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the oxford handbook of LANGUAGE EVOLUTION oxford handbooks in linguistics The Oxford Handbook of Applied Linguistics Second edition Edited by Robert B Kaplan The Oxford Handbook of Case Edited by Andrej Malchukov and Andrew Spencer The Oxford Handbook of Cognitive Linguistics Edited by Dirk Geeraerts and Hubert Cuyckens The Oxford Handbook of Comparative Syntax Edited by Gugliemo Cinque and Richard S Kayne The Oxford Handbook of Compounding Edited by Rochelle Lieber and Pavol Sˇtekauer The Oxford Handbook of Computational Linguistics Edited by Ruslan Mitkov The Oxford Handbook of Compositionality Edited by Markus Werning, Edouard Machery, and Wolfram Hinzen The Oxford Handbook of Field Linguistics Edited by Nicholas Thieberger The Oxford Handbook of Grammaticalization Edited by Heiko Narrog and Bernd Heine The Oxford Handbook of Japanese Linguistics Edited by Shigeru Miyagawa and Mamoru Saito The Oxford Handbook of Laboratory Phonology Edited by Abigail C Cohn, Ce´cile Fougeron, Marie Hoffman The Oxford Handbook of Language Evolution Edited by Magggie Tallerman and Kathleen R Gibson The Oxford Handbook of Language and Law Edited byLawrence Solan and Peter Tiersma The Oxford Handbook of Linguistic Analysis Edited by Bernd Heine and Heiko Narrog The Oxford Handbook of Linguistic Interfaces Edited by Gillian Ramchand and Charles Reiss The Oxford Handbook of Linguistic Minimalism Edited by Cedric Boeckx The Oxford Handbook of Linguistic Typology Edited by Jae Jung Song The Oxford Handbook of Translation Studies Edited by Kirsten Malmkjaer and Kevin Windle the oxford handbook of LANGUAGE EVOLUTION Edited by M A G G I E TA L L E R M A N and KATHLEEN R GIBSON Great Clarendon Street, Oxford ox2 6dp Oxford University Press is a department of the University of Oxford It furthers the University’s objective of excellence in research, scholarship, and education by publishing worldwide in Oxford New York Auckland Cape Town Dar es Salaam Hong Kong Karachi Kuala Lumpur Madrid Melbourne Mexico City Nairobi New Delhi Shanghai Taipei Toronto With offices in Argentina Austria Brazil Chile Czech Republic France Greece Guatemala Hungary Italy Japan Poland Portugal Singapore South Korea Switzerland Thailand Turkey Ukraine Vietnam Oxford is a registered trade mark of Oxford University Press in the UK and in certain other countries Published in the United States by Oxford University Press Inc., New York # Editorial matter and organization Maggie Tallerman and Kathleen R Gibson 2012 # The chapters their several authors 2012 The moral rights of the authors have been asserted Database right Oxford University Press (maker) First published 2012 All rights reserved No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, without the prior permission in writing of Oxford University Press, or as expressly permitted by law, or under terms agreed with the appropriate reprographics rights organization Enquiries concerning reproduction outside the scope of the above should be sent to the Rights Department, Oxford University Press, at the address above You must not circulate this book in any other binding or cover and you must impose the same condition on any acquirer British Library Cataloguing in Publication Data Data available Library of Congress Cataloging in Publication Data Data available Typeset by SPI Publisher Services, Pondicherry, India Printed in Great Britain on acid-free paper by MPG Books Group, Bodmin and King’s Lynn ISBN 978–0–19–954111–9 10 Contents Preface and Acknowledgements List of Figures List of Tables List of Contributors Introduction: The evolution of language Maggie Tallerman and Kathleen R Gibson xi xiii xiv xv PA RT I I N S I G H T S F RO M C O M PA R AT I V E A N I M A L B E H AV I O U R Introduction to Part I: Insights from comparative animal behaviour Kathleen R Gibson and Maggie Tallerman 39 Language or protolanguage? A review of the ape language literature Kathleen R Gibson 46 Primate social cognition as a precursor to language Robert M Seyfarth and Dorothy L Cheney 59 Cooperative breeding and the evolution of vocal exibility Klaus Zuberbuăhler 71 Gesture as the most flexible modality of primate communication Frans B M de Waal and Amy S Pollick 82 Have we underestimated great ape vocal capacities? Katie Slocombe 90 Bird song and language Peter Slater 96 Vocal communication and cognition in cetaceans Vincent M Janik 102 vi contents 10 Evolution of communication and language: insights from parrots and songbirds Irene M Pepperberg 109 11 Are other animals as smart as great apes? Do others provide better models for the evolution of speech or language? Kathleen R Gibson 120 PA RT I I T H E B I O LO G Y O F L A N G UAG E EVO LU T I O N : A NATO M Y, G E N E T I C S , A N D N E U RO LO G Y 12 Introduction to Part II: The biology of language evolution: anatomy, genetics, and neurology Kathleen R Gibson and Maggie Tallerman 13 Innateness and human language: a biological perspective W Tecumseh Fitch 133 143 14 Evolutionary biological foundations of the origin of language: the co-evolution of language and brain Szabolcs Szamado and Eoărs Szathmary 157 15 Genetic inuences on language evolution: an evaluation of the evidence Karl C Diller and Rebecca L Cann 168 16 Not the neocortex alone: other brain structures also contribute to speech and language Kathleen R Gibson 176 17 The mimetic origins of language Merlin Donald 180 18 Evolution of behavioural and brain asymmetries in primates William D Hopkins and Jacques Vauclair 184 19 Towards an evolutionary biology of language through comparative neuroanatomy Wendy K Wilkins 198 contents 20 Mirror systems: evolving imitation and the bridge from praxis to language Michael A Arbib 21 Cognitive prerequisites for the evolution of indirect speech Frederick L Coolidge and Thomas Wynn 22 The anatomical and physiological basis of human speech production: adaptations and exaptations Ann MacLarnon vii 207 216 224 PA RT I I I T H E P R E H I S TO RY O F L A N G UAG E : W H E N A N D W H Y D I D L A N G UAG E EVO LV E ? 23 Introduction to Part III: The prehistory of language: When and why did language evolve? Kathleen R Gibson and Maggie Tallerman 239 24 Molecular perspectives on human evolution Rebecca L Cann 250 25 The fossil record: evidence for speech in early hominins Bernard A Wood and Amy L Bauernfeind 258 26 The genus Homo and the origins of ‘humanness’ Alan Mann 273 27 The Palaeolithic record Thomas Wynn 282 28 Musicality and language Steven Mithen 296 29 Linguistic implications of the earliest personal ornaments Francesco d’Errico and Marian Vanhaeren 299 30 Inferring modern language from ancient objects Rudolf Botha 303 viii contents 31 Natural selection-itis David Lightfoot 32 The role of hominin mothers and infants in prelinguistic evolution Dean Falk 33 Infant-directed speech and language evolution Bart de Boer 34 Displays of vocal and verbal complexity: a fitness account of language, situated in development John L Locke 313 318 322 328 35 Tool-dependent foraging strategies and the origin of language Kathleen R Gibson 340 36 Gossip and the social origins of language Robin I M Dunbar 343 37 Social conditions for the evolutionary emergence of language Chris Knight and Camilla Power 346 PA RT I V L AU N C H I N G L A N G UAG E : T H E D EV E LO P M E N T O F A LINGUISTIC SPECIES 38 Introduction to Part IV: Launching language: the development of a linguistic species Maggie Tallerman and Kathleen R Gibson 353 39 The role of evolution in shaping the human language faculty Stephen R Anderson 361 40 The origins of meaning James R Hurford 370 41 The origins of language in manual gestures Michael C Corballis 382 contents 42 From sensorimotor categories and pantomime to grounded symbols and propositions Stevan Harnad ix 387 43 The symbol concept Terrence W Deacon 393 44 Words came first: adaptations for word-learning Robbins Burling 406 45 The emergence of phonetic form Michael Studdert-Kennedy 417 46 The evolution of phonology Peter F MacNeilage 423 47 The evolution of morphology Andrew Carstairs-McCarthy 435 48 What is syntax? Maggie Tallerman 442 49 The origins of syntactic language Derek Bickerton 456 50 The evolutionary relevance of more and less complex forms of language Andrew Carstairs-McCarthy 469 51 Protolanguage Maggie Tallerman 479 52 The emergence of language, from a biolinguistic point of view Cedric Boeckx 492 PA RT V L A N G UAG E C H A N G E , C R E AT I O N , AND TRANSMISSION IN MODERN HUMANS 53 Introduction to Part V: Language change, creation, and transmission in modern humans Maggie Tallerman and Kathleen R Gibson 505 subject index Kenyanthropus 264, 265 keyboard, use of in ape language research 51–2, 86, 355, 383, 453, 481 kin selection 246–7, 324, 326–7 KMN-WT15000 274–5, see also Homo erectus; Homo ergaster; Nariokotome knapping 283–4, 289, 292 KNM-ER3733 274–5, see also Homo erectus; Homo ergaster Koko, gorilla 48, 50, 54 Kyaaro, parrot 110, 113–4 La Chapelle-aux-Saints, France 243, see also Neanderthal Lana, chimpanzee 51, 54 language: acquisition 9, 10, 17, 19, 20–1, 69, 110, 113, 144, 151, 154, 182, 213, 219, 222, 297, 324–5, 357, 366, 380, 403, 426, 464, 481, 496, 507, 535–6, 562, 590, 592, 621–5, 627, 630–1, 635–6, 638 adapting to become learnable 13, 14, 16, 325, 354, 361, 367, 509, 587, 597–604, 617, 636 and brain, coevolution of 9, 165–6 and fitness 12, 21, 165, 328–39, 354, 363, 459, 603–4, 615 and social grooming 217, 249, 344–5, 430 and timescales 12–14, 590–1 artificial 477–8, 602, 622–4, 634 as an autapomorphy 2–6 as an evolving system 14, 627, 631 as a saltation 21, 25, 30 amodal nature of 183, 208, 432 biological basis of 199, 201, 627 capacities, see language capacities change, see language change cognitive/developmental relationships 248 compared with animal communication systems 1–6, 10–12, 15, 25, 31, 362, 442, 445, 451, 452 complex 310, 327, 519 complexity, see language complexity comprehension, see language comprehension contact 13, 573–80 creation 35, 505, 507, 508, 511, 537, 542–4 design features of 2–4, 6, 24, 542 developmentally adaptive 328–39 diversity 11, 22, 173, 300, 493, 494, 500, 508, 560, 561–3, 570, 572, 618, 637 emotional regulation by 223 faculty, see language faculty function 11–12, 15, 23–5, 500, 633 genesis 9, 516 handedness, relationship to 138–9, 184–5 749 in children 14, 16, 17 information sharing 18, 248, 335, 341, 345, 373, 379, 382, 441, 554, 631, 633 innate mechanisms of, see language, acquisition; innate language faculty; Language Acquisition Device Monocategoric Grammar 472, 475–8 multicomponent perspective on 155 modern 9, 34, 220, 299, 302, 303–12, 337, 429, 463, 472, 479, 505, 514, 518, 520, 521, 526, 542, 568–9 non-configurational 448–9 of thought 23–4, 396, 498 ‘organ’ 199, 201, 459, 497, 500–1, 627–8, 630 phonemic 124, 310 processing 16, 19, 20, 62, 75, 111, 141, 165, 167, 199, 219, 404, 458, 461, 462, 507, 510, 530, 533, 534–7 prerequisites for 3, 12, 32, 34, 88, 233, 323, 371, 383, 459, 461, 486, 536 second 309, 323, 473–4, 481, 539, 543, 560, 562, 574 shift 35, 505, 508, 538, 575, 577–9 sign, see sign(ed) languages syntactic 35, 245, 301, 302, 304–7, 310, 381, 456–68 tonal 136–7, 172, 174, 363–4 transmission 3, 10, 12, 13–14, 16, 33, 35, 157, 183, 224, 225, 320, 325, 347–8, 355, 390, 394, 424, 427, 430, 431, 432, 434, 486, 505, 507, 508–10, 535, 540, 559, 560, 561–2, 583, 585, 590–604, 609–10, 617–18, 627, 631–3, 636, 638 typology 558, 636, 638 universals 15, 17, 19, 20, 22, 202–3, 353, 367–8, 404, 424–5, 442–3, 445, 460, 461, 473, 476, 483, 493, 591, 593–4, 601–3, 627, 628, 629, 636 usage-based approaches to 18–19, 21, 528, 638 vocabulary size in 5–6, 10, 99, 163–4, 213, 218, 222, 247, 319, 337, 357, 374, 380, 406, 413–4, 419–20, 637 vs animal communication systems 4–5, 10–12, 15 Language Acquisition Device 9, 17, 426, 593 language capacities: early modern humans 559 great apes 67–76, see also ape language studies Homo erectus 242 Homo habilis 268 Neanderthals 58, 243–5, 300, 302, 480 language change 13, 35, 134, 174, 465–6, 488, 505, 506–8, 510, 514, 518, 526, 528, 534, 535, 573–80, 616, 619, 629, 631, 638 and language evolution, similarities 314, 518, 542, 631 750 subject index language change (cont.) and technological innovation 163 rate of 134–5, 162–3 Plio-Pleistocene 134, 167 language complexity 97, 301, 327, 412, 519, see also coevolution of language and brain as driver of biological evolution 325 language comprehension 42, 189, 213, 218–9, 222, 268, 338, 356, 404, 410–12, 485, 609 by apes 41, 42, 47–8, 51–2, 122, 356, 411, 443, 453–4 by dolphin 44, 106, 122 by sea lion 44 language faculty 2, 6, 7, 8, 9, 11, 12, 13, 14, 16, 17–18, 21–3, 25, 30–2, 33, 34, 35, 155, 240, 242, 244–6, 313–6, 325, 354, 357, 358, 359, 361–9, 437, 439, 451, 464, 466, 470, 475, 477, 480, 484, 492–501, 505, 506, 508–9, 510–11, 541, 544, 591, 592, 593, 629, see also innate language faculty dating of 8, 30, 31, 34, 240, 242, 244–5 language game model 602, 608–9, 610, 617–19 language-specific adaptations 614 language-specific biases, patterns, principles and mechanisms 17, 18, 19, 20, 22, 232, 395, 404, 443, 601, 630, 636, 638 languages in the vocal modality discussed/ illustrated, see also sign(ed) languages; American Sign Language of the Deaf: Afrikaans 543 Buriat 532, 533 Chinese 385, 445, 457, 483, 520, 522–6 English 3, 20, 162, 366, 378, 402, 410, 423, 435, 436, 439, 440, 444, 445, 447, 449, 450, 451, 454, 455, 465, 481, 513, 516–7, 529, 531–2, 534, 553, 568, 632 Esperanto 477–8 Evenki 579 French 437, 438, 465, 489, 513, 514–5, 632 German 439, 444, 513, 556 Latin 437, 438, 513, 632 Piraha˜ 309, 358, 450–1, 474–5, 478 Portuguese 465 Riau Indonesian 309, 358, 445, 471–2, 474, 478 Sakha 579 Saramaccan 465 Seselwa 465 Tok Pisin 540 Warlpiri 449 langur 73 lark 99 laryngeal descent, see hyolarygneal descent larynx 94, 128, 129, 212, 226, 227, 385, 417–18, 427, 430 articulating cartilages of 225, 233 descent of in animals 229, 365 descent of in development 229, 247 descent of in human phylogeny 26, 27, 141–2, 229–30, 243, 365 position of 141, 228, 243, 270–1 temporary descent of 229 valvular/non-valvular 233–235 last common ancestor: African apes and humans 330 apes and humans 7, 13, 32, 42, 46, 57, 62, 88, 205, 210, 246, 250, 254 early hominins and humans 142, 171, 233, 240, 256, 277, 298 great apes and humans, behaviour/cognition of 7, 13, 32, 42, 57, 62, 210, 246 monkeys, apes, and humans 62, 63, 205, 210, 231, 254 panins and humans 7, 32, 39–40, 60, 61, 67, 83, 88, 170, 185, 210, 250, 254, 261–2, 363, 484, 499 lateral fissure, see Sylvian fissure lateralization, see asymmetry; handedness; orofacial asymmetry learning 19, 29, 35, 106, 110, 134, 135, 138, 144, 147, 178, 179, 181, 209, 213, 242, 243, 249, 332, 366, 388–91, 393, 507, 536, 582, 584, 586, 588, 590–1, 628, 633, 638 and innateness 10, 17, 19, 20, 59, 75, 143–56, 368, 414, 444, 458, 470, 506, 510, 559, 563, 592, 600, 601, 603–4, 621, 625, 633 and instinct 15, 133–4, 144–6, 147–8, 155, 563 and language 9, 16, 24, 45, 47, 113, 130, 157, 165, 167, 177, 179, 210–11, 213, 247, 248, 319–20, 325, 342, 354, 366–7, 374, 380, 383, 396, 406, 407, 415, 433, 440, 481, 509, 510, 545, 549, 554, 560, 562, 563, 581, 587, 590, 592–4, 595, 596–600, 602–3, 603–4, 605, 608, 609, 610, 617–18, 619, 627, 629 Bayesian model of 600–2 biases 17, 18, 510, 584, 585, 586, 594, 600–2, 604, 624, 634, 636 in lexigram-using apes 50–3, 54 mechanisms 14, 17, 19, 20, 144, 147, 154, 368, 388, 506, 510, 591, 593, 618, 621–2, 625, 634–5 of bird song 10–11, 33, 99–101, 116–17, 121, 125–6, 147, 179, 334, 362, 365 of calls in birds 10, 101, 179 of lexicon 69, 213, 355, 356, 406–16, 443, 486 sequential 20, 634–5 statistical 14, 19, 506, 509–10, 621–5, 634 subject index vocal 10, 12, 33, 44, 48, 54, 80, 93–4, 99–101, 103, 104, 106, 108, 110, 112, 116–8, 121, 126, 128, 129, 130, 136, 179, 337, 355, 365, 383, 486, 490, 612 learning bottleneck 583, 584, 585, 586, 595, 597–600, 601, 603, 609 legisign 397–8, 400, 402 lemur 231, 254 lesser apes 32, 76, 190, 227, 228, 233, 235, 254, 255, 297, 452, see also gibbon; siamang Levallois 290, see also prepared core techniques lexical envelope 359, 497–9 lexicalization 203, 497, 498, 499 lexicon 4, 6, 23, 213–14, 336, 355, 356, 401, 406–7, 413–14, 415, 416, 418, 420, 421, 443–5, 451, 479, 487, 509, 520–1, 523, 529, 538, 541, 546, 575, 606, 607–9, 609–10, 619, 630 lexigram(s) 41, 50–5, 56, 121, 122–3, 374, 378, 453, 454, 481 life history 314, 331 changes in hominin 27, 338 limbic control 383 linearization, see word order linguistic change, see language change linguistic diversity 11, 22, 173, 300, 493, 494, 500, 508, 560, 561–2, 563, 570, 572, 618, 637 linguistic expressions 370, 378, 522, 524 spatial nature of 202–3, 513 linguistic fossils 410, 438, 481, 482, 539, 540–1, 564, 634 linguistic universals, see language universals linguistics: evolutionary 16, 34, 505, 508, 540, 543, 583, 588, 591 functionalist 20–1, 367, 493 generative 13, 18, 21, 248, 459, 474, 493–7, 510–11, 528, 529, 593, 626–7, 630 lips 93, 112, 117, 128, 129, 178, 225, 226, 231, 234, 411, 417–8, 428, 429, 430, 433 lizards 159, 160, 161 loris 231 LRC (Language Research Center, Georgia State University) 50–1, 52, 53 Lucy, chimpanzee 49–50, 54, 56 Lucy, fossil hominin 241, 264, see also Australopithecus afarensis lumpers 241–2 lunate sulcus 139, 200, 204, 270 lyrebird 97 macaques 42, 62, 170, 191, 208, 212, 220, 254, 376, 418 crab eating 191 751 Japanese (Macaca fuscata) 61, 194, 320 rhesus (Macaca mulatta) 61, 139, 187, 191, 193, 194, 195, 419 Machiavellian intelligence 348 maintainable tools 292 manual gestures, see gestures marmoset/common marmoset/pygmy marmoset 42, 75, 79–80, 128, 138, 187, 191, 195 Matata, bonobo 52, 481, 482 material culture 347 relationship to symbolic and syntactic language 301–2, 311, 544 meaning (linguistic) 2, 4, 5–6, 24–5, 41, 67, 77, 97, 106, 125, 207, 211, 214, 219, 301, 306–9, 354, 355, 362, 370–81, 382, 387–92, 394–6, 402, 406–16, 427, 440–1, 443, 445–9, 452, 454, 462, 466, 472, 477, 480–1, 483, 487, 489–90, 509, 513, 514, 529–32, 534, 536, 539, 541, 542, 544, 547, 551, 583–7, 600, 604, 608, 622, 633 conceptual 354, 370–7, 489 flexibly defined 43, 84–5, 115, 485 pragmatic 25, 354, 370, 377–9, 435–6, 440, 446, 454, 480, 507, 633–4 symbolic 85, 89, 301, 305, 306–7, 310–11, 355, 382 meat-eating 129, 241, 275, 285, 287, 342, 349 and brain size 230, 275, 287 and stone tools 27, 28, 57, 275, 285, 341 by Homo erectus 29 megadont 259, 260, 265, 266, see also hominin fossils, megadont archaic memory 2, 16, 65, see also working memory capacity 30, 209, 218, 219, 222, 288, 291, 294, 629 declarative 135, 177 episodic 372–3 and hippocampus 135, 177–8 in apes 53, 121 in birds 124, 373 prospective/retrospective 374 semantic 372, 373, 544 Mendelian genetics 418 and Darwinian evolution 145 mental constructional skills 47, 55, 58, 243 mental representation 24, 42, 66, 108, 329, 347, 354, 371, 374, 376, 377 of predator classes in monkeys 74 Merge (linguistic) 18, 22, 30, 359, 451, 461, 462–3, 473, 483, 498, 544, 630 Michael, gorilla 50 microcephalics/microcephaly 136, 137, 161–2, 169, 172–3, 179 752 subject index microcephalics/microcephaly (cont.) and ASPM 136, 169, 172–3 brain organization vs brain size in 137, 169, 173, 179 language ability in 137, 169, 173 microcephalin (MCPH1): and brain development 161, 172, 173 and genetic drift 172 and natural selection 172 global frequency of variants 173 recent selection or genetic drift 136, 161–2 tone languages and 172–4 microliths 30 Middle Palaeolithic 279, 291 Middle Stone Age 31, see also Mousterian symbolic behaviour in 303–12 migration 63, 107, 170, 252, 257, 280, 341, 473, 508, 539, 560, 566, 573, 577, 578–80, 629 mime 180, 181, 182, 210–11, 211–12, 214, 249, 341, 384, 385, 387–92, 393, 479, 552 mimesis 138, 180–3, 208, 242 mimetic act 180 mimicry 42, 53, 64, 129, 130, 168, 209 definition of 110 mind-reading, see theory of mind Minimalist Program 18, 30, 359, 459, 461, 473, 494, 495, 496–7, 498, 499, 501, 630–1 mirror neurons 117, 118, 139, 140, 182, 207–15, 220, 384, 391, 419, 430, 433, 605–6 mirror self-recognition 106, 121, 123 mitochondrial DNA 8, 31, 239–40, 251–2, 253–4, 255, 256, 257, 576–7, 578–9 mockingbird 97 modality, language 12, 34, 39, 67, 82–9, 95, 96, 110, 130, 139, 329, 355, 380, 391, 432, 434, 479, 486, 507, 545–51, 555–7 gestural or manual 84, 355, 507, 545–57 vocal or oral 82, 95, 355, 391, 545–6, 550, 556–7 modality-neutral 139, 200, 204, see also amodal mode technologies 283–8, see also Oldowan and butchery 284–5 and vegetable food 284–5 distribution of 283–4, 284–5, 286–7 mode technologies 242, 285–8, see also Acheulean mode knapping 292, see also Mousterian and Homo heidelbergensis 289 model(s): animal, of language evolution 45, 88, 110, 116–7, 120–30, 155 avian, of vocal learning 110, 117–19 of human evolution, constrasting 240 modelling: computational 215, 357, 417, 418, 420–2, 488, 581, 583, 596–600, 601, 603, 610, 621, 634 experimental 602–3 mathematical 10, 155, 320, 582, 600–2, 603 robotic 10, 35, 505, 508, 605–11 using embodied agents 10, 509, 605–11 Model-Rival (M/R) method 113–14 modern human(s) 31, 58, 134, 208, 243, 245, 277, 281, 294–5, 301–2, 303–4, 341, 495, 559, see also anatomically modern humans; Homo sapiens cognition 294–5 division of labour 293, 294 fire usage 31, 293 foraging on fish 293, 340, 341 language capacity in 34, 165, 168, 219, 244–5, 330, 495, 540–1, 544, 558–9, 572, 637 multicomponent tools 244, 292, 294 subsistence 293 modern synthesis 145, 501 Mojo, Tatu, and Dar, chimpanzees 48 monkey 4–5, 42, 43, 44, 63, 66, 68–70, 71, 83–4, 86, 89, 90–1, 95, 99, 115, 116–17, 118–19, 126, 127, 137, 138, 139, 140, 182, 186, 187, 191, 194, 195–6, 202–5, 206, 209, 210, 212, 231, 254, 318, 329, 331, 345, 372, 384, 408, 418, 430, 433, 442, 452–3, 485, 486, see also baboon; blue monkey; Campbell’s monkey; capuchin; colobine; colobus monkey; Diana monkey; gelada; langur; last common ancestor, monkeys, apes, and humans; macaques; marmoset; New World monkey; Old World monkey; putty-nosed monkey; spider monkey; squirrel monkey; tamarin; vervet monogenesis of language 558–72 morphologization 439, 477 morphology: biological 4, 139, 160, 262, 263, 264–5, 268, 270, 271, 279, 280 erosion in 632 linguistic 3, 22, 24, 357–8, 362, 403, 435–41, 444, 445, 471, 522–3, 541, 554, 565–6, 602, 632 morphosyntactic features 25, 445, 506, 516, 521, 524, 526 mortuary treatment at Sima de los Huesos, Spain 290 mother and child interactions 122, 128, 246–8, 319, 323–4, 326, 327, 334 motherese 246–7, 296–7, 319, 320, see also infant-directed speech motor control 536, 636 subject index of articulators 77, 80, 129, 178 of beak 116 of hand 178 over vocalization 72, 77, 80 motor cortex 202, 204 and grasping 204, 220 motor neurons 118, 178–9 in brain stem, spinal cord 137, 178 motor sequencing behaviours 136 motor skills 123, 138, 249, 356, 391 motor theory of speech perception 382, 384, 385 Mousterian 291, 298 hafting in 243–4, 291 movement, see displacement, syntactic moving target: adapting to 158–9, 167, 637 and natural selection 158, 162, 167, 458, 629 MSA, see Middle Stone Age multi component tools 244, 292, 294 multimodal communication 87 bonobos vs chimpanzees 87–8 Multiregional model of hominin evolution 240, 278, 280 music/musicality 30, 386, 430, 499, 631, 632 and language, coevolution of 296–8 and language, common neural control 297–8 hierarchical nature of 126, 296, 491 mutation 60, 136, 166, 168–75, 177, 245, 250–2, 253, 255, 314, 385, 499, 563, 607 for language or speech 9, 14, 21, 23, 33, 137, 142, 168–75, 257, 302, 356, 410, 458, 459, 562, 615 naming insight 356, 380 Nariokotome, Kenya 274, see also Homo erectus; Homo ergaster natural selection 60, 157–67, 252, 257, 333, 418 and language evolution 21, 22–3, 25, 72, 79, 162–5, 220, 246, 247, 313–17, 334, 335–6, 338, 354, 363, 366, 368–9, 409, 410–11, 412, 413–14, 415, 458, 459, 500, 628, 629, 637 and nature/nurture debate 18, 145, 152, 157, 500–1, 506, 509–10, 595, 603–4, 627 overuse of 313–17 navigation 374, 606, 607 Neanderthal, see also La Chapelle-aux-Saints, France; archaic Homo sapiens: and Aurignacian 300–2 characteristics of 269–70, 278, 279, 280, 281, 291, 384 cognition 291 distribution of 240, 243, 277, 279–80 753 DNA 171, 239–40, 243, 245, 250–1, 254, 256–7, 279–80, 302 fossils and human fossils 142, 233, 239, 240, 243, 259–60 group size 291 hafting and Mousterian technology 58, 243–4, 291 hunting 291 hyoid bone 141–2, 233, 243, 245 interbreeding with modern humans 257, 280 language capacity in 58, 231, 233, 234, 243–5, 298, 301–2, 386, 480 pigment use by 291 ornaments 300–2 tool use by 243–4, 245, 256 vertebral/thoracic canal 142, 232–3, 245, 270 vocal tract 231, 243, 298 negation marker (linguistic) 409, 454, 514–5 neocortex 136, 141, 176–9, 342, 345 chimpanzee/gorilla 177 components of 176 neocortical expansion 137, 176, 178, 342 and group size 29, 345, 348 neocortical ratio 177, 179, 345 Neolithic 163 neural mechanisms 42, 117, 207–8, 215, 376 involved in speech 62, 171, 208, 212, 313, 628 neural plasticity 135, 136, 142, 355, 404, 464 neuroanatomical structure(s), see asymmetry (neural); arcuate fasciculus; amygdala; anterior cingulate cortex; avian neuroanatomical structures; basal ganglia; brain growth and development; brain stem; Broca’s area; Brodmann’s areas; cerebellum; cortex; dorsolateral prefrontal cortex; F5; frontal lobe; fronto-cerebellar pathway; fronto-parietal circuitry; Heschl’s gyrus; hippocampus; hypoglossal nerve; inferior frontal gyrus; inferior parietal lobe; intraparietal sulcus; lunate sulcus; motor cortex; neocortex; occipital lobe; pallium; parietal cortex/lobe; parietal-occipitaltemporal junction (POT); peri-Sylvian region; petalia; planum temporale; posterior parietal cortex (PPC); prefrontal cortex; premotor area; spinal cord; subcortical connectivity; superior longitudinal fasciculus; superior temporal gyrus; Sylvian fissure; temporal lobe; thoracic spinal cord; Wernicke’s area neuronal production 135 neuronal pruning 135 754 subject index New Caledonian crow 124–5, see also Betty New World monkeys 59, 78, 196, 254, see also capuchin; marmoset; platyrrhine; spider monkey; squirrel monkey; tamarin niche construction 134, 142, 354, 368 nightingale 97 Nim Chimpsky, chimpanzee 47, 49, 53, 54, 467 non-verbal communication 2, 307 nouns 13, 25, 320, 378, 395, 402, 436, 438, 439, 443, 445, 454, 463, 464, 474, 483–4, 486, 506, 515, 519, 520, 529, 532, 534, 542, 565, 567, 568, 624 novel vocalizations 42, 108, 125, 127, 128, 207, 210, 337 great ape 12, 32, 43, 48, 53, 94, 109, 129, 130, 138, 210, 383, 430, 486 parrot 108, 109, 111, 112, 123 object permanence 123, 371–2, 486 observational priming 209, 210 occipital lobe 139, 176 ochre(s) 31, 307, 308 Old World monkey 42, 59, 62, 63, 68, 80, 191, 196, 254, see also baboon; blue monkey; Campbell’s monkey; catarrhine; colobine; colobus monkey; Diana monkey; gelada; langur; last common ancestor of monkeys, apes, and humans; macaque; putty-nosed monkey; rhesus monkey; vervet Oldowan tools 276, 283, 298, see also mode technologies Omo Kibish, Ethiopia 250, 280, see also anatomically modern humans orang-utan: anatomy, neuroanatomy 32, 191, 192 behaviour, cognition 41, 43, 46, 50, 61, 72, 83, 84, 93, 94, 120, 340 phylogeny, genetics 254 ornaments, see also beads: and language evolution 299–302 disappearance of 300 language and symbolism, relationship to 8, 277–8, 301–2 Neanderthal 300 personal 30, 31, 299–302 symbolic culture, relationship to 30, 299, 305, 311 syntactic language, relationship to 301–2, 305 orofacial asymmetry 187–8, 195, 196 orofacial control 170 in voluntary vocalization 212 orofacial dyspraxia 136, 171, 177 orofacial praxis 170, 302 Orrorin tugenensis 261–2 oscine songbirds 99, 115, 116, 117–8, 362, 365, see also songbirds Out of Africa Model 240, 256–7, 276, 277, 278, 280, 281, 292, 508, 562–3, 566 pair bonds 12, 126, 245 and foraging 342 Palaeolithic 282–95, see also Upper Palaeolithic palaeoanthropology pallium / pallial structures 116, 178–9 Pan 57, 83, 255, 262–4, 486, see also bonobo; chimpanzee; panins panins 7, 14, 239–40, 258–9, 261, 262 pant-hoot, chimpanzee 86, 277 acoustic flexibility 75–6 dialects 43, 75–6 Panzee, chimpanzee 53, 372 parallels 3, 4, 5, 185, 246, 395, 625 avian, ape, human 10, 62, 72, 113, 115, 116–17, 118–19, 125–6, 365 Paranthropus boisei 266, 267, 274, see also Australopithecus boisei parental selection 334, see also kin selection parietal cortex/lobe 139–41, 164–5, 176, 203, 219 and mimesis 128 and object identity 203–4, 208 and theory of mind 220–1 evolutionary expansion of 176, 182, 204, 219, 269 parietal-occipital-temporal junction (POT) 139–40, 141, 199–200, 201–2, 204, 205, 206 parrot 101, 129, see also African Grey parrot; Alex; Alo; Griffin; Kyaaro parsimony principle 40, 59–60, 61, 62 parsing behaviour 54, 66, 112, 113, 214, 316, 319, 419, 539, 609 passive construction (linguistic) 25, 447, 450, 459, 488, 506, 519, 533, 553 Peking man 277, see also Homo erectus; Zhoukoudian pelvic narrowing and earlier birth 318, 331 perceptuo-motor system 421, 608, 635, 637 peri-Sylvian region 199 petalia 269, 270 phenotype 14–15, 34, 134, 145, 147, 149, 315, 330, 331, 332, 333, 338, 354, 363, 364, 418, see also genotype and phenotype phenotypic plasticity 15, 34, 134, 152, 155, 158, 167, 354, 511 Phineas T Gage 221 Phoebe, dolphin 122 phonation 225–6, 232, 355, 357, 385, 428 flexible 61 subject index phoneme(s) 3, 6, 125, 126, 136, 225, 336, 421, 423, 433, 452, 551, 635 anatomy and physiology of 3, 225–6 production by Alex (African Grey parrot) 111–13, 123 phonemic language 123, 310 phonetic 428, 462, 487, 607, 623, 624 alphabet 225, 400, see also International Phonetic Alphabet behaviour 329, 425 control 128, 329, 339 distinctions 112, 228, 356, 425, 487–8 flexibility 329, 535 reduction 522, 531, 534 structure 417–22, 490 phonological gesture 357, 417–22 phonological segmentation 14, 111, 421, 507, 551, 552–6, 622–3 by Alex (African Grey parrot) 111–13 phonological storage capacity 140, 218 and intelligence 218 and language 218–19 and syntax 219, 220 phonological syntax phonological systems 4, 368, 406, 486, 499 phonology 19, 22, 24, 246, 339, 356–7, 367–8, 369, 407, 410, 412, 414–16, 417, 436, 440, 443, 444, 481, 486, 490, 496, 541, 577, 637 articulatory 418, 420 as amodal adaptation 183, 432 phonotactics 3, 113, 622 phylogenetic split: between hominins and panins 26–7, 42, 57, 83, 184–5, 239–40, 255, 484, 486 between Neanderthals and modern humans 171, 240, 251, 256, 291 phylogeny 14, 59, 152, 155, 214, 246, 357, 418, 427–31, 445–6, 472, 496 primate 59–60, 246 pidgins 9, 11, 35, 309, 400, 460, 462, 464, 470, 481, 507, 537–44, 631 pigment use 30, 31, 290, 291, 300, 341 pinnipeds 128, 129 Pit of Bones, see Sima de los Huesos, Spain planning 23, 107, 181, 221, 242, 354, 374 and executive functions 140, 218, 222, 248, 294 long range, in modern humans 248, 294, 374 planum temporale 189–90, 201 plastic token languages 41, 50–5 plasticity 75, 499, see also brain plasticity; developmental plasticity; neural plasticity; phenotypic plasticity; vocal plasticity behavioural 134, 333, 464 755 Plato’s problem 359, 493, 496 platyrrhine 254, see also New World monkey play 49, 61, 72, 78, 85, 88, 111–12, 129, 180, 182–3, 187, 196, 211, 320, 334, 485, 608, 609 and mimesis 180, 180–3 playback experiments: with cetaceans 104 with primates 62, 64, 65, 67, 74–5, 76, 91, 92, 94 pointing 122–3, 378, 391–2, 393, 401–2, 403, 485 comprehension of, dolphin 106 in apes 20, 122–3, 347, 378, 485 polygenesis of language 558–72 polysemy of the terms ‘instinct’ and ‘innate’ 146–7, 152 population 13, 29, 99–100, 134, 153, 158, 161, 163, 172, 185–6, 189, 191, 193–6, 252, 300, 332, 341, 559, 561–3, 566, 594, 612–13, 617, 618 contact 13, 240, 252, 256, 278, 279–80, 505, 508, 573–80 population-specific communicative behaviours 48, 75–6, 77, 86, 94, 105, 126, 127 posterior parietal cortex (PPC): and grasping 204–5 and spatial cognition 139–40, 203 POT, see parietal-occipital-temporal junction poverty of the stimulus 19, 20, 357, 359, 366–7, 426, 496, 595, 630, see also Plato’s problem practice 29, 116, 138, 146–7, 181, 183, 213, 418, 531, see also rehearsal vocal, in Alex (African Grey parrot) 111–12 pragmatics 113, 324, 354, 521, 542, 633, 634, 636 prairie dog 5, 127 praxis 170, 207–15, 302 preadaptation 27, 129, 130, 180, 181, 183, 379, 383, 440, 496, 634 predicate (linguistic) 206, 376, 387, 390, 403, 488, 514, 539 prefrontal cortex 138, 182, 219, 221, 269, 303, 384 prehistoric humans 303 pre-language 24, 34, 35, 359, 479–91, 508, 558, 559–60, 561, 62–3, 572, see also protolanguage premotor area 138, 204 and laryngeal representation in macaque 212 and mirror neurons 182, 208, 220 prepared core techniques 243, 290 pre-syntactic principles 29, 34, 481–2 primate calls 25, 42, 62, 90–95, 297, 355, 383, 485–6, 487–9 in humans 444, 481 Principles and Parameters model 18, 21, 493–4, 500, 593, 614 756 subject index procedural learning 29, 138, 177, 179, 181, 242, 243, 249, 342 production, see also vocal production: linguistic 6, 42, 165, 167, 218, 227, 268, 269, 356, 362, 404, 410–2, 458, 484, 485, 635 of vocalization in non-human primates 60–3, 68–9, 71–3, 77–8, 86, 93–5, 110, 129, 185, 187, 195, 212, 453, 457 rates for sound 227 prognathism 278 propositional structures/thought 6, 66, 67–8, 363, 376–7, 391 propositions (linguistic) 24, 69–70, 308, 376–7, 378, 387–92, 447, 485, 487, 489, 540, 541 prosimian 59, 186, 254, see also lemur; tarsier; loris prosody 112, 183, 319, 423, 499 evolution of 183 proto-concepts 354, 371, 374, 486 protogrammar 453, 454, 541 protolanguage 14, 34–5, 44, 122–3, 125, 141, 163–4, 165, 359, 440, 445–6, 460, 461, 462, 467, 472, 479–91, 498, 542, 544 ancestral 32, 483–4, 486, 508, 558 and beads 302 capacity in modern humans 481–3 compositional 359, 480–4, 507, 539, 602 context specificity of 163 dating of 26–7, 29, 57–8, 239–40, 508 evolution of 13, 40, 210–12, 213–14, 241, 242, 247–8, 319, 320, 342, 359, 451, 462, 507, 539, 541 gestural 41, 164, 479 holistic 359, 487–90, 583, 602 in apes 17, 26, 32–3, 41, 46–58, 120–5, 130, 136, 355–6, 467, 481–2, 484–7 in children 464 lexical, see protolanguage, compositional musical 359, 490–1 non-compositional 596 origins of 479–80 vocabulary size of 163–4 protosign 211, 212, 214, 460 protospeech 211, 212, 214 protosyllable 357, 428, 488 protowords 210, 358, 451, 460, 480–1, 483, 485, 486, 487, 489–90 Proto-World 367, 558, 561, 564, 566, 569, 570–1 provisioning 246, 249, 342 of young 29, 341–2 PTBD hypothesis, see putting the baby down hypothesis and motherese punctuated equilibrium 25, 314 putting the baby down hypothesis and motherese 319–21 putty-nosed monkey 42, 76–7, 452 pyow vocalization 42–3, 76–7, 452–3 pyow–hack combinations 76–7, 452–3 and meaning 43, 452–3 Qafzeh, Israel 280, 299, see also anatomically modern humans quantal speech 228, 230, 234, 270 quantifiers (linguistic) 402 quantity discrimination 115, 548 by Alex (African Grey parrot) 110–11, 114, 123 range expansion 276–7, 293, 341, 562–3, 578 by Homo erectus 278, 280, 287 by Homo sapiens 292 rapid evolution 136, 159–60, 257, 509, 614 in guppies 159–60 in humans 134–5, 162 in lizards 159–60 raspberry sounds 94, 138, 188, 195 raven 124 recursion 4, 6, 22, 219–20, 221, 223, 301, 302, 309, 315, 337, 358, 363, 450–1, 452, 458–9, 474, 583, 614, 630 and combinability 18, 220, 362, 451 recursive syntax 115–6, 307–8, 309, 364, 499, 604 lacking in non-human primates 54–5, 58, 67 reduction: in genetic diversity 579 in pidgins 542 of jaw and face in hominins 230–1 phonetic/phonological 18, 461, 522, 531, 534 reference 73–5, 113, 127, 358, 371, 377, 457, 458, 463, 485–6, 509, 515 displaced, see displaced reference functional 5, 91–2, 485 iconic 391, 393–4, 399 indexical 378, 393–4, 397, 399, 400, 402 symbolic 5, 41, 329, 355, 393–405, 485 referential communication 32, 44, 108, 125, 126–7, 128, 241, 246, 341–2, 355, 371, 403, 404, 410, 445, 463, 542, 606 about absent objects, by apes 120 about absent objects, by dolphins 106 and food calls 5, 43 in apes 43, 92, 120, 347 in language-trained apes 41, 51, 111 in monkeys 91 of artifical signals 607–9 referential vocalization 5, 44, 85, 91, 125, 127, 128, 355, 403, 410 subject index by Alex (African Grey parrot) 110–12 by great apes 43, 48, 91–2, 93 learned 108, 110 rehearsal 180, 181, 182 relative clause 453, 472, 474, 519 reliable tools, production by Homo sapiens 292, 294 ‘reliably developing’ concept 151–2, 153, 154 replacement models of later human evolution 240, see also Out of Africa model representation, embodied 180–3 reproductive success 78, 315, 330, 332, 333, 338, 347, 368 respiratory control 141, 178, 225, 226–7, 228, 232–5 inferred from fossils 242 reverse engineering 9–10, 590 rhesus monkey, see macaque, rhesus Rico, dog 122 Rinnie, orang-utan 50 Rocky, sea lion 122 roles, social 56, 606 comprehension of 92 rook 124 rule of law 349 Sahelanthropus tchadensis 261–2, see also hominin fossils, possible Sally Ann test, see false belief tests same/different concept 380 comprehension of 51 comprehension of by Alex (African Grey parrot) 123, 375 Sarah, chimpanzee 51, 54 scaling relationships 314 scavenging 29, 123, 275, 304, 340, 342, 480 Schoăningen, Germany 29, 290, 292 scrub jay 124, 373 sea lion 10, 44, 69, 121, 122, 125, 128, 129, see also Rocky seasonal migrations and foraging 341 segment, see consonant; vowel segmentation 14, 40, 111, 421, 507, 551, 552–7, 622–3, 624, 625 vocal, by Alex (African Grey parrot) 112–13 selective attention/selective inhibition 140 self-embedding 4, 450, 451 self-organization 361, 421, 429, 506, 509, 612–20 definition of 613 semantic compositionality 4, 43, 66, 380, 400, 452–3, 472, 480, 484, 487–8, 489, 583, 596–7, 600, 603, 609, 618, 633, 637 semantic roles 449–50 757 sensitive period 10–11, 116, 126, see also critical period sensorimotor induction 355, 387–92 sentence comprehension 411, 453, 539, 609 in sea lion, dolphin 122 sentences 6, 42, 47, 51, 54, 56, 66, 69, 70, 114–5, 122, 125, 126, 137, 213, 216, 219, 308, 319, 329, 337, 362, 367, 370, 373, 374, 377, 396, 400, 401, 402, 403, 411, 423, 435, 436, 446–9, 454, 456, 461, 463, 467, 472, 474, 477, 506, 519, 524–5, 551, 553, 555, 575, 618, 623, 624, 630, see also clauses complex 47, 55, 58, 214, 308–9, 464 sequential articulation 170, 243, 637 sexual selection 246, 247, 327, 336, 338 and vocal behaviours 97, 247, 335, 490 sexual signals and language evolution 349 shared attention 68, 69, 220 shared cognitive-cultural networks 181 shared goals/intentions 19–20, 356, 370, 379, 485–6 in apes 55–7, 354–5, 485–6 shell 311 beads see beads tick 304–5 shellfish: collecting 31, 341 foraging on by modern humans 28, 293, 340 shells-to-beads inference 311 Sherman, chimpanzee 41, 52, 54, 56 short-term memory capacity 209, 219 siamang: anatomy, neuroanatomy 32, 227 phylogeny, genetics 254 Sierra de Atapuerca, Spain 30, 277, see also Gran Dolina, Spain; Sima de los Huesos, Spain hyoid bone from 233 sign (in Peirce’s and/or Saussure’s sense) 393–405 sign(ed) languages 26, 135–6, 164, 323, 329, 367, 382, 384, 409–10, 417, 432–4, 544, 545–57 Al-Sayyid Bedouin Sign Language 535, 554, 556 American Sign Language, see American Sign Language of the Deaf emergent 9, 35, 507, 508, 545, 552–6 Italian Sign Language 384 Nicaraguan Sign Language 11, 14, 320, 507, 535, 553–4, 631 Sign Language of the Netherlands 551 structure of 3, 432–4, 507, 551 signature whistles 44, 104, 122 758 subject index signs, gestural: combinations, in apes 41, 48–50, 53, 54, 120–1, 453 deceptive usage, by apes 50, 86 invention of, by apes 120–1, 129, 130 usage, in apes 43, 47, 54, 55, 71–3, 82–9, 95, 120, 122, 185–7, 210, 214, 347, 355, 378, 383, 485, 560 Sima de los Huesos, Spain; mortuary treatment in 290 simple recurrent network 634 Simpler Syntax 21 single origins model, see Out of Africa model singularist approaches 315 skills 207, 209, 211, 214, 607 rehearsal of 13, 29, 180, 181 refined 181, 183 stone tool-making 138, 181, 243, 249, 284 skull shape 189 in hominin evolution 200, 229, 230, 243, 261, 266, 275, 276, 278, 279, 331 slings, baby 246, 318 social awareness in non-human primates 73 social behaviour 78, 217, 282, 285, 290, 293–4 in Homo erectus 287–8 social bonding, see social hypothesis; gossip social cognition 291 and language evolution 33, 59–70 baboon 41–2 social grooming 129, 249, 343–4 social group size 290, 293, 332, 341, 344–5, 348 and language evolution 29, 217, 249, 294, 342, 344–5, 429–30 social hypothesis 249 of language origins 343–4 vs tool-making hypothesis 344 social intelligence 67, 124, 172 and group size 348 in dogs 122 in hyenas 123 social knowledge in non-human primates 63–5, 66, 67 social learning 128, 210–11, 383, 536, 581 by Homo erectus 29 in cetaceans 105, 106 social mind 347 social relationships 101, 126, 127, 345, 346, 372, 373, 401 representation of 66, 67, 205–6 soft palate 128, 226, 270, 428, 430 song: avian, see bird song flycatcher 115, 117 ‘syntax’ 115, 452 three-wattled bellbird 117 hierarchically structured 4, 44, 116, 126, 128, 296 humpback whale 44, 103–4, 126 repertoire size, avian 97–9, 100, 101, 115, 126, 344, 362 song complexity, avian 99, 101, 118, 337, 452 song learning: benefits of 99–101, 337 bird 10–11, 12, 44, 96–101, 115, 116, 117–8, 121, 125–7, 128, 146, 171, 334, 337 evolution of 101 mate choice and 96, 97, 337, 490 neighbour relationships and 98–9, 410 whale 44, 102, 103–5, 126, 410 song territories 44, 96–7, 100, 128 songbird 10, 12, 79, 99–101, 109–19, 125–6, 334, 362, 365–6, 490, 496, 604, see also oscine songbird; suboscine sonority 3, 426, 433 sound, advantages of 96, 385, 391 sound articulation 228–32 sound sequences 3, 77, 224, 227 speed of 227, 232 spandrel 7, 21, 25, 26, 34, 246, 316, 499, 501 sparrow: song 97–8, 100 rufous-collared 100 white-crowned 97 spatial cognition/intelligence 29, 203–4, 222, 243, 288, 291 spatial functions/information and language 105, 139–40, 202–6, 218, 242, 243 spatial terminology 438–9, 513 spears 30, 55, 58, 242, 243, 291, 341 and Homo erectus 242, 287 and Homo sapiens 58, 243, 282 from Schoăningen, Germany 29, 290, 292 spectacled parrotlet 101 speech: advantages/disadvantages of 479, 545–6, 548, 549–50 anatomy of 212, 224–35, 357, 423, 426, 430, 433, 533, 615 child-directed, see infant-directed speech comprehension of by apes 48 distinguishing features of 6, 125–7, 224, 225–7, 383, 432 errors 428, 431, 616 evolution of 7–8, 9, 27, 40, 43–4, 89, 120–30, 139, 177, 183, 211, 214, 216–23, 232–5, 241, 245, 246, 247, 248, 258–72, 297, 302, 418–19, 427–31, 466–7, 479–80, 490, 559, 615 subject index evolutionary fitness, marker of 336 in children 222–3, 229, 335, 419, 548 in infants 14, 77–8, 80, 128, 329, 357, 418–9, 428, 431 infant-directed, see infant-directed speech nature of 343–4, 377, 462, 466–7, 469, 490, 546, 552, 622 particulate 355, 357, 417, 419–20, 430 perception 24, 110, 313, 319, 382–3, 384, 385, 427, 431–2, 560 physiology of 4, 12, 29, 33, 62, 83, 94, 138, 140–1, 171, 174–5, 176–9, 189, 270, 342 production 4, 14, 24, 26, 83, 110, 208, 224–35, 269, 329, 365, 417, 427–32, 559, 629 quantal 230, 234, 270 speech articulation 229, 230–1, 365 neural control of 232 speech sounds 228, 319, 324, 382–3, 396, 616–7 anatomy of 430, 616–7 range and variation of 226–7, 234, 241, 365 spider monkey 192 spinal cord 60, 137, 142, 178, 261, 270 and respiratory control 141, 178, 232, 270 splitters 241–2 spontaneous communications by apes 48–50, 54, 86, 94, 372, 378, 481 squirrel monkey 192, 372 starling 115–6, 452 stimulus enhancement 110, 209 Stone Age see Palaeolithic; Upper Palaeolithic; Middle Palaeolithic; Neolithic; Middle Stone Age Stage X in language evolution 439, 440 stone tools 27, 28, 57, 181, 241, 242, 276, 277, 285, 287, 292, 293, 479, 480 earliest 275, 282–3 structure dependence 353, 447 subcategorization (linguistic) 443–4, 466, 480–1, 482, 483, 609 subcortical connectivity 179 subglottal pressure 232, 233, 235 subjacency condition 18, 315, 316, 337, 368, 459 subject, grammatical 18, 25, 67, 206, 219, 387, 390, 438, 443, 447–50, 455, 456–7, 458, 477, 482–3, 532, 551, 565, 631–2 suboscine 115, 117–8, 119 superior longitudinal fasciculus 199 superior temporal lobe 140–1, 189, 219 supralaryngeal vocal tract, see vocal tract 4, 129, 225, 228, 229, 270 SVTH:SVTV ratio 228, 229, 230 swallowing 228, 229, 230–1, 233 759 Swartkrans, South Africa 268, 271, 287 syllable 3, 112, 115, 226, 232, 357, 417, 421, 423, 426, 428, 429, 432–3, 452, 488, 525, 530, 596, 602, 616, 617, 622, 623, 624, 635, see also protosyllable combinations 3, 43 in bird song 97–8, 116, 126 in infant speech 14, 334–5, 418, 419–20, 429 Sylvian fissure 139, 190–2, 196, 200, 204 symbol 5, 13, 24, 47, 301, 305–12, 354, 355, 362, 371, 387–92, 393–405, 407, 409, 433, 484–5, 539, 540, 563, 597, 605 and dogs 122 and dolphins/sea-lions 125 ape usage of 7, 41, 51, 54, 86, 111, 121, 213, 355, 383, 408, 453, 481 definition of 54, 301, 394 in archaeological record 298, 299–302, 303–12 Neanderthal use of 291 symbol grounding problem 24, 354–5, 387–92, 395, 605 symbolic: behaviour 52, 298, 299–301, 303–12, 562 cognition/thought 30, 31, 34, 244, 277, 281, 295, 301, 498, 559 culture 301–2, 307, 309, 349 instruction 355, 389–92 language 171, 301, 309, 310, 460, 468, 469, 541 meaning 89, 301, 306–7, 355, 370–1, 382 nonlinguistic codes 401 reference 5, 41, 393–5, 398, 399–405, 485 symbolic communication 8, 84–6, 88, 298, 327, 334 symbolic theft hypothesis 609 symbolism 8, 40, 41, 243–4, 249, 277–8, 294, 303, 342, 400 symbols-to-syntax inference 305–9, 310–11 syntactic: acquisition 464, 622–5 categories 445, 472, 475, 476, 638 change 515, see also grammaticalization constructions/processes 203, 214, 218, 324, 337, 338, 367, 381, 400–4, 412, 438, 445–50, 472, 476, 481–3, 488, 575, 632, 634, 638 language 13, 15, 17–18, 21–2, 25, 29, 34–5, 47, 206, 219, 245, 248, 301–2, 303–12, 358–9, 381, 410, 413, 415–6, 426, 442–55, 456–68, 480, 482, 507, 509, 510, 520, 537, 539 potential in animal species 47–55, 67, 95, 97, 105–6, 121–4, 127, 452–5, 481–7 structure 22, 203, 438, 464, 472, 482, 539, 543, 551, 618, 623, 637 760 subject index syntax: and vocal complexity 338 complex 219, 247, 324, 337–8, 406, 415, 540 dependencies in, see dependency linear 16, 24, 308, 366, 442, 446–7, 449, 456, 482, 496, 499, 506 origins and evolution of 310, 358, 359, 437, 442, 452, 456–68, 479–91, 606, 609–10, 618 precursors of 66, 77, see also pre-syntactic principles recursive language, see recursion; recursive syntax Talking Heads model 608, 609, 610 tamarin 229, 372 Tania, chimpanzee 49 tarsier 231, 254 teeth 7, 128, 226, 331, 332 in fossil hominins 27, 40, 129, 241, 252, 260, 261, 262, 263, 264, 265, 266, 267, 268, 269, 274, 276, 279, 287 temporal lobe 135, 139, 141, 189, 191, 193, 194, 195, 219, 230 and event perception 182 tense (linguistic) 67, 385, 436, 439, 445, 449, 471, 477, 506, 515–16, 518–19, 522–3, 529, 532, 632 thematic relations 202–3, 204–5, 449–50, 539, see also semantic roles and event perception 202–5 theory of mind 16, 106–7, 108, 140, 220–3, 354, 378, 499, see also false belief tests and avian cognition 124 and ornaments 244, 301, 308 and tool-making 243 in Alex (African Grey parrot) 114 in dolphin 108 in non-human primates 55–6 language evolution, relationship to 68–70, 244, 301, 308 mirror self-recognition test for 106 third factor effects 21, 359, 496, 497, 500 thoracic spinal cord and respiratory control 29, 141–2, 270 throwing 140, 200, 202, 248, 341 thrush 99 timescale(s) 12–14, 159, 590, 591, 602 token ‘languages’ 41, 50–1, 54–5 tone languages 172–4, 363–4, 471 tongue 77–8, 117, 128, 129, 137, 178, 225, 226, 228–31, 234, 235, 270, 357, 385, 417–8, 419, 421, 426, 428, 429, 430 and diet 231, 235 human vs non-human primate and other mammals 77–8, 137, 228, 230–1 innervation in fossils 231 neural control of 116–7, 128, 137, 178 parrot 129 role of in primate articulation 77 tool-dependent foraging 340–2 tool-making / tool use: and hierarchical structure 126, 213 and mimesis 180–3, 249 and neural control 138, 140, 202 as evidence in language evolution 8, 300–02, 303–12 avian 124–5 by dolphins 121 by great apes 7, 41, 52, 56, 58, 93, 121, 195, 213, 379 by modern humans 248 by other mammals 123, 129, 245 composite 8, 30, 244–5 earliest evidence of 275, 282–3, 479 foraging-related 248, 340–2 hominin 27–9, 57, 163, 165, 181, 200, 241–3, 274, 275–7, 282–95, 298, 341, 343, 480 language, relationship to 246, 343, 355, 385, 495 Neanderthal 256, 279 tool-making hypothesis 248–9, 344 transmission: bottleneck, see bottleneck in transmission cultural, or social, or traditional 3, 10, 13, 14, 16, 35, 86, 99–100, 120, 157, 183, 320, 347, 394, 486, 505, 506, 508–10, 535, 560, 562, 583, 590–604, 610, 617–18, 627, 631–3, 636, 638 genetic 253–4 horizontal 13–14, 320 traps as evidence for advanced cognition 244, 292, 294 triadic communication 377, 485 Trinil, Java 275, see also Homo erectus Turkana, Kenya 264, 265, 266, 267, 274 Twin Rivers, Zambia, pigment use at 290 understanding 2, 12, 41–2, 55–6, 57, 64–5, 66, 69, 72, 78, 105–6, 113, 114, 117, 121, 123, 182, 408–12, 415, 453–4, 481, 485, 498, 560, 633, see also theory of mind in dolphins 105 in non-human primates 2, 41, 55–6, 57, 64–5, 69, 72, 114, 121, 182, 411, 453, 481 in parrots 69, 113, 123 others’ goals, intentions, and knowledge 41, 55–6, 57, 64–5, 66, 69, 78, 105, 121, 123, 182 subject index uniformitarianism 300, 302, see also anti-uniformitarianism Universal Grammar 9, 17–22, 170, 213, 314–16, 359, 385, 426, 432, 444, 451, 458, 468, 493–6, 499–501, 506, 510–11, 539, 540, 593, 621, 625, 626–31, 633, 636–8 ‘toolkit’ view of 17, 22, 451 Upper Palaeolithic 8, 30–1, 163, 244, 279, 293–4, 341 art 8, 94 beads in situ European development of 279 tools 30–1, 279 verb(s): at Stage X and/or in pre-language 439, 480, 484, 542 changes in and/or grammaticalization of 162, 463, 466, 488, 506, 515, 516, 519, 532, 533 constructions using 529, 531 grammar of in Chinese 522–6 in protolanguage 446, 483–4, 486, 506, 542 learning 319, 464 morphology 435, 436 open and/or lexical classes of 436, 445, 483 semantic and/or syntactic classes of 443, 444, 480, 482, 530, 624 serial constructions 465, 466 verbal complexity 328–39, see also vocal complexity vertebral canal in fossil hominins 232–3, 243, 270 vertical transmission of information 13, 320, 594 vervet (Chlorocebus aethiops) 4, 42, 61, 62, 75, 115, 127, 297, 375, 403, 485 visuospatial sketchpad 140, 218, 222 vocabulary 4, 5–6, 10, 13, 20, 24, 34, 69, 99, 106, 163–4, 169, 207, 213, 218, 222, 247, 319, 320, 337, 357, 374, 380, 406–16, 419–20, 440, 443–5, 454–5, 462, 466, 473, 475, 480–1, 484–7, 490, 540, 541, 570–1, 623, 637 learning, see word learning Proto-World 570–1 size 5–6, 63–4, 337, 374, 380, 407, 413–4 size and relationship to rate of language evolution 163–4, 357, 419–20 storage of 6, 34, 219, 412–14, 487 protolanguage 462, 466, 480–1, 484–6 vocal behaviour 71–3, 79–80, 90, 112, 113, 121, 130, 136, 329, 335 and audience effects, see audience effects social consequences, awareness of, baboons 73 vocal capacities 29, 40, 41–4, 90–5, 243, 490 great ape 29, 41–3, 90–5 monkey 42 761 vocal combination 117 vocal communication 2, 60, 67, 68, 73, 88–9, 90–5, 101, 102–5, 107–8, 109, 115, 117, 118–9, 125, 128, 195, 202, 224, 297, 332, 344, 433, 490, 560 advantages of 385, 391 dialogue-like, in bonobos 88 non-human primate 71–81, 88–9, 90–5, 202, 297, 490 vocal complexity 329, 336–9, 344 as fitness clue 329, 338 vocal control 12, 80, 121, 328–30, 486 vocal displays 336–9 vocal flexibility 41, 68, 71–80, 108, 118, 126, 129, 227, 329, see also phonetic flexibility alarm calls 75, 485 comprehension 71 evolution of 78–80 great ape 32, 43, 93–5 in pinnipeds 129 non-human primate 12, 42, 71–80, 383 vocal imitation 12, 13, 29, 43, 68, 100, 122–3, 129, 329, 365, 408, 418–20, 486, 490, 619 avian 100, 109–11, 129, 179 dolphin 122 non-human primate 12, 43, 53, 94, 486 vocal labelling of external events, primate 75 vocal learning 10, 12, 33, 44, 48, 54, 106, 108, 116–18, 126, 136, 337, 365, 486 and FOXP2 136 avian 10–11, 44, 99–101, 110, 115, 116–18, 121, 126, 129–30, 178–9, 334 bottlenose dolphin 44, 104, 108 humpback whales 103 mammalian 129 non-human primate 48, 75, 93–4, 99, 329, 355, 365, 383, 386 selective advantages of 337, 490 sexual selection and 337, 490 signature whistles, dolphins 104 vocal ornateness 329 vocal plasticity, see vocal flexibility; vocal learning vocal praxis 302 vocal production 4, 6, 14, 24, 26, 42, 60–3, 68–9, 71–80, 83, 87, 93–5, 110–13, 116, 129, 177, 187, 195, 208, 212, 224–35, 269, 270–1, 336–9, 356, 364, 365, 382–3, 417–22, 423–34, 484, 559 constrained/limited 60, 63, 355, 383–4 flexible 61, 68–9, 71–80, 93–5 vocal signals 75, 77, 79, 84–6, 87 visual interactions, relationship to 79 762 subject index vocal tract: adapted for speech 4, 32, 225–35 anatomy 141–2, 225–35, 270–1 and Broca’s aphasia 171 and diet 141 and speech production 225–35, 357, 380, 417–8, 420–1, 430–1, 559 articulation, speed of 227 evolution of 225, 229–30, 270–1, 479–80, 559–60 fossil evidence of 229–30, 270 hominin 7, 479 human infant 77, 420 human vs primate 4, 26, 32–3, 77, 141, 68, 227 innervation of 231 mammalian 225, 228 modern human 7–8, 16, 141–2, 201, 225–6 motor control of in relationship to vocal repertoire 130 Neanderthal 298 parrot 111 two-tubed 228–9, 270–1 vocal usage 61–2 vocalization(s), see also acoustic signals; calls; vocal signals: and infant-directed speech 320 and social grooming 249, 429–30 as semantic signals 409, 412, 415 avian 44, 101, 109, 111–13, 115, 116, 118, 125–6, 129, 130, 138–9, 140, 179 brainstem control of 137, 178, 212 control of 4, 12 evolution of 319, 391, 615 homologues in earlier hominins 229–30, 271 in human infants and children 246, 334, 357 in language evolution 13, 77, 128 innate 115, 487 language-specific 489 mammalian 44, 60, 104, 108, 122, 129, 225 non-human primate 12, 41–2, 43–4, 58, 61–70, 73–8, 84, 85–7, 89, 90–5, 115, 187, 194–6, 210, 211–12, 214, 227, 297, 329, 330, 383, 430, 452–3, 484 novel 12, 32, 42, 43, 48, 94, 108, 109, 112, 128, 129, 383, 430, 486 properties of in humans 224, 225–32, 393, 407, 485–7 referential 5, 43, 44, 48, 85, 91, 92, 108, 110–12, 125–8, 355, 391, 403 role of FOXP2 in 171, 174 rule-governed 4, 442, 452 volitional control 42, 76 vowel: acoustic properties of 417, 420 as compared with formatives of sign 433 as reproduced by parrots 110–13 harmony 438 in infant-directed speech 297, 319, 323 patterns 624 production 226, 228, 231, 357, 417–22, 426, 428, 429 production by human infants 419, 429 production by Neanderthals 243 systems 16, 228, 323, 325, 420, 432, 474, 509, 617, 619 vowel-like formants / sound sequences 62, 77, 110 Waddington’s concepts 134, 152, 155, 354 warbler 97–8, 99, 100 warrant / warrantedness condition 305–7, 309–10, 311 Washoe, chimpanzee 47, 48–50, 54, 481 weaning 213, 245 and relationship to language learning 331, 332, 334 Wernicke’s area 167, 185, 189, 199, 201 whale 10, 44, 102–8, 410, 490, see also cetacean humpback 44, 103–4, 126 whale song 4, 11, 44, 103–4, 126, 410, 442, 452, 490 wh-constructions 402 whistle 107, 117, 225 nonsignature, dolphin 104 orang-utan 94 signature, dolphin 44, 104, 122, 128 Whorfian hypothesis 633 windows approach to language evolution 32, 463–6, 467, 505, 540, 544 word learning 5, 69, 145, 213, 319, 329, 354, 355, 356–8, 387, 389–90, 402–3, 406–16, 418–20, 428–9, 444–6, 453, 454, 468, 480–2, 489, 506, 510, 559, 608, 614, 622, 624, 633 in animals 44, 48–54, 106, 122, 123, 374, 446, 453, 489 word(s): boundaries 510, 622 class 214, 436, 445–6, 447, 464, 484, 488, 506, 513–22, 526 closed-class 436, 438–9, 445, 454, 472 comparison with primate calls 485–6 content see word, lexical subject index ‘defective’ 410, 481, 539 functional 13, 444–5, 454, 483, 506, 513, 515, 529–30, 533, 542, 544 half-life of 162 lexical 9, 13, 438, 443, 446, 453, 454, 471, 477, 483, 488, 506, 513, 515–6, 523, 526, 529–30, 533, 542, 573–4, 575, 631, 632, 633, 634 open-class 438, 445 order 20, 24, 44, 47, 54, 122, 177, 402–3, 411, 414, 448–9, 453–5, 456, 466, 467, 482, 499, 507, 510, 541, 564, 575, 609, 623 segmentation of 14, 111, 510, 560, 621–5 working memory 30, 140, 177, 217–8, 219, 221–2, 242, 244, 288, 291, 342 and tool production 288, 291, 294 763 executive functions, synonymous with 140, 218, 244, 294 wren 99 X chromosome 252, 576 Y chromosome 8, 252, 253, 254, 576–7, 578, 579 zebra finch 97, 171, 365 Zhirendong, China 240, 278, see also anatomically modern humans Zhoukoudian, China 277, see also Homo erectus Zinjanthropus, see also Paranthropus boisei; Australopithecus boisei 266 .. .the oxford handbook of LANGUAGE EVOLUTION oxford handbooks in linguistics The Oxford Handbook of Applied Linguistics Second edition Edited by Robert B Kaplan The Oxford Handbook of Case... and the conceptual foundations of linguistic theories He is the author of twelve books, including Unravelling the evolution of language (Elsevier, 2003) He was the organizer of the Cradle of Language. .. 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