods and men emphasized hunting. Nuclear family or­ ganization and a newly developed division of labor might have led in turn to modern notions of kinship and descent that promote economic and political cooperation among individuals and groups. he emergence

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of modern hunter-gatherer social organization could thus have stimulated rapid population growth, and larger, denser populations could in turn explain the accelerating pace of technological innovation, the proliferation of symbols (art), and other novelties that mark the archaeological record ater 50 ka. his kind of explanation is logically coherent, but it fails in at least one crucial respect—it does not explain why social relations changed when they did or why they changed at all. Population growth could underlie the timing, if it is assumed that growth came irst and that behavior changed about 50 ka when popula­ tion density crossed a critical threshold that forced social reorganization. A major objection to this idea is that the advanced behavior presumably appeared irst in Africa, and archaeology suggests that African popula­ tions were shrinking, not growing, when it emerged. In southern Africa, where Blombos Cave is said to anticipate the new behavior by 75 ka, populations crashed within 15 ky aterward. he archaeological indica­ tions for small African numbers around 50 ka are consistent with genetic analyses, cited above, which suggest that the recent African population from which all living people derive included no more than 10,000 breed­ ing adults. Under these circumstances, it is at least as plausible to tie the ba­ sic behavioral shit, which may have included major changes in social organization, to a neurological change that launched the fully modern human ability to manipulate culture as an adaptive mechanism. A neural change, in turn, requires no special explanation since it would follow from the kind of fortuitous but highly advantageous mutation that must underlie all macroevolutionary change. he neural hypothesis in fact follows directly from the notion that selection for larger and presum­ ably more sophisticated brains was a vital aspect of human evolution long before the origin of modern humans and from the observation that earlier advances in human behavior, from the Oldowan to the Acheulean to the Middle Paleolithic, corresponded broadly to changes in brain size and probably also in brain organization. It is also in keeping with the discovery that genes involved in brain development evolved especially rapidly in the human line ater it split from the chimpanzee line 4–6 Ma. Fossils and genetics imply that the last crucial neural change occurred in Africa, but an African origin is not central to the argument. he underly­ ing mutation could have occurred in Europe, in which case, I myself and readers of this book would be Neanderthals contemplating the strange people who used to live in Africa.

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SOURCES: climate, modern human expansion, and Neanderthal contraction (Finlayson 2004; Mellars 2006b); century­to­millennial climate variability during glaciations before the last (Jouzel et al. 2007; Martrat et al. 2007); possible social or demographic causes for cultural change 50 ka (Kuhn and Stiner 2006; Sofer 1990a, 1994); modern hunter­gatherer social organization (Marlowe 2005); rapid evolu­ tion of a gene involved in human brain development (Pollard et al. 2006)

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Language and the Evolution of Modern Human Behavior Assuming for argument’s sake that a neural change explains the appearance of advanced (fully “modern”) behavior, it is reasonable to suppose that the change promoted the fully modern capacity for rapidly spoken phonemic speech, that is, for “fully vocal language—phonemicized, syntactical, and ininitely open and productive” (Milo and Quiatt 1993, 579). his inference follows not simply from the way living humans use pho­ nemic language to communicate but, even more, from the way they use it cognitively to model both nature and culture. Perhaps most important in the present context is that modern humans probably need fully modern language to ask the hypothetical “what if ” questions that underlie the modern capacity for innovation. Archaeological evidence for this capac­ ity burgeons ater 50 ka, and its appearance could explain why human itness—the ability to survive and reproduce—increased simultaneously. At least three lines of evidence can be brought to bear on the evolu­ tion of linguistic ability—historical linguistics, the anatomy of the vocal apparatus, and the genetic basis for language. Historical linguistics is rel­ evant if it is accepted that there are lexical similarities (that is, cognates or homologous words) among all modern languages and that these im­ ply a common linguistic ancestor or “mother tongue.” Only a minority of historical linguists recognize such lexical similarities, and they have not developed a method for estimating the age of the mother tongue. However, it would probably be undetectable if it existed before 50 ka, and it could thus represent the language of the African population from which all later people descend. Exceptional linguistic diversity in Africa may also place language origins there. he two previous chapters touched on the anatomy of the vocal ap­ paratus and particularly on the inferred position of the larynx (or voice box) in Neanderthals. he primitive position, which is retained in the apes and in human infants, is for the larynx to reside high in the neck. Such a placement sharply reduces the likelihood of choking since it al­ lows food and air to descend along separate digestive and respiratory tracts. Newborn humans run little risk of choking while they feed, but the danger is greatly enhanced between one and a half and two years of age when the larynx begins to drop and the digestive and respiratory tracts partially merge. his potentially dangerous arrangement is expli­ cable only if it confers a strong selective counteradvantage, and the most plausible one is that it enlarges the supralaryngeal space and thus allows humans to produce a wider range of sounds. Among these sounds are the vowels and consonants that separately or together characterize all known languages and that are essential for the production and decod­ ing of rapidly articulated speech. he larynx does not fossilize, but its adult position corresponds to the lexion or uparching of the adult cra­

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nial base. Modern human adults have highly lexed cranial bases that restrict the space available for the larynx, forcing it downward in the neck. In contrast, the only three Neanderthal skulls (from La Chapelle-auxSaints, Saccopastore, and Guattari [Circeo]) on which basicranial lexion can be assessed have relatively lat bases. his might mean that the Ne­ anderthals had a more highly placed larynx and a more limited ability to produce crucial sounds. It might also mean that they could not have beneited from a neural change that enhanced spoken language. In this, they would have difered from near­modern humans like those from Skhul and Qafzeh Caves, Israel, whose uparched basicraniums imply a fully modern laryngeal position. It is thus tempting to conclude that the Neanderthals were incapable of fully modern language and that this partly or wholly accounts for their extinction. However, the inluence of basicranial lexion on the position of the larynx is controversial, and laryngeal position is probably at least as closely tied to the form of the hyoid (or tongue) bone that lies just above the larynx in the throat. Two Neanderthal hyoids are known— from Kebara Cave, Israel, and El Sidrón Cave, Spain—and the previous chapter noted that they are essentially modern in form. So are two ad­ ditional hyoids from the much older proto­Neanderthals of Atapuerca SH, and the implication may be that adult laryngeal position was the same in Neanderthals as it is in living humans. In addition, even if basi­ cranial lexion closely relects laryngeal position, skulls that are as old as or older than the Neanderthals exhibit an essentially modern degree of basicranial lexion. his is true not only of the well­known near­modern skull 5 from Skhul Cave, Israel, but also of the much older fossil skulls from Petralona, Greece, and Broken Hill (Kabwe), Zambia, discussed in chapter 5. hus, even if we assume that only an uparched basicranium implies a lowered larynx and that a lowered larynx is essential to modern language, a change in basicranial lexion could not explain the modern human expansion 50 ka. If enhanced linguistic ability stimulated the ex­ pansion, the enhancement must have been in the brain and not in the vocal apparatus. he identiication of genes that bear on speech and language is just beginning, and only one is well­known so far. his is FOXP2, which codes for a transcription factor or regulatory protein that governs the embryonic development of the basal ganglia and other subcortical ele­ ments of the neural circuits involved in speech and language. Individuals who inherit a damaged (mutant) version of the FOXP2 gene exhibit spe­ ciic speech and comprehension deicits, and it was the presence of such individuals in three successive generations of a British family, known anonymously as KE, that allowed geneticists to pinpoint the importance of FOXP2 to speech and language. Analysis of variability in noncoding

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segments of the FOXP2 gene (that is, segments that do not afect the amino­acid composition of the FOXP2 transcription factor) revealed an unusual number of rare variants (alleles). he most parsimonious expla­ nation for the excess is that natural selection recently swept the current form of FOXP2 to ixation, which means that ater this form arose, those who possessed it enjoyed a substantial reproductive and survival advan­ tage. he advantage could have been an enhanced capacity for language, and for present purposes it would be crucial to know when the selective sweep occurred. Initial calculations, based on FOXP2 diversity in living humans, placed the sweep within the last 200 ky, ater the Neanderthal and mod­ ern human lineages had diverged, However, as noted above,modern FOXP2 has now been found in two Neanderthal bones from El Sidrón Cave, Spain, and if the fossil FOXP2 is authentic (as opposed to mod­ ern human contaminant), the selective sweep must have occurred more than 400 ka, in a population that was ancestral to both the Neanderthals and modern humans. Conceivably, this population authored the more sophisticated stone tools that marked the shit from the early to the late Acheulean discussed in chapter 5. It will be recalled that this shit oc­ curred in Africa perhaps 700–600 ka and that late Acheulean Africans, equipped with more sophisticated stone technology, then expanded their range to Europe. Previous chapters assigned the people involved to the species Homo heidelbergensis. However, even if it is clear that FOXP2 cannot bear on the much more recent modern human expansion from Africa about 50 ka, it still exempliies research that could since clinical investigations have isolated numerous other genes that are probably relevant to communication and cognition. he problem remains to determine their precise function and to overcome the probability that complex behaviors like language de­ pend on the interaction of many genes. It may also be that if a crucial genetic change occurred 50 ka, it was not in a gene or genes that encode a protein that afects cognition or communication, but in one or more noncoding genes that modulate the function of coding genes, and such a regulatory change might be particularly diicult to isolate and date. Dat­ ing in this context means determining whether natural selection is likely to have afected any candidate genes around 50 ka. Assuming that ongoing research reveals other genes that deeply in­ luence modern human communication or cognition and that the Ne­ anderthal nuclear genome becomes available, it should be possible to determine if Neanderthals and modern humans shared these genes. If not, the diference would provide circumstantial support for the neural hypothesis of modern human origins, for it would show that behaviorally relevant genetic change continued even ater the modern human lineage had emerged. An expectation to the contrary is implicit in hypotheses

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that link the modern human expansion to strictly social or demographic factors.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: language and culture (Bickerton 1990); the mother tongue (Ruhlen 1994a, 1994b, 1996); lar­ ynx position in humans and other primates (Aiello and Dean 1990, 236–243; Laitman 1985; Lieberman 2008, 64–65), efects on sound production (Lieberman 1991, 1992; Lieberman et al. 1992), and rela­ tion to basicranial lexion (Houghton 1993; Laitman 1985; Laitman et al. 1979); Neanderthals and proto­Neanderthal hyoids (Arensburg et al. 1989, 1990; Martínez et al. 2008); basicranial lexion in fossil humans (Franciscus 1995); inluence of FOXP2 on speech and language (Groszer et al. 2008; Lai et al. 2001; Lieberman 2006, 2007b; Vargha­Khadem et al. 2005); molecular evolution of FOXP2 (Coop et al. 2008; Enard et al. 2002; Zhang et al. 2002); recovery of FOXP2 from Neanderthal bones (Krause et al. 2007a); genes that regulate brain size and behavior and that might have played important roles in human brain evolution (Dorus et al. 2004); importance of noncoding regulatory genes (Carroll 2003); genes and cognition (Fisher 2006); possible recovery of the entire Neanderthal nuclear genome (Green et al. 2006)

Challenges to the Neural Hypothesis for Modern Behavioral Origins he greatest strength of the neural hypothesis is that it parsimoniously explains the burst of behavioral innovation that ushers in fully modern humans and that surely explains their spread. Its most obvious weakness is that, on present knowledge, it cannot be explored independently in fossils since the putative change was in brain organization, not size, and fossil skulls provide little or no irm evidence for brain structure. Nean­ derthal skulls, for example, difer conspicuously in shape from modern ones, but they were as large or larger, and the diference in skull shape does not imply a signiicant diference in brain function. No one doubts an essential link between the human brain and uniquely human behav­ ior, but paleoneurologists disagree on many key details of brain evolu­ tion, and the details are in any case not detectable in the fossil record. he only obvious work­around is to identify and analyze genes that bear on communication and cognition, as discussed in the previous section. Beyond the present diiculty of testing the neural hypothesis, it faces at least two other objections that are more strictly archaeological. Both are founded in the rich archaeological record of Europe, and in essence both concern the correlation between Neanderthals and Mousterian (Middle Paleolithic) artifacts on the one hand and between anatomically modern people and Upper Paleolithic artifacts on the other. he irst objection goes to the heart of the assumption that Neanderthals were biologically precluded from behaving in a fully modern way, and it is the more diicult objection to dismiss. he second concerns the possibil­ ity that advanced (fully modern) behavior did not appear abruptly with the advent of the Upper Paleolithic roughly 40 ka but evolved gradually aterward. SOURCES: skull shape and brain function (Holloway 1991b); human brain evolution (Deacon 1992, 1998; Falk 1992a, 1992b; Holloway 1998)

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Were Neanderthals Fundamentally Incapable of Fully Modern Behavior?

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As outlined here, Out of Africa postulates that the Neanderthals were replaced because they could not compete culturally with their modern human successors. he argument is bolstered over most of Europe by the relatively abrupt nature of the replacement. At many sites, Cro­Magnon/ Upper Paleolithic occupations overlie Neanderthal/Mousterian layers with no evidence for a major break in time or for any transition between the two, suggesting the replacement took only decades or, at most, cen­ turies. Demographic modeling shows that a 1% or 2% rise in Neanderthal mortality would have been suicient to extinguish Neanderthal popu­ lations within a thousand years, and the Cro­Magnons might have in­ duced such a rise simply by excluding the Neanderthals from essential resources. To accept this possibility, however, we must assume there was little or no gene low or cultural exchange between Neanderthal resi­ dents and Cro­Magnon invaders. As discussed in the previous chapter, some paleoanthropologists see fossil evidence for hybridization, but the indications are not compelling, and modern and fossil DNA so far argue strongly against it. Assuming that it was possible, it might have been largely precluded by a biologically grounded behavioral gulf between Neanderthals and Cro­Magnons. If this is accepted, the Cro­Magnon invasion of Europe would have difered fundamentally from the historic European invasion of the Americas or Australia, where the indigenes and invaders clearly had the same biological capacity for culture and in­ terbreeding was rampant. However, there is a signiicant problem with the idea that the Nean­ derthals could not behave like moderns. his is the occasional discovery of artifact assemblages that comprise a blend of Neanderthal/Mousterian and Cro­Magnon/Upper Paleolithic artifact types. At some sites such “mixed” assemblages may have been created when excavators inadver­ tently merged the contents of adjacent Mousterian and Upper Paleolithic layers, but this possibility is usually dismissed for sites of the Châtelper­ ronian Culture in central and western France and adjacent northern Spain. As discussed in chapter 6, Châtelperronian stone­artifact assem­ blages generally combine typical Mousterian sidescrapers, denticulates, and backed knives with numerous characteristic Upper Paleolithic end­ scrapers and burins. At one site, the singular Grotte du Renne at Arcy­sur­ Cure in the Paris Basin, typical Châtelperronian stone artifacts are ac­ companied by carefully shaped bone artifacts and by bone beads and pendants (ig. 6.60 in the previous chapter). he stone and bone artifacts were recovered from occupation loors with patterned arrangements of postholes, mammoth tusks, limestone plaques, and hearths that probably mark the positions of ancient huts (ig. 6.61 in the previous chapter). By themselves, the stone artifacts might be ambiguous, but the bone artifacts,

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ornaments, and highly structured loors point unequivocally to the Upper Paleolithic. Remarkably, as indicated in chapter 6, an associated human temporal fragment and twenty­nine isolated teeth exhibit Neanderthal (as opposed to Cro­Magnon) features. Neanderthal authorship of the Châtelperronian is implied even more surely at La Roche à Pierrot Rock­ shelter, Saint­Césaire, west­central France, where a partial Neanderthal skeleton was directly associated with typical Châtelperronian stone tools. Châtelperronian layers have been dated variously by radiocarbon, by thermoluminescence, and by correlation to regional or global climate stratigraphy. he results are somewhat inconsistent, but as discussed in chapter 6, a reasonable inference now is that the Châtelperronian existed for a few millennia between perhaps 45 and 36 ka. It was during this period that the earliest undeniable Upper Paleolithic culture or culture complex, known as the Aurignacian, appeared widely in southeastern, central, and western Europe. As summarized below, the Aurignacian is marked by multiple highly formalized, distinctive Upper Paleolithic stone­ and bone­artifact types and by a variety of art objects, including human and animal representations. At most sites where the Aurignacian and the Mousterian occur together, the Aurignacian immediately over­ lies the Mousterian, and in the version of Out of Africa favored here, the Aurignacian is a plausible artifactual manifestation of the Cro­Magnon invasion. Physically the makers of the early Aurignacian are poorly known, but sparse, fragmentary fossils from France and more numer­ ous and complete ones from Moravia, Czech Republic, suggest they were fully modern (rather than Neanderthal). How, then, to explain the Arcy Châtelperronian? he last chapter presented three possible alternatives. he least popular is that relatively crude excavation methods inadvertently mixed items from late Châtelper­ ronian layers that lacked bone artifacts and ornaments with items from overlying early Aurignacian layers that had them. his could explain why ity years ater Arcy was excavated, alone among the eighteen or so com­ monly accepted Châtelperronian sites, it has provided a wide variety of well­made Châtelperronian bone artifacts and ornaments. Aurignacian admixture could also explain why the Arcy Châtelperronian ornaments are so Aurignacian­like. A second possibility, also not widely accepted, is that the Arcy Châtelperronian documents an independent, Neander­ thal invention of the Upper Paleolithic. his interpretation fails to ex­ plain why the Arcy Châtelperronian remains unique and further why its well­made bone artifacts and ornaments appeared just as Aurignacian invaders were introducing similar items to the neighborhood. his leads to the third and probably most popular possibility—that Arcy relects cultural difusion from Cro­Magnon Aurignacians to Châtelperronian Neanderthals, before the Neanderthals inally succumbed. But even if this third alternative appears credible, it begs one fundamental question:

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If the Upper Paleolithic way of doing things was clearly superior and Neanderthals could imitate it (that is, they were not biologically precluded from doing so), why did they not acculturate more widely, with the result that they or their genes would have persisted much more conspicuously into Upper Paleolithic times (ater 40 ka)? here is no com­ pelling answer, and if the Arcy Châtelperronian was not created during excavation, it remains a puzzle whose solution is important for closure on Out of Africa.

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SOURCES: demographic modeling and Neanderthal extinction (Zubrow 1989); proposed fossil evidence for Neanderthal/Cro­Magnon interbreeding (Trinkaus 2005) and genetic evidence against (Krause et al. 2007a; Noonan et al. 2006); Châtelperronian stone artifacts (Harrold 1983, 1988, 1989); Châtelperronian bone artifacts and ornaments (Baier and Julien 1990; d’Errico et al. 1998b; Farizy 1990a, 1990b, 1994; Leroi­Gourhan 1965a; Taborin 1990, 2002; White 2001a); Arcy Châtelperronian human remains (Bailey and Hublin 2006; Hublin et al. 1996); Saint­Césaire human remains (Lévêque et al. 1993); earliest Aurignacian in Europe (Mellars 1993, 1996, 2006b; Straus 1993–1994); early Auri­ gnacian art and ornamentation (Bahn 1994; White 1989); early Aurignacian human remains (Bailey and Hublin 2005; Churchill and Smith 2000b; Frayer 1986; Gambier 1989; Hublin 1990; Smith et al. 1989); excavation methods at Arcy­sur­Cure (Schmider 2002; White 2001a); the Châtelperronian as an independent Upper Paleolithic development from the Mousterian (Zilhão and d’Errico 1999) or as a product of difusion from the Upper Paleolithic (Demars and Hublin 1989; Harrold 1983, 1989; Klein 1973b)

Does the Upper Paleolithic Truly Represent an Abrupt Departure?

he Upper Paleolithic contrasts with the Mousterian in many ways, of which the most oten cited is the widespread Upper Paleolithic emphasis on stone lakes whose length was at least twice their width. Archaeolo­ gists distinguish such elongated lakes as “blades.” Most Mousterian peo­ ple produced very few blades, and at Mousterian sites where blades are common, they were made mainly by a variant of the Levallois technique that many Mousterian knappers also used to produce lakes. In contrast, Upper Paleolithic people developed specialized techniques to produce blades regularly and consistently. In general, compared with Mousterian lake technology, Upper Paleolithic blade production provided more cutting edge from a given stone core, and among the irst Upper Paleo­ lithic people it may have helped to conserve scarce raw material. Later Upper Paleolithic people probably produced blades mainly for historical reasons, however, as a part of their cultural heritage. Besides emphasizing blades, most Upper Paleolithic people routinely manufactured stone­tool types that tend to be rare and crudely made (“atypical”) in Mousterian assemblages. he most commonly cited Upper Paleolithic types are probably endscrapers (elongated lakes or blades with smooth, continuous retouch on the edge opposite the striking plat­ form) and burins (lakes or blades from which a second smaller lake or blade (a burin spall) was struck along one edge, leaving a scar at an abrupt angle to the ventral surface of the parent) (ig. 7.7). Burins and

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657 FIGURE 7.7. Top: a burin and an endscraper on a blade, tool types that are conventionally said to distinguish the upper Paleolithic from the preceding mousterian. Middle and bottom: bifacial points, backed blades, and carinate (or keeled) endscrapers, types that singly or together especially distinguish the upper Paleolithic.

cm

dihedral burin

endscraper on a blade

leaf-shaped points with flat, invasive, bifacial retouch

backed blades

carinate endscrapers

endscrapers do characterize most Upper Paleolithic assemblages, but as truly diagnostic Upper Paleolithic types, they are probably surpassed by carefully made leaf-shaped points with lat invasive bifacial retouch; backed or truncated pieces on which a lateral edge or end has been methodically dulled or blunted; and carinate (keel-shaped) and nose-ended scrapers on which the presumed working edge has been formed by removing numerous small thin blades (“bladelets”). Singly or in combination, inely made leaf­shaped points, backed or truncated elements, and carinate or nose­ended scrapers distinguish many Upper Paleolithic industries, and they are correspondingly rare in Mousterian ones.

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Further, unlike Mousterians, Upper Paleolithic people commonly cut, carved, polished, or otherwise shaped bone, ivory, and antler into a wide variety of formal artifact types. hese include not only pieces that were probably projectile points, awls, punches, needles, and so forth but also nonutilitarian items that are clearly interpretable as art objects or items of personal adornment. As noted in the previous chapter, unequivocal art and decorative items are essentially unknown in a Mousterian context. In general, Upper Paleolithic people made a much wider variety of readily distinguishable artifact types than their predecessors did, and Upper Paleolithic industries or “cultures” varied far more in time and space. Finally, as detailed below, in virtually every conceivable category of recoverable items—graves, house ruins, ireplaces, and such—Upper Paleolithic examples are commonly far more elaborate than Mousterian ones. In all the ways that the Upper Paleolithic difers from the Mouste­ rian, the Mousterian difers much less from earlier cultures, and it thus seems reasonable to conclude that the Upper Paleolithic represents a quantum change from everything that went before. It has sometimes been suggested that an “early Upper Paleolithic” antedating 25–20 ka should be distinguished from a “late Upper Paleo­ lithic” aterward and that only the late Upper Paleolithic was signiicantly diferent from the Mousterian. In this view, the early Upper Paleolithic was transitional from the Mousterian and did not depart radically from it. he case is perhaps clearest for Cantabrian Spain (north of the Picos de Europa), where there was a dramatic increase in the number of sites ater about 20 ka, accompanied by an artistic lorescence and by sig­ niicant changes in the location of settlements and in hunting patterns. However, direct radiocarbon dating at Chauvet and Cosquer Caves now shows that Aurignacians or other early Upper Paleolithic people were producing spectacular wall art in nearby France more than 27 ka. French Aurignacian sites like Abri Blanchard, Abri Castanet, and La Souquette, antedating 30–28 ka, are also among the most proliic sources of Up­ per Paleolithic pierced animal teeth, shaped ivory and steatite beads, and other personal ornaments. Moreover, even if northern Spain and neighboring parts of western Europe appear quiescent before 20 ka, parts of central and eastern Europe were not. he cultural elaboration indicated by sites like Vogelherd, Hohlenstein­Stadel, Geissenklösterle, and Hohle Fels in southwestern Germany, Stratzing/Krems­Rehberg (Galgenberg) and Willendorf in Austria, Dolní Věstonice, Pavlov, and Předmostí in the Czech Republic, or Sungir’ in Russia, all antedating 20 or even 30 ka, rivals any later developments in Spain or France. More­ over, it could be argued that the “late” Upper Paleolithic of central Eu­ rope began with a cultural slump since site density and richness declined signiicantly about 20 ka and recovered to early Upper Paleolithic stan­ dards only ive or six millennia later.

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For the most part, the shiting locus of especially abundant, rich, or elaborate Upper Paleolithic sites probably relects the vicissitudes of Last Glaciation climatic change, which sometimes favored one area, some­ times another. It is surely signiicant that a major climatic inlection oc­ curred about 20 ka, when the Last Glaciation ice sheets reached their maximum. Following this time, the Scandinavian ice sheet on the north and the Alpine ice sheet on the south advanced to within 600 km of each other across central Europe. he intervening ice­free area became a frigid near­desert, and this surely explains why much of central Europe was so sparsely populated between roughly 20 and 14 ka. Paradoxically, however, from a human perspective the Last Glacial Maximum probably had its most dramatic efect in Africa, where people were able to reoc­ cupy areas they had largely abandoned before 40 or 50 ka, both in the north and in the south. he reason was a signiicant increase in regional precipitation. Together with temperature variation, similar, if less dra­ matic long­term moisture luctuations probably also contributed to local long­term population and cultural luctuations in Last Glacial Europe. his is not to say that all the known Upper Paleolithic traits were present from the very beginning or that all subsequent elaboration (or decline) was due directly to climate. Some of the speciic cultural inno­ vations discussed below unquestionably were developed ater the Upper Paleolithic began, and they difused more or less widely depending on geographic constraints, their local utility, and so forth. Undoubtedly some did promote local or regional population increases or cultural lores­ cences, but none were as fundamental as those that distinguish the Upper Paleolithic from everything before it. In fact, to the extent that an “early” Upper Paleolithic can be distinguished from a “late,” the apparent difer­ ences are less than those between many historic hunter­gatherer cultures, and nowhere is the early Upper Paleolithic truly a link or evolutionary transition between the Mousterian and the late Upper Paleolithic. In sum, from the characteristics that have been listed and that will be discussed further below, one can argue that the Upper Paleolithic and its African counterpart, the Later Stone Age, signal the most fundamental change in human behavior that the archaeological record may ever reveal, barring only the primeval development of those uniquely human behav­ iors that made archaeology possible. Excepting the puzzle posed by the association of Neanderthals with the late Châtelperronian at Arcy­sur­ Cure, the strong correlation between Upper Paleolithic artifacts and modern human remains clearly suggests that it was the modern human physical type that allowed the Upper Paleolithic (and all subsequent cul­ tural developments).

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SOURCES: blade manufacture—Mousterian (Kozlowski 1990a; Mellars 1996), Upper Paleolithic (Bordaz 1970); Upper Paleolithic stone­tool types (Kozlowski 1990a; Sonneville­Bordes and Perrot 1954, 1955, 1956a, 1956b); division of the Upper Paleolithic into early and late—general (Clark 1997c; Lindly and

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Clark 1990; Straus 1994) and in Cantabrian Spain (Straus 1977, 1990b, 1992; Straus and Heller 1988); early Upper Paleolithic wall art (Bahn 1995–1996) and ornaments (White 1989) in France; early Upper Paleolithic cultural elaboration in southwestern Germany (Conard 2003, 2005; Conard et al. 2003; Conard and Floss 2001; Hahn 1993; Hahn et al. 1985), Austria (Felgenhauer 1959; Neugebauer­Maresch 1988), the Czech Republic (Klíma 1963; Sofer et al. 1993; Svoboda 1994; Svoboda et al. 1994), and Russia (Bader 1978; White 1993); decline in site density and richness in central Europe beginning 20 ka (Hahn 1976; Kozlowski 1990c; Kozlowski and Kozlowski 1979; Otte 1990b; Roebroeks et al. 1992a; Svoboda 1990; Svoboda et al. 1994; Weniger 1990); glacial climatic change and Upper Paleo­ lithic elaboration (Guillien and Laplace 1978); Last Glacial Maximum human occupation in northern Africa (Camps 1974; Wendorf et al. 1979) and southern Africa (Deacon and hackeray 1984; Klein 1994)

Cultural Variability Previous chapters have emphasized that before 40 ka—that is, before the Upper Paleolithic and comparable cultural manifestations had com­ pletely supplanted earlier ones—vast areas were characterized by re­ markably uniform artifact assemblages that difered from one another mainly in the relative abundance of the same basic artifact types. In ad­ dition, artifactual change through time was painfully slow: basic assem­ blage types lasted tens or even hundreds of thousands of years. Ater 40 ka, however, the general pattern changed radically. Like­aged artifact assemblages from neighboring regions oten difered qualitatively, and within single regions the pace of artifactual change accelerated sharply. he greatly enhanced tendency to spatial variability is clearly illustrated by the divergence between the Upper Paleolithic of Europe, western Asia, and northern Africa on the one hand and the contemporaneous Later Stone Age (LSA) of sub­Saharan Africa on the other. Unlike their Middle Paleolithic and Middle Stone Age antecedents, whose separation relects scholarly tradition rather than artifactual content, the Upper Pa­ leolithic and the LSA difered artifactually from their very beginnings. he punched blades and varied burins that are a hallmark of the Upper Paleolithic never seem to have been an important element in the LSA. Instead, LSA people more commonly produced a range of scrapers and other retouched forms on lakes and lake blades. Some LSA retouched artifacts broadly resemble MSA ones, but they are commonly much smaller, and in the better known later LSA industries, postdating 20 ka, they are more standardized. In fact, to the extent that the LSA recalls the Upper Paleolithic, it is in relatively abstract features like greater artifact standardization, greater assemblage variation through space, accelerated assemblage change through time, and of course the routine production of formal bone artifacts and of art objects or items of personal deco­ ration. he sum suggests a common mindset that difered qualitatively from the mindset of earlier peoples. Within Africa, the Upper Paleolithic tendency to cultural diversity is particularly conspicuous in the Nile Valley, where diferences in stone­

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FIGURE 7.8. Typical aurignacian split-base bone points, “pendants,” and chipped stone artifacts. The split-base points (redrawn after bernaldo de Quiros Guidotti [1982], 98) come from the cave of el Castillo, northern spain. The remaining artifacts (redrawn after González echegaray [1980], 36, 39, 153) come from the nearby cave of el Pendo.

keeled or carinate endscrapers

perforated deer canine

661

imitation perforated deer canine in soapstone

endscrapers on Aurignacian blades

multiple dihedral burins 0

5 cm

split-base bone points

artifact types through time and space reveal no fewer than six cultures between 40–35 and 17 ka. hese include the Shuwikhat Industry and the Fakhurian, Kubbaniyan, Idfuan, Halfan, and Gemaian complexes, some of which existed in diferent parts of the Nile Valley at the same time. Within Europe, the highly diverse nature of Upper Paleolithic cultures is amply demonstrated by the numerous Upper Paleolithic sites of the east European plain, many of which have provided artifact assemblages that are unique and that cannot be assigned to an industry or culture repre­ sented at other sites, even in the same region. he dramatic acceleration in change through time is perhaps best exempliied by the classic Upper Paleolithic sequence of southwestern France. his was established long ago by careful excavations in caves such as La Ferrassie and Laugérie­Haute and has been repeatedly con­ irmed by meticulous modern excavations, including, above all, those

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FIGURE 7.9. a range of Gravettian (Perigordian) stone artifacts. backed bladelets are especially characteristic. The ones here (redrawn after bernaldo de Quiros Guidotti [1982], 145) come from the Gravettian deposits of Cueva morín, northern spain. The remaining pieces (redrawn after González echegaray [1980], 113) come from Gravettian levels in the nearby cave of el Pendo.

backed bladelets

Font Robert point 0

dihedral burin 5 cm

multiple mixed burin

denticulate

at the Abri Pataud. he sum shows that in the period between 40–35 and 11 ka, France harbored a remarkable succession of Upper Paleolithic industries, comprising most fundamentally (from older to younger) the Aurignacian, Gravettian (or Perigordian), Solutrean, and Magdalenian. Each industry (or culture) was characterized by speciic artifact types that are rare or unknown in the others. hus the Aurignacian, which intruded France from the east by 36 ka and was present until perhaps 28 ka, was distinguished by large blades with invasive, overlapping (“sca­ lar”) retouch on the lateral edges, “beaked” burins, nosed and keeled (carinate) endscrapers, and distinctively shaped bone or antler points, the most famous being those with split bases (ig. 7.8). he Gravettian spanning the period, between about 28 and 21 ka and extending eastward to European Russia and southward to Italy and Ibe­ ria, was marked especially by numerous small, narrow, parallel­edged, oten pointed, steeply backed blades (ig. 7.9). In western Europe later Gravettian people made characteristic tanged or stemmed points, while their central and eastern European contemporaries produced shouldered (or highly asymmetric stemmed) forms. Aurignacian­type bone points were absent throughout, and the principal bone artifacts were “awls,” “punches,” and other presumably domestic implements, accompanied by well­made bone art objects and items of personal adornment. he Solutrean, present between about 21 and 16.5 ka and essentially conined to France and Spain, was characterized above all by inely made foliate (leaf­shaped) stone points of various shapes and sizes (ig. 7.10). Some forms are regionally restricted, implying that local Solutrean groups

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FIGURE 7.10. solutrean artifacts from la riera Cave, northern spain (redrawn after straus and Clark [1986], 99, 103, 107). The solutrean is distinguished especially by various kinds of well-made unifacial and bifacial points.

shouldered points

fragments of laurel-leaf points

dihedral burin 0

5 cm

663

willow-leaf point

endscraper on a retouched flake

denticulated bladelets

developed their own particular styles. Finally, the Magdalenian, between about 16.5 and 11 ka, restricted initially to France and found later also in northern Spain, Switzerland, Germany, Belgium, and southern Britain, was distinguished primarily by a sophisticated bone and antler technol­ ogy that resulted in points, harpoons, and other implements and weapons (ig. 7.11), sometimes elaborately incised or decorated. As in the Solu­ trean, regional diferences in artifact form can be used to deine Mag­ dalenian subcultures. Some German Magdalenian sites may provide the oldest known evidence for domestic dogs. he late Upper Paleolithic of Europe seems to have been particu­ larly diverse, and while various kinds of Solutreans and Magdalen­ ians occupied the north and west, people who continued to emphasize Gravettian­like backed bladelets lived to the south and east in Italy, Hun­ gary, Bulgaria, Greece, and the former Yugoslavia. he artifact assem­ blages these people produced are oten lumped as “Epi­Gravettian.” he division between two major cultural groupings probably originated dur­ ing the last glacial maximum, around 20 ka, when much of central Eu­ rope became uninhabitable and Upper Paleolithic populations were split between refuges on the southwest and southeast. Within any given region, each major Upper Paleolithic industry not only tended to replace its predecessor far more quickly than earlier in­ dustries did, but artifactual change within each industry was also far more conspicuous. hus archaeologists can readily recognize distinctive

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FIGURE 7.11. magdalenian bone and stone artifacts from el Juyo Cave, northern spain. The magdalenian is distinguished above all by a wide variety of well-made bone and antler artifacts, including many decorated or artistic pieces.

El Juyo

needle

antler beveled-base “spear point”

antler “awl”

deer canine “pendants”

antler beveled-base “spear point”

dihedral burin pierced shell “beads” (string added)

burin on the corner of a snapped blade 0

bird bone tube Figure 7.11

borers

5 cm

retouched backed microblade microblade

subdivisions within the Aurignacian, Gravettian, Solutrean, and Magdalenian. he number and content of the subdivisions varies from region to region, thereby complicating formal deinition, but this only under­ lines the remarkable internal diversity of the Upper Paleolithic. Some of the extensive spatial and temporal variability that charac­ terized artifact assemblages ater 40–35 ka was undoubtedly functional, relecting the fact that some people began to manufacture new or dis­ tinctive artifacts because they had a new or distinctive purpose in mind. Much of the variability was probably stylistic, however, relecting cul­ turally diferent ways of doing the same thing. Although it may seem peculiar to us that material items should suggest so little cultural dif­ ferentiation before 40–35 ka, the amount that characterizes the succeed­ ing period is reminiscent of more recent history. he extensive cultural variability of the past 40–35 ky almost certainly required the existence of modern people, with their seemingly ininite capacity for innovation.

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SOURCES: LSA stone artifacts (Deacon 1984; Mitchell 2002; Wadley 1993); Nile Valley Upper Paleo­ lithic (Phillips 1994; Wendorf and Schild 1989); east European Upper Paleolithic (Anikovich 1992; Hofecker 1986, 1988, 2002; Klein 1969a, 1973b; Sofer 1985, 1989b, 1990b); Upper Paleolithic sequence of southwest France (Laville et al. 1980; Rigaud and Simek 1990; Sonneville­Bordes 1963, 1973); Abri Pataud sequence (Mellars et al. 1987; Movius 1975, 1977); later Gravettian stone points (Otte 1990b); regional Solutrean stone artifacts (Straus 1990b); dogs in the German Magdalenian (Weniger 1989); late Upper Paleolithic variability in Europe (Gamble 1986; Kozlowski et al. 1992; Montet­White 1994; Mussi 1990a; Runnels 1995); Last Glacial European refuges (Hublin 1998b); subdivisions of major Up­ per Paleolithic industries (Straus 1987, 1995)

The Late Paleolithic his section describes various aspects of culture in the period between 40–35 and 12–10 ka, when early anatomically modern people had com­ pletely replaced their predecessors. his period encompasses the later part of the Last Glaciation, and it corresponds to the timespan of the European Upper Paleolithic, as it is conventionally deined (ig. 7.12). For simplicity’s sake, the people who lived in this period will be referred to here as “late Paleolithic” to include both Upper Paleolithic people in Eu­ rope and their early­modern counterparts elsewhere in the world, par­ ticularly the contemporaneous LSA people of Africa. Economy

Although the emphasis here is on the diferences between late Paleo­ lithic (early­modern) people and their predecessors, economically they were broadly similar. All Paleolithic people lived entirely by hunting and gathering wild resources. Only in the transition from the Pleistocene to the Holocene, between roughly 11 and 9 ka, is there irm evidence for a signiicant change, when some people, particularly in southwestern Asia, began to domesticate animals, plants, or both. Even these people, however, probably continued to depend on wild resources for centuries, if not millennia. Ethnohistorically, most hunter­gatherers relied more on gathered, mostly vegetal foods than on lesh from hunting. Undoubtedly, late Pa­ leolithic people also depended heavily on plants, but plant residues are seldom preserved in Paleolithic sites. Table 7.5 lists ive exceptional sites, where charring and dense (anaerobic) deposits facilitated preservation, particularly at Ohalo II, or where water, sometimes chemically charged, was successfully used to “loat” of large numbers of seeds from the sedi­ mentary matrix (Franchthi Cave, Dolní Vĕstonice II, and El Juyo Cave). Ohalo II stands out from the others for the sheer volume of plant remains it has provided and for the remarkable abundance of wild grass seeds. hese demonstrate intense prehistoric interest in cereal grasses 10–12 ky before some were domesticated. Starch grains from barley and perhaps wheat that remained on a ground stone artifact conirm that wild cereal seeds were processed for food. Still, preservation conditions like those

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FIGURE 7.12. Climate and cultural stratigraphy after 186 ka. The “late Paleolithic,” as described in the text, occupied the later part of the last Glaciation, between 40–35 and 10 ka. This corresponds to the timespan of the european upper Paleolithic.

O-isotope stages

666

ka

1 11

10

20

Rough Indication of Average Temperature cold

warm

Glaciations & Interglacials Present Interglacial

Traditional French Stages

European Cultural Stratigraphy

Holocene

Neolithic, etc. Mesolithic

Würm IV Würm III/IV

2

24

Upper Paleolithic

Würm III 30

west 40

3

Last Glaciation

east

Würm II/III

50

57

Würm II

60

4 71

70

80

5a

90

5b

100

5c

110

5d

120

5e

Würm I/II

Last Interglacial

Würm I

Mousterian (= Middle Paleolithic)

Riss/Würm

127 130

6

Penultimate Glaciation

Riss

186

at Ohalo II are rare, and it is understandable that archaeologists have probably overstressed the importance of hunting in Paleolithic economies. he overemphasis is probably most misleading for low­latitude sites where gatherable plant foods were probably always more abundant relative to game. It is less misleading for middle­ and upper­latitude sites where game animals were relatively more numerous, particularly during glacial intervals. Game was especially abundant at middle latitudes in late Paleolithic Eurasia because glacial cold and aridity favored the formation of grassy steppes over vast areas where forests prevailed during interglacials, includ­ ing the Present or Holocene Interglacial. he principal steppe species were gregarious ungulates such as the woolly mammoth (Mammuthus primigenius), reindeer (Rangifer tarandus), bison (Bison priscus), horse (Equus prezwalski), and saiga antelope (Saiga tatarica) that could never have pros­

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TABLE 7.5. Late Paleolithic sites with remains of plants that people probably introduced. At El Juyo and Franchthi

Caves, rodents or birds could also have been partly responsible. Estimated ages are in uncalibrated radiocarbon years. Site

Age

Plant Remains

References

Ohalo II. Israel

19 ka

>90,000 charred plant remains from 142 taxa, including grasses, acorns, almonds, pistachios, wild olives, raspberries, wild igs, and wild grapes. he grasses included both wild barley and wild wheat.

(Nadel et al. 1994; Nadel et al. 2004; Piperno et al. 2004; Weiss et al. 2004)

Franchthi Cave, Greece

22–10 ka

Seeds from deposits > 13 ka may have been naturally introduced; those in deposits < 13 ka come mainly from wild lentils, vetches, pistachios, and almonds that were probably intro­ duced by people.

(Hansen 2001; Hansen & Renfrew 1978)

Dolní Vĕstonice II

26 ka

Charred remnants of roots or tubers and a seed “loated” from a hearth

(Mason et al. 1994)

El Juyo Cave, Spain

14–13 ka

853 identiied seeds or other plant parts, including fragments of acorns, hazel nuts, and raspberry pits.

(Freeman et al. 1988)

Niah Great Cave, Borneo

27–9 ka

Charred nut and fruit tissues and starch grains from yams and sago.

(Barker 2002; Barker et al. 2003; Barker et al. 2007)

pered in forested environments. Reindeer reached the far south of France, and some even penetrated the Pyrenees to northern Spain. In France, in keeping with independent evidence for especially cold climate, reindeer were particularly common during the early Aurignacian (before 30 ka) and again during the Solutrean and Magdalenian (between roughly 20 and 11 ka). heir bones dominate many French Magdalenian sites, includ­ ing such well­known ones as Verberie, Pincevent, Laugérie­Haute, La Madeleine, and Isturitz. Because of regional environmental diferences, other species prevail in contemporaneous sites elsewhere—for example, red deer (Cervus elaphus) at La Riera, El Juyo, Tito Bustillo, and other sites in northern Spain. and bison at Amvrosievka, Bol’shaya Akkarzha, Zolotovka I, and other sites on the south Russian plain—but everywhere late Paleolithic people emphasized large gregarious herbivores. In contrast to large herbivores, large carnivores such as lions, hye­ nas, and bears rarely occur in late Paleolithic archaeological sites, though one or more large carnivore species coexisted with people everywhere in Eurasia and Africa. At Istállóskö Cave in Hungary, Bacho Kiro Cave in Bulgaria, and other central or southeastern European early Upper Paleolithic (Aurignacian) cave sites, cave bear bones abound, but these are spatially or stratigraphically separate from the human occupational debris, and they probably result from natural deaths during hibernation.

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Human population growth and more intensive human use of caves probably extinguished the cave bear over most of its range in the later Upper Paleolithic (by 25 ka), but in general, Upper Paleolithic people and large carnivores probably avoided one another since lose­lose was the most likely result of confrontation. Large herbivores provided not only food but also critical raw mate­ rials, including hides, sinew, and, most obviously, bone, antler, or ivory. Late­Paleolithic people used bone, antler, and ivory to fashion a wide variety of implements and art objects, and they employed larger pieces as weights, supports, and other structural elements in their dwellings. Pat­ terned arrangements of large bones constitute key evidence for dwellings at the Gravettian sites of Krakow­Spadzista Street B in Poland, Pavlov I and Milovice in the Czech Republic, and at Mezhirich, Yudinovo, Mezin, and other Upper Paleolithic sites in Ukraine and Russia. In parts of cen­ tral and eastern Europe where trees were especially scarce, the people even used fresh bone for fuel, judging by the large amount of bone ash and charred bone fragments in their ireplaces. Like Native Americans on the American Great Plains, they probably also burned the dried dung of large herbivores. Although it is probable that late Paleolithic people exploited local resources more efectively than earlier people did, irm evidence remains elusive. In western Europe perhaps the best indication is that, per unit time, Upper Paleolithic sites are much more numerous than Mousterian ones, implying that Upper Paleolithic populations were larger and denser. he principal alternative—that Upper Paleolithic people simply moved camp more oten—seems unlikely since Upper Paleolithic sites tend to be richer and more extensive than Mousterian ones. Since Upper Paleolithic and Mousterian people lived under broadly similar conditions, Upper Paleolithic people could have been more numerous only if they used local resources more eiciently. heoreti­ cally, this should be relected in contrasts between Upper Paleolithic and Mousterian food refuse, but systematic studies of the ungulate re­ mains that dominate everywhere so far show no consistent diferences in prey species choice, skeletal part representation, butchering methods, or mortality proiles. he problem may be that Upper Paleolithic people were distinguished primarily by greater hunting success, grounded in more advanced technology, and the analytic methods that archaeologists apply to bone assemblages do not measure the success rate. At the mo­ ment, the clearest diference is that Upper Paleolithic people seem to have ished and fowled more oten, and more sophisticated technology could again be the reason. Upper Paleolithic sites tend to contain more ish and bird bones, and stable carbon and nitrogen analysis of human bones implies that some Upper Paleolithic people consumed ish and aquatic birds routinely, while no Neanderthals did. Increased reliance on

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ish may especially account for late Upper Paleolithic lorescence in parts of northern Spain, southwestern France, and southern Russia, where an­ nual salmon runs probably provided knowledgeable hunter­gatherers with an unusually rich and reliable resource. It has sometimes been suggested that some Upper Paleolithic people were unique or advanced in their tendency to specialize on just one or two herbivorous species, such as reindeer in northwestern and southwestern France or red deer in northern Spain (ig. 7.13). However, some Mouste­ rian levels in southwestern France are dominated by a single species, for example, by reindeer at Combe Grenal and Les Pradelles (Marillac), by aurochs (Bos primigenius) at La Borde, and by bison at Mauran. One or two species also strongly dominate some Mousterian sites elsewhere, for example, bison at Wallertheim and reindeer at Salzgitter­Lebenstedt in Germany; horses at Zwoleń in Poland; wild asses, saiga antelope, or both at Starosel’e, Kabazi II and other sites in Crimea (Ukraine); and bison at Rozhok I, Sukhaya Mechetka, and Il’skaya on the south Russian plain (locations in ig. 6.2 in the previous chapter). More fundamentally, in virtually all cases where bones of a single species predominate at an Upper Paleolithic or earlier site, the species could also have dominated the ancient environs of the site, and the bones therefore need not demonstrate human hunting specialization or prefer­ ence. he extraordinary abundance of reindeer at late Upper Paleolithic sites in southwestern France is a case in point since it almost certainly relects climatic deterioration that all but eliminated other herbivores. Finally, it is not necessarily true that people hunted or even ate the spe­ cies whose bones are most common at a site. his is most obvious with respect to the woolly mammoth, whose bones sometimes dominate open­air Upper Paleolithic sites in central and eastern Europe. In most cases the bones appear to have been used extensively as building mate­ rial, as fuel, or as raw material for artifact manufacture, and they may simply have been scavenged from long­dead animals. his possibility is directly implied at the artifactually unique late Upper Paleolithic site of Mezin (Ukraine), where chemical analysis shows that the mammoth bones composing a “ruin” came from individuals that probably died de­ cades apart. It is also suggested at the eastern Gravettian site of Krakow­ Spadzista Street B (Poland), where many of the mammoth bones used to build three structures were conspicuously gnawed by carnivores. At many other sites it may be indicated by diferences in supericial bone weathering, although diferential exposure of bones ater site abandon­ ment might produce the same result. As discussed in chapter 6, the extreme southern tip of Africa has provided perhaps the best evidence for a late Paleolithic advance in re­ source exploitation. In brief summary, unlike their MSA predecessors, local LSA people living under similar environmental conditions actively

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horse

red deer

roe deer

bison

El Juyo 4

79/3

3925/44

178/4

94/4

El Juyo 6/7

31/1

2627/78

85/4

32/2

El Juyo 8/9

34/3

1289/17

76/5

25/1

Altamira 2

3/1

972/15

19/3

25/2

0

50%

boar

ibex

polecat

lion

El Juyo 4

40/2

1/1

El Juyo 6/7

47/4

1/1

El Juyo 8/9

2/1

fox

wolf

bear 10/2 1/1

2/1

Altamira 2

1/1

hedgehog

1/1

22/4

3/1

10/2

1/1

17/3

1/1

2/1

FIGURE 7.13. abundance of various mammal species in layers 4, 6/7, and 8/9 at el Juyo Cave and in layer 2 at nearby altamira Cave, northern spain. The associated artifacts belong to the local “lower” magdalenian, bracketed between roughly 15 and 13 ka. The bars represent the percentages of each species in each layer, as measured by the minimum number of individuals (mni) necessary to account for the bones of each species. The numbers accompanying the bars are the number of specimens assigned to each species divided by the mni. The chart illustrates the abundance of red deer and the rarity of large carnivores, which are characteristic aspects of most magdalenian sites in northern spain.

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ished and fowled, and they obtained dangerous game such as bufalo and wild pigs much more oten. In addition, on average, LSA shellish and tortoises are signiicantly smaller than their MSA counterparts, and the most economic explanation is that LSA pressed harder on the spe­ cies because LSA populations were larger and denser. LSA advances in resource exploitation were probably due mainly to technological inno­ vations, including the development of ishing and fowling gear and also of snares or weapons that reduced the hunters’ exposure to danger. he contrasts are stark, but unfortunately it is not yet clear that they charac­ terize the very earliest LSA people, whose sites are locally rare, probably because they date from a time (between 60 and 30 ka) when hyperaridity had greatly reduced human numbers. It has occasionally been suggested that very late Paleolithic Europeans managed or domesticated reindeer and even the horse, but compelling bony evidence is available only for the dog (Canis familiaris). By ethno­ historic analogy, late Paleolithic people probably used dogs as sentinels, hunting partners, drat animals, or even food. Analyses of mitochondrial DNA show that dogs derive exclusively from the gray wolf (Canis lupus) and that the original domestication event or events occurred in Eurasia, perhaps in eastern Asia at about 15 ka. Determining the exact time and place is complicated by the likelihood of persistent backcrossing between wolves and dogs. To begin with, dogs difered from wolves primarily in a shortening and broadening of the muzzle, and the oldest skulls that unambiguously exhibit this characteristic come from the Eliseevichi I Epigravettian site in the Desna Valley, Russia. Radiocarbon dates bracket Eliseevichi I between 17 and 13 ka. German Magdalenian sites dated to roughly 14 ka have provided less securely identiied dog bones. Among them, a partial mandible from Oberkassel is the most persuasive. Dogs are well­documented at widely scattered European, west Asian, and North American sites that postdate 10 ka. SOURCES (also table 7.5): domestication of animals and plants in western Asia (Bar­Yosef and Meadow 1995); mammalian fauna of the Eurasian glacial steppes (Guthrie 1990, 1996); reindeer abun­ dance and glacial climate in France (Boyle 1990); French sites with abundant reindeer (Audouze 1987; Audouze et al. 1989; Bahn 1983; Bouchud 1959; Delpech 1975, 1983, 1989); Spanish sites with abundant red deer (Altuna 1972, 1992; Clark and Straus 1983; Straus 1992; Straus and Clark 1986); south Russian sites with abundant bison (Hofecker 1986; Hofecker et al. 1991; Klein 1973b; Sofer 1990b); cave bear bones in central European Aurignacian sites (Allsworth­Jones 1990; Kozlowski 1982, 1990b); extinc­ tion of the cave bear (Anderson 1984; Estévez 2004); bones as Upper Paleolithic construction material in Poland (Kozlowski 1983; Kozlowski et al. 1974), the Czech Republic (Klíma 1963; Oliva 1988), and Ukraine and Russia (Gladkih et al. 1984; Hofecker 1986; Klein 1969a, 1973b; Sofer 1985, 1989b); Upper Paleolithic use of animal bone for fuel (Villa et al. 2002); Mousterian and Upper Paleolithic site num­ bers per unit time (Clark and Straus 1983; Mellars 1973, 1982; Straus 1977); similarity between Mouste­ rian and Upper Paleolithic sites in ungulate prey (Gaudzinski 1998; Grayson and Delpech 2003; Steele 2004); increased ishing and fowling in the Upper Paleolithic (Pettitt et al. 2003; Richards et al. 2001); possible Upper Paleolithic specialization on one or two ungulate species—Upper Paleolithic (Mellars 1973, 1989b; Straus 1977; White 1982a), Mousterian sites where one species dominates—Combe Grenal (Bordes and Prat 1965; Chase 1986, 1989), Les Pradelles (Costamagno et al. 2006), La Borde (Jaubert

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et al. 1990), Mauran (Farizy and David 1992; Farizy et al. 1994), Wallertheim (Gaudzinski 1992, 1995), Salzgitter­Lebenstedt (Butzer 1971; Gaudzinski and Roebroeks 2000; Tode et al. 1953), Zwoleń (Gautier 1989; Schild et al. 1988), Starosel’e, Kabazi II et al. (Burke 1999a, 1999b, 2006; Burke et al. 1999; Chabai et al. 1995; Klein 1969b); Rozhok I, Sukhaya Mechetka, and Il’skaya (Hofecker 1987; Hofecker et al. 1991); reindeer abundance and climate in southwestern France (Rigaud and Simek 1990); Paleolithic scavenging of mammoth bones at central and east European sites—general (Klein 1973b; Sofer 1985, 1990b), at Mezin (Pidoplichko 1969), at Krakow­Spadzista Street B (Kozlowski and Kubiak 1972); advanced late Paleolithic resource exploitation in southern Africa (Klein 1994; Klein and Cruz­Uribe 1996); possible late Paleolithic domestication of reindeer or horse (Bahn 1983); genetic origins of the domestic dog—general (Vilà et al. 1997), and in Eurasia (Leonard et al. 2002), perhaps eastern Asia (Savolainen et al. 2002); dog bones from Eliseevichi I (Sablin and Khlopachev 2002) and Oberkassel (Benecke 1987)

Technology

Late Paleolithic people were far more inventive than their predecessors, and they made technological innovations at an unprecedented pace. It is these innovations that are particularly stressed here. Many remain impre­ cisely dated, and even when dates are available it seems unlikely that the oldest known occurrences were actually the irst. For this reason, and be­ cause speciic dates are not crucial to the main point, the discussion will treat the entire late Paleolithic period 40–10 ka, as a single unit. Figure 7.14 outlines the known chronology of major technological innovations in four major regions of Eurasia. he outline is inevitably tentative since fresh excavations are likely to push back the known ages of major inno­ vations in various places. In general, late Paleolithic artifact assemblages contain a much wider range of recognizable artifact types than do earlier ones, suggesting that late Paleolithic people were engaged in a wider range of activities. hese clearly included shaping formal bone, ivory, and antler artifacts, and they probably also involved manufacturing more tools for the creation of other tools rather than for immediate use as hide scrapers, projectile points, and so on. To judge by the small size or the shape of many late Pa­ leolithic stone artifacts and by the form of many bone pieces, the people probably also manufactured many more composite tools, that is, imple­ ments combining separate pieces of stone, bone, or other materials. Un­ fortunately, because the composite tools were held together mainly by perishable glues, leather thongs, and so forth, few have survived intact. Although Mousterians and even earlier people had adapted to gla­ cial climates in central and western Europe, late Paleolithic people were the irst to inhabit the harsh environments of easternmost Europe and northern Asia (Siberia) (ig. 7.15), where the winters were exceptionally long and cold even during interglacials. Upper Paleolithic innovations for dealing with intense cold probably included both better clothing and better housing. Winter clothing almost certainly incorporated fur, and east European Upper Paleolithic sites including, for example, Mezin, Mezhirich, Eliseevichi I, Avdeevo, and Kostenki XIV have provided the

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WESTERN EUROPE

CENTRAL EUROPE

12

bow and arrow

14

fishhooks domesticated dog harpoon heads

16 18 20 22 24 26

spearthrower self-barbed points stone lamps with handles eyed needles

28 30

38

notational systems? wind instruments fat-burning stone lamps pigment recipes split-base points

40

bone awls

32 34 36

42 44

EASTERN EUROPE

SOUTHERN SIBERIA

harpoon heads fishhooks

cordage, basketry & textiles fired clay & kilns storage pits eyed needles

hunting traps domesticated dog mammoth bone houses

throwing darts

tailored clothing cordage fat-fueled bone lamps large winter houses w/ cold-storage pits tailored clothing heated shelters eyed needles bone shovels

heated shelters eyed needles

bone points snares? fire dills? bone awls

bone awls bone points

split-base points

673 FIGURE 7.14. The chronology of major late Paleolithic technological innovations in eurasia, as established by archaeology (modified after Hoffecker [2005b], fig. 4). The interpretation of some items is controversial, and this pertains above all to sequentially marked bones that may represent counting or notational (information-storage) systems (d’Errico et al. 2003) or even lunar calendars (Marshack 1972b). Future excavations will undoubtedly show that the chart underestimates the time when some innovations appeared.

oldest known evidence for systematic fur trapping. Each site contains an unusual number of wolf or arctic fox bones, and the bones tend to occur as either whole or nearly whole skeletons lacking the paws or as articulated paw skeletons, occurring separately. he implication is that the people removed the feet with the skins, as modern trappers oten do, and then discarded the skinned carcasses. he “awls,” “punches,” and other pointed bone objects that appear in even the earliest Upper Paleolithic sites could have been used to sew skins together, and Kostenki 15 on the Russian plain and Tolbaga in south­ central Siberia have provided the oldest known eyed needles, dated be­ tween 35–30 ka and 35–28 ka, respectively. he next oldest example is from the eastern Gravettian site of Předmostí, Czech Republic where it was made about 26 ka. Specimens that postdate 19–18 ka have been found across Europe. he existence of sophisticated, well­tailored cloth­ ing is directly documented in some Upper Paleolithic burials, especially those found at the site of Sungir’, 192 km northeast of Moscow. Here, dis­ colored soil and strings of ivory beads surrounding, girdling, and paral­ leling the skeletons of three people buried between 26 and 19 ka suggest the details of fur or leather garments, comprising a cap, a shirt, a jacket, trousers, and moccasins. he beads and other objects found with the skel­ etons were apparently sewn on the clothing as decorations or fasteners. Moccasins or other supportive footwear could further explain why the Sungir’ people had slim (nonrobust) toes relative to the overall ro­ busticity of their legs. An equally slender middle toe may indicate even

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FIGURE 7.15. siberia, showing the approximate locations of the mousterian and upper Paleolithic sites mentioned in the text (adapted from vasil’ev [1993], fig. 1). Mousterian and most early/middle Upper Paleolithic sites are restricted to temperateSiberia, below55°N. Late Upper Paleolithic sites are also common in the subarctic and arctic regions, above 55°N.

▲ ▲ Yana Berelekh ▲ Mayorych

Ikhine ▲

Ezhantsy ▲ ▲Kukhtuj I. Ust’-Mil’ ▲ IN ▲ Diring Yuriakh AL Verknetroitskaya H K ▲ Dyuktaj SA Aldan R.

SIBERIA

Ob ’ R.

R.

▲

i R. ese Yen

Afontova Gora

Ir tys h R.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

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. Angara R ●

Le na

674

▲Mamakan ▲

▲ ● ● ▲ ▲ ▲ Mal’ta, ▲ ▲▲ ▲ Buret’ L. BAIKAL ❑▲ ▲ Kokorevo Tolbaga ▲▲ ▲ ▲ ■❑ ●●■ ■▲ ■ ▲ ▲ ● ▲ ▲ ❑ ■ Malaya ▲ ▲ ❑■■ Syya ▲ ■ ● ▲ Kara-Bom ■■ Maloyalomanskaya

Denisova, Okladnikov

CHINA

MONGOLIA

❑ Mousterian (> 40 ky ago) ■ Early Upper Paleolithic (40–30 ka) ● Middle Upper Paleolithic (30–20 ka) ▲ Late Upper Paleolithic (20–10 ka)

0

200 km

older footwear for the modern people who inhabited Tianyuan Cave, near Zhoukoudian, northern China, about 35 ka. A molecular clock (ge­ netic) estimate of 72 ± 42 ka for the time when the human body louse (Pediculus humanus corporis) diverged from the head louse (P. h. capitis) may imply yet greater antiquity for tailored clothing. he body louse feeds on the skin but lives in close­itting clothing, and it evolved from the head louse, which lives and feeds exclusively on the scalp. Like their predecessors, late Paleolithic people oten occupied rock­ shelters or cave mouths in regions where these were available, such as southwestern France and northern Spain. Meticulous modern excavations like those at Cueva Morín and El Juyo in northern Spain show that they sometimes built walls or otherwise modiied natural shelters to make them more habitable. Over much of their range, however, late Paleolithic people did not have access to caves, and even where they did, they un­ doubtedly also camped in the open air. By analogy with historic foragers, they surely built wood or brush huts at many open­air sites, but most de­ composed long ago in the ground. he 19­ky­old lakeshore encampment at Ohalo II, Israel, provides a rare glimpse of what is usually missing. Ohalo II is a classic late Paleolithic site in every predictable respect, including numerous well­made bone artifacts, abundant shell beads, some from species that had to be imported from the Mediterranean coast, 55 km to the west, large numbers of bones from small ish that the people caught in the nearby lake, and a relatively elaborate burial of an adult male. he deposits are virtually unique, however, because they

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have been waterlogged since they formed, and they consequently preserve a wide array of plant remains. he presence of thousands of grass seeds, acorn fragments, and other plant remains has already been noted, for they directly document what archaeologists usually only suspect— that late Paleolithic foragers relied heavily on plant foods. he relevant point here is that the plant fossils cluster with numerous lint artifacts and fragmented animal bones in oval depressions, 10–20 cm deep, 2–5 m across, and 5–13 sq m in area. In all, six such depressions have been par­ tially or wholly excavated, and in addition to plant food debris, each con­ tains thick fragments of tamarisk, willow, and oak branches that could have framed the roof of a now­collapsed structure. here are also smaller twigs and stems from other plants that could have illed the gaps between branches. Finally, the loor of the most thoroughly investigated depres­ sion preserved a spread of loosely interwoven grass stems surrounding a difuse expanse of ash. he grass stems have been interpreted as bedding (mattress) material around a central ireplace, and the sum suggests that each depression marks a brush hut, the oldest so far on record. he huts were apparently all destroyed by ire, and the charring helped to preserve plant tissues. Ohalo II remains unique, but many European late Paleolithic sites provide indications of structures that were remarkably substantial by comparison to ones known or inferred from earlier times. For the most part, only the foundations survive because the superstructures were presumably made mainly of wooden poles, hides, and other perish­ able organic materials. he most spectacular examples come from the harsh open plains of central and eastern Europe where they variously comprise large, artiicial depressions, regular arrangements of postholes, patterned concentrations of large bones or stone blocks (serving as con­ struction material), sharply deined concentrations of cultural debris, or a combination of these features. Between perhaps 18 and 13 ka, late or Epi­Gravettian people built houses largely of mammoth bones (igs. 7.16 and 7.17), and the remnants have been found at no less than fourteen sites, from Milovice (Czech Republic) on the west to Kostenki (Russia) on the east. he structures were commonly accompanied by deep pits that penetrated the frozen subsoil and that served at least in part to re­ frigerate lesh and bones intended for fuel. Wood was rare or absent, and fresh bone was probably the best available substitute. he various types of “ruins” generally border, encircle, or cover patches of ash and charcoal that mark ancient ireplaces for heating and cooking (ig. 7.18). Some of the ireplaces are more complex than any ear­ lier ones and have intentionally corrugated loors or small ditches lead­ ing out from the central ash accumulation. Modern experiments show that these modiications increase oxygen low and thus produce a hotter lame. he eastern Gravettian people who occupied Kostenki 1, Russia,

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FIGURE 7.16. Top: roughly circular concentration of large bones believed to mark the base of a ruined hut at the upper Paleolithic (epigravettian) site of mezin (complex 1) in ukraine. radiocarbon dates on associated mammoth teeth and shells scatter between 29.7 and 15.1 ka, but the artifacts support an age near 15.1 ka (hoffecker 2002). Below: hypothetical reconstruction of the hut, showing the bones used as weights to hold down skins stretched over a wooden framework (redrawn after boriskovskij [1958], fig. 1).

0

5m

Mezin Complex 1

concentration of large bones

about 25 ka also appear to have heated or lit their homes with mammal­ fat lamps fashioned from the hollowed­out heads of mammoth femurs. Other late Paleolithic Europeans, especially in southwestern France, fash­ ioned fat­fueled lamps from limestone and sandstone slabs. he oldest examples antedate 30 ka, and ones that postdate 20 ka are sometimes decorated and furnished with handles. As noted below, some French spec­ imens were surely used to illuminate the cave walls on which late Paleo­ lithic people painted and engraved. For the most part, we do not know exactly what late Paleolithic peo­ ple did with their individual artifacts, but it is safe to assume that they used them mainly to obtain or process foods and raw materials. Given the resemblance of stone burins to modern metal engraving tools by the same name, it seems likely that many were used to shape and incise bone, ivory, or antler. Stone endscrapers were probably used to process hides, perhaps particularly in the initial removal of hair and fat. Endscrapers from the Magdalenian site of Verberie in the Paris Basin exhibit polishes and striations that abundantly conirm hide working, and many also exhibit wear from friction in bone handles. Spatulate bone “burnish­ ers” may also have been used for hide processing, perhaps particularly for smoothing or for spreading sotening agents or pigment. Enigmatic bone objects found at many sites may have been trips or other parts of compound traps for snaring foxes, hares, and other valuable fur­bearing animals.

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FIGURE 7.17. Top left: Plan view of a circular heap of mammoth bones used to construct a hut at the upper Paleolithic (epigravettian) site of mezhirich in ukraine about 15.6 ka. Top right: distribution of artifacts, fragmentary animal bones, and other debris beneath the bone heap. Bottom left: Conjectural reconstruction of the hut (adapted from Gladkih et al. [1984], 165). in total, excavations at mezhirich have uncovered four round or oval mammothbone ruins from huts that may have been occupied simultaneously, although probably mainly in winter (soffer et al. 1997). The bones came from at least 149 mammoths, but the people could have obtained them from animals that died naturally.

heap of mammoth bones

hypothetical partial reconstruction

hearths

6m

0

Mezhirich

flint nodules & fragments of granite

pieces of amber

flint artifacts bone artifacts bits of charred bone pieces of red ocher

scatter of artifacts and other cultural debris beneath bone heap

N

hearths

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FIGURE 7.18. Top: Plan of a roughly rectangular, 30-cm-deep depression dug by the upper Paleolithic (epigravettian) inhabitants of Pushkari 1 in ukraine, showing the internal distribution of hearths and large bones. radiocarbon suggests the site was occupied about 17 ka (hoffecker 2002). Below: reconstruction of a hypothetical three-part structure that may have covered the depression (redrawn after boriskovskij [1958], fig. 3).

3m

Pushkari I hearths

margin of depression with large bones

Some bone objects retain wear or use traces like those on historic battens, weaving sticks, mat needles, and net spacers that were used to manufacture textiles or cordage from plant ibers. Such objects span the entire Late Paleolithic across Europe, but their interpretation is partic­ ularly clear in 27–26­ky­old Moravian (Czech) Eastern Gravettian (or “Pavlov Culture”) sites noted below, where ired clay objects preserve impressions of textiles, cordage, and nets. Textile or cordage impressions also occur on much rarer ired clay objects at other European late Paleo­ lithic sites, ranging in age from 24 to 12.5 ka. he unusual waterlogged deposits at Ohalo II, Israel, preserved actual fragments of twisted­iber cordage, dated to 19 ka. Many pointed stone and bone artifacts must have been used to tip spears or arrows whose perishable wooden shats have long since van­ ished. Points still embedded in animal bones provide conirmation–for example, a lint point in a wolf skull at the Eastern Gravettian site of Dolní Věstonice in the Czech Republic, another in a reindeer vertebra

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at the late Magdalenian-like “Hamburgian” site of Stellmoor in northern Germany, and a bone point in a bison scapula at the Siberian late Paleolithic site of Kokorevo I. Like other formal bone artifacts, bone points were essentially a late Paleolithic innovation, and their variety constitutes a further indication of late Paleolithic creativity. Some, including the very earliest examples from Europe, have split or beveled bases (ig. 7.8) that would have made hating easier. Others that are especially common in Siberia bear longitu­ dinal grooves or slots into which sharp stone bits were probably inserted to produce a more ragged wound (ig. 7.19). Points with the inserts still in place have been found at several sites, including Kokorevo I mentioned above. On some points the grooves were too broad and shallow to ac­ cept inserts, and they may have been runnels to promote bleeding in a wounded animal. Finally, among all the known point types, among the most striking are those with barbs. Possible self­barbed points—slender double­pointed antler spearheads that could have been mounted so that one end served as a projectile tip and the other as a barb—are known from northern Spain by 20 ka, and more compelling objects that re­ semble historic harpoon heads occur across Eurasia ater 15 ka. Both the self­barbed points and the harpoons could have been used to spear ish, and the harpoons could also have been used to capture reindeer or other mammals crossing a stream. Arguably, the manufacture of bone harpoon heads can be traced to the people who occupied the Katanda sites in the eastern Democratic Republic of the Congo, before 60 ka. However, as discussed in the sec­ tion on nonstone artifacts in chapter 6, the dating at Katanda is contro­ versial, and the oldest bone harpoon heads in Africa are perhaps those from Ishango, also eastern Democratic Republic of the Congo, dated to ca. 25 ka, or those from various north and east African sites that were occupied mainly ater 12–10 ka. Bone points are only one of many late Paleolithic novelties that en­ hanced hunting eiciency. For acquiring large, mobile game, the most important new items were surely projectile weapons, including particu­ larly the spear­thrower and the bow and arrow. he spear­thrower, oten known by its Aztec name as the atlatl, is a bone or wooden rod hooked at one end to accommodate the dimpled or notched nonpointed end of a spear or dart shat (ig. 7.20). With the shat resting against the rod, the rod extends a person’s arm so that the spear can be thrown harder and farther. Bone spear­throwers were being used in southwestern France by 18 ka. he antiquity of the bow and arrow is more diicult to establish be­ cause the most diagnostic parts were made of perishable materials. How­ ever, small sharp stone bits, well­fashioned stone points, or small backed bladelets closely similar to ones that tipped arrows later on occur from at

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FIGURE 7.19. Grooved antler artifacts from the siberian upper Paleolithic site of Kokorevo i, dated to approximately 13–14 ka (redrawn after abramova [1982], fig. 1). The grooves probably held microblades, some of which were still present in one piece (left center). The artifacts probably served mainly as projectile points, and the broken piece in the upper left-hand corner was found stuck in a bison shoulder blade. The piece on the far right is less pointed than the others and may have been a knife handle. The grooved antler and microblade technology is well documented in Siberia only after 18 ka, but it may have been present earlier.

stone inserts

0

5 cm

antler "knife handle" antler projectile points

least 20 ka in various parts of Eurasia and Africa. Once the bow and ar­ row were invented, the combination would obviously have difused very rapidly, and a sharp increase in the abundance of tiny backed bladelets in both Africa and Eurasia about 21–20 ka may signal the spread. More securely, blunt­ended bone rods strikingly similar to historic or proto­ historic arrow linkshats or foreshats circumstantially place the bow and arrow in southern Africa by 20 ka or a little later. However, the oldest un­ equivocal evidence, consisting of fragmentary wooden bows or arrows, comes only from late Magdalenian or related sites, dating between 12 and 10 ka in France and northern Germany. Late Paleolithic sites in Europe have also provided the oldest known ishhooks, dated to roughly 14 ka, while objects interpreted as “ish gorges”—shaped bone slivers resembling double­pointed toothpicks (ig. 7.21)—occur in South African sites that are only slightly younger. Fish­

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0

5 cm

Mas d’Azil ing with spears, traps, and so on had probably been underway for some time before 14 ka, to judge by ish bones found in earlier sites. In sum, late Paleolithic people were obviously very resourceful, and it seems likely that by 12–10 ka they had devised the entire range of tech­ nology observed among historic hunter­gatherers. he Eastern Gravet­ tian people who occupied the Czech sites of Dolní Věstonice I and II, Pavlov I, Předmostí, and Petrkovice 27–26 ka even discovered that clay or ine silt mixed with water and other materials (temper) and heated to 600–800°C, hardens into a much more durable material. he sites have provided more than 10,000 ire­hardened clay fragments, including some that may be from linings of baskets, others that may represent daub used in hut construction, and more than 3,700 that were parts of animal and human igurines. Some fragments retain human inger or palm prints, and a small number preserve unique impressions from woven fabrics, nets, cordage, and basketry that shed additional light on late Paleolithic ingenuity. he greatest number of hardened pieces (more than 6,700) oc­ curred at Dolní Věstonice I, which also preserved two walled structures

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bone points

FIGURE 7.21. bone artifacts from the lsa layers of nelson bay Cave, south africa (after deacon [1984], 176). formal bone artifacts are rare or absent in msa sites, and where they occur, many could represent undetected intrusions from lsa levels. by ethnographic analogy, the nelson bay lsa “fish gorges” were probably baited and attached to lines to catch many of the fish represented in the same layers. In preceding MSA sites, objects interpretable as fishing gear are absent and fish bones are correspondingly rare.

cut and polished with snapped end

bone spatulates

bone spatulates “fish gorges”

bone spatulate

0

“fish gorges” 5 cm bone bead multiringed bone tubes

bone pendants

that probably represent kilns. Roughly 15 ky later, about 13–12 ka, inal Pa­ leolithic, semisedentary foraging people in the lower Amur Basin (Rus­ sian Far East) and nearby Japan independently rediscovered ired clay technology, which they used to manufacture the world’s oldest known ceramic vessels. SOURCES: overviews of late Paleolithic technology (Bordaz 1970; Bordes 1968; Clark 1967b; Coles and Higgs 1969; Hofecker 2005b; Sonneville­Bordes 1973); east European fur­trapping sites (Hof­ fecker 1986; Klein 1973b; Sofer 1985); eyed needles at Kostenki XV and Tolbaga (Hofecker 2005b), Předmosti (Sofer 2000), and later European sites (Schmider 1990; Stordeur­Yedid 1979; Straus 1990b); Sungir’ and tailored clothing (Bader 1978; Kuzmin et al. 2004; Pettitt and Bader 2000); Sungir’ and Tianyuan footwear (Trinkaus 2005d; Trinkaus and Shang 2008); divergence of clothes lice from body lice (Kittler et al. 2003); structural modiication at Cueva Morín (Freeman and González Echegaray 1970) and El Juyo (Barandiarán et al. 1985; Freeman et al. 1988), structural remnants from Ohalo II (Nadel et al. 2004; Nadel and Werker 1999) and central and east European sites (Hofecker 1986, 2002;

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Klein 1969b, 1973b; Kozlowski 1990c; Kozlowski et al. 1974; Kozlowski and Kubiak 1971, 1972; Sofer 1985, 1989b; Svoboda 1993, 1994); lamps at Kostenki I (Hofecker 2005b) and in western Europe (de Beaune and White 1993); wear on Verberie endscrapers (Rots 2005); wear traces from weaving, etc. (Sofer 2004); European late Paleolithic objects with textile impressions (Sofer et al. 2000); Ohalo II cordage (Nadel et al. 1994); points stuck in animal bones at Dolní Věstonice (Klíma 1963), Stellmoor (Schild 1984), and Kokorevo I (Abramova 1964); antiquity of self­barbed points (Pokines and Krupa 1997); antiquity of harpoons in Europe (Julien 1982; Peterkin 1993; Straus 1993) and in Africa (Brooks et al. 1995; Phillipson 2005; Robbins et al. 1994, 2000); oldest known spear­thrower (Cattelain 1989); arrow linkshats or foreshats in southern Africa (Deacon 1984); oldest known arrows (Tyldesley and Bahn 1983); oldest known ishhooks (Julien 1982); east (Czech) Gravettian ired clay objects (Vandiver et al. 1989) and textile impressions (Adavasio et al. 1996, 2001; Sofer 2000; Sofer et al. 1998); early ceramic vessels in the Amur Basin and Japan (Dolukhanov et al. 2002; Kuzmin and Orlova 1998; Reynolds and Barnes 1984; Teruya 1986)

Social Organization

he archaeological record sheds little light on late Paleolithic social or­ ganization, except perhaps for greater contact or interchange between groups than ever before. In Europe and northern Asia, some Upper Pa­ leolithic sites contain “luxury” items such as amber and seashells that had to be imported or traded from hundreds of kilometers away. Even the stone used to make tools was sometimes transported over great distances. Perhaps the most spectacular examples come from very late Upper Paleolithic (terminal Pleistocene) sites on the great plain of north­ central Europe, which demonstrate that particularly desirable types of lint were routinely carried 100–200 km and even more. here are also earlier examples, however, such as those from the “Spitsyn” set of Upper Paleolithic sites near Kostenki in European Russia, from the very early Aurignacian levels of Bacho Kiro, Bulgaria, and from the Eastern Gra­ vettian occupations at Pavlov and Dolní Věstonice, Czech Republic. he Spitsyn people who lived near Kostenki before 32 ka imported almost all their lint from 150–300 km away; the early Aurignacians who occupied Bacho Kiro by 40 ka carried more than 50% of their lint at least 120 km; and the Eastern Gravettians who inhabited Pavlov and Dolní Věstonice roughly 27 ka imported more than 90% of their lint over a distance of 120 km. Not all late Paleolithic people routinely moved “luxury” or “ba­ sic” goods over such great distances, but before the late Paleolithic, no one did. he wider intergroup contacts that such long distance move­ ments may imply could have been both cause and efect of the enhanced cognitive and communication abilities that are probably implied by late Paleolithic art. In the absence of direct evidence for social organization, the physi­ cal and material cultural similarities between late Paleolithic people and many historic hunter­gatherers suggest that late Paleolithic populations were similarly organized or, perhaps more precisely, that they enjoyed broadly the same range of social structures as their historic counterparts. Some late Paleolithic people occupying more marginal environments in

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which resources were limited and population density was low probably lived in small egalitarian bands of related families, like many of the historic hunter-gatherers of Australia and of the dry interiors of western North America and southern Africa. Based on historic observations, average band size was probably around thirty, and individuals oten moved between neighboring bands. During the day, the band usually divided into smaller foraging groups that reunited at night. Individual bands commonly moved camp in response to seasonal variation in the location of key foods. Related bands speaking the same language formed what might be called a tribe, but the tribe was strictly a social or linguistic unit, not a political one. At the other extreme, some late Paleolithic people who inhabited rel­ atively rich settings that supported much denser populations may have formed “ranked” societies like those of the historic hunter­gatherers of the American Paciic Northwest. In such societies, the cohabiting group might oten have exceeded 200 individuals, and it probably moved rela­ tively rarely. A cadre of hereditary chiefs may have coordinated many activities, including food acquisition and distribution, rituals and cer­ emonies, trade, and even warfare. Relatively complex social organization has been posited particularly for the very late Paleolithic (ca. 16–11 ky old) Magdalenian people of southwestern France and northern Spain, whose sites are especially numerous, rich, and closely packed. It may also have characterized some of the Gravettian and Epi­Gravettian (ca. 26–14 ka) peoples of Ukraine, Belarus, and Russia, whose sites contain elaborate structural “ruins” and impressive food storage pits. Whereas the simpler societies of the late Paleolithic may have difered little in ba­ sic organization from Middle Paleolithic and even earlier ones, the more complex societies that characterized some late Paleolithic people prob­ ably had no earlier counterparts.

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SOURCES: Upper Paleolithic transport of luxury items (Harrold 1988; Mussi and Roebroeks 1996; Weniger 1989, 1990); stone raw material transport in north­central Europe (Schild 1984), at Kostenki (Hofecker 1986; Klein 1969a; Sofer 1985, 1989b), Bacho Kiro (Kozlowski 1990a), and Pavlov and Dolní Vĕstonice (Svoboda 1994); historic hunter­gatherer social organization (Marlowe 2005); social organization in the east European Gravettian and Epi­Gravettian (Sofer 1985, 1989a)

“Ideology”: Art and Graves

he thoughts, ideas, beliefs, and values of late Paleolithic people are not preserved in the archaeological record, but their art and their graves pro­ vide the irst clear evidence for ideological systems like those of historic people. he art has been summarized and analyzed many times, and it may be divided between two basic categories—wall art, comprising paintings and engravings on rock surfaces, and portable or home art, comprising items that occur alongside other artifacts in the ground.

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Paintings and engravings weather of rock surfaces that are exposed to the elements, and late Paleolithic wall art is therefore conined almost entirely to deep caves. he irst examples were discovered in 1879 by Marcelino Sanz de Sautuola in Altamira Cave in the Cantabrian region of northern Spain, but for many years their great antiquity and authen­ ticity were disputed. Full acceptance came only in the early 1900s, af­ ter important additional examples were found nearby in southwestern France. More than 150 caves with late Paleolithic paintings or engrav­ ings have now been identiied in Franco­Cantabria (ig. 7.22), making it by far the richest region in the world. In other regions with apparently suitable caves, late Paleolithic wall art is either rare or absent, suggesting it was not an aspect of local culture. But it is also possible that, like di­ verse historic or late­prehistoric hunter­gatherers throughout the world, many late Paleolithic peoples produced wall art on exposed rock surfaces where it no longer survives. It is both puzzling and fortunate that, almost uniquely, the late Paleolithic artists of Franco­Cantabria oten chose cave walls as a basic medium. For more than a hundred years ater Franco­Cantabrian cave art was irst discovered, it was assigned to the Paleolithic largely because it depicted mammoth, bison, reindeer, wild horses, and other region­ ally or globally extinct animals. Stylistic comparisons with animal ig­ ures carved from or engraved in bone, antler, or ivory suggested that the inal Paleolithic (16–11 ky old) Magdalenians were the main artists at famous caves like Lascaux, Les Trois­Frères, Niaux, and Altamira, but until recently direct dating was impossible. his was true even though the black pigment used in many paintings was based on charcoal that is amenable to the radiocarbon method. he problem was that the amount of carbon necessary for a conventional radiocarbon date would require the destruction of whole paintings or even of multiple paintings. Now, with the advent of the accelerator radiocarbon method, it is possible to date charcoal fragments the size of a pin prick (half a milligram), whose removal does little or no damage, and this shows that the Magdalenians indeed produced most if not all the remarkable art at Niaux, Altamira, El Castillo Cave, and other Franco­Cantabrian sites. In addition, however, accelerator dating has also shown that some French cave art substantially antedates the Magdalenian. he most spec­ tacular examples are at Cougnac Cave (southwestern France), where paintings of the extinct Irish elk have been bracketed between 25 and 20 ka; at Cosquer Cave near Marseilles, where stenciled or negative handprints (produced by blowing paint over a human hand) and naturalistic animal paintings have been ixed at roughly 27 and 18.5 ka, respectively; and at Chauvet Cave in the Ardèche Valley (south­central France), where natu­ ralistic paintings have been dated to about 31 ka. Cosquer Cave contains

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BRITAIN 0

GERMANY

200 km

BE

LG I

UM Hohlenstein-Stadel

Atlantic Ocean

Geissenklösterle Vogelherd

FRANCE SWITZERLAND Jovelle Las Chimeneas (El Castillo) La Covaciella

AL

Micolón Altamira Santimamiñe

POR TUG

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ChaP Ter s even

Rouffignac

Vacheresse, Ebbou, Lascaux Gabillou Le Colombier, Oulen, Cougnac Les Deux-Ouvertures, La Mouthe Peche Merle Tête-du-Lion, Le Figuier Les Trois-Frères & Chabot Gargas Le Portel Bayol Fontanet La Baume-Latrone Réseau Clastres Niaux Cosquer

ITALY

Chauvet

SPAIN

Parpalló

Mediterranean Sea FIGURE 7.22. approximate locations of the main decorated caves in france and spain and of caves in Germany that have provided some of the oldest known ivory figurines. The overwhelming majority of decorated caves occur in northern (Cantabrian) Spain and southern France, which together comprise the Franco-Cantabrian region. Exploration continues to reveal new paintings or engravings in previously known caves, and a previously unknown decorated cave is found every year or two. The insets show a lion figurine from the early Aurignacian deposits of Vogelherd Cave, dated to roughly 32 ka, a painted rhinoceros from Chauvet Cave that probably dates to about 31 ka, and a red deer from Las Chimeneas Cave (El Castillo) that dates to 14–13 ka. (Base map and rhinoceros modified after Chauvet et al. [1995], 13; Vogelherd figurine after Hahn [1993], fig. 3; and Las Chimeneas deer after Straus [1995], fig. 9).

numerous inger tracings in the once­sot cave wall that are thought to be coeval with the stenciled hands, which they sometimes overlie, and both Cosquer and Chauvet caves contain engraved animals whose species identity and style imply broad contemporaneity with the painted ones. At Chauvet the artists were presumably Aurignacians, and at Cosquer the earliest ones could have been Aurignacians or Gravettians and the later ones were probably Solutreans. At both sites, subject matter and style further support a pre­Magdalenian age for the art. his is particularly true at Chauvet, where rhinoceroses, lions, mammoths, and bears oc­

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cur far more oten than in well­known Magdalenian art, in which horse, bison, deer, and ibex generally dominate. he focus at Chauvet recalls the prominence of rhinoceroses, bears, mammoths, and lions among the seventeen Aurignacian ivory igurines found in the caves of Vogelherd, Geissenklösterle, and Hohlenstein­Stadel, southwestern Germany (ig. 7.22). he ivory statuettes are as old as the Chauvet paintings, or older, and as already indicated, they join Chauvet in underscoring the gulf be­ tween even the early Upper Paleolithic and the Mousterian. Much supposed Paleolithic cave art from outside the Franco­ Cantabrian region is diicult to authenticate, but Fumane Cave, north­ western Italy, has provided six or more ocher­painted fragments of limestone roof or wall that became embedded in early Aurignacian lay­ ers, dated between 36 and 32 ka. he fragments mainly bear lines that may have formed parts of larger paintings, but one exhibits the partial, styl­ ized outline of a four­legged mammal and a second shows what may be a crude human igure, perhaps with an animal head. he Fumane paint­ ings conirm that even the earliest Upper Paleolithic people produced representational art. Scanning electron microscopy, X­ray difraction, and other advances in technology that can reveal the physicochemical composition of inini­ tesimal pigment samples have now shown that Franco­Cantabrian artists oten mixed iron and manganese oxide or charcoal pigments with miner­ als that served as paint extenders or with plant oils that served as binders. he identiication of distinct pigment recipes has illuminated the prob­ able order in which paintings were executed, and together with radiocar­ bon dating, pigment variation among paintings suggests that successive generations of artists may have visited a single cave for centuries or even millennia. Replication experiments have shown further that the artists could have applied paint in a variety of ways, including not only brushing but also spitting in the manner of some historic Australian Aborigines. If the techniques of the artists have become clearer, however, the meaning or purpose of their art remains mysterious. Perhaps the most se­ cure inferences can be drawn from historic hunter­gatherers, who rarely produced art for its own sake. Instead they embedded their art in other aspects of culture, where it variously functioned to enhance hunting suc­ cess, to ensure the bounty of nature, to illustrate sacred beliefs and tradi­ tions (perhaps on ritual occasions), or to mark the territorial boundaries of an identity­conscious group. Conceivably much Paleolithic wall art symbolizes or encodes the social structure or worldview of its makers, and like the much more recent rock art of southern Africa, some could register the visions of shamans or medicine men in the trance state. he deeply cultural (vs. strictly artistic) meaning of much Franco­Cantabrian art is probably relected in its location not only in caves but sometimes deep within them, in chambers or passages that were diicult to reach.

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To penetrate the darkness, the artists used wooden torches or burned animal fat or oil on limestone and sandstone slabs. Splotches of possible torch charcoal from attempts to expose unburned wood underneath occur on the walls of Chauvet Cave, and shallow sculpted rock bowls that broadly resemble historic Inuit (Eskimo) oil lamps have been found in several French Magdalenian sites. Some even retain chemical traces of vegetal wicks and of the mammal fats that served as fuel. By its mode of occurrence, portable or home art, including items of personal adornment, is much easier to date. he ivory igurines from Vogelherd, Hohlenstein­Stadel (ig. 7.23), Geissenklösterle, and Hohle Fels in southwestern Germany, occur in early Aurignacian layers that have been securely dated by radiocarbon to 32 ka or before. An early Au­ rignacian layer, at Galgenberg Hill (Stratzing/Krems­Rehberg), Austria, also dated to about 32 ka, has provided an equally sophisticated igurine in serpentine (ig. 7.24), and if we didn’t know better, we might assume that as a group, the igurines represent the culmination of a multimil­ lennial learning curve. In fact, they have no antecedents, and the same is true of early Aurignacian ornaments, dated at various sites to 32 ka or before. hey comprise meticulously shaped ivory or sot stone beads and numerous carefully perforated animal teeth that were probably pendants or beads. At Brassempouy, France, the teeth include four human speci­ mens on which the roots were perforated or circumincised for hanging. Together with perforated human teeth from Aurignacian horizons at La Combe and Isturitz, also France, the Brassempouy examples are the old­ est known human remains to be modiied for ornamental or perhaps ritual purposes. he practice of ornament production almost certainly originated in Africa, and it was noted previously that early LSA sites (table 7.4), ante­ dating 30 or even 40 ka, have produced carefully made ostrich eggshell beads resembling those that some Africans still produced historically. Africa is also justly famous for its rock art, but this was mostly done on exposed surfaces where it would last a few hundred or a few thou­ sand years at most. Deep caves that might preserve much older wall art are rare, but deposits dated between 27.4 and 19 ka at Apollo 11 Cave, Namibia, have provided wall fragments on which painted animals show that representational art also has great antiquity in Africa. Like the stone artifacts that accompany them, art objects and per­ sonal ornaments vary signiicantly in form through time and space. For example, in Europe well­produced naturalistic engravings of animals are concentrated in Magdalenian sites (16–11 ka), whereas human igurines occur most commonly in Gravettian levels (28–21 ka) (ig. 7.25). Like the wall art, the portable art is impossible to interpret precisely, but little of it was probably done for its own sake. Some enigmatic engraved or incised objects may have been gaming pieces, and others were perhaps counting

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689 FIGURE 7.23. mammoth ivory “lion-man” from an early aurignacian layer at holhenstein-stadel, southwestern Germany (redrawn from an original by Joachim hahn in Clottes [1996], 280). The object had disintegrated in the ground and had to be painstakingly reconstructed. it illustrates the remarkable artistic ability of even the earliest upper Paleolithic europeans, 32 ka or before.

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Hohlenstein-Stadel “lion-man” or recording devices, even lunar calendars. Flute-like bone tubes with evenly spaced perforations from the early Aurignacian layers at Geissenklösterle and Isturitz probably functioned as lutes or pipes. he three examples from Geissenklösterle—two on swan bones and one labori­ ously carved in mammoth ivory—are irmly dated to 33 ka or before, and together with the Isturitz specimens, they are the oldest widely accepted musical instruments known so far. Later examples from Isturitz suggest a musical tradition that persisted for perhaps 20 ky. Many animal igurines could be the totemic symbols of kinship groups, and the human igurines obviously could represent deities or spirits. Most are highly stylized, lacking facial features or details of the hands and feet. Many, known popularly as “Venus igurines” (igs. 7.24, 7.26, also ig. 5.62), have exaggerated buttocks and breasts, leading to speculation that they were fertility symbols or depictions of earth mother goddesses. Whatever the case, with the rest of the art, they clearly imply

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FIGURE 7.24. The early aurignacian “venus” from Galgenberg hill, near Krems, austria (drawn by Kathryn Cruz-uribe from a photograph). The figurine was carved from green serpentine, and it may depict a dancing woman, left hand in the air, right arm bent with hand on hip, and left breast protruding in profile. The artist who produced it was among the earliest on record, yet the quality of craftsmanship has never been exceeded.

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Galgenberg “Venus” that late Paleolithic people not only were physically identical to their living descendants but possessed basically the same cognitive and communication facilities, including languages that were as complex as any historic ones. Enhanced cognition and communication were surely crucial to many late Paleolithic activities, particularly to ones that depended upon within- and between-group cooperation, and insofar as the art implies the fully modern mind, it helps to explain why other aspects of late Paleolithic culture appear so advanced. Graves are of course known from Mousterian sites, and late Paleolithic people were thus not the irst to bury their dead. However, late Pa­ leolithic sites are the earliest to contain undoubted multiple burials. he probable Aurignacian skeletons, excavated at the famous Cro­Magnon rockshelter, southwestern France, probably came from a communal grave, and some truly spectacular examples have been found at East­ ern Gravettian (“Pavlov Culture”) open­air sites in the Czech Republic, radiocarbon­dated to 27–26 ka. hese Eastern Gravettian cases include a young woman and two young men, possibly her brothers, buried to­

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Last Glaciation coastline

Scandinavian ice sheet

Gagarino

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et

e ice sh

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Grimaldi single figurine multiple figurines

gether at Dolní Věstonice and a veritable graveyard at Předmostí. Here, excavations near the end of the nineteenth century uncovered a large (4 × 2.5 m) oval pit that was covered by limestone slabs and mammoth bones and that contained the skeletons of eighteen individuals of various ages and both sexes. Even richer cemeteries, containing dozens of indi­ viduals, have been found in terminal or epi­Paleolithic (ca. 15–10 ky old) cave sites such as Taforalt and Afalou­Bou­Rhummel in northwestern Africa (locations in ig. 7.2), where they are associated with artifacts of the Iberomaurusian Industry. Equally important, recall from the previous chapter that Mouste­ rian graves tend to be very simple, with no certain indication of a burial ritual or of the inclusion of valued items or “grave goods.” Many Upper Paleolithic graves are also relatively simple, but others are much more elaborate, and individuals were oten buried with special bone, shell, or stone artifacts (ig. 7.27). Clusters of perforated seashells occurred with the skeletons from the early Aurignacian layers at the Cro­Magnon Shel­ ter, and clusters of pierced shells or animal teeth, dense concentrations of ocher, or both characterize numerous other Upper Paleolithic graves, notably including the multiple examples in the Aurignacian and Gravet­ tian layers of the Grimaldi Caves and of Arene Candide Cave (Italian Riviera). Two neonates buried at the Eastern Gravettian site of Krems­ Wachtberg, eastern Austria, 26–27 ka were laid on a bed of red ocher,

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FIGURE 7.25. approximate locations of the upper Paleolithic Gravettian sites that have provided female figurines or engravings (redrawn after Champion et al. [1984], fig. 3.19). The Gravettian complex spanned Europe from Portugal to Russia between about 28 and 21 ka, and similarities in stone artifacts, figurines, personal ornaments, burial style, and so forth imply it sprung from a common center, perhaps in central or eastern Europe. The map also illustrates the maximum extent of the last glacial ice sheets in Europe and corresponding changes in coastlines (dotted lines mark present coastlines).

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FIGURE 7.26. mammoth ivory “venus” figurines from Kostenki 1, layer 1, Russia (redrawn after Abramova [1962], plate III). The associated artifacts belong to the Eastern Gravettian Culture, which dates mainly between 24 and 21 ka in Russia and Ukraine. The Kostenki specimens closely resemble figurines from like-aged sites across Europe in the exaggeration of female characteristics and in the casual portrayal of the head and limbs. Figure 5.62 illustrates a closely similar figurine from a French Gravettian site.

cm

and their burial pit, 40 cm long, was covered by a mammoth scapula supported by a tusk. More than thirty ivory beads accompanied one of the bodies, and the sum contrasts sharply with the simple graves of infant Neanderthals, known, for example, from Roc de Marsal, France, Mezmaiskaya Cave, Russia, and Dederiyeh Cave, Syria. he child’s skeleton from Lagar Velho, Portugal, mentioned in the previous chapter as a possible Neanderthal/Cro­Magnon hybrid, fur­ ther illustrates the diference between Mousterian and Upper Paleolithic graves since it was identiied as Gravettian even before it was directly dated. his is because the bones were stained red from pulverized ocher and they were associated with a shell pendant and four pierced deer ca­ nines like those recovered previously from Gravettian burials in Italy and Moravia (Czech Republic). he child had been laid out on its side, fully extended in the manner that characterizes Gravettian burials else­ where but never Mousterian ones. he excavators believe that the ocher originated from a hide shroud (which explains how it came to coat all the bone surfaces), the pendant hung around the neck (it occurred near the

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N bone “hide burnisher”

tip & handle of a “bone knife”

bone needle mound of yellow clay

pierced arctic fox teeth

mandible

flint artifacts

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pierced arctic fox teeth

693 FIGURE 7.27. Plan of a burial pit dated to roughly 30 ka at Kostenki Xv on the don river in european russia (redrawn after rogachev [1957], fig. 56). The pit contained the remains of a six-to-seven-year-old child, whose reconstructed skull is illustrated in figure 7.3. The excavators believe the child’s body was seated, head on knees, on a 4–6 cm-high mound of bright yellow clay within a structure that subsequently collapsed inward. The collapse caused the head to separate from the semidecomposed postcranium. Probable grave goods include more than seventy flint artifacts (blades, flakes, ten endscrapers, and a borer), three well-made bone artifacts (identified as a knife, a hide burnisher, and a needle), and more than 150 arctic fox teeth, each pierced through the root. The teeth may have been part of a headdress that disintegrated when the skull separated from the body.

60 cm

neck vertebrae), and the deer canines were part of a headdress (they were closely associated with what remained of the skull, which a bulldozer had unfortunately shattered before excavation.) Radiocarbon dating of charcoal from immediately below the skeleton and of animal bones from immediately above it, bracket it between 24.9 and 23 ka, squarely within the later part of the Gravettian era. he actual age is assumed to be closer to the 24.9 ka date since the charcoal lay in a pit that may have formed part of the burial ritual. As discussed in the last chapter, most specialists believe the child was fully modern, if perhaps unusually stocky, and few accept the idea it relects hybridization many millennia earlier. An equally old or older grave at Sungir’, Russia, mentioned above for its evidence of clothing, may best illustrate the extraordinary con­ trast between some Upper Paleolithic burials and all Mousterian ones. he Sungir’ grave was dug into permanently frozen subsoil (permafrost)

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sometime between 26 and 19 ka and contained the extended skeletons of two children, one arguably male and the other female, placed head to head. he putative male was covered with 4,903 beads whose arrange­ ment suggests they were attached to closely itting clothing. In addition, there were 250 perforated arctic fox canines placed as if they had been attached to a belt at the waist, an ivory animal pendant on the chest, an ivory pin (perhaps a fastener) near the throat, a large ivory mammoth igurine under the let shoulder, a highly polished human femoral shat packed with red ocher on the let side, and a 2.4­m­long ivory “lance” on the right side. he putative female was covered and surrounded by 5,374 beads or bead fragments that were also probably attached to clothing, and there was an ivory pin at the throat, several small ivory “lances” on both sides, and on one side, two perforated antler “wands,” one of them decorated with rows of shallow drilled holes. Experimentation suggests that the beads alone required thousands of hours to manufacture. he nearby burial of an adult male was equally elaborate, and the sum sug­ gests a burial ritual and perhaps notions of an aterlife similar to those recorded among historic hunter­gatherers. In contrast to Mousterian graves, some Upper Paleolithic ones, such as those at Předmostí and Krems­Wachtberg, were covered by large rocks or bones, perhaps to complete a ritual or to prevent wolves or hyenas from exhuming the bodies. Like earlier sites, however, late Paleolithic ones of­ ten contain isolated scraps of human bone, and carnivore exhumation of burials could be responsible. Alternatively, disarticulated bones might sometimes relect complex burial practices that deliberately dissociated the skull from the postcranium or separated various postcranial elements from each other. he most persuasive example is probably from the 22­ ky­old late Perigordian (or Protomagdalenian) layer of the Abri Pataud, France, where the skull of a iteen­to­eighteen­year­old woman was po­ sitioned among three stones about 4 m away from a cluster of postcranial bones that may include hers and those of her newborn child. In yet other instances, including the polished human femur in the double grave at Sungir’ and intentionally shaped or perforated human bones and teeth from Isturitz, Saint­Germain­la­Rivière, and other French Upper Paleo­ lithic sites, isolated elements may represent trophies or heirlooms. Finally, the possibility exists that some disarticulated bones repre­ sent food debris, but compelling Paleolithic evidence for it is rare and limited so far to the much more ancient sites at Atapuerca GD (Spain), dated to 800 ka (chap. 5), and Moula­Guercy (France), dated to roughly 110 ka (chap. 6). Cannibalism has been more commonly observed at prehistoric agriculturalist sites like Fontbrégoua Cave (France), dated to about 6 ka, and various Anasazi Pueblo sites in the American South­ west, dated between roughly 1,100 and 700 years ago. he evidence at each site comprises numerous disarticulated human bones that were

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demonstrably broken, burned, cut, or otherwise treated like the bones of other animals and that cannot be easily attributed to mortuary practices, carnivore exhumation, or postdepositional disturbance. So far, no late Paleolithic site has provided comparable bones, and like historically observed hunter-gatherers, late Paleolithic people probably rarely if ever practiced cannibalism. For the most part, like their historic counterparts, they probably lived in small bands that had far more to lose than to gain from hunting their neighbors.

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SOURCES: summaries of Upper Paleolithic art (Abramova 1967; Bahn 1995–1996; Bahn and Vertut 1988; Breuil 1952; Conkey 1981, 1983, 1987; Freeman et al. 1987; Graziosi 1960; Guthrie 2005; Kozlowski 1992a; Leroi­Gourhan 1965b; Sieveking 1979; Ucko and Rosenfeld 1967; White 1986a, 2003a); authen­ ticity of Altamira (Bahn and Vertut 1988; Straus 1992); rarity of cave art outside of Franco­Cantabria (Bar­Yosef 1994a); radiocarbon dating of Franco­Cantabrian art to the Magdalenian (Bahn 1994, 1995–1996; Chauvet et al. 1995; Lorblanchet 1994; Valladas et al. 1992); pre­Magdalenian cave art—at Cougnac Cave (Lorblanchet 1993, 1994), Cosquer Cave (Clottes and Courtin 1993, 1995; Clottes et al. 1992), and Chauvet Cave (Chauvet et al. 1995; Clottes et al. 1995; Genty et al. 2004); Chauvet vs. typical Magdalenian cave art (Clottes 1996); Fumane Cave (Broglio et al. 2006; Mellars 2006b); composition of Paleolithic paint (Clottes 1993; Clottes et al. 1990; Lorblanchet 1994; Lorblanchet et al. 1990); experi­ ments to replicate Paleolithic paintings (Lorblanchet 1991); interpretation of southern African rock art (Lewis­Williams 1981, 1982; Lewis­Williams and Dowson 1988; Lewis­Williams and Loubser 1986); Magdalenian lamps (de Beaune and White 1993); early Upper Paleolithic beads, pendants, and other possible ornaments (Gamble 1986; Hahn 1977, 1993; Hofecker 1986; Kozlowski 1982; Movius 1969b; Sonneville­Bordes 1973; White 1989); Brassempouy human tooth pendants (Henry­Gambier et al. 2004); Apollo 11 paintings (Wendt 1976); possible Paleolithic counting devices (d’Errico and Cacho 1994; d’Errico et al. 2003) or lunar calendars (Marshack 1972a, 1972b, 1991b); lutes from Geissenklösterle (Conard 2005) and Isturitz (Buisson 1990); persistence of a musical tradition at Isturitz (d’Errico et al. 2003); communal graves or cemeteries—at Cro­Magnon (Lartet 1868), Dolní Věstonice (Alt et al. 1997; Formicola 2007; Klíma 1987), Předmostí (Absolon and Klima 1977; Smith 1982, 1984; Svoboda 2008), and Iberomaurusian sites (Camps 1974; Lubell 2001); detailed description of Dolní Vĕstonice graves and human remains (Trinkaus and Svoboda 2006); variation in Upper Paleolithic graves, from simple (Belfer­Cohen and Hovers 1992; Straus 1990a; Svoboda and Vlček 1991) to elaborate (Harrold 1980); burials from the Grimaldi Caves (Formicola et al. 2004; Mussi 1990a; Petit­Maire et al. 1971), Arene Candide (Pettitt et al. 2003), Krems­Wachtberg (Einwögerer et al. 2006), Lagar Velho (Zilhão 2001b), and Sungir’ (Formicola and Buzhilova 2004; Pettitt and Bader 2000; White 1993); Pataud human remains (Movius 1975; Villa 1992); human bones as Upper Paleolithic heirlooms (Sonneville­ Bordes 1973; Villa 1992); cannibalism—among historic hunter­gatherers (Arens 1979), at Fontebrégoua Cave (Villa 1992; Villa et al. 1986), and Anasazi sites (White 1992; White and Folkens 2000)

Mortality and Disease

he vital statistics of late Paleolithic people can be only crudely estimated because there are so few skeletons whose age and sex can be established and because the skeletal sample represents a mishmash of catastrophic and at­ tritional mortality. However, a composite sample of seventy­six Eurasian Upper Paleolithic skeletons from various times and places, supported by a remarkable series of 163 skeletons from the terminal Pleistocene Iberomaurusian levels of Taforalt in Morocco, suggest that the common mortality pattern resembled that of most later prehistoric and historic hunter­gatherers. he conclusions must be tentative because available methods for assessing age at death from human skeletons are notoriously

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imprecise, and underestimation of adult age is probably common. With this caveat in mind, the composite Eurasian/North African late Paleolithic sample suggests that child mortality was high, women oten died before age forty, probably because of the risks associated with childbear­ ing, and men commonly died before sixty or even ity. Recall from chap­ ter 5, however, that a survey of mortality through time, beginning with the australopiths, concluded that late Paleolithic people were far more likely than anyone before to survive into older adulthood. he conclu­ sion is questionable, in part because the survey did not acknowledge diferences among fossil samples in the underlying causes or agents of mortality, but at least tentatively, it still suggests that late Paleolithic hu­ man groups contained more older people. heir presence could have en­ hanced group survival, partly because they retained knowledge on how to respond to occasional, unusual crises, and perhaps even more because they provided direct economic support. Among Hadza hunter­gatherers in northern Tanzania today, young women are oten freed to have ad­ ditional children much sooner because their mothers or aunts, beyond childbearing age themselves, actively provision their daughters’ ofspring. Such economically advantageous “grandmothering” may have initially selected for the long postreproductive lifespans that now distinguish human from nonhuman ape females, and the greater availability of ec­ onomically active older women could help explain why late Paleolithic pop­ ulations were so much larger (denser) than earlier ones. Unlike Neanderthal skeletons, late Paleolithic ones rarely show evi­ dence of serious accidents or disease, suggesting that late Paleolithic culture provided a far more efective shield against environmentally in­ duced trauma. Skeletal anomalies that may reveal cause of death are par­ ticularly rare but include conspicuous lesions, perhaps caused by a severe fungal infection, on the skull, mandible, pelvis, and femur of the famous Old Man of Cro­Magnon (actually probably only in his late forties) and dental abscessing that may have produced a fatal septicemia (blood in­ fection) in the iteen­to­eighteen­year­old woman buried about 22 ka at the Abri Pataud, very near Cro­Magnon. Her dental infection was prob­ ably tied to the eruption of aberrant, supernumeary teeth that partially destroyed her normal right upper molars. his abnormality has not been observed in any other Upper Paleolithic people, most of whom had com­ paratively healthy teeth, probably because their diets included few foods that encouraged caries or plaque formation. Additional late Paleolithic skeletal anomalies that probably relect cause of death have been reported from the cave sites of Rochereil in France and Romito in Italy. At Rochereil, a child in a Magdalenian grave had a bulging forehead and other features that suggest hydrocephaly (a normally fatal excess of cerebrospinal luid in the skull). An artiicial perforation that was probably intended to provide relief may have been

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the immediate cause of death. At Romito, an adolescent male in a late Epi-Gravettian or “Romanellian” grave apparently sufered from a kind of genetic dwarism (acromesomelic dysplasia) that certainly reduced life expectancy under Paleolithic conditions. he Romito dwarf is dated to 12–11 ka and antedates the next oldest instance by at least 5 ky. It is the only case of dwarism yet recorded among prehistoric, stone age hunter­ gatherers. Other known late Paleolithic skeletal abnormalities, reported mainly from Czech and French sites, were variably debilitating but probably not fatal. Like the ones already cited, none indicate epidemic diseases, which were probably rare until the greater population densities promoted by the development of food production, beginning 12–10 ka. he Czech exam­ ples include bone deformation of the let temporomandibular joint, sug­ gesting partial facial paralysis, in one woman buried at Dolní Věstonice and a shortened, deformed right leg accompanied by pronounced spinal curvature to the let (scoliosis) in a second. he French cases include fused cervical vertebrae (cervicoarthrosis) in skeletons from Cro­Magnon, Chancelade, and Combe­Capelle; a healed skull fracture, bone lesions or degeneration implying a permanently dislocated let shoulder, and a lat­ erally deviated right big toe (hallux valgus) in the skeleton from Chance­ lade; and an asymmetric sacrum, relecting lateral curvature of the spine, in the one skeleton from Combe­Capelle. By modern analogy, the oc­ currence of cervical fusion may mean that some older Upper Paleolithic people were relatively sedentary, literally remaining seated much of the time. Similarly, if modern people are guides, the laterally deviated big toe of the Chancelade skeleton may relect poorly itting footwear, though other explanations are possible. Skeletal evidence for deliberate injury is also rare, probably because, like most ethnographically recorded hunter­gatherers, late Paleolithic ones rarely engaged in warfare or mass violence. In some instances, such as the healed fractures on male skulls from Chancelade and Dolní Věstonice, the cause could have been accidental, and in many oth­ ers bone fracturing or crushing probably occurred ater death, in the ground. Prominent examples of damage that was once believed to be an­ temortem but that was probably postdepositional include the fractured female skull (individual 2) from the Cro­Magnon site and the four frac­ tured or crushed skulls from the Upper Cave at Zhoukoudian. But even if evidence for violence is rare, it does exist—for example, at the Grimaldi Caves in northern Italy, where an (Aurignacian?) child was buried with a projectile point embedded in its spinal column; at Wadi Kubbaniya near Aswan in Egypt, where a young adult male, buried perhaps 25–20 ka, had a healed parry fracture on the right ulna, a stone chip embedded in the let humerus, and two blades (projectile armatures?) in the abdomi­ nal cavity; and above all in an extraordinary terminal Pleistocene (ca.

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14–12 ky old) cemetery near Jebel Sahaba in Sudanese Nubia. Nearly half the ity­nine individuals exhumed here either had unhealed antemor­ tem skeletal injuries or had stone artifacts lodged in or near their bones. hey provide a remarkably graphic exception to the stated generalization that mass violence is rare among hunter­gatherers. Both the Old Man of Cro­Magnon and the man buried at Chancel­ ade were probably too disabled to fend for themselves, and their survival shows that Upper Paleolithic people, like Neanderthals before them and historic people later, cared for their old and sick. Such care need not have been entirely charitable or unselish since older people in hunter­ gatherer societies commonly possess vital knowledge and experience. But care in a more emotional, abstract sense not known for the Nean­ derthals may be indicated for the Dolní Věstonice woman with the de­ formed temporomandibular joint. Both the face engraved on an ivory fragment and the face of a sculpted clay head found nearby droop on the let side, just as hers probably did. hey were produced 27–26 ka, and they could represent the oldest known portraits.

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SOURCES: mortality in European Upper Paleolithic and Moroccan terminal Pleistocene populations (Vallois 1961); survival to older adulthood in fossil hominins (Caspari and Lee 2004); contribution of older Hadza women to children’s diets (Hawkes et al. 1997a); grandmothering and long postre­ productive lifespans in humans (Hawkes et al. 1998); lesions on the skeleton of the Old Man of Cro­ Magnon (Dastugue 1982); Abri Pataud dental infection (Dastugue and de Lumley 1976; Legoux 1975); hydrocephaly at Rochereil (Dastugue and de Lumley 1976; Vallois 1971); dwarism at Romito (Frayer et al. 1988); skeletal pathologies at Dolní Vĕstonice (Klíma 1962, 1963, 1987; Trinkaus et al. 2006); pathologies in late Paleolithic French skeletons (Dastugue and de Lumley 1976); skeletal evidence for deliberate injury—general (Roper 1969), at Chancelade (Dastugue and de Lumley 1976) and at Dolní Věstonice (Svoboda and Vlček 1991); postdepositional skeletal damage at Cro­Magnon (Dastugue 1982) and Zhoukoudian Upper Cave (Pei 1939b); skeletal evidence for violence at Grimaldi (Dastugue and de Lumley 1976), Wadi Kubbaninya (Wendorf et al. 1986), and Jebel Sahaba (Anderson 1968; Wendorf 1968)

Late-Paleolithic Population Expansion Previous sections noted that in Europe and southern Africa, late Pa­ leolithic peoples appear to have been much more abundant than their predecessors. At least equally important, late Paleolithic people greatly extended the geographic range of humankind by colonizing the eastern­ most part of Europe (in what are today Ukraine, Belarus, and European Russia), central and northern Siberia (Asiatic Russia), the Americas, and Australasia (Australia and neighboring islands). Easternmost Europe

Mousterian and earlier sites occur mostly, if not entirely, on the west­ ern and southern margins of easternmost Europe, in spite of extensive archaeological reconnaissance and the kind of intensive commercial ac­ tivity that leads to site discovery. herefore, as each year passes it seems

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Mamontovaya Kurya (1000 km) Garchi I Zaozerʼe Sungirʼ MAXIMUM EXTENT OF LAST GLACIATION ICE

0 a Ok

Timonovka, Karachizh

Khotylevo, Negotino, Betovo, Korshevo

Eliseevichi, Yudinovo Pogon, Pushkari tʼ ya ip Pr Mezin

Don

Chulatovo

KIEV

Ataki, Korman, Molodova

Kostenki

Sukhaya Mechetka

Northern Donets

Mezhirich

Amvrosievka

Dnep r

Zolotovka

Muralovka

t Pru

AN HI AT RP CA

Beglitsa Rozhok, Nosovo

Bolʼshaya Akkarzha Dn e

Volga

a

Ketrosy

Gagarino

Avdeevo

n Des

Kulichivka

200 km

MOSCOW

ba Ku

str

NS MT

CRIMEA

Syurenʼ1& Kabazi Staroselʼe BLACK SEA

Ilʼskaya

Zaskalʼnaya Kiik-Koba

CASPIAN SEA

nʼ

Kudaro

CAUCASUS MTNS

Dmanisi

FIGURE 7.28. approximate locations of major Paleolithic sites in eastern europe (the western part of the former soviet union) (modified after Hoffecker [1987], fig. 1). Squares mark key Upper Paleolithic sites; circles mark earlier sites. Note the position of the Sungir’ site at 56°N, 40°E, 192 km east of Moscow. In addition to a large assemblage of classic Upper Paleolithic artifacts, it provided the remains of eight fully modern humans. Four came from graves where they were associated with numerous ornaments and other special objects. Skeletons in two of the graves have been directly dated to between 26 and 19 (radiocarbon) ka (Kuzmin et al. 2004; Pettitt 2000a). Upper Paleolithic people were the first to live so far north and east, whether during a glaciation or an interglacial.

increasingly likely that Mousterian and earlier people simply could not occupy most of eastern Europe because of the harsh continental climate that prevailed even during interglacials. Upper Paleolithic people obviously did not ind the climate an insuperable problem, and the Sungir’ site (56°N), dated between 26 and 19 ka, is spectacular proof that even early Upper Paleolithic people could inhabit areas of permafrost (where the subsurface never thawed) north and east of present­day Moscow (ig. 7.28). During a relatively mild interval between 32 and 24 ka within the Last Glaciation, yet earlier Upper Paleolithic people settled 1,400 km fur­ ther east and north at Garchi I (59°N) and Zaozer’e (58°N) in the western foothills of the Ural Mountains that separate European Russia from Si­ beria. During an earlier, comparably mild interval between 42.5 and 36

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ka, some may have penetrated another 1,000 km north to Mamontovaya Kurya (66°N) on the Arctic Circle. Mamontovaya Kurya has provided seven stone artifacts, a mammoth tusk with apparent stone­tool marks, and 123 mammal bones from a small excavation at the base of a river bluf. Radiocarbon­dated bones and OSL dates on sediments place the accumulation between 38 and 34 ka, but an enlarged artifact sample, ad­ ditional occurrences, or both may be necessary to conirm such ancient human presence in the Arctic. A striking feature of the earliest east European Upper Paleolithic sites is their artifactual variability, and with rare, mostly arguable exceptions, none can be assigned to the early Aurignacian Culture. Remember that the early Aurignacian represents the oldest undeniable manifestation of the Upper Paleolithic over most of central and western Europe. If, as seems increasingly likely, the early east European Upper Paleolithic ap­ proaches or even exceeds the 40­ka age of the earliest Aurignacian, the implication could be for a yet earlier Upper Paleolithic that gave rise to both. he reason for such early divergence between east and west is ob­ scure, but it may relate to the fact that over most of eastern Europe, early Upper Paleolithic people encountered no humans, whereas in central and western Europe early Upper Paleolithic (Aurignacian) people in­ vaded areas that the Neanderthals (Mousterians) had long inhabited. he conspicuous internal diversity of the earliest European Upper Paleolithic is a further reminder of how much even the earliest Upper Paleolithic difered from the far more monotonous Mousterian that preceded it.

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SOURCES: Garchi I, Zaozer’e, and Mamontovaya Kurya (Pavlov et al. 2001, 2004); early Upper Paleo­ lithic variability in eastern Europe (Anikovich 1992; Hofecker 1988, 2002)

Siberia

he southwestern corner of Siberia enjoys a cool temperate climate like that of adjacent west­central Asia, and it was likewise occupied by Mous­ terians before 40 ka. So far, their artifacts have been excavated from ive caves and three open­air sites. Levallois lakes are common, and the as­ semblages difer in no essential respect from Levallois­rich Mousterian assemblages that have long been known in Europe and western Asia. Stone raw materials were obtained locally, and bone artifacts are lacking, even where bone is well­preserved. Seven isolated teeth from Denisova and Okladnikov Caves and mitochondrial DNA in a subadult humerus and femur from Okladnikov Cave indicate that the people were Nean­ derthals living at or near the northeastern edge of their range. Radiocarbon dates from the Kara­Bom open­air site show that a typical blade­and­burin Upper Paleolithic industry replaced the Mous­ terian in southwestern Siberia at or before 40 ka. Roughly twenty addi­ tional sites in southern Siberia (south of 55°N), extending from the Altai

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Mountains on the southwest to just beyond Lake Baikal in the center, have provided similar early Upper Paleolithic assemblages that date between 40 and 30 ka (site locations in ig. 7.15). Human teeth at Malaya Syya and Maloyalomanskaya Cave imply that the artifact makers were anatomically modern. In addition to typical Upper Paleolithic stone im­ plements, the people produced points, awls, retouchers, and other for­ mal artifacts in antler, bone, or ivory. At Tolbaga and perhaps other sites, they also let stone rings that probably mark hut bases, together with stone­lined hearths and probable storage pits. So far, the early Upper Paleolithic is undocumented in southeastern Siberia, but its presence by 35–30 ka is implied by the coeval colonization of Japan. Most likely this occurred via a landbridge from Siberia to Sakhalin Island, which was the northern end of a long peninsula that also comprised the four main Jap­ anese Islands (Hokkaido, Honshu, Shikokou, and Kyushu) during much of the Last Glaciation. he femur and tibia of a six­year­old child from a layer radiocarbon­dated to >32 ka at Yamashita­cho Cave, Okinawa, show that the earliest Japanese were anatomically modern. Southern Siberian sites that date between 30 and 20 ka are assigned to the local “middle Upper Paleolithic.” he rich Mal’ta open­air site is the leading example. It was occupied sometime between 25 and 20 ka, and both its elaborate artifact inventory and its probable structural remnants broadly recall those at contemporaneous east European Upper Paleolithic sites. Blades, frequently made into burins and endscrapers, dominate the stone artifacts. Bone, ivory, and antler artifacts abound, and they include utilitarian pieces like awls, points, “handles,” and nee­ dles, together with numerous pendants or beads and a variety of animal and human igurines (ig. 7.29). Dense, patterned clusters of cultural de­ bris may mark hut bases (ig 7.30), and if so, they imply semipermanent camps where the people processed parts of mammoth, reindeer, wild horses, bison, hares, and other species for food or for raw material, cloth­ ing, and parts of structures. A grave with the remains of two children and some elaborate artifacts with which they were buried completes the complex (ig. 7.31). In its particulars, Mal’ta difers from any known Eu­ ropean Upper Paleolithic sites, but the contrast implies only a diference in ethnicity not in fundamental lifestyle. It further underscores the bur­ geoning cultural diversity that characterized the late Paleolithic and that separates it from what went before. Almost all Siberian Middle Upper Paleolithic sites occur in temper­ ate Siberia, south of 55°N, but the Yana Rhinoceros Horn Site site at 71°N, 500 km north of the Arctic Circle, suggests middle Upper Paleolithic people penetrated the Arctic, at least briely. he Yana site occurs in a region of permanently frozen ground, and the artifacts and associated animal bones have been recovered mainly on the surface below frozen de­ posits from which they eroded. he bones come from mammoth, woolly

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FIGURE 7.29. nine of thirty human figurines excavated at the Mal’ta site, southcentral Siberia (redrawn after Abramova [1967], 161). Similar figurines have also been found at the nearby site of Buret’. They were produced sometime between 25 and 21 ka, and they are conspicuously female, like the broadly contemporaneous or somewhat earlier “Venus figurines” from Europe. However, they differ from their European counterparts in key features, including their greater linearity, the tendency to show breasts and other bodily features in outline rather than relief, and the presence of facial features and hair or headdresses.

Mal’ta cm

rhinoceros (Coelodonta antiquitatis), horse, bison, musk ox (Ovibos moschatus), reindeer, wolverine (Gulo gulo), lion (Panthera leo), wolf (Canis lupus), Arctic fox (Alopex lagopus). brown bear (Ursus arctos), and Don hare (Lepus tanaiticus), and specimens that have been artifactually shaped or that exhibit butchering marks have provided radiocarbon dates centered on about 27 ka. he artifactually modiied items include two spear foreshats in ivory and a remarkable foreshat in rhi­ noceros horn, preserved by the frozen sediments. By historic analogy, a stone point in a bone holder was mounted on one end of each foreshat, which enabled a hunter quickly to replace a point that broke or that be­ came embedded in an animal. he Yana foreshats are the oldest known, and they conirm the technological ingenuity of late Paleolithic people.

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limestone slabs

reindeer antlers

before removal of reindeer antlers m

hearth

FIGURE 7.30. remnants of a structure in horizon 2 at the mal’ta site, southcentral siberia (redrawn after Gerasimov [1958], figs. 19 and 20; [1961], fig. 58). Mal’ta was occupied sometime between 25 and 20 ka, and it contained perhaps thirteen other similar “ruins.” These varied somewhat in size and shape, but they generally shared: (1) an artificially depressed floor 50–70 cm below the surrounding surface; (2) limestone slabs, large bones, or both positioned so that they could have anchored a wooden framework; (3) a profusion of reindeer antlers that could have been interwoven on wooden poles to support a skin covering; (4) one or more hearth pits in the floor; and (5) cache or storage pits dug that were dug into the sides of depressed floor.

after removal of reindeer antlers

hypothetical, partial reconstruction

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FIGURE 7.31. The fragmentary skeleton of a three-tofour-year-old child from a grave at the siberian upper Paleolithic site of mal’ta (redrawn after Gerasimov [1935], fig. 36). The poorly preserved remains of an infant occurred in the same grave (Alekseev 1998). Radiocarbon places the grave at about 21 ka (Kuzmin 1997). Like many other late Paleolithic graves, the one at Mal’ta contained special artifacts that may represent funerary offerings, the deceased’s personal belongings, or both.

diadem

braincase dentition

bracelet necklace

upper limb fragment vertebrae & ribs

limestone slabs

retouched blade bird figurine

bone “button”

lower limb fragments

N borer

blade

bone point

blade

he Yana site may imply that Middle Upper Paleolithic people were distributed throughout subarctic and arctic Siberia (north of 55°N), but excepting Yana itself, virtually no sites are known there before 20 ka. he most stunning claim to the contrary comes from Diring Yuriakh at 61°N in the Lena River Basin, where TL dating shows that eolian (windblown) sands that overlie more than 4,000 laked stones accumulated between 370 and 260 ka. he dating is intriguing, but the laked pieces are not clearly artifactual. he next oldest artifacts may come from alluvial de­ posits at Ezhantsy, Ikhine I and II, and Ust’­Mil’ II, also in the Lena Basin. In each case radiocarbon dates on wood fragments or stratigraphic cor­ relations to radiocarbon­dated deposits elsewhere bracket the artifacts between 35 and 20 ka. he dating is questionable, however, partly be­ cause the artifacts are few and they may have been displaced from higher

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120O 70O

140O

180O

160O

140O

120O

160O

ARCTIC OCEAN

60O

La Ice uren Sh tide ee t

Yana Berelekh

Bluefish Caves

BERING LAND BRIDGE Broken Mammoth Nenana Valley

50O

ALASKA Healy Lake

Co Ice rdille Sh ran ee t

SIBERIA

SEA OF OKHOTSK Ushki Lake

BERING SEA

0

PACIFIC OCEAN

1000 km

FIGURE 7.32. northeastern siberia and northwestern north america, showing the extent of the bering landbridge in relation to the modern land configuration and the maximum extent of the Last Glaciation ice sheets. Some important archaeological sites that may bear on the initial colonization of the Americas are also shown. Arguably, the oldest are the Bluefish Caves (Yukon) (Ackerman 1996; Morlan 1987), where putative bone artifacts have been radiocarbon-dated to nearly 25 ka. Less controversial are Berelekh (Yakutia, Siberia) (Mochanov and Fedoseeva 1996) and Ushki Lake (Kamchatka Peninsula, Siberia) (Dikov 1996; Goebel et al. 2003), which were occupied by 13 ka, and sites in the Tanana Basin, the Nenana Valley, and at Healy Lake (all central Alaska), which were occupied between 12 and 11 ka (Hoffecker and Elias 2003; Holmes 2001; West 1996a). At Broken Mammoth in the Tanana Basin, the artifacts are accompanied by numerous animal bones that probably represent kitchen debris (Holmes 1996). The first human penetration of the landbridge may have occurred only about 12 ka, when trees that could supply fuel reappeared after an absence of 20 ky or more. (Base map modified after Szathmary [1993], fig. 1).

levels, partly because some of the dated wood fragments could have been reworked from older sediments (this could explain some stratigraphic inversions in the dates), and partly because there are inconsistencies in the published stratigraphic descriptions. Seemingly conirmatory radio­ carbon dates on bone are equally problematic because people ater 20 ka probably sometimes obtained bones from older, natural bone accumula­ tions, like the mammoth “cemeteries” that occur near Berelekh and other sites. Such an origin could explain the dated Yana bone artifacts, but it is obviously unlikely for the dated bones that exhibit butchery marks. Human occupation of central and northern Siberia between 20 and 10 ka is uncontroversial and has been well­documented at Verkhne­ troitskaya, Kukhtuj, Berelekh, Mayorych, Ushki Lake, and Dyuktaj Cave (locations in igs. 7.15 and 7.32). Arguably, occupation began only about 18 ka, following the exceptionally cold and dry conditions of the the Last Glacial Maximum when people may have been forced to abandon Siberia

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almost entirely. However, radiocarbon dates from a handful of sites suggest that some populations may have found a way to cope before this. Sites that represent the Siberian “late Upper Paleolithic,” postdating 20–18 ka, occur in all three major climatic zones—temperate, subarc­ tic, and arctic. At least twenty have been excavated, and they tend to share microblades struck from polyhedral “wedge­shaped” (Gobi) cores and grooved or slotted antler points into which the microblades were inserted. Similar, broadly contemporaneous artifacts from adjacent re­ gions, including Japan, suggest a distinctive northeast Asian late Paleo­ lithic cultural zone. he artifact assemblages from individual Siberian sites or layers within sites are mostly small, and structural remnants, where they occur, suggest relatively simple, transitory housing. he sum may imply a highly mobile lifestyle focused on migratory herds of rein­ deer, bison, and other large mammals. Considered in its entirety, from 40–35 to 10 ka, the Siberian late (or Upper) Paleolithic difered from its west Asian and European counter­ parts in detail, but it shared with them the routine manufacture of bone, ivory, and antler artifacts; the presence of readily identiiable, oten spec­ tacular art objects and personal ornaments; the construction of dwell­ ings; relatively elaborate burial of the dead; signiicant stylistic variation through time and space; and so forth. In general, Siberian late Paleolithic people seem to have lived very much as did their European contempo­ raries, subsisting largely on gregarious herbivores that had become espe­ cially numerous and widespread under glacial climatic conditions.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: Siberian Mousterian (Vasil’ev 1993); identiication of Neanderthal teeth at Denisova and Okladnikov Caves (Turner 1990b); Neanderthal mitochrondrial DNA in subadult bones from Oklad­ nikov Cave (Krause et al. 2007b); Kara­Bom (Brantingham et al. 2001; Derev’anko and Markin 1998; Goebel 1999; Goebel et al. 1993); Siberian Upper Paleolithic between 40 and 30 ka (Dolukhanov et al. 2002; Germonpré and Lbova 1996; Goebel and Aksenov 1995; Kuzmin 1997; Vasil’ev 1993); colo­ nization of Japan (Ikawa­Smith 1986; Ono et al. 2002; Reynolds 1985; Reynolds and Barnes 1984); Yamashita­cho Cave (Trinkaus and Ruf 1996); Mal’ta (Medvedev 1998); Yana RHS (Pitul’ko et al. 2004); Diring Yuriakh—pro (Waters et al. 1997) and con—(Kuzmin 1997); possible occupation sites older than 20 ka in subarctic and arctic Siberia (Dolitsky 1985; Hopkins 1985; Kuzmin 1994, 1997; Kuzmin and Tankersley 1996; Larichev et al. 1992; McBurney 1976; Morlan 1987; Yi and Clark 1985); Siberian Upper Paleolithic between 20 and 10 ka (Dolitsky 1985; Klein 1971; Larichev et al. 1990, 1992); occupation of Siberia at the Last Glacial Maximum—con (Goebel 1999, 2002; Hofecker 2005a), pro (Vasil’ev et al. 2002); Siberian late Upper Paleolithic artifacts and lifestyle (Goebel 2002)

The Americas

For consistency with prior sections, the geologic ages presented below are based on uncalibrated radiocarbon dates, and they are likely to be 2–3 ky younger than true calendar ages, for reasons discussed in chapter 2. he diference is important for establishing correspondences between events dated by radiocarbon and by other numeric methods, but it is inconse­ quential in the span of human evolution, which is the focus of this book.

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Both genes and physical characteristics imply that historic Native Americans derive overwhelmingly from northeast Asians. Native Americans and northeast Asians share the same unique mtDNA and Ychromosome haplotypes, and they are strikingly similar in the derived “Mongoloid” form of their skulls, featuring especially broad, short braincases; broad, lat faces with high, frontally directed cheekbones; narrow noses; and high frequencies of “Sinodont” dental traits. hese include strong tendencies for the upper incisors to have shovel­shaped crowns, for the upper third molars to be unusually small, for the lower irst molars to have an extra (third) root, and for the lower second molars to exhibit ive (vs. four) cusps. “Sinodonty” stands in contrast to “Sundadonty,” a less specialized dental pattern that characterizes southeast Asians and their Polynesian relatives, all of whom tend to have nonshoveled upper incisors, large upper third molars, and so forth. he Sinodont/Sunda­ dont split occurred only ater 30 ka, and this may place a lower limit on the time when the Americas were irst colonized. he oldest known American skeletal remains, dated between 11 and 8 ka, are less clear in their implications for Native American ori­ gins. Skulls that recall those of historic south Asians, the Ainu people of northern Japan, or Polynesians dominate the small North American sample, while skulls that are more like those of indigenous Australians or sub­Saharan Africans dominate the larger South American assemblage, recovered mainly from caves in the Lagoa Santa region of central Brazil. he accompanying dentitions are sometimes Sinodont and sometime Sundadont. he famed skeleton of “Kennewick Man” from the Columbia River Gorge, Washington State, illustrates the common contrast with re­ cent Native Americans since the skull initially suggested that the owner was not a Native American, but a radiocarbon date of roughly 8.4 ka demonstrated that it was, just a very early one. Following a long struggle about who should have custody, the Kennewick skeleton is now available for scientiic analysis. he variation in early Native American cranial and dental form does not mean that Native Americans originated variously in southern Asia, Australia, or Africa since the shared Mongoloid traits that linked historic Native Americans and northeast Asians appeared in both populations only ater 8–7 ka. As discussed below, this discovery and the observation of ancient American skulls and teeth may instead imply a complex suc­ cession of population dispersals from northeastern Asia. Determining when these began depends mainly on archaeological evidence in north­ eastern Siberia, in the Americas, and in the region of the Bering Strait, where Siberia and North America approach each other most closely. he growth of glaciers requires moisture as well as cold, and dur­ ing late Paleolithic times Siberia was mainly too dry for large glaciers to form. However, the huge ice sheets of Europe and especially North

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America locked up so much water that sea level fell by up to 140 m, ex­ posing wide swaths of land at the margins of the continents. A dry land­ bridge up to 1,000 km wide formed across the Bering Strait (ig. 7.32), and Alaska and the adjacent ice­free areas of northwestern Canada be­ came an extension of Siberia, largely separated from the rest of North America by the thick ice sheets that covered most of Canada. he last glacial ice sheets began to melt about 14 ka, but the land connection was not fully severed until about 10 ka. Most archaeologists agree that the irst Americans were simply Siberian Upper Paleolithic people who like the saiga antelope, the yak (Bos grunniens), and other north Asian spe­ cies naturally extended their range eastward across the landbridge. he timing of the extension is not established, but ater many years of ex­ ploration and commercial activity on both sides of the landbridge, the oldest unequivocal sites all postdate 13–12 ka. Older sites may be absent because the landbridge was sparsely vegetated between 25 and 14 ka and supported few large herbivores. Perhaps even more important, wood (for fuel) was absent until about 13 ka, when climatic amelioration al­ lowed trees to recolonize sheltered valleys. he earliest known Alaskans let behind endscrapers, sidescrapers, distinctive small teardrop­shaped and triangular bifacial points, and other artifacts that have been assigned to the Nenana Complex. his was irst identiied at the Walker Road and Dry Creek sites in the Nenana Valley of central Alaska, and it has been radiocarbon­dated to between 12 and 11 ka. It has no obvious northeast Asian counterpart, but it was succeeded between 11 and 10 ka by the Denali Complex, which includes wedge­shaped cores, lanceolate points, and other items that closely recall Siberian artifacts postdating 20–15 ka. he Alaskan archaeology may imply two early west­to­east migra­ tions, perhaps corresponding to two of the movements that linguists, skeletal biologists, and some geneticists have postulated. More spe­ ciically, Nenana and Denali could represent, respectively, the original speakers of the two oldest American linguistic stocks: Amerind, wide­ spread throughout the Americas, and Na­Dene (Athabaskan), conined largely to Alaska and western Canada with outliers in the southwestern United States. Speakers of the third major stock, Eskaleut (or Eskimo­ Aleut), probably arrived more recently and settled almost exclusively in the Arctic. At the moment, the most serious objection to this scenario centers on the integrity of the Amerind linguistic stock. Many authorities believe that this is an artiicial conglomeration of unrelated languages and that such great linguistic diversity would require tens of thousands of years to develop, assuming it stemmed from a single source. Alterna­ tively, the heterogeneity of the putative Amerind phylum might imply multiple west­to­east migrations ater 12 ka.

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he antiquity of human occupation south of Alaska has been hotly debated for decades. Based primarily on estimates for the last shared mtDNA ancestor of Native Americans, geneticists have oten postulated human arrival between 30 and 15 ka, but if there were multiple migra­ tions, the last shared ancestor may have lived in eastern Asia rather than the Americas. Many, probably most, archaeologists see little or no com­ pelling evidence for human presence in the Americas before 12–11 ka, while some stress sites where stone artifacts or other humanly modi­ ied objects may be much older. Most such sites are problematic, and their claimed antiquity variously depends on absolute dates that are in­ consistent with each other or with other stratigraphic evidence; on the crude nature of the artifacts; on laked stones whose artifactual quality is oten arguable or whose stratigraphic context is not clear; on charred wood, earth, or bones that could have been burned naturally; or on other questionable observations. As a result, the “shelf­life” of each claim has averaged less than ten years, but fresh examples continue to surface, and there is always the possibility that one will be valid. Recent claims for the rockshelter of Toca do Boquerião da Pedra Furada in northeastern Brazil, for the streamside site of Monte Verde in south­central Chile, and for the Paisley 5 Mile Point Caves, south­central, Oregon, have received especially wide attention. At Pedra Furada, a stratigraphically consistent series of radiocarbon dates implies human occupation from before 48 ka until roughly 6.1 ka. he cultural origin of the laked stones and of well­delineated hearths dated between 10.4 and 6.1 ka is not disputed, but the evidence for older occupation is more problematic because it is based primarily on crudely laked quartzite cobbles and on dispersed charcoal. he cobbles origi­ nated from a cemented layer (or conglomerate) in the clif face approxi­ mately 100 m above the shelter, and the laked examples might thus be “geofacts,” created when cobbles weathered out naturally and struck the hard ground below. Such a process is unlikely to have mimicked human laking very oten, but the laked specimens were selected from a vastly larger number of unlaked ones. Occasional natural brushires nearby could similarly explain the dispersed charcoal, and Pedra Furada may already have joined the long list of dubious claims referred to above. At Monte Verde, eight battered or crudely laked stone artifacts, three naturally fractured cobbles that show traces of use, and iteen nat­ urally fractured cobbles that were apparently carried to the locality but not necessarily used were found deeply buried in riverine sands. Char­ coal from the same level nearby has been radiocarbon­dated to more than 33 ka. Basically similar but more numerous naturally fractured or crudely laked cobbles also occur at a higher level, dated to about 12.5 ka, where they are associated with bones and hide fragments from at

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least seven gomphotheres (mastodon-like proboscideans), a wishboneshaped mound of sand, gravel, and animal fat, parts of many diferent edible or medicinal plants drawn from a wide range of environments, and a mass of logs and branches, all preserved in dense, boggy deposits. he excavator believes that the logs and branches mark the bases of twelve small, adjacent rectangular huts. At the Paisley Caves, the oldest human occupation is marked by cop­ rolites (fossil feces) that have been directly dated to 12.3 (radiocarbon) ka. Based on form alone, the coprolites might derive from a nonhuman spe­ cies, especially perhaps a canid, and three specimens have provided canid mtDNA. However, six have also provided distinctively Native American mtDNA (from haplogroups A and B), and it is primarily for this reason that they are thought to imply ancient human occupation. he excava­ tors discount the possibility that the Native American mtDNA could have intruded the coprolites, perhaps when later occupants of the caves urinated inside, but there are no stone tools or other occupational debris to support human, as opposed perhaps to canid, occupation at 12.3 ka. In the rarity or absence of indisputable cultural refuse, the Monte Verde and Paisley Cave sites difer from numerous like­aged (or older) sites in Eurasia and Africa and also from sites that postdate 12 ka in the Americas. he famous Folsom open­air site, New Mexico, illustrates a key younger site. Excavations at Folsom in 1927–1928 repeatedly pro­ duced carefully crated, luted bifacial lanceolate points among abundant bones of an extinct bison. he lutes (elongated scars proceeding from the base on both faces) would have facilitated mounting on wooden spear shats, and the spears were presumably used to kill the bison. he artifact­bone association, observed in place by acknowledged authori­ ties, was undeniable and it was the irst to demonstrate that people had entered the Americas before the end of the Pleistocene. he Folsom site itself is now known to date to about 10.5 (radiocarbon) ka. Its geomor­ phic/paleontological context showed archaeologists where to seek simi­ lar occurrences, and by 1940, the Folsom result had been replicated at a dozen other localities. he occupations of Monte Verde at 12.5 ka and the Paisley Caves at 12.3 ka are not only older than Folsom, they are as old as or older than any other widely accepted archaeological site in the Americas, including sites on the American side of the Bering landbridge, and if both sites are accepted at face value, their practical and theoretical implications are profound. Archaeologists will be hard­pressed to explain either why, in contrast to contemporaneous Eurasians and Africans, Native Americans seem to have been so rare in the millennia preceding 12 ka or, conversely, why, in contrast to Native Americans before 12 ka, contemporaneous Eurasians and Africans let hundreds of rich, indisputable sites with well­ made artifacts and an abundance of other unmistakable cultural debris.

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here is also the problem that Last Glacial migrants from Siberia to Alaska would have found it diicult or impossible to spread farther south in North America. Canada was almost completely covered by two great ice sheets, the Cordilleran on the west and the Laurentide on the east, and if people arrived in Alaska before 25 ka, they could have moved south only through a relatively narrow, inhospitable ice­free corridor in western Canada. Alternatively, if they arrived only between 25 and 13 ka, during the peak of Last Glacial cold, continuous ice would have blocked their dispersal southward. If they were seafarers with the ability to hunt from boats, they might have been able to travel down the north­ west Paciic Coast beginning 17 ka, but archaeology on both sides of the Bering Straits records such people only ater 5 ka, and for others, move­ ment south would have become practical only beginning 13 ka, when the ice sheets began to pull apart. Together with the simultaneous devel­ opment of somewhat warmer, moister climate in the Beringian source region, this could help explain why the oldest irm and universally ac­ cepted archaeological sites south of the Canadian/American border date from only about 11 ka. hese sites are commonly assigned to the Clovis Complex of the Paleo­Indian Tradition, which extended from southern Canada as far south as Panama. It is especially well­known at carefully excavated sites in the western United States (ig. 7.33). he characteristic artifacts of the Clovis Complex were bifacial, concave­based, lanceolate, luted projectile points that were larger and usually less carefully crated than subsequent Folsom points. Clovis as­ semblages also include endscrapers, sidescrapers, large bifacial imple­ ments, occasional burins (known as “gravers” in North America), and bone or ivory implements that recall late Paleolithic examples in Eurasia. he Clovis Complex provides a plausible origin for its Folsom successor, but its own origins are obscure. Arguably, it stemmed from the Nenana Complex of Alaska, which may be two or three centuries older and con­ tains bifacial points that difer from Clovis ones mainly in smaller aver­ age size and lack of luting. Clovis people did not penetrate South America, but other groups reached the very tip of the continent by 11 ka. he earliest South Ameri­ cans made a range of regionally diverse projectile points that imply either a long­standing pre­Clovis occupation or a very rapid diferen­ tiation from a Nenana­ or Clovis­like base. At present both alternatives may seem improbable, but a rapid diferentiation is less so, and it would imply that people colonized South America not in a smooth wave, but in a series of skips and jumps that quickly isolated populations in pockets of especially favorable habitat. he ecological shock of human arrival, perhaps combined with dra­ matic climatic change in the transition to the present interglacial, may explain why North America lost thirty­ive large mammal genera, more

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Nenana, AK

Broken Mammoth, AK

Wally’s Beach, AB Colby, WY Mill Iron, MT Lange-Ferguson, SD Hell Gap, WY Kanorado, KS

East Wenatchee, WA Indian Creek, MT Paisley Caves, OR Anzick, MT Sheaman, WY Bonneville Estates, NV Union Pacific, WY Arlington Springs (Santa Rosa Island), CA Dent, CO Lehner, AZ Murray Springs, AZ Blackwater Draw, NM

Shawnee-Minisink, PA Paleo Crossing, OH Jake Bluff, OK Sheriden Cave, OH Domebo, OK Cactis Hill, VA Sloth Hole, FL Aubrey, TX Lubbock Lake, TX

cm

0

2000 km

Clovis sites other sites of roughly Clovis age

Monte Verde, Chile Cerro Tres Tetas, Cuevas Casa del Minero, & Piedra Museo, Argentina Fell’s Cave, Chile

FIGURE 7.33. The main in situ occurrences of Clovis Paleo-indian artifacts and of widely accepted like-aged archaeological sites in the americas (map modified after Waters and Stafford [2007], fig. 1; Clovis point redrawn after Wormington [1964], fig. 68). The most distinctive Clovis artifacts are carefully crafted bifacial, concave-based projectile points with one or more basal flutes (elongated flake scars) proceeding from the base on both faces. Accelerator radiocarbon dates from eleven key sites bracket the Clovis Culture tightly between 11.05 and 10.8 ka (Waters and Stafford 2007), and Clovis constitutes the oldest widely accepted presence for human presence in North America south of the Canadian/U.S. border. Sites of the Nenana Complex in Alaska are perhaps three centuries older, and they might provide the base from which Clovis later developed. North American sites that lack characteristic Clovis artifacts but that are roughly the same age may imply that Clovis and like-aged traditions differentiated from an earlier, pre-Clovis base. The most significant non-Clovis sites are perhaps Mill Iron (Montana) and Hell Gap (Wyoming) assigned to the Goshen Complex and dated to nearly 11 ka. The Clovis culture never penetrated South America, but equally early sites with distinctive projectile points are known from its very tip. Fells Cave, Chile, radiocarbon-dated to 11 ka, is perhaps the best known, and together with three like-aged sites in southwestern Argentina, it may also imply a pre-Clovis occupation of the Americas. Monte Verde, also Chile, may document such an occupation at 12.5 ka, but its archaeological nature is controversial, as discussed in the text.

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than 70% of its total, between perhaps 12 and 10 ka. Six continued to exist elsewhere, but the remainder vanished entirely. An even larger number apparently disappeared from South America at about the same time. he lost “megafauna” included mammoths, horses, and native camels whose brief association with Paleo­Indians of the Clovis Complex is now well­ established. he possibility that the Paleo­Indians caused or contributed to large mammal extinctions is a potent reminder of the level of hunting­gathering competence that late Paleolithic peoples had probably achieved. Similar, though less extensive, extinctions occurred at or near the Pleistocene/ Holocene transition in Eurasia and Africa, where people perhaps deliv­ ered the coup de grâce to species whose numbers and distribution had already been reduced by environmental change. Environmental change alone is an inadequate explanation for the American, Eurasian, and Afri­ can extinctions since the extinct species had survived the similar change that occurred at the end of the Penultimate Glaciation, roughly 130 ka. What diferentiated the end of the Last Glaciation most clearly was the presence of more advanced hunter­gatherers, whose behavioral innova­ tions and capabilities have been emphasized here.

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SOURCES: radiocarbon calibration and the earliest Americans (Fiedel 2002); cranial resemblances between historic Native Americans and east Asians (Howells 1973a, 1995); Sinodonty and Sundadonty (Turner 1985, 1987, 1989, 1990a, 1995a); early Native American skeletal remains (Jantz and Owsley 2001; Lahr 1995; Lahr and Foley 1998; Neves et al. 2004; Neves and Pucciarelli 1998; Powell and Neves 1999; Steele and Powell 1993); Kennewick Man (Chatters 2001; Dalton 2005; Holden 1999; Huckleberry et al. 2003; Taylor et al. 1998); oldest sites in Beringia (Dumond 1982; Hofecker 2005a; Hofecker et al. 1993; Hofecker and Elias 2003, 2007); vegetation on the Bering landbridge (Colinvaux 1996; Col­ invaux and West 1984; Guthrie 1990); Nenana Complex—artifacts (Goebel 1999; Goebel et al. 1991) and age (Hofecker 1996; Hofecker et al. 1993; Powers and Hofecker 1989); Denali complex (West 1996b); migrations across the Bering Straits—implied by language (Greenberg 1987, 1996; Greenberg et al. 1986). by genes (Eshleman et al. 2003; Hey 2005; Kitchen et al. 2008; Schurr and Sherry 2004; Szathmary 1993; Torroni et al. 1992, 1993, 1994; Wallace et al. 1985; Zegura 1987; Zegura and Karafet 2004); integrity of the Amerind lingustic stock (Nichols 1990); debate over antiquity of people south of Alaska (Bray 1988; Martin 1987; Meltzer 1995); problematic American sites >12 ka (Dincauze 1984; Fiedel 2002; Lynch 1990; Owen 1984); Pedra Furada—pro (Guidon 1989; Guidon and Arnaud 1991; Guidon and Delibrias 1986) and con (Meltzer et al. 1994); Monte Verde—pro (Dillehay 1984, 1987, 1989, 1997; Dillehay et al. 1982; Dillehay and Collins 1988; Meltzer 1993a, 1997; Meltzer et al. 1997; Morlan 1990) and con (Fiedel 1999; Haynes 1999); Paisley 5 Mile Point Caves (Gilbert at al. 2008); Folsom site (Meltzer 2005); ice­free corridor (Jackson and Duk­Rodkin 1996; Rutter 1980; Wright 1991); possible movement of the irst Americans along the Paciic Coast (Dillehay et al. 2008; Goebel et al. 2008); age of the oldest widely accepted American archaeological sites (Haynes 1993); Clovis Complex (Haynes 1980, 1982, 1984; Meltzer 1993b, 1995); accelerator radiocarbon dating of Clovis and like­aged sites (Waters and Staford 2007); Nenana­Clovis connection (Goebel et al. 1991; Hofecker et al. 1993); Clovis contemporaries in South America (Dillehay 1999; Dillehay et al. 1992; Meltzer 1995); early projectile point traditions in South America (Haynes et al. 1997b; Roosevelt et al. 1996); early occupation of southernmost South America (Miotti and Salemme 2003); terminal Pleistocene mam­ mal extinctions in the Americas (Kurtén and Anderson 1980; Martin 1984), in Eurasia (Stuart 1991; Vereshchagin and Baryshnikov 1984), and in Africa (Klein 1984); Paleo­Indians responsible for ter­ minal Pleistocene extinctions—(Fiedel and Haynes 2004; Martin 1967; Mossiman and Martin 1975), not responsible (Graham and Lundelius 1984; Grayson and Meltzer 2003; Guthrie 1984); mammalian extinctions and human evolution (Klein 2000)

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Australia, New Guinea, and Tasmania (Sahul)

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Unlike the Americas, Australia has not been connected to another continent since the late Cretaceous, roughly 70 Ma. Lower sea levels dramati­ cally enlarged the Malay Peninsula, fusing it with Sumatra, Java, Borneo, Bali, and smaller Southeast Asian islands to form a subcontinent paleo­ geographers call “Sunda Land,” while Tasmania and New Guinea were similarly joined to Australia to produce a supercontinent called “Sahul Land.” However, Sunda and Sahul always remained separate (ig. 7.34). Travelers between the two could have island hopped, but some open­sea travel remained unavoidable, minimally including at least one voyage of 70–90 km and three others of 30 km. Such distances required boats that could maintain buoyancy for several days, and arguably only late Paleolithic people, with the fully modern ability to innovate, could have designed them. he boats themselves probably perished long ago, and in any case the sites where remnants might persist are now inaccessible on the drowned continental shelves of Sunda and Sahul. Until the mid­1960s most archaeologists thought that people colo­ nized Sahul only ater 10 ka, but it is now clear that they arrived much earlier. here is a striking parallel with the investigation of early Ameri­ can colonization, for once the accepted late date in Sahul was breached, it was rapidly pushed back. By the early 1980s, a date of 40 ka was widely accepted, and there are presently more than ity sites scattered through­ out Sahul that unequivocally document human presence between 40 and 10 ka. Debate today centers on claims for signiicantly older sites. In the mid­1990s, TL dates on artifact­bearing sands at Jinmium Rock­ shelter, Northern Territory, Australia, briely appeared to support hu­ man presence between 176 and 116 ka. TL at Jinminum also placed sands containing an exfoliated, engraved wall fragment between 75 and 58 ka, 20 ky earlier than the oldest well­documented European rock art. How­ ever, the radiocarbon and OSL methods subsequently showed that the entire Jinmium deposit postdates 10 ka. Jinmium now excluded, the old­ est proposed Australian sites are the Malakunanja II and Nauwalabila I rockshelters, also Northern Territory, that contain artifact­bearing sedi­ ments that luminescence places between 60–50 ka, and the open­air ar­ chaeological sites at Lake Mungo, New South Wales, that luminescence, ESR, and U­series dates place as early as 62 ka. he Lake Mungo sites are particularly notable because they have also provided the oldest known Australian human remains, discussed below. Dates of 60–50 ka for Malakunanja II and Nauwalabila I are widely accepted, but their bearing on human occupation is questionable since termite activity is likely to have displaced dated artifacts downward from deposits that formed less than 40 ka. he problem at Lake Mungo centers on a conlict between luminescence estimates that could sup­ port occupation at 60 ka and ones that would imply a time nearer 45

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present-day land sea bottom exposed at time of maximum glaciation

Tabon

Malakunanja Lene Hara, & Nauwalalabila Jerimalai & Yombon & Misilil Matja Kuru (New Britain) (Timor) Buang Merabak Bobongara & Matenkupum (Huon) (New Ireland) Kilu (Buka)

Moh Kwieh Niah

Ngandong

SUNDA

Wajak

715 FIGURE 7.34. map of sunda and sahul, showing the approximate locations of the sites mentioned in the text (modified after Bellwood [1998], fig. 1). Dark gray indicates new land that would be exposed by a 200 m drop in sea level. Even with a drop of this magnitude, substantial stretches of open sea would separate Sahul from Sunda, implying that the earliest human inhabitants of Sahul arrived by boat. Based on present knowledge, they arrived between 45 and 40 ka. Stars mark the oldest well-documented sites.

Liang Bua (Flores) Jinmium

SAHUL

GRE8

Laura South

Mandu Mandu Carpenter’s Gap & Riwi Allen’s Cave Devil’s Lair most likely routes from Sunda to Sahul 0

600

1200 km (at Equator)

L. Mungo & Willandra Lakes L. Tandou Talgai

Wyrie Swamp Keilor

Kow Swamp, Coobool Creek, Nacurrie & Cohuna

ka. Radiocarbon dates and the stratigraphic position of the archaeological remains generally support the younger luminescence estimates. A comprehensive review of the evidence at Malakunanja II, Nauwalabila I, Lake Mungo, and other less celebrated localities that could antedate 45 ka, suggests that none certainly do. he same review concludes that the oldest securely dated sites in Sahul are Allen’s Cave, Berang Mera­ bak, Carpenter’s Gap, Devil’s Lair, GRE 8, Bobongara Point (Huon Pen­ insula), Lake Mungo, and Riwi, all of which document human presence between 45 and 40 ka. In theory, the diversity of Australian Aboriginal mtDNA could help pinpoint the timing of initial colonization, but it is consistent with an en­ try at either 60 or 45 ka. However, only an entry at 45 ka meshes with Out of Africa at roughly 50 ka. Entry at 60 ka or before, if this is eventually established, would force revision in the Out­of­Africa model. We would

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have to accept a local origin for the modern Australian Aborigines, a dispersal of modern humans from Africa that proceeded irst along the south Asian (Indian Ocean) coast and only later from there to Europe, or two separate dispersals—an earlier one along the south Asian coast, perhaps partly by boat about 60 ka, and a later one northward and then westward around the Mediterranean Basin that brought modern hu­ mans to Europe roughly 20 ky later. he known mtDNA and Y­chromosome haplotypes of Australian Aborigines preclude their local origin since all the haplotypes fall within the founder mtDNA and Y­chromosome clades associated with a mod­ ern human exodus from Africa between 70 and 50 ka. he mtDNA of some south Asians may support a 60­ka dispersal along the south Asian coast, but the timing lacks archaeological and fossil support, and it would mean that the dispersing population must have crossed the Bas­el­Mandeb Strait at the southern end of the Red Sea. A more north­ erly land route would have required passage through the Sinai Desert into what is now Israel, and archaeology shows that fully modern hu­ mans arrived in Israel only ater 50 ka. he Bas­el­Mandeb route cannot be ruled out, but it would also require rapid adaptation to desert condi­ tions on the Arabian Peninsula, and it seems unlikely. In sum, evidence external to Sahul suggests it was colonized no earlier than 50 ka. he time when people irst entered Sahul aside, their seafaring abil­ ity is clear, and by 40 ka some had even reached the oceanic islands of New Britain and New Ireland to the east of New Guinea. Minimally this required two 30­km hops from New Guinea to New Britain and a third voyage of similar length from New Britain to New Ireland. he early col­ onists did not stop at New Ireland, and by 28 ka they had reached Buka, the northernmost island in the Solomon chain. Travel to Buka from New Ireland required either a single direct voyage of 175 km or several smaller, island­hopping trips of up to 50 km each. Fish bones igure prominently in the most ancient sites on both New Ireland and Buka, and ishing may thus be added to seafaring as a marker of local behavioral modernity. Assuming that Sahul was irst colonized by boat from Sunda, the most likely entry points were in the northwest, where coastal environ­ ments closely resembled those of Sunda. To begin with the settlers may have spread mainly along the shore, but they had occupied much of the Australian interior by 25–20 ka. heir artifacts, their hunting­gathering way of life, and, on occasion, even their rock art are now well­documented throughout Sahul, from New Guinea on the north to Tasmania on the south. he earliest Australian laked stone artifacts are remarkably infor­ mal, and they have been assigned to a loosely deined Core­Tool­and­ Scraper Tradition that persisted basically unchanged until roughly 4 ka. Similar stone artifacts occur widely in southeast Asia in late Pleistocene

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and early Holocene deposits, and where they are found alone, the behavioral modernity of the makers can be questioned. However, at several Australian sites the laked stones are accompanied by such advanced behavioral markers as formal bone artifacts, art, complex burials, or a combination of these. A layer that has been radiocarbon-dated to roughly 32 ka at Mandu Mandu Rockshelter, Western Australia, contains artiicially modiied cone shells that are among the oldest known beads anywhere, and the remnant of a rock painting dated to at least 25 ka at Laura South, Cape York Peninsula, rivals European Upper Paleolithic paintings in its antiquity. In addition, stone artifacts that date from at least 40 ka at Bobon­ gara on the Huon Peninsula, northeastern Papua New Guinea, include distinctive large ax­shaped lakes that were intentionally grooved or “waisted” around the middle. Similar tools are known from other New Guinea sites dated to roughly 25 ka, and smaller, morphologically similar lakes with systematically ground edges occur in layers dated between 24 and 20 ka in northern Australia. he grooving or waisting could have simpliied mounting on wooden handles, and the various pieces might then have been used to chop or to work wood. he watercrat by which Sahul was colonized indirectly imply sophisticated working of wood, bamboo, or related materials, but just as on other continents, on Sahul, prehistoric wooden artifacts have rarely survived. A prominent excep­ tion occurs at Wyrie Swamp, South Australia, where dense peat deposits preserved a range of wooden artifacts, including the oldest­known boo­ merangs, dated to 10–9 ka. Among the best­documented and certainly most informative early Australian sites are those at Lake Mungo, one of thirteen now­dry lake basins in the Willandra Lakes district of western New South Wales. As already noted, the dating of the Lake Mungo sites is controversial, but luminescence and 14C together suggest that people exploited shellish, ish, emus, marsupials, and probably plants around the Willandra Lakes from 50–45 ka until ater 40 ka, when climatic change caused the lakes to shrink and eventually disappear. Lake Mungo is best known for parts of three anatomically modern human skeletons labeled Mungo 1–3 that probably date to around 40 ka. Arguably, the skeletons are too incom­ plete for sex determination, although Mungo 1 was initially identiied as female and Mungo 3 as male. Mungo 1 was perhaps 20–25 years old at time of death, and the body had been partially cremated before the bones were intentionally fragmented and placed in a small, shallow pit. he oc­ currence is the oldest known cremation. Mungo 2, found with Mungo 1, was too fragmentary and too poorly represented to analyze. Mungo 3 was an aged adult who had been laid out in a shallow grave and liber­ ally sprinkled with red ocher before burial. European Upper Paleolithic

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people sometimes used ocher in the same way, but the Mungo case is older than any known European one, and the ocher is notable not only for its abundance, but for its probable origin 200 km from the site. Curiously, compared with the historic Australian Aborigines, the Lake Mungo people, together with possibly contemporaneous or some­ what younger ones from Keilor, Victoria, and Lake Tandou, New South Wales, possessed relatively high­vaulted, thin­walled, smooth­browed, spherical skulls with relatively lat faces, whereas the people who lived at Kow Swamp, Coobool Creek, and Nacurrie, northern Victoria, between perhaps 14 and 9 ka and at the probably contemporaneous sites of Co­ huna, Victoria, and Talgai, Queensland, had exceptionally rugged skulls with relatively low vaults, thick walls, lat and receding foreheads, strong browridges, and projecting faces. A similar, arguably even more rugged morphology characterizes a cranial vault known as Willandra Lakes Hominid 50 that eroded from ancient lake deposits just north of Lake Mungo and that may be closer in age to the gracile Mungo specimens. he range of variation is extraordinary and may indicate that Australia was colonized more than once, by very diferent people. A very late in­ cursion, perhaps about 3–4 ka, is suggested indirectly by a major turn­ over in stone­artifact types across the Australian continent and more directly by the coeval introduction of the dingo, a semidomesticated dog widely associated with the historic Aborigines. he Lake Mungo people may derive from a population represented by broadly contemporaneous skeletal remains from Niah Great Cave on Borneo, Tabon Cave on Palawan Island in the Philippines, and further aield, from Liujiang and Ziyang in China. he origin of people like those represented at Kow Swamp, Coobool Creek, and Nacurrie is much less clear, although a fragmentary skull from Moh Kwieh Cave, hailand, could belong to the same group. It has been dated to 25 ka, and it could imply that people like those at Kow Swamp were once widespread in Sunda. Human remains that date to 10 ka and later imply that people similar to the Australian Aborigines lived throughout southeast Asia until perhaps 7 ka when rice farmers from the north and east progres­ sively encroached on them. Based on skull form, the Kow Swamp people and their Australian contemporaries might then be regarded as a stage between Willandra Lakes Hominid 50 and the historic Aborigines in a line that began in Sunda with classic Indonesian Homo erectus and ran through the Ngandong (Solo) people of Java. Remember from previ­ ous chapters that bovid teeth associated with the Ngandong fossils have been bracketed between 53 and 27 ka by the ESR and mass­spectrometric U­series methods. As noted previously, mtDNA from the Mungo 3 skeleton initially seemed to imply an ancient, now­extinct mtDNA lineage, but the result is now more commonly thought to relect modern contamination. In

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contrast, apparently autochthonous mtDNA suggests broad genetic continuity between people like those at Kow Swamp 14–9 ka and the historic Aborigines. he possibility of a much older evolutionary link between Ngandong and Kow Swamp–Coobool Creek–Nacurrie cannot be dis­ counted, but there are at least three arguments against it. First, on present knowledge, the Kow Swamp–Coobool Creek–Nacurrie morphology ap­ parently postdates the more gracile morphology of Lake Mungo, probably by a substantial interval. Second, while the Kow Swamp skulls may appear archaic, the postcranial bones are morphologically modern in every de­ terminable respect. his is particularly clear for the Kow Swamp femurs, which lack the thick cortical bone and peculiarities of shat shape that mark the femurs of archaic Homo. hird, sophisticated morphometric analysis employing several key cranial, facial, and dental measurements does not in fact diferentiate the Kow Swamp skulls strongly from their Lake Mungo predecessors, and their diference from later prehistoric and historic Aboriginal skulls could simply relect their greater average size and robusticity. If the smaller size and reduced robusticity of later people relect relaxed selection under the mild conditions of the Holocene, the especially robust Kow Swamp morphology might relect a local response to the especially harsh conditions of the millennia surrounding the Last Glacial Maximum, 20–18 ka. his was a time of great aridity that dramati­ cally reduced human populations throughout Australia. he origin of the Kow Swamp morphology and, by extension, its less robust historic suc­ cessor might be resolved by the recovery of fossils irmly dated between those of Lake Mungo and Kow Swamp, but Aboriginal objections to the study of prehistoric human remains may preclude this. Like the peopling of the Americas, the peopling of Sahul may be linked to a wave of large vertebrate extinctions. hese involved all nine­ teen marsupial species whose body weight exceeded 100 kg, twenty­two of the thirty­eight species whose body weight was between 10 and 100 kg, three large reptiles, and the ostrich­sized lightless bird, Genyornis newtoni. None of the extinct forms have so far been found at an archaeologi­ cal site where their bones are unequivocally associated with artifacts, but luminescence and U­series dates from twenty­eight sites across Australia and New Guinea suggest that many disappeared more or less simultane­ ously sometime between 51 and 40 ka. Direct dating of eggshell places the last appearance of Genyornis at 50±5 ka. Eight Tasmanian archaeo­ logical caves that have provided the largest humanly collected faunal as­ semblages in Sahul, including more than 600,000 bones, imply that the extinctions occurred before the caves were irst occupied about 34 ka. he assemblages include bones from marsupials that no longer reside locally, indicating diferent (more grassy) vegetation, but no bones from the especially large, totally extinct species that occur in older or less well dated contexts elsewhere.

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Unlike the American extinctions, the Australian ones did not occur at a time of radical climatic change, and the extinct species apparently lourished under the arid conditions of the Australian interior. he sum may pinpoint people in their disappearance. he indigenous Australian fauna included few large predator species, and the herbivores could have been particularly vulnerable to the sudden appearance of an exceptionally potent one. Alternatively, people could have precipitated the extinctions by iring the landscape, which would probably have ad­ versely afected plants that were vital to many indigenous herbivores. he historic Aborigines oten set brushires to aid movement or to con­ centrate game, and stark pollen changes, concentrations of charcoal, or both in ancient sediments suggest the practice has deep roots. Genyornis and most of the large extinct Australian marsupials were browsers, and the historic vegetation across much of the Australian interior was ire­ adapted desert scrub in which they probably could not have prospered. Stable isotopes in eggshell of the Australian emu (Dromaius novaehollandiae) from three scattered Australian localities spanning the last 140 ky suggest the desert scrub may have replaced more nutritious, drought­ resistant vegetation mosaics about 50–45 ka, and early human iring could have been the cause. Human involvement in extinctions remains as debatable in Sahul as it is in the Americas, however. A stronger case will require irmer estimates of when the extinctions took place, together with persuasive stratigraphic associations between artifacts and bones of the extinct species.

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SOURCES (also table 7.1): minimum voyages from Sunda to Sahul (Birdsell 1977); investigation of human antiquity in Sahul (Jones 1979, 1989, 1990, 1992; Lourandos 1987; O’Connor 1995; horne 1980; White and Habgood 1985; White and O’Connell 1979, 1982); Jinmium—original TL dating (Bahn 1996; Fullagar et al. 1996) and corrected dating (Roberts et al. 1998); antiquity of 60–55 ka for Malakunanja II (Roberts et al. 1990) and Nauwalabila I (Bird et al. 2002; Roberts et al. 1994b) and reasons for likely younger age (Allen 1994; O’Connell and Allen 1998); Lake Mungo dating—overview (Grün 2006, 38–41), to 62 ka (Grün et al. 2000; horne et al. 1999) or 50–45 ka (Bowler et al. 2003; Gillespie and Roberts 2000; Olley et al. 2006); oldest archaeological sites in Sahul at ca. 45 ka (O’Connell and Allen 2004, 2007); coalescence of Australian mtDNA haplotypes (van Holst Pellekaan et al. 2006); mtDNA, Y chromosomes, and the recent African origin of Aboriginal Australians (Hudjashov et al. 2007); genetic evidence for early­modern human dispersal along the south Asian coast (Forster and Mat­ sumura 2005; Macaulay et al. 2005); initial occupation of New Britain (Pavlides and Gosden 1994), New Ireland (Allen 1994; Allen et al. 1988; Leavesley 2005; Leavesley et al. 2002), and Buka (Wick­ ler and Spriggs 1988); initial occupation of Sahul along the coasts (Bowdler 1977, 1990); Sahul sites older than 10 ka (Allen and Holdoway 1995; Bowdler 1992; Cosgrove et al. 1990; Davidson and Noble 1992; Jones 1979, 1989, 1992; White and Habgood 1985; White and O’Connell 1982); Huon Peninsula (Groube et al. 1986; Roberts 1997); early Australian stone artifacts (Holdoway and Stern 2004); Mandu Mandu (Morse 1993); Laura South (Watchman 1993); Wyrie Swamp (Luebbers 1975); Lake Mungo sites and graves (Bell 1991; Bowler et al. 1970, 1972; Bowler and horne 1976; Oyston 1996); Mungo sex determination (Brown 2000; horne and Cornoe 2000); Willandra Lakes Hominid 50 (Hawks et al. 2000; Stringer 1998; horne 1984); successive colonizations of Australia (Jones 1992; horne 1977, 1980); cultural change in Australia 3–4 ka (Bowdler 1989; Jones 1989); southeast Asian distribu­ tion of Australian (Australo­Melanesian) phenotype before 6–7 ka (Bellwood 1998; Matsumura and Pookajorn 2005); cranial continuity from H. erectus to Kow Swamp–Coobool Creek–Nacurrie (Hab­ good 1985, 1989; horne and Wolpof 1981; Wolpof 1985a; Wolpof et al. 1984); antiquity of Ngandong

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(Swisher et al. 1996); Lake Mungo and Kow Swamp mtDNA (Adcock et al. 2001); likelihood that Lake Mungo mtDNA is not authentic (Cooper et al. 2001; Smith et al. 2003); Kow Swamp femurs (Kennedy 1984a); morphometric similarities between Lake Mungo and Kow Swamp skulls (Brown 1987) and diferences due to size and robusticity (Brown 1993); Aboriginal objections to the study of prehistoric human remains (Mulvaney 1991); extinct Australian megafauna (Horton 1984; Murray 1984); tim­ ing of Australian megafaunal extinctions (Roberts et al. 2001); extinction of Genyornis (Miller et al. 1999b); lack of totally extinct species in ancient Tasmanian sites (Cosgrove and Allen 2001; Cosgrove et al. 1990); arid adaptations of extinct Australian vertebrates (Prideaux et al. 2007); Aboriginal burn­ ing and extinctions (Jones 1989); stable isotopes in ancient emu eggshell (Miller et al. 2005b); lack of stratiied archaeological sites with extinct megafauna (Cupper and Duncan 2006; White 1996c) and possible counterexamples (Johnson 2005; Trueman et al. 2005); uncertainty on the causes of Austra­ lian megafaunal extinctions (Wroe and Field 2006)

Conclusion Together, the fossil and archaeological records suggest that the modern physical form evolved before the modern capacity for culture. From a strictly behavioral (archaeological) perspective, the earliest anatomically modern or near­modern people were not signiicantly diferent from their nonmodern predecessors and contemporaries, and this probably explains why they were conined to Africa for thousands or even tens of thousands of years. It was only sometime between 60 and 40 ka, when anatomically modern people developed the fully modern capacity for culture, that they were able to spread widely through Eurasia. he im­ portance of their unique behavioral (cultural) capabilities is underlined by the apparent rapidity of their spread as well as by subsequent cultural change and diversiication. Although the basic human form did not change signiicantly in the ensuing 40 ky, the evolution of culture accel­ erated dramatically. Plainly it was culture and not body form that pro­ pelled the human species from a relatively rare and insigniicant large mammal 40 ka to a geologic force that threatens the very existence of so many other species today. However, if the broad outline of modern human origins is now clear, many details remain to be established. A rapid replacement of nonmod­ ern humans is unequivocal only in Europe, particularly western Europe, but even here the evidence is mainly artifactual not physical. Additional skeleton remains of the earliest modern Europeans are needed to con­ irm their African origin beyond a shadow of a doubt. In eastern Asia, the rapidity and even the dating of the replacement is much less clear, and only a much enlarged fossil and archaeological record can conirm the hypothesis that modern invaders essentially extinguished nonmod­ ern residents, as in Europe, and that they rarely if ever interbred with their predecessors. Finally, if we accept that modern human behavior provided the competitive advantage that allowed modern humans to spread from Africa, it remains uncertain what promoted behavioral ad­ vance. Did it follow strictly on social, economic, or technological change,

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as most specialists believe, or was it sparked by a neurological change that fostered fully modern cognitive ability? he issues that require an enlarged fossil and archaeological record will not be resolved quickly, but fresh research in behavioral genetics and the extraction of nuclear DNA from Neanderthal and contemporaneous nonmodern human fos­ sils may soon allow a decision on whether genetic change could have sparked the modern human diaspora.

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Addendum: Homo floresiensis In October 2004 and October 2005, a mainly Australian­Indonesian team announced the 2003–2004 recovery of distinctive human remains from Liang Bua Cave on the island of Flores, eastern Indonesia. he principal ind was the skull and partial skeleton of a diminutive individual called Liang Bua (LB) 1, but the excavations produced elements from at least two and perhaps as many as eight additional individuals. he minimum number—three including LB1—depends on elements that must repre­ sent diferent individuals—two mandibles with the let third premolars (P3’s) in place and an isolated let P3. he maximum number—nine in­ cluding LB1—depends on the assumption that the stratigraphy has been accurately assessed and that bones in diferent layers come from difer­ ent individuals. he dentition of LB1 shows it was a full adult, but its femoral length implies it stood only about 1 m tall, and its body mass has been estimated at between 16 and 29 kg. he remains of other individuals imply compa­ rably small size, and the endocranial capacity of LB1 was only about 400 cc, just above the lower limit for adult australopiths. In some features, including the long, low shape of the LB1 braincase and the absence of a “chin” (mental eminence) on both mandibles, the fossils suggest a min­ iature Homo erectus. In others, including the great length of the arms relative to the legs or the structure of the wrist, LB1 was arguably more australopith­like, while in still others, including the size and proportions of the teeth, it could pass for Homo sapiens. he mix of primitive, derived, and unique features convinced the discoverers to create a new species, Homo loresiensis, nicknamed the “hobbit” ater a mythical creature in J. R. R. Tolkien’s epic fantasies. he discoverers suggested that H. loresiensis descended from H. erectus, which arrived on Flores and then dwarfed in response to the small available living area (about 14,000 sq km). In advance, derivation from H. erectus might seem unlikely since Flores was always separated from other islands to the west and thus from the southeast Asian mainland by at least 19 km of open water. Only people with seaworthy boats could have reached it. Recall from chapter 5, however, that Mata Menge and

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other sites dated to roughly 800 ka on Flores have provided fractured rocks that may be artifacts, and if they are, they imply that H. erectus had the seafaring ability. he initial colonization event would have been unusual, even unique, however, since recurrent ones would have inter­ rupted the dwaring process. he stratigraphic context and associations of the H. loresiensis fos­ sils have been described only briely, but radiocarbon, U­series, ESR, and luminescence dates suggest a duration from before 38 ka to 12 ka. he key specimen, LB1, dates from about 18 ka. he surrounding de­ posits contained laked stone artifacts and animal bones. he artifacts are mostly simple lakes, but they include “points,” “perforators,” blades, and bladelets that are said to imply technological convergence on Homo sapiens. he animal bones come mainly from two large species of moni­ tor lizard (Varanus) and a dwarf form of Stegodon (an extinct east Asian relative of the elephants). hey include pieces that were cut or charred and that therefore imply butchery and cooking. he overlying deposits contain remains of H. sapiens, which also occur in other parts of the cave where they might be as old as those of H. loresiensis. Flores is on one of the likely routes that H. sapiens took from Sunda to Sahul perhaps 45 ka (ig. 7.34), and deposits dated to roughly 38 ka at Jerimalai Shelter and to 30–35 ka at Lene Hara and Matja Kuru 2 caves on the island of Timor, yet closer to Sahul, contain stone artifacts that may resemble those from the upper levels of Liang Bua. he Matja Kuru 2 assemblage also includes simple shell beads, and the Timor artifact makers are assumed to have been H. sapiens, like the earliest occupants of Sahul. hus, from 45–30 ka forward, H. loresiensis and H. sapiens probably shared Flores and per­ haps even Liang Bua. As an alternative to dwaring in H. erectus, the discoverers of H. loresiensis have suggested it might represent the end point of an ancient and otherwise unknown, small­bodied lineage of Homo. his could ex­ plain both its small size and peculiarities of the LB1 postcranium like exceptionally long arms and short legs. Observers have largely ignored this alternative possibility, probably because it seems unlikely that such a lineage could have reached Flores while remaining undetected else­ where. Skeptics have focused instead on what they see as indications of abnormal development in LB1 and on the possibility that it represents a small­bodied modern human alicted with a severe growth disorder. Such disorders can afect both body and brain, leading in the brain to microcephaly, a pathological condition relected in abnormally small size and usually also subnormal function. Microcephaly is more likely than dwarism to explain LB1’s small brain because population­level re­ ductions in average body size within recent mammalian species (includ­ ing H. sapiens) commonly result in only moderate reductions in brain

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size. hus, if H. loresiensis descended from H. erectus and H. erectus were subject to the usual mammalian scaling constraints, even a dwarf form would have had a signiicantly larger brain than LB1. In addition, if the stone tools from Liang Bua imply that H. loresiensis converged behaviorally on H. sapiens, it did so with a brain that was far smaller than the brain of any normally functioning individual of H. sapiens. Since, key functional components of the brain scale closely to brain size in living primates, the implication would be that scaling in H. loresiensis departed from the general primate pattern. From a be­ havioral perspective, this means that 1 cc of brain in H. loresiensis could not have been functionally equivalent to 1 cc of brain in either modern humans or modern chimpanzees. Ancient DNA might resolve the validity of H. loresiensis, but DNA preserves well only in dry conditions, and the Liang Bua deposits are exceptionally damp. In the absence of relevant DNA, the separation H. loresiensis from H. sapiens will probably be questionable so long as there is only one skull. Validation will ultimately depend on the discovery of additional skulls, especially ones whose stratigraphic context and asso­ ciations are more clearly speciied. Even if H. loresiensis is eventually upheld, however, it will be as a curiosity, and its existence would not alter the basic pattern of human evolution outlined in this book.

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

SOURCES: initial description of H. loresiensis (Brown et al. 2004) and its stratigraphic context (Mor­ wood et al.2004, 2005); anatomical arguments for separation from Homo sapiens (Argue et al. 2006; Falk et al. 2005, 2006, 2007; Gordon et al. 2008b; Lahr and Foley 2004; Larson et al. 2007; Tocheri et al. 2007) or for pathology, including dwarism and microcephaly, in an otherwise modern human (Hershkovitz et al. 2007; Jacob et al. 2006; Martin et al. 2006; Obendorf et al. 2008; Richards 2006; Weber et al. 2005); Timor archaeology (O’Connor 2007; O’Connor et al. 2002; Veth et al. 2005); scal­ ing of functional brain components to overall brain size in living primates (Conroy and Smith 2007)

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SYnOPSIS: anaTOmY, BehaVIOr, anD mODern human OrIGInS

8

he previous chapters have outlined ive apparent phases in human evolution: 1.

2.

3.

An australopith (a.k.a. australopithecine) phase between roughly 4.5 and 2.6 million years ago (Ma), when hominins (or humans broadly understood) were distinguished from apes primarily in their lower limbs, which facilitated habitual bipedal locomotion. he australo­ piths were remarkably apelike in brain size and in many aspects of their dentition, trunk, and upper limbs, and like chimpanzees, they probably did not rely on technology for survival. From everything we know about them, if they had survived to the present, we would probably call them bipedal apes. Sparse fossils support biomolecular indications that their last shared ancestor with chimpanzees lived be­ tween 8 and 5 Ma. A phase initiated by the emergence of the genus Homo and the ap­ pearance of the oldest archaeological sites roughly 2.5 Ma. he earli­ est representatives of Homo, dating between 2.5 and 2 Ma are poorly known, but by 2–1.9 Ma, their descendants were distinguished from the australopiths by larger brains and by smaller premolars and mo­ lars. he oldest archaeological sites are clusters of informally laked stones and fragmentary animal bones that imply a greatly increased reliance on technology and carnivory. Brain enlargement, stone lak­ ing, and carnivory probably coevolved, and the use of laked stones and a growing emphasis on animal lesh probably explain the de­ crease in cheek tooth size. A phase marked by the broadly coincidental emergence of Homo ergaster and the Acheulean Hand Ax Tradition roughly 1.7 Ma. In distinction from previous hominins, H. ergaster approximated living people in its nearly exclusive reliance on bipedalism (as opposed to a

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ChaP Ter eiGhT

FIGURE 8.1. a working phylogeny linking the human species discussed in the text. Au. = Australopithecus; P. = Paranthropus; H. = Homo).

Ma

Ma

0

0

0.5

H. neanderthalensis

H. sapiens

0.5 H. heidelbergensis

1.0

H. erectus (China)

H. erectus (SE Asia)

1.0 1.5

1.5 H. ergaster

2.0 H. rudolfensis

2.5

P. robustus P. boisei

2.0

H. habilis

2.5

Au. garhi P. aethiopicus

3.0

3.0

Au. africanus

3.5

3.5 Au. afarensis

4.0 4.5

4.

5.

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Au. anamensis Ar. ramidus

4.0 4.5

mix of bipedalism and tree climbing), in its body size and proportions, and in its reduced level of sexual dimorphism. It was the irst hominin species to invade truly arid, highly seasonal environments, and sometime between 1.7 and 1.4 Ma, it became the irst to colonize Eurasia. A phase marked by the appearance of Homo heidelbergensis and the probably simultaneous shit in Africa from the early to the late Acheu­ lean Tradition about 700–600 ka. H. heidelbergensis equaled or ex­ ceeded living humans in body mass, and it approached living humans in average brain size. Equipped with late Acheulean artifacts, it spread to Europe, where it was probably the irst hominin to gain a perma­ nent foothold. Its European representative was ancestral to the Nean­ derthals, and its African representative evolved into modern humans. A phase signaled by the emergence and spread of fully modern hu­ mans from Africa about 50 ka (thousand years ago).

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Ma 0.00.01

0.05

Middle Paleolithic/ Upper Paleolithic/ Middle Stone Age Later Stone Age

s Y noP s is : anaTom Y , b ehavior , and moder n hum an or iGins Ma 0.00.01

dihedral burin ivory needle

FIGURE 8.2. some common artifact types in the main culture-stratigraphic units discussed in the text. The individual drawings are not to scale.

figurine

endscraper on a blade beveled-base antler point

borer

bone pendants 0.05

simple, convex sidescraper

Levallois flake

Mousterian point

double, straight sidescraper

simple, straight sidescraper

notch

denticulate

backed knife

0.25

0.25

Lower Paleolithic/ Early Stone Age Oldowan Acheulean

1.7

727

hand ax

cleaver

pick

acute-edged flake

steep-edged flake

flake scraper

discoid 1.7

acute-edged flake

steep-edged flake

discoid

flake scraper

core scraper

chopper

hammerstone

2.5

polyhedron

anvil 2.5

he australopiths, the genus Homo, and H. ergaster probably all emerged irst in equatorial Africa, and each may have evolved abruptly, in a punctuational speciation event sparked by global climatic change. H. heidelbergensis and fully modern humans also appeared irst in Africa, and their emergence may have been equally sudden, but it was not clearly linked to climate change. his chapter summarizes the evidence behind each phase or event, with special emphasis on the emergence and spread of modern humans. Arguably this was the most signiicant event that paleoanthropology will ever reveal, and its recency means it has let an extraordinary wealth of fossil and archaeological evidence. Figure 8.1 presents the working phylogeny that underlies this sum­ mary, while igure 8.2 illustrates the basic artifactual changes to which it refers. Figure 8.3 summarizes two highly conspicuous trends referred to

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millions of years ago

FIGURE 8.3. Top: The encephalization Quotient (eQ) in hominins versus time, showing that long-term increase in brain size in Homo was not due simply to increased body size. for each species, eQ is the ratio of actual brain weight divided by the brain weight that would be predicted from body mass in extant hominoids. The sloping line roughly describes the general trend. Bottom: The megadontia Quotient (mQ) versus time in hominins, showing that megadontia irst increased among the australopiths and then declined in Homo. for each species, mQ is the ratio between the summed occlusal areas of P4, m1, and m2 and the sum predicted from body mass in extant hominoids. The sloping lines roughly approximate the trends before and after the appearance of Homo. The data are from mchenry (1994b) and mchenry and Cofing (2000), which explain more precisely how eQ and mQ were calculated. fossils that have been reported since the diagrams were constructed would not alter the basic trends.

0

encephalization quotient

5.5

4.0 3.5

1.0

1.5

2.0

2.5

3.0

4.0

neanderthalensis 0.15-0.1 Ma (early sapiens & neanderthalensis) 0.55-0.2 Ma (heidelbergensis & late erectus)

habilis

robustus

3.0

ergaster rudolfensis africanus

boisei

2.0

3.5

observed brain volume expected volume

extant & Upper Paleolithic sapiens Skhul/Qafzeh

5.0 4.5

0.5

2.5

megadontia quotient

1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44

ChaP Ter eiGhT

afarensis common chimpanzee

1.5 3.0

boisei

observed occlusal area expected area

2.5 robustus

2.0

africanus habilis

afarensis

rudolfensis

1.5

anamensis

1.0 0

ergaster

extant sapiens common chimpanzee

0

0.5

1.0

1.5

2.0

2.5

3.0

3.5

4.0

millions of years ago in the text: a tendency for average brain size to increase through time, even when increases in body size are considered, and a tendency for cheek tooth size irst to increase (in the australopiths) and then to decrease (in Homo). Chapters 4–7 provide the supporting references.

The Australopiths Most genetic studies indicate that chimpanzees and people are more closely related to each other than to any other living species, and if a constant rate of genetic divergence is assumed, protopeople probably split from protochimpanzees only between 8 and 5 Ma. he geographic range of the chimpanzees and the locations of the oldest unequivocal hominin fossils show that the split occurred in tropical Africa, and three sparsely known fossil species—Sahelanthropus tchadensis (Chad), Ardip-

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ithecus kadabba (Ethiopia), and Orrorin tugenensis (Kenya)—may now document it. All three date from between 7 and 5 Ma, and each exhibits cranial or dental characters that suggest hominin (as opposed to great ape) ainities. he species samples share few parts in common, and they could represent a single species or segments of a single evolving lineage. Assuming that they represent separate species and that bipedalism is the ultimate litmus test for hominin status, O. tugenensis is the strongest candidate for hominin statusso far because its femur resembles that of later bipedal species. If future inds demonstrate that either S. tchadensis or A. kadabba were also bipedal, the implication might be that hominins diversiied only shortly ater they appeared.Aan alternative possibility would be that closely related ape species evolved bipedalism indepen­ dently near 6 Ma as they adapted to the thinning of forests and the spread of woodlands in tropical and subtropical Africa. In this case bipedalism could no longer be the deining character of the hominins. he oldest unequivocal hominin fossils come from the Middle Awash and Gona, Ethiopia, where they date between roughly 4.5 and 4.3 Ma. hey have been assigned to the species Ardipithecus ramidus, and they reveal creatures that were strikingly chimpanzee­like in many aspects of their dentition, including their relatively thin dental enamel. heir arms were strongly muscled, with elbows that could have been locked to aid in tree climbing. As presently described (chap. 4), A. ramidus is linked to later hominins mainly by the incisor­like form of its canines and by the forward position of the foramen magnum on the base of the skull. Leg bones that await description are said to show that it was bipedal, but per­ haps less completely so than its more human­like successors. Associated animal remains indicate that it frequented relatively moist, wooded en­ vironments, and it probably resembled chimpanzees in diet and in many aspects of its behavior and social organization. If Ardipithecus ramidus was the only hominin between 4.5 and 4.3 Ma, then about 4.2 Ma it evolved abruptly into Australopithecus anamensis, which is known primarily from Kanapoi and Allia Bay, northern Kenya, and from the Middle Awash, Ethiopia. A. anamensis resembled both chimpanzees and A. ramidus in its relatively large, sexually dimor­ phic canines and in other aspects of its dentition and jaws. Unlike A. ramidus, it anticipated later australopiths in its relatively thick dental enamel and broadened molars. It is the oldest known hominin for which a described leg bone unambiguously demonstrates bipedalism. Like all the australopiths, however, it retained powerful apelike arms, and it probably mixed bipedalism with a signiicant amount of tree climbing. About 3.8–3.7 Ma, Australopithecus anamensis was succeeded by the much better known species, Australopithecus afarensis. Fossils of A. afarensis have been found at no less than seven sites in Ethiopia, Kenya, and Tanzania, but the most signiicant specimens come from Laetoli,

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Tanzania, and especially from Hadar (including Dikika), Ethiopia. he oldest representatives appear to have difered little from A. anamensis, and additional inds may suggest that A. anamensis should be sunk into A. afarensis. Like A. anamensis, A. afarensis was remarkably apelike in all known parts except the lower limbs. It is the oldest hominin species for which a complete skull is known, and this exhibits many chimpanzeelike features, including strong forward projection of the face and jaws, an exceptionally broad and lat cranial base, and a braincase that was small both absolutely and in relation to the size of the face and body. A partial skeleton from Hadar, various isolated lower limb bones, and fossil foot­ prints at Laetoli show that A. afarensis walked bipedally in broadly the same manner as living people, but its upper limb skeleton shows that it also retained an apelike ability to climb trees. Its canines were smaller than those of chimpanzees, but they remained sexually dimorphic, and this may imply that it was chimpanzee­like in its social organization. A. afarensis is usually considered ancestral to all later hominin spe­ cies, but the details are unresolved. In the view that is tentatively accepted here, at or shortly ater 2.9 Ma, it founded three distinct lineages: one that produced Australopithecus africanus, a second that led to Paranthropus (the robust australopiths), and a third that resulted in Homo. A. africanus is known exclusively from South Africa, where it probably existed between 2.8 and 2.3 Ma. In its body, it was arguably even more apelike than A. afarensis, and its habitat included trees that it may have climbed for food and refuge. It had larger molars than A. afarensis, and these were mounted in jaws that could exert great force between the upper and lower cheek rows. In some individuals, the anterior parts of the tempora­ lis muscles met at the top of the skull and promoted a weak sagittal crest along the midline. In its large molars and in the craniofacial structures to which the jaw muscles attached, A. africanus resembled Paranthropus. In its relatively unpneumatized cranial base, arched forehead, reduced sub­ nasal prognathism, and other features, it also resembled Homo. Strictly on morphology then, it is a plausible common ancestor for both the ro­ bust australopiths and Homo, and the second edition of this book tenta­ tively favored this assessment. here was the problem then that Australopithecus (here Paranthropus) aethiopicus, dated between 2.7 and 2.3 Ma in eastern Africa, also provided a plausible ancestor for the robust australopiths. It had even more massive cheek teeth than A. africanus, a well­developed sagittal crest, and the “dish­shaped” face that is oten regarded as a robust aus­ tralopith hallmark. It could be eliminated from robust australopith an­ cestry only by assuming that it evolved similar craniodental features in parallel, as it came to emphasize the same hard or grit­encrusted foods that the robust australopiths may have preferred. Chapter 3 pointed out

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that such parallelism (or convergence)—the evolution of similar features in genetically similar taxa adapting to similar circumstances—occurred repeatedly in primate evolution before the emergence of hominins. As noted above, future research may even show that bipedalism—the deining character of the hominins—evolved more than once in closely related apes responding to shared environmental change. he possibility that Australopithecus africanus was ancestral to both Paranthropus and Homo, and by extension that A. aethiopicus represents an extinct hominin side branch, cannot be ruled out, but it became less likely following the description of A. garhi in 1999. A. garhi is known mainly from a skull dated to 2.5 Ma in the Middle Awash, Ethiopia, but it may also be represented by fragmentary dentitions dated between 2.7 and 2.3 Ma in deposits near Lake Turkana. It was thus a contemporary of both A. africanus and A. aethiopicus. It closely resembled A. afarensis in the shape of its braincase, in its marked subnasal prognathism, and in the relatively large size of its incisors and canines. It difered most conspicuously from A. afarensis in the extraordinary size of the molars and premolars, which approximated the largest known specimens of Paranthropus. However, as the molars wore, they retained their cuspal topography, in contrast to the molars of Paranthropus, which generally wore lat. In further contrast to Paranthropus, in which huge cheek teeth were associated with small front teeth, in A. garhi the rear teeth and front teeth were both large. Among all the known australopiths, based on the pattern of molar wear and on the size relations of diferent teeth, A. garhi anticipates Homo most closely. he same deposits that produced the skull of A. garhi provided hom­ inin limb bones, and if these are assumed to represent A. garhi, it may have combined relatively long humanlike legs with long, powerful ape­ like forearms. A similar coniguration has been tentatively inferred for earliest Homo. he A. garhi site also produced three animal bones with stone­tool cut marks, and the Gona site, about 90 km to the north, has provided the oldest known stone artifacts in deposits that date to 2.6–2.5 Ma, broadly coeval with A. garhi. In sum, from everything we know or can hypothesize about A. garhi, it provides a credible ancestor for Homo, and the case is probably stronger than for A. africanus. A. africanus could still be ancestral to Paranthropus, but if the split between the Homo and Paranthropus lineages occurred in eastern Africa, then by geography, morphology, and temporal position, Paranthropus (ex­Australopithecus) aethiopicus is a better candidate. Future discoveries may force a reevalu­ ation, but the current bottom line is the scheme in igure 8.1, where A. garhi is on the line leading to Homo, P. aethiopicus is ancestral to the later robust australopiths, and A. africanus represents a lineage that became extinct without issue.

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Whatever the origins of Paranthropus, its coexistence with Homo ater 2.5 Ma is not in dispute. By 2 Ma, Paranthropus had itself diferen­ tiated between a “hyperrobust” form, P. boisei, in eastern Africa and a somewhat less robust form, P. robustus, in southern Africa. Arguably P. boisei and P. robustus were simply geographic variants of a single wide­ spread species. Both were “robust” strictly in their massive cheek teeth and in the associated craniofacial structures that relect the power of the jaw muscles. In average body size they were no larger than other aus­ tralopiths, and they were signiicantly smaller than living humans. hey retained the basic australopith structural plan in which small braincases and a somewhat apelike upper body were mounted on legs adapted for habitual bipedalism. hey may have been no more carnivorous than chimpanzees, and they probably did not lake stone, though they may have used pebbles, sticks, and other objects for the same simple tasks that chimpanzees sometimes do. Fossils of Paranthropus actually outnumber those of Homo in depos­ its that antedate 1.5 Ma, but they are rarer in later deposits, and Paranthropus probably became extinct near 1 Ma. Its demise may have resulted from global climatic change that produced drier, more seasonally vari­ able conditions throughout its range, from unsuccessful competition with the evolving genus Homo, or from the interaction of both factors.

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Homo habilis and the Oldest Archaeological Sites Meager fossils show that Homo was present in eastern Africa between 2.4 and 2.0 Ma, but it is well­represented only in deposits that postdate 2–1.8 Ma. Most authorities assign all fossils of very early Homo to a single species, H. habilis, which difered from the australopiths primarily in its larger brain and in its reduced cheek teeth. he increase in brain size was particularly striking since it occurred without any signiicant increase in body size. here is the complication, however, that both endocranial volume and cheek tooth size varied greatly within H. habilis. Some indi­ viduals had large skulls and large australopith­sized teeth, while others had much smaller australopith­sized skulls and relatively small cheek teeth, similar in size to those of the succeeding species, Homo ergaster (or early African H. erectus). his variability might imply extreme sexual dimorphism, or it might mean that H. habilis was actually two species: H. habilis (in the narrow sense) for the individuals with smaller brains and teeth, and H. rudolfensis for those with larger brains and teeth. Chapter 4 accepted the hypothesis of extreme sexual dimorphism, mainly to avoid the inconvenient need to speculate on how two contemporaneous spe­ cies of early Homo might have difered in behavior and ecology. In the phylogeny that is tentatively presented here (ig. 8.1), H. habilis narrowly understood is taken as the stem species for later Homo, and H. rudolfen-

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sis is placed to the side. his admittedly begs an important issue, and the phylogeny may mask an even more complex situation, for it can be argued that neither H. habilis nor H. rudolfensis constitutes a morphologically suitable ancestor for later Homo. We would then have to contemplate a third contemporaneous species of earliest Homo, perhaps an early representative of H. ergaster, which is unambiguously documented in the fossil record only ater 1.8–1.7 Ma. he brain expansion that marks H. habilis (or the variants into which it may ultimately be split) is notable not only in itself but also because it coincided broadly with the appearance of the oldest known archaeo­ logical sites. hese are dated between 2.6 and 1.7 Ma, and they comprise clusters of stone artifacts and fragmentary animal bones that provide the earliest nonanatomical evidence for human behavior. he artifacts are usually grouped in the Oldowan Industrial Complex, named for Olduvai Gorge, where Oldowan artifact assemblages were irst thoroughly de­ scribed. In general, Oldowan tools include a range of sharp lakes and the cores or core (“pebble”) tools from which the lakes were struck (ig. 8.2). Oldowan stone­working technology was primitive by later standards, and individual pieces are notoriously diicult to assign to dis­ crete types. Still, the Oldowan Complex relects an ability to lake stone that living chimpanzees probably cannot acquire, and the artifacts and associated fragmentary animal bones demonstrate a commitment to ar­ tifact manufacture and to carnivory beyond anything known in apes. hey show further that, by 2.6–2.5 Ma, at least one hominin species had developed the uniquely human habit of accumulating garbage at favored spots on the landscape. A causal link between brain expansion, cheek tooth reduction, and stone­tool manufacture follows from the assumption that early Homo (or its immediate ancestor) produced most if not all of the earliest stone tools, and it would be weakened if it could be shown that Paranthropus manufactured a signiicant number. his possibility cannot be evaluated directly, but it has been suggested by a strong similarity in thumb form between Paranthropus robustus and Homo, including H. habilis. humb form in Homo enables the precision grip, while thumb form in chimpan­ zees promotes the power grip. humb bones of A. afarensis indicate that the earliest australopiths had a chimpanzee­like power grip. If natural selection for the precision grip or dexterity in stone lak­ ing drove the development of the human thumb, then it is reasonable to argue that P. robustus made at least some early stone tools. However, P. robustus/P. boisei and early Homo coexisted for perhaps a million years, from before 2 Ma until 1.2–1 Ma. During this long period, stone artifacts changed signiicantly, but evidence exists for only one evolving artifact tradition, not two. Additionally, there is no obvious rupture in the ar­ chaeological record at the time that P. robustus/P. boisei became extinct

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at 1.2–1 Ma. he same kinds of artifacts were made before and ater, when only Homo survived. he most economical conclusion then is that Homo produced most if not all of the earliest stone tools. Conceivably P. robustus applied its precision grip to food processing or to some other tool­ using activity besides stone knapping. Alternatively, the thumb bone that suggests the precision grip in P. robustus may actually have come from early Homo, which is represented in the same deposit (at Swartkrans Cave, South Africa), but by many fewer diagnostic craniodental frag­ ments.

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Homo ergaster and the Initial Colonization of Eurasia In most current accounts of human evolution, Homo habilis (or one of its variants) is said to have evolved into Homo erectus approximately 1.8–1.7 Ma. he oldest specimens of H. erectus include a nearly complete skull, a second partial skull, several lower jaws, and some postcranial bones from East Turkana in northern Kenya, a skull and an associated partial skeleton from West Turkana, a skull from Swartkrans Cave in South Af­ rica, and at least three of the reported skulls from Dmanisi, Georgia. he various skulls difer from those of H. habilis in several features, includ­ ing especially the presence of a conspicuous supraorbital torus (brow­ ridge) across the top of the orbits, thicker skull walls, and a yet larger brain (greater endocranial volume). Much of the increase in brain size, however, may relate to a concomitant increase in average body size, es­ pecially in females. In the conventional scheme, H. habilis is assumed to antedate H. erectus, but two fossils from Koobi Fora (East Turkana) and one from Olduvai Gorge suggest that H. habilis survived long ater 1.7 Ma, to per­ haps 1.45 Ma. In addition, at Dmanisi, the three described skulls that closely match those of east African early H. erectus are accompanied by a fourth that closely resembles skulls of east African H. habilis in the narrow sense. All the Dmanisi skulls are now commonly dated to 1.7 Ma, and their contemporaneity is diicult to reconcile with an evolutionary sequence from H. habilis to H. erectus. A possible explanation is that the Dmanisi skulls record the evolution of H. habilis into H. erectus and that H. erectus subsequently spread from Asia to Africa. However, this makes no sense if H. erectus was actually present in eastern Africa be­ fore 1.7 Ma and if H. habilis persisted there until 1.45 Ma. he apparent contradictions could stem partly from incorrect dates in eastern Africa, at Dmanisi, or in both places, but whatever the cause, they will probably not be resolved quickly. he type skulls of H. erectus come from China and Java, and they arguably difer from those of early African and Georgian H. erectus in having somewhat larger, lower, latter, and more angular braincases, yet

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thicker skull walls, thicker brow ridges from top to bottom, and ear and nasal specializations that could justify placing the African and Georgian fossils in a separate, more primitive species, for which the name Homo ergaster has been proposed. he distinction of H. ergaster from H. erectus is admittedly subtle, and it will require careful, continuous reevaluation as the fossil record grows. However, H. ergaster is provisionally accepted here (chap. 5) as the stem species for all later humans, including H. erectus narrowly understood (in the Far East) and other forms of primitive Homo in Africa and Europe (ig. 8.1). Limb bones and inner ear structure show that Homo ergaster (or “early African H. erectus,” if preferred) was the irst human species to rely almost exclusively on bipedalism, in contrast to the australopiths and even H. habilis, which depended on a mix of bipedalism and tree climb­ ing. H. ergaster was also the irst human species in which body weight and stature increased to approximately their modern values, and it may have been the irst in which males and females difered in size no more than in living people. In extant higher primates, species that are more sexually dimorphic tend to have polygynous mating systems in which males compete vigorously for females, while less dimorphic species tend to have monogamous mating systems in which males and females pair for long periods. Decreased dimorphism in H. ergaster may thus mark the beginnings of a distinctively human pattern of sharing and coop­ eration between the sexes, preiguring the social organization of historic human hunter­gatherers. he appearance of H. ergaster in eastern Africa coincided broadly with the appearance of more sophisticated stone artifacts 1.7–1.6 Ma, and the connection is probably more than coincidental. Among the novel ar­ tifacts, the most conspicuous forms are the shaped bifacial tools known as hand axes (ig. 8.2). Moreover, in distinction from their Oldowan predecessors, the hand ax makers also knew how to strike large lakes, sometimes exceeding 30 cm in diameter, on which they oten made hand axes. Together, hand axes and the associated advances in laking technol­ ogy are hallmarks of the Acheulean Industrial Tradition, named for St. Acheul in northern France, where the complex was irst identiied in the nineteenth century. More sophisticated stone technology, essentially modern body form, and implicit changes in cognition and social organization enabled H. ergaster to expand its range into previously unoccupied environments. It became the irst human species to invade truly arid, highly seasonal en­ vironments in Africa, and it was also the irst to disperse from Africa to Eurasia. he timing of this dispersal is unsettled, and new claims ap­ pear regularly. At the moment, the oldest widely accepted claim is for Dmanisi, Georgia, where fossils of H. ergaster/H. habilis that were just mentioned occur with crudely laked stones in deposits that may have

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formed as much as 1.77 Ma. he dating depends primarily on radiopotas­ sium and paleomagnetic readings that place a basal basalt and overlying luvial deposits within the Olduvai Normal Subchron, between 1.96 and 1.77 Ma. he fossils and artifacts, however, originate from hollows within the luvial deposits, and the sediments that ill the hollows exhibit re­ versed magnetism, which means they must have formed ater 1.77 Ma. hey could have formed just ater 1.77 Ma or up to several hundred thou­ sand years later, and the morphology of the human fossils may present the strongest case for a time near 1.77 Ma. Elsewhere in Eurasia, dates that may support the 1.77­My estimate for Dmanisi have come from two localities in Java, where radiopotas­ sium analysis of volcanic extrusives places H. erectus fossils at 1.8–1.6 Ma, and from the Nihewan Basin in northern China, where paleomagnetic readings bracket a series of artifact horizons between 1.66 and 1.1 Ma. he Javan dates are problematic because the stratigraphic relationship of the dated samples to H. erectus fossils remains unclear, but if the dates are accepted, they could imply again that H. erectus evolved in Asia and subsequently spread to Africa. he dates might also help to explain an archaeological peculiarity—the Acheulean hand ax tradition seems never to have penetrated the Far East, and if H. erectus was present in Java 1.8 Ma, its ancestor could have arrived there before hand axes were invented in Africa. he Nihewan paleomagnetic ages are intriguing, but if people actu­ ally penetrated northern China as much as 1.66 Ma, they should have let traces on the way, perhaps above all in central or southern Asia, and until such traces are discovered, the Nihewan ages should probably be regarded as tentative. For the moment, if the Javan radiopotassium dates and the Nihewan paleomagnetic readings are held in abeyance, nei­ ther Java nor China provides compelling evidence that people reached eastern Asia before about 1 Ma. Hominin fossils and artifacts together suggest that east Asian populations ater 1 Ma followed a diferent evo­ lutionary trajectory than their African, west Asian, and European con­ temporaries. Arguably, east Asian fossils imply two diferent trajectories: one in southeastern Asia, where classic H. erectus morphology persisted from 1 Ma until perhaps 50 ka and a second in China where people by 200 ka combined massive uninterrupted browridges, keeled, lat, receding fron­ tal bones, low vault heights, and other features of classic H. erectus with larger, more rounded braincases, less massive faces, and other advanced features that mark H. sapiens. he Chinese skulls broadly recall those of Afro­European H. heidelbergensis, considered in the next section, and the similarity could imply that H. heidelbergensis extended eastward to China (but not to Java). Alternatively, it could imply that Chinese H. erectus and H. heidelbergensis evolved similar characters in parallel.

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Parallel evolution is tentatively favored here, partly because it is more parsimonious and partly because the Chinese archaeological record so far reveals no evidence for a population intrusion. In addition, the later Chinese fossils tend to resemble Chinese H. erectus in a handful of features, including a short maxilla, lat, horizontally oriented cheekbones, a very broad nasal bridge, strong shoveling of the upper incisors, and small third molars (M3s). Artifacts are too poorly known in China and especially in Java to de­ termine if biological divergence may have been accompanied by cultural divergence, but artifacts are well enough known, particularly in China, to show that ancient east Asians never manufactured typical Acheulean hand axes, even if they sometimes produced tools that appear equally sophisticated. It has usually been assumed that Europe was occupied as early as eastern Asia, but many possible European sites that antedate 500 ka are problematic, either because the dates are insecure or because the evi­ dence for human presence depends on laked stones that may be natural in origin. Until recently the oldest known European human fossils and artifacts appeared to date from about 500 ka, and some specialists argued that Europe was colonized only about this time. It now appears, however, that southern Europe (south of the Alps and the Pyrenees) was occu­ pied between 1.1 and 0.8 Ma. he most compelling evidence comprises a fragmentary hominin mandible and associated artifacts dated by paleo­ magnetism and associated mammalian fossils to 1.1 Ma at Atapuerca SE, Spain; numerous fragmentary hominin fossils and associated artifacts dated by paleomagnetism to 0.8 Ma or before at nearby Atapuerca GD; and a hominin skull putatively dated by radiopotassium to 0.9–0.8 Ma at Ceprano, Italy. At Atapuerca GD and perhaps also at Atapuerca SE, the earliest oc­ cupations may have been conined to interglacial periods, and even dur­ ing interglacials, people may have failed to penetrate northern Europe until ater 600 ka. hirty­two laked stones picked from ancient river channel deposits at Pakeield, southeastern England, may indicate spo­ radic occupation of northern Europe as early as 700 ka. However, a compelling case will require localities like those that postdate 600–500 ka, where artifacts and animal bones are tightly stratiied together and where some of the bones exhibit stone­tool marks.

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Homo heidelbergensis and the First Permanent Settlement of Europe In Africa, the Acheulean hand ax tradition comprises two stages—an early one when the hand axes were relatively thick, weakly trimmed, and relatively unsymmetrical, and a later one when they were oten much

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thinner, far more extensively trimmed, and much more symmetrical in both plan and edge view. Late Acheulean artifact assemblages also tend to contain a wider range of well-made lake tools that anticipate those of the succeeding Middle Stone Age. he transition from the early to the late Acheulean is imperfectly dated, but a reasonable inference now is that it occurred 700–600 ka. Late Acheulean hand ax makers then expanded to Europe, where they were probably the irst people to gain a permanent foothold, particularly in the north. A small number of hu­ man fossils, oten only weakly dated, suggest that the earliest European hand ax makers closely resembled their African contemporaries, and at least tentatively, the Europeans and Africans can be assigned to the same species, for which the name Homo heidelbergensis is available. In times past, the same fossils were oten referred to “early” or “archaic Homo sapiens,” but this obscures their evolutionary implications because only the African examples belong to people who probably lie near the origin of modern H. sapiens. Artifact assemblages remained similar between Africa and Europe ater 500 ka, while so far as we know, the Far East continued on its own distinctive course. About 250–200 ka, Acheulean (Early Stone Age/ Lower Paleolithic) assemblages were widely replaced in both Africa and Europe by Middle Stone Age/Middle Paleolithic assemblages emphasiz­ ing a reined lake technology, usually without hand axes (ig. 8.2). By 250 ka, however, the human form had come to difer markedly between the two continents, and by roughly 150 ka Europe was occupied exclu­ sively by the highly distinctive Neanderthals, whereas Africa was inhab­ ited by people who looked far more like living humans. he evolution of the Neanderthal lineage in Europe is particularly well­documented, beginning with the people whose bones accumulated at Atapuerca SH at least 530 ka. European fossils dated between roughly 400 and 150 ka suggest that Neanderthal cranial features did not evolve as an integrated complex but accumulated piecemeal and in variable combinations. he pattern suggests a process driven mostly by random drit (vs. natural selection) in a lineage whose numbers and distribution changed dramatically when glacial conditions replaced interglacial ones and vice versa. Many fewer fossils document the evolution of the modern human lineage in Africa, and their dating is oten insecure. However, no Afri­ can fossils that likely date between 250 and 100 ka exhibit Neanderthal specializations, and both individually and collectively, they far more closely resemble living humans. Ot­cited examples include three skulls from Herto, Ethiopia, that may date from as much as 160 ka, and much more fragmentary fossils from Klasies River Main, South Africa, that are irmly dated between 115 and 70 ka. he especially abundant and com­ plete modern or near­modern human remains from Skhul and Qafzeh

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Caves, Israel, well-dated to between 110 and 80 ka, can be added to the African list since they accumulated at a time when “Ethiopian” mam­ mal species indicate that Africa had expanded ecologically to include the adjacent, southwestern margin of Asia. As a group, the key African fossils (including those from Skhul/Qafzeh) reveal people with relatively short, high braincases overhanging the face in front, in contrast to the long, low braincases and forwardly mounted faces of the Neanderthals. It is this fossil diference that most strongly supports the now­famous Out­of­Africa theory, according to which modern humans spread from Africa about 50 ka to replace the Neanderthals and other equally archaic humans in Eurasia.

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The Out-of-Africa Hypothesis and the Evolution of Human Behavior he Out­of­Africa hypothesis for modern human origins might better be called Out of Africa 3, since it concerns the pattern of human evolution ater Out of Africa 1, the widely accepted initial human dispersal from Africa before 1 Ma, and Out of Africa 2, the less celebrated dispersal of late Acheulean people from Africa roughly 600 ka. However, for simplic­ ity’s sake here, in keeping with present convention, Out of Africa 3 will be known simply as Out of Africa. Genetics have played the leading rule in its scientiic acceptance, but at base, it is grounded in the fossil record, which shows that human populations diverged morphologically between Africa, Europe, and the Far East, especially ater 500 ka. From this time onward, there were at least three evolving human lineages (ig. 8.1): Homo sapiens in Africa, H. neanderthalensis in Europe, and H. erectus in the Far East. he European lineage is the best documented, and as noted above, it is marked by the accretion of Neanderthal craniofacial features, culmi­ nating in the classic Neanderthals between 190 and 130 ka. Physiologi­ cal adaptation to recurrent cold probably explains why the Neanderthals were distinguished by especially broad trunks and short distal limbs. he pertinent African fossil record is much less complete than the European one, but it tentatively suggests a similar evolutionary pattern driven mostly by random genetic drit, in which the craniofacial char­ acters of living people accumulated piecemeal between 250 ka or before and 50 ka. he Far Eastern record is the most sketchy, and as noted pre­ viously, it may actually comprise two distinct evolutionary trajectories: one in southeastern Asia that suggests continuity within Javan Homo erectus from perhaps 1 Ma until 50 ka, and a second in China that may indicate evolution from classic Chinese H. erectus between 1 Ma and 500 ka to populations that, by 200–100 ka, combined classic H. erectus cra­ nial features with ones that recall more ancient Afro­European Homo heidelbergensis.

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As presently revealed, the Chinese fossil record is perhaps the biggest impediment to unqualiied acceptance of Out of Africa, but it is actually not so much contradictory as it is poorly known. China has provided only a handful of fossils—most notably those from Dali, Jinniushan, and Maba—that probably postdate classic Homo erectus, and none of these is precisely dated. In this light, the real problem with the Chinese fossil record is that its bearing on Out of Africa is uncertain. A fuller record may show that, like European H. neanderthalensis, archaic Chinese populations (advanced H. erectus?) were replaced by immigrant African H. sapiens 50–40 ka, or it might suggest that local populations and African immigrants interbred. In the latter instance it might even reveal a dispersal from Africa to the Far East long before 50 ka. If in­ terbreeding (gene low) were demonstrated, the species distinction be­ tween invaders and locals could not be maintained. It is the growing evidence against interbreeding in Europe and the abrupt replacement of the Neanderthals there that most strongly support the separate species labels for the Neanderthals and modern humans. he skeletal remains of early modern Europeans sometimes exhibit features that could imply interbreeding with Neanderthals, and the strong­ est recent claims concern an early modern individual from Portugal and three from Romania. In each case, however, the supposed Neanderthal traits are swamped by others that are indisputably modern, and the indi­ cations for hybridization are problematic. he occurrence of a diagnos­ tic Neanderthal feature—the suprainiac fossa—on one of the Romanian fossils has been contested, but the more general problem is the muted expression of the supposed Neanderthal characters and their unknown heritability. It is especially unclear if they could be expected in individu­ als that lived long ater the putative hybridization event. he fossils for which Neanderthal characters have been proposed mainly postdate this event by at least 100 generations. he heritability of genes is unproblematic, and ancient DNA has now joined the genetics of living humans to argue that Neanderthals and early modern Europeans rarely if ever interbred. Since Neanderthal mi­ tochondrial (mt) DNA was irst reported in 1997, it has been extracted from iteen additional Neanderthal bones, representing at least thir­ teen diferent individuals, scattered throughout the Neanderthal range, from Spain on the west to south­central Siberia on the east. he fossil mtDNA underscores the divergent evolution of the Neanderthals, and like modern human mtDNA, it is roughly ive times less variable than the mtDNA of the African great apes. he providing Neanderthals died mainly, if not entirely, in the middle of the Last Glaciation (early OIS 3), between roughly 59 and 40–35 ka, and their limited mtDNA diversity suggests a recent climatically forced bottleneck of the kind whose re­ currence earlier on could help explain the mosaic accumulation of Ne­

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anderthal craniofacial features. More important from the perspective of this book, the absence of Neanderthal mtDNA from the living human gene pool implies that Neanderthal females either did not participate in interbreeding or participated so rarely that their mtDNA was lost by chance long ago. he recent recovery of the Y chromosome from two Neanderthal bones suggests a parallel situation with regard to males: Neanderthal males either did not mate with modern human females or the mating events were so infrequent that Neanderthal Y chromosomes failed to survive to the present. he DNA of the earliest modern Europeans might yet reveal occasional interbreeding, but testing so far, involving well-preserved bones from seven individuals who lived in western and central Europe mostly between 35 and 22 ka, has produced no Neanderthal mtDNA. Recent advances in DNA extraction and sequencing promise to pro­ vide the entire Neanderthal genome, and if the same procedures can be applied to fossils of “late Homo erectus,” they could provide the most compelling check on Out of Africa. Admittedly, however, even if fresh fossil DNA conirms what available fossil DNA implies, it would not demonstrate the species distinctions postulated here. In fact, given the relatively short interval over which the proposed species diverged, they are likely to have remained genetically compatible, and according to the model that this book advocates, behavioral separation would account for the absence or rarity of interbreeding when modern human invaders came into contact with nonmodern Eurasians. An obvious objection to Out of Africa is the failure of modern or near­modern humans to expand from Africa immediately ater they ap­ peared, by 100 ka or before. Instead they seem to have been conined to Africa until roughly 50 ka, and it is even possible that they were replaced by Neanderthals on the southwest Asian margin of Africa (in what is now Israel) roughly 80 ka. Archaeology provides a partial answer to the apparent dilemma. he people who inhabited Africa between 100 and 60–50 ka may have been physically modern or near­modern, but they were behaviorally very similar to the Neanderthals and other nonmod­ ern humans. he relatively full African and European archaeological records show a distinct rupture 50–40 ka, when the Middle Stone Age (MSA) in Africa and the broadly similar Middle Paleolithic in Europe gave way to the Later Stone Age (LSA) and Upper Paleolithic, respec­ tively. It is only LSA and Upper Paleolithic sites, postdating ater 50–40 ka, that commonly provide material residues that are indistinguishable from those of many later prehistoric and historic hunter­gatherers. Some novel features that mark the archaeological record beginning 50–40 ka can be found in the bulleted list below (chaps. 6 and 7 provide the details). In the view that this book espouses, these features are not

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isolated traits that deine “modern” behavior, but related outcomes of the innovative burst behind the Out of Africa expansion.

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• Substantial growth in the diversity and standardization of artifact types. • Rapid increase in the rate of artifactual change through time and in the degree of artifact diversity through space. • First routine shaping of bone, ivory, shell, and related materials into formal artifacts (“points,” “awls,” “needles,” “pins,” etc.). • Earliest appearance of incontrovertible art and personal ornamentation. • Oldest undeniable evidence for spatial organization of camp loors, including elaborate hearths and the oldest indisputable structural “ruins.” • Oldest evidence for the transport of large quantities of highly desir­ able stone raw material over scores or even hundreds of kilometers. • Earliest secure evidence for ceremony or ritual, expressed both in art and in relatively elaborate graves. • First evidence for human ability to live in the coldest, most conti­ nental parts of Eurasia (northeastern Europe and northern Asia). • First evidence for human population densities approaching those of historic hunter­gatherers in similar environments. • First evidence for ishing and for other signiicant advances in hu­ man ability to extract energy from nature. he most signiicant novelty is oten taken to be the burgeoning of unequivocal art and personal ornamentation because this suggests a ca­ pacity for abstract or “symbolic” thought. However, it is impossible to demonstrate that this capacity did not exist tens of thousands of years ear­ lier, when it was expressed only occasionally, as for example, at Blombos Cave, discussed in chapter 6 and again below. he diference at 50–40 ka is that unequivocal art, ornamentation, and other advanced behavioral markers were closely tied to a dramatic increase in human numbers, nota­ bly including the Out­of­Africa expansion. In short, they appear to have been part of a package that signiicantly enhanced human itness—the ability to survive and reproduce—and it is in this sense that they signal true evolutionary as opposed to mere historical change. Ater the ini­ tial lowering of advanced behavioral markers, local historical and envi­ ronmental conditions and the vagaries of preservation inluenced their presence or expression, and many sites lack them. here are even places, perhaps above all Tasmania, where they were signiicantly more conspic­ uous prehistorically than they were ethnohistorically. he fundamental point, however, is that their variable occurrence ater 50–40 ka contrasts sharply with their near uniform absence before, and it is this diference that signals something special.

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he beginnings of the LSA and the Upper Paleolithic lie beyond the practical lower bound of the radiocarbon dating method, and methods that can provide older ages are oten more reliable in theory than in practice. his is particularly true of optically stimulated luminescence, thermoluminescence, and electron spin resonance, in large part because they require an accurate estimate of the radiation dose to which buried items were subjected, and either they must assume a constant rate when sediments oten imply otherwise or they must make unveriiable site-speciic assumptions about how the rate may have varied through time. he calculated ages are most compelling when they closely match results from radiocarbon or other methods that do not require site-speciic assumptions, and in the instances that matter most to this book, such matches are not possible. here is the additional complication that the earliest LSA is still poorly known and thus weakly dated. Still, with these caveats in mind, the available dates suggest that the archaeological markers of advanced behavior appeared irst in Africa, probably between 50 and 45 ka, that they spread to western Asia and eastern Europe between 45 and 40 ka, and that they reached western Europe only between 40 and 36 ka. he geographic sequence is plainly what Out of Africa would predict. he advanced behavioral traits that blossom ater 50 ka imply the fully modern capacity for innovation that underlies culture in the nar­ row anthropological sense. he next section briely explores the thorny problem of explaining why modern innovative ability developed when it did, but the most fundamental point is that it provided the competitive advantage that allowed fully modern humans to replace their nonmod­ ern contemporaries. he time when advanced behavioral markers ap­ peared in China and adjacent southeastern Asia remains unsettled, and most artifact assemblages that are thought to date between 50 and 10 ka difer little from older, even much older, assemblages. In fact, however, the relevant archaeological record is slim and inadequately dated, and if neighboring regions are considered—especially Siberia and Sahul (the glacial supercontinent that included Australia, New Guinea, and Tasma­ nia)—archaeological change is conspicuous at 45–40 ka. Much fresh ar­ chaeological research will be necessary to determine whether the pattern in China and southeastern Asia difered from the pattern in Africa and western Eurasia.

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Some Problems with Out of Africa Out of Africa is the most reasonable and parsimonious explanation of the available fossil and archaeological data, but previous chapters pro­ vided some contrary observations, and these cannot simply be ignored. Some specialists also believe that proponents of Out of Africa have in­ advertently imposed their intellectual preconceptions on contrary or

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ambiguous data, and this leads them to reject Out of Africa in advance. Carried to its logical extreme, however, this perspective precludes any decision on Out of Africa, barring the unlikely development of data collection procedures that do not require advance assumptions or expectations. New intellectual frameworks or paradigms may yet prove helpful, but the aim of this section is to reiterate some problems with Out of Africa that are more evidentiary than epistemological.

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What explains the relatively abrupt florescence of advanced behavior (the modern capacity for Culture) 50 ka? he most popular answer is

probably that it followed on some momentous social or demographic change. Social change, for example, could have involved the initial development of the nuclear family as the fundamental productive unit and together with this, the division of labor by sex and age that characterized all historic hunter-gatherer societies. Nuclear family organization and a newly invented division of labor could in turn have led to the modern notions of kinship and descent that promote economic and political cooperation among individuals and groups. Population growth could have followed, and larger, denser populations could in turn explain the accelerated innovation that marks the archaeological record ater 50 ka. his kind of explanation is logically coherent, but it fails in at least one crucial respect—it does not explain why social relations changed when they did or why they changed at all. Population growth could underlie the timing, if it is assumed that growth came irst and that behavior changed about 50 ka when popula­ tion density crossed a critical threshold that forced social reorganization. However, even if we ignore the need to explain what drove population growth, an explanation that invokes it must confront the problem that the advanced behavior presumably arose in Africa, and archaeology suggests that African populations were shrinking not growing when it appeared. In southern Africa, proxies for human population density described in chapter 6 imply that density remained low and more or less constant between 120 and 60 ka. Shortly ater 60 ka, people became archaeologically all but invisible, probably because hyperaridity in the middle of the Last Glaciation sharply reduced their numbers. A similar pattern appears to have characterized northern Africa. Eastern Africa was more mesic 60–50 ka, and people may have remained more numer­ ous there, but even when this is considered, the archaeological indica­ tors for African population size about 50 ka are consistent with genetic analyses, summarized in chapter 7, which suggest that the African popu­ lation from which all living people derive included no more than 10,000 breeding adults. Given what we know or don’t know about social and demographic change, it then becomes at least as plausible to tie the basic behavioral

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shit 50 ka to a fortuitous mutation that promoted the fully modern brain. his proposal follows on two commonly accepted observations: that natural selection for more efective brains largely drove the earlier phases of human evolution and that the relation between morphologi­ cal and behavioral change shited abruptly about 50 ka. Before this time, morphology and behavior appear to have evolved relatively slowly and more or less in tandem, but ater this time morphology remained rela­ tively stable while behavioral (cultural) change accelerated rapidly. What could explain this better than a neural change that promoted the ex­ traordinary modern human ability to innovate? his is not to say that the Neanderthals and their nonmodern contemporaries had apelike brains or that they were as biologically and behaviorally primitive as yet earlier humans. It is only to suggest that an acknowledged genetic link between morphology and behavior in yet earlier people persisted until the emer­ gence of fully modern ones and that the postulated genetic change 50 ka fostered the uniquely modern ability to adapt to a wide range of natural and social circumstances with little or no physiological change. Most archaeologists dismiss the neural hypothesis out of hand, per­ haps because they have been trained to distrust biological explanations for cultural change. In addition, some specialists seem to feel that a bio­ logical explanation relects unwitting bias against the Neanderthals and other nonmodern humans. Ideological issues aside, however, the main problem with a neural explanation has long been that it cannot be tested with fossils. Earlier on in human evolution, a link between behavioral and neural change can be inferred from conspicuous increases in aver­ age brain size, but humans virtually everywhere had achieved modern or near­modern brain size by 200 ka. Any neural change that occurred around 50 ka must thus have been in organization, and fossil skulls pro­ vide only speculative evidence for brain structure. Neanderthal skulls, for example, difer dramatically in shape from modern ones but were just as large if not larger, and on present evidence it is not clear that the diference in form implies a signiicant diference in function. A link between form and function becomes especially unlikely if, as suggested above, random genetic drit was primarily responsible for the diference in form. Fossils may never allow an independent test of the neural hypoth­ esis, but there is now the possibility that genetics could. Clinically ori­ ented investigations have identiied numerous genes that probably bear on communication and cognition, and if their function can be more precisely determined, it may be possible to determine if one or more underwent strong selection at roughly the time of the modern human expansion from Africa. In addition, once the Neanderthal nuclear ge­ nome becomes available, it will become possible to assess the extent to which Neanderthals and modern humans shared behaviorally relevant

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genes. If diferences are found, they would indirectly support the neural explanation, for they would suggest that behaviorally relevant genetic change occurred even ater the modern human lineage had emerged. An expectation to the contrary is implicit in hypotheses that link the modern human expansion to strictly social or demographic factors.

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Is it really true that advanced behavioral markers appear widely only about 50–40 ka? With regard to art and ornamentation, for example,

virtually all specialists agree that they become commonplace only ater 50 ka and that older examples are both rare and crude. he most widely cited older examples are abstract engravings and perforated shells dis­ cussed below, and these occur in only a small fraction of the sites where they might be expected. Representational (igurative) art objects and in­ tentionally shaped ornaments (beads or pendants) appear only at sites that postdate 50 ka. However, even if authorities agree that 50 ka marks a sharp change in course, they disagree sharply on what the change means. To some, the rarity and simplicity of supposed art before 50 ka implies that modern cognitive abilities were present but were weakly or infrequently expressed before 50 ka, while to others (including myself), it suggests that the fully modern capacity for culture may have appeared only about this time. Some of the extremely rare art objects that antedate 50 ka are prob­ ably younger intrusions that even the most careful excavation cannot de­ tect, while others are probably the result of human or natural actions that will inevitably, on rare occasions, mimic crude human attempts at art. In this regard, credible claims for art or other advanced behavioral markers before 50 ka must involve sizable numbers of conspicuously patterned objects from carefully documented contexts. As discussed at length in chapter 6, with this criterion in mind, two sites—Blombos Cave, South Africa, and the Katanda open­air (riverside) site cluster, Democratic Republic of the Congo—probably present the most serious obstacles to the neural explanation for Out of Africa favored here. At Blombos, the objects include a pigment (ocher) lump on which an abstract pattern has been incised, forty­one perforated tick shells interpreted as beads, and twenty­nine indisputable bone artifacts, all ixed by luminescence dating of sand grains and heated stone artifacts between 84 and 74 ka. At Ka­ tanda, the objects comprise eight whole or partial barbed bone points and four other well­made bone artifacts, dated to between 90 and 60–70 ka by luminescence on surrounding sands and by Electron Spin Resonance on associated mammal teeth. he incised pattern on the Blombos pigment lump comprises a se­ ries of six­to­seven crude X’s framed by additional incised lines (ig. 6.46), and many observers accept the sum as crude art. Some of the other 8,000 plus pigment lumps from Blombos exhibit incised lines that could

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represent incomplete or less successful stabs at similar abstract expression, and scored lines have been observed on lumps from other broadly contemporaneous Middle Stone Age (MSA) sites. Everywhere, however, MSA people ground or scraped pigment to obtain powder, and there is the possibility that they scored it to test its quality or to roughen its surface before grinding. Conceivably MSA people used the powder to make paint, perhaps for application to human skin, but the disposition of powder on ancient stone artifacts and modern experimentation indicate it was an important ingredient in the glue used to attach stone bits to wooden handles or shats. his is the only function that archaeological and experimental observations conirm, and it is reasonable to ask, given the large number of pigment lumps at Blombos, Klein Kliphuis, and other sites, whether scoring for practical purposes might not occasionally produce overlapping marks that appear deliberately patterned. he Blombos engraving would be more persuasively artistic if it were widely replicated or, of course, if it were less abstract and it suggested a human or animal igure. In addition, whatever its meaning, it also cannot bear specially on Out of Africa since the Neanderthals who occupied Pech de l’Azé Cave 1, France, also incised engraved patterns on pigment lumps. he perforated Blombos tick shells may be beads, but the case de­ pends mainly on the need to explain why people would bring home such tiny shells otherwise. he perforations are less conspicuously artiicial than they are on much later, LSA shell beads, and the basic shape of the tick shells remained unaltered. he same is true of thirteen putative tick shell beads dated by luminescence to roughly 82 ka at Pigeon Cave (Taforalt), Morocco. Unlike the proposed Blombos beads, some of the Pigeon Cave specimens were abraded and initially choked with beach sediment, and the circumstances of their excavation have been only weakly described. hey are less compelling than their Blombos coun­ terparts, and they might have been ignored altogether, if the Blombos inds had not already been publicized. he key point is that neither the Blombos nor Pigeon Cave beads are deliberately shaped, and so far, the oldest unequivocally shaped beads postdate 50 ka. he oldest artiicially shaped beads are in ostrich eggshell, some of which date to beyond 40 ka at early LSA sites in eastern Africa. Shaped beads are also well­known from European Upper Paleolithic sites that postdate 40 ka. he Blombos bone artifacts may be divided between two groups— twenty­six that were shaped mainly by use and three that were more conspicuously fashioned in advance and that closely resemble pol­ ished LSA bone projectile points. he pieces that were shaped by use recall the smoothed and pointed antelope horncores and bone splinters dated to more than 1 Ma at the Swartkrans, Sterkfontein, and Drimolen Caves, South Africa (chap. 4), and their cognitive implications need be no greater. he three that resemble polished LSA projectile points are

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another matter, for if their stratigraphic position has been correctly assessed, they could imply LSA-like hunting proiciency. One was initially associated with three radiocarbon readings on charcoal near 2 ka, but if these dates are discounted and the MSA associations of all three are accepted, the Blombos mammalian assemblage might be expected to exhibit the same elevated proportion of more dangerous ungulates that distinguish LSA assemblages from MSA assemblages elsewhere in the region. It doesn’t, and in this connection, the large average size of the associated shells and tortoise bones imply that the Blombos people were much less numerous than LSA people living under similar conditions. he bottom line is that whatever the cognitive or cultural implications of the Blombos artifacts, they do not appear to have conferred an LSA­ like itness (reproductive and survival) advantage. his may explain why they did not spread from Blombos and why they did not prompt an Out­ of­Africa expansion. he Katanda barbed points may be seen to parallel the Blombos pol­ ished points, but they are far more elaborate, and if they are genuinely older than 60 ka, they would more surely imply modern cognition long before Out of Africa occurred. his would raise the question again, how­ ever, of why they remained geographically isolated and why their use did not promote human population increase. he answer may be that they are actually much younger than proposed, and this might be checked by direct radiocarbon dating. hey are much fresher looking than the heav­ ily abraded mammalian teeth that have been partly used to date them, and the next oldest similar points in eastern Africa date from 25 ka or less. Most postdate 10 ka.

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Were Neanderthals fundamentally incapable of fully modern behavior?

As outlined here, Out of Africa postulates that the Neanderthals were replaced because they could not compete culturally with their modern uman successors. If cognitive limitations explain their inability, the limi­ tations might have created a behavioral barrier to interbreeding between Neanderthals and modern humans. he idea that the Neanderthals were not competitive is bolstered over most of Europe by the relatively abrupt nature of their replacement. At most sites, where Cro­Magnon/Upper Paleolithic occupations directly overlie Neanderthal/Middle Paleolithic layers, there is no indication for a substantial break in time or for any transition between the two, and it is reasonable to infer that in any given region the replacement took a relatively short time, a few centuries or perhaps even a few decades. However, there is the occasional discovery of artifact assemblages that include subequal numbers of Neanderthal/Middle Paleolithic and Cro­Magnon/Upper Paleolithic artifact types. As described in chapter 6, the most compelling examples come from western France and northern

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Spain, where they have been assigned to the Châtelperronian Industry. he Châtelperronian probably began around 45 ka, and it persisted until perhaps 37–36 ka, when it was truncated by the early Upper Paleolithic Aurignacian Industry. Human fossils from La Roche á Pierrot Rockshel­ ter, Saint­Césaire and the Grotte du Renne, Arcy­sur­Cure, indicate that Châtelperronian people were Neanderthals. In contrast, early Aurigna­ cians were anatomically modern people whose biological roots probably lay in Africa. If only stone tools were involved, the Châtelperronian might be re­ garded as the inal stage of the local Mousterian of Acheulean Tradition Type B (with numerous backed knives) that converged on the Upper Paleolithic, and the early Châtelperronian may have been just that. How­ ever, at Arcy­sur­Cure, late Châtelperronians not only produced a mix of typical Middle and Upper Paleolithic stone artifacts, they also manufac­ tured typical Upper Paleolithic bone tools and personal ornaments, and they modiied their living space in a characteristically Upper Paleolithic fashion. his might mean that they borrowed Upper Paleolithic cultural traits from Aurignacian neighbors, but even if this is accepted, it begs one fundamental question: if Upper Paleolithic culture was clearly su­ perior and Châtelperronian Neanderthals could imitate it (that is, they were not biologically precluded from behaving in an Upper Paleolithic way), why didn’t the Neanderthals acculturate more widely, with the re­ sult that they or their genes would have persisted much more conspicu­ ously into Upper Paleolithic times (ater 40–35 ka)? he Arcy Châtelperronian counters other evidence for a behavioral diference between Neanderthals and Cro­Magnons, and it is reason­ able to ask whether excavation could have created it. he excavations were conducted between 1948 and 1963, and the methods were excellent for their time. However, hindsight suggests that they would not meet present standards, particularly in a deposit that was stratigraphically complex, and it is possible that the excavators inadvertently failed to separate a typical Châtelperronian occupation without ornaments and well­made bone tools from an overlying early Aurignacian occupation that had both. It has been argued that the Arcy Châtelperronians em­ ployed distinctive methods to manufacture ornaments and other special pieces, but a comprehensive comparative analysis shows that both the manufacturing methods and the inal products it comfortably within the early Aurignacian range. he Arcy example underscores the extent to which archaeological interpretation depends on excavation quality. Outsiders must usually take quality for granted, but even the most careful, experienced exca­ vators may fail to detect mixture between occupations in a complex cave like the Arcy Grotte du Renne. his reinforces what should be an archaeological maxim: the irst discovery of a unique or unexpected

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assemblage should be regarded as a possible accident, and even the second could be a coincidence. In general, only repeated, independent discoveries establish a pattern that merits serious behavioral interpretation. Nearly ity years ater the Arcy Châtelperronian was irst reported, it remains nearly unique. Only the Châtelperronian layers at Quinçay Cave, southwestern France, provide meager corroboration, in the form of four pierced animal teeth. Well-excavated Châtelperronian layers elsewhere, for example, at Grotte XVI, also southwestern France, lack Upper Paleolithic– like ornaments and bone tools, and among the eighteen or so commonly accepted Châtelperronian sites, Grotte XVI illustrates the rule. At least until the Arcy Châtelperronian is independently replicated then, it need not imply fundamental similarity in behavioral capability between Neanderthals and Cro-Magnons.

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Conclusion Since the 1910s, fossil and archaeological discoveries have suggested that fully modern (Cro­Magnon) immigrants replaced the Neanderthals in Europe. Fossil and archaeological support for an abrupt replacement grew stronger in succeeding decades, but it became particularly compel­ ling from the late 1980s onward, when new dates and fresh fossils dem­ onstrated that near­modern or modern humans had appeared in Africa by 100 ka; new fossils (especially those from the Sima de los Huesos, Ata­ puerca, Spain) documented the accretional evolution of the Neanderthal lineage in Europe; genes showed that all living humans share a common African ancestor that existed ater the Neanderthal and modern human lineages had diferentiated; and DNA extracted from the bones of Nean­ derthals simultaneously underscored their divergent evolutionary his­ tory of the Neanderthals and showed that they rarely if ever interbred with modern humans. Some of the new (and old) evidence is ambiguous, circumstantial, or even contradictory, but if the study of human origins were a jury trial, the verdict would surely be that modern humans, origi­ nating in Africa, swamped or extinguished the Neanderthals. he jury would probably also accept that a behavioral transformation accounted for modern human success, but they might deadlock on whether so­ ciodemographic or genetic change underlay the transformation. In fact, of course, human origins research difers from a jury trial in that no verdict need ever be inal, and new evidence and new jury members are always welcome. It seems increasingly unlikely now that the verdict on the extinction of the Neanderthals will be reversed, but fu­ ture juries, faced with new evidence, may decide diferently on a range of other controversial issues, including, for example, the relation between climatic change and speciation early on in human (hominin) evolution, the number and relationships of early human species, and the number

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and timing of ancient human dispersals from Africa. hese matters are inherently more diicult to resolve than modern human origins be­ cause they center on much older time periods for which the record is far sparser. hey may never be resolved as completely as modern hu­ man origins, but uncertainty stimulates data collection, and it is fresh data—expanding genetic studies and new fossils and artifacts from well­ documented contexts—that have eliminated all reasonable doubt in the century­old controversy over the fate of the Neanderthals.

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