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This page intentionally left blank The Evolution of Language Language, more than anything else, is what makes us human It appears that no communication system of equivalent power exists elsewhere in the animal kingdom Any normal human child will learn a language based on rather sparse data in the surrounding world, while even the brightest chimpanzee, exposed to the same environment, will not Why not? How, and why, did language evolve in our species and not in others? Since Darwin’s theory of evolution, questions about the origin of language have generated a rapidly growing scientific literature, stretched across a number of disciplines, much of it directed at specialist audiences The diversity of perspectives – from linguistics, anthropology, speech science, genetics, neuroscience, and evolutionary biology – can be bewildering Covering diverse and fascinating topics, from Kaspar Hauser to Clever Hans, Tecumseh Fitch provides a clear and comprehensible guide to this vast literature, bringing together its most important insights to explore one of the biggest unsolved puzzles of human history w tecumseh fitch is Professor of Cognitive Biology at the University of Vienna He studies the evolution of cognition and communication in animals and man, focusing on the evolution of speech, music, and language He is interested in all aspects of vocal communication in terrestrial vertebrates, particularly vertebrate vocal production in relation to the evolution of speech and music in our own species The Evolution of Language w tecumseh fitch CAMBRIDGE UNIVERSITY PRESS Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo, Delhi, Dubai, Tokyo Cambridge University Press The Edinburgh Building, Cambridge CB2 8RU, UK Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521859936 © W Tecumseh Fitch 2010 This publication is in copyright Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press First published in print format 2010 ISBN-13 978-0-511-67747-2 eBook (NetLibrary) ISBN-13 978-0-521-85993-6 Hardback ISBN-13 978-0-521-67736-3 Paperback Cambridge University Press has no responsibility for the persistence or accuracy of urls for external or third-party internet websites referred to in this publication, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate Dedicated to my father Contents List of figures [page ix] Acknowledgments [xi] Introduction [1] section the lay of the land Language from a biological perspective [13] Evolution: consensus and controversy [35] Language [73] Animal cognition and communication [143] section meet the ancestors Meet the ancestors [205] The LCA: our last common ancestor with chimpanzees [234] Hominid paleontology and archaeology [250] section the evolution of speech The evolution of the human vocal tract [297] The evolution of vocal control: the neural basis for spoken language [338] 10 Models of the evolution of speech and phonology [364] section evaluating phylogenetic models of language evolution 11 Historical overview: Western theories of language origin before Darwin [389] viii Contents 12 Lexical protolanguage [401] 13 Signs before speech: gestural protolanguage theories [433] 14 Musical protolanguage [466] 15 Conclusions and prospects [508] Glossary [513] Appendix: species names [519] References [521] Author index [605] Subject index [607] Species index [611] References Trubetskoy, N S (1939/1969) Grundzăuge der Phonologie/Principles of Phonology (Berkeley, CA: University of California Press) Tsai, L S and Maurer, S (1930) “‘Right-handedness’ in white rats,” Science 72, pp 436–438 Tulving, E (2002) “Episodic memory: From mind to brain,” Annual Review of Psychology 53, pp 1–25 Tulving, E and Thomson, D M (1973) “Encoding specificity and retrieval processes in episodic memory,” Psychological Review 80, pp 352–373 Turing, A M (1952) “The chemical basis of morphogenesis,” Philosophical Transactions of the Royal Society of London, Series B 237, pp 37–72 Tutin, C E G (1979) “Mating patterns and reproductive strategies in a community of wild chimpanzees (Pan troglodytes schweinfurthii),” Behavioral Ecology and Sociobiology 6, pp 29–38 Tyack, P L and Clark, C W (2000) 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180, 185–201, 237, 324, 403, 416, 425, 431, 452, 457, 461 Chomsky, Noam, 20f, 32ff, 57, 62, 79f, 87–117, 401, 406f, 497, 505 Clayton, Nicky S., 145, 147, 151, 160, 229, 480 Condillac, E B., 339, 400, 436, 438f Corballis, Michael, 145, 153, 179, 288, 434, 443, 446, 451, 459, 461, 464 Darwin, Charles, 16, 25, 31, 34–71, 89f, 92, 144, 149, 154, 177, 181, 187, 200, 208, 221, 249, 251f, 284, 290, 298, 309, 321, 328, 335, 340, 343, 362, 384, 401, 414f, 439 Deacon, Terrence, 34, 71, 91f, 93, 168, 177, 198, 227, 246, 248, 278, 280, 286, 289ff, 347, 349f, 354f, 370ff, 378, 380, 401, 404, 420ff, 428, 433, 460, 467 Dennett, Daniel C., 71, 89, 109, 135f, 190 Donald, Merlin, 82f, 88, 266, 298, 309, 399, 403, 431, 433, 474, 495, 505 Dunbar, Robin, 156, 187f, 195, 198, 228, 231, 368, 378, 415–428, 467, 486 Emmorey, Karen, 437–465 Enard, Wolfgang, 278, 359f, 478 Falk, Dean, 248, 253, 286f, 332, 477, 492f, 504 Fitch, W Tecumseh, 21, 42, 57, 99, 109f, 115, 180, 183, 195f, 199, 237, 248, 299f, 303, 306, 308f, 311, 314ff, 319, 321ff, 326, 328–331, 335f, 338, 340f, 354, 368, 375, 424f, 428, 431, 469, 474f, 477, 479, 489f, 492f, 495 ´ T., 23, 86, 88, 118f, 401 Givon, Goldstein, Louis, 96f, 102, 374f, 447, 467f, 499 Goodall, Jane, 154, 180, 184, 234, 237f–243 Gould, Stephen Jay, 21, 35, 42, 49, 51f, 57f, 64ff, 210, 284f, 342, 405 Grafen, Allen, 40, 42ff, 196–199 Grice, Paul, 131, 133f Hamilton, William D., 41, 44, 199, 415f, 425, 427, 490 Hare, Brian, 158f, 457 Harnad, Steven, 16, 122 Hauser, Kaspar, 73, 74f, 343, 354, 405 Hauser, Marc D., 21, 57, 144, 146, 149, 152, 165, 171, 180, 187–198, 237, 299f, 306, 311, 326, 354, 375, 403 Hayes, Cathy, 13, 15, 163, 166, 237, 286, 346, 352, 372 Hewes, Gordon, 16, 25, 266, 390, 400, 433f, 439–465, 466f Hockett, Charles, 16, 18f, 26, 35, 93f, 97, 142, 201, 400, 467, 469 Holloway, Ralph, 153, 155, 259, 278, 282, 284–291 Hrdy, Sarah B., 242f, 246, 248, 268 Hurford, James R., 16, 33f, 77, 90, 122, 124, 172, 403, 415, 417, 427, 447, 455f, 484, 501 Jackendof, Ray, 15, 16, 18, 23, 57, 76f, 86f, 96, 115, 117, 122f, 142, 170, 372, 403f, 407, 410ff, 420ff, 424, 474 606 Author index Jespersen, Otto, 16, 345, 470, 466, 474, 481490, 494498, 500, 503f Jăurgen,Uwe, 177f, 181, 347ff, 350ff, 355, 368, 371, 460 Kirby, Simon, 15, 33f, 91f, 378, 380, 383, 430, 496, 501–504 Kuhl, Patricia K., 99, 146, 326f, 343 Ladd, D Robert, 27, 96, 292, 503 Laitman, Jeffrey T., 309, 318, 331, 333 Lamarck, Jean, 36, 38, 47 Langer, Susanne K., 17, 47, 144 Lenneberg, Eric H., 74, 79f, 87, 101, 175, 177, 279, 285, 344, 402 Lewontin, Robert C., 15, 21, 60, 64f Lieberman, Daniel, 260f, 322, 331 Lieberman, Philip, 20, 101, 117, 298, 306ff, 326, 336, 355, 365f, 370, 374f, 401, 403, 407, 431, 451, 467, 496 Lindblom, Bjorn, 94, 195, 300, 368, 381f, 445, 466–507 Locke, John L., 242, 247, 289, 345, 372, 396f Lorenz, Konrad, 27, 31, 68, 80, 194, 384 MacLarnon, Ann, 335f MacNeilage, Peter F., 218, 288, 305, 355, 366–370, 375f, 431, 438, 467 Markman, Ellen, 125, 128ff Marler, Peter, 31, 75, 99, 149, 161, 165, 180, 183, 187f, 191, 301, 326, 335, 339–342, 345ff, 471, 485 Maynard Smith, John, 21, 43f, 50f, 57f, 62, 71, 190, 195–199, 209, 212f, 385, 416 McGrew, William C., 154f, 238ff, 246, 262, 265 Merker, Bjorn, 470 Miller, Geoffrey, 40, 100, 108, 146, 208, 248, 301, 326, 419, 428, 431, 490f Mithen, Steven, 246, 260, 266, 335, 368, 470, 474, 481, 486ff, 492, 495ff, 504f Măuller, F Max, 31, 389, 392, 394ff, 470, 472, 478, 482, 494 Newmeyer, Frederick, 20, 57, 65, 76, 86, 106, 403, 407 Nowak, Martin, 87f, 102, 107, 109 Ohala, John J., 303, 305, 323, 368, 374 Păaaă bo, Svante, 361 Pepperberg, Irene M., 81, 147, 152, 157, 165f, 168ff, 345, 370 Pfungst, Otto, 143 Pinker, Steven, 16, 18, 20, 23, 31, 35, 46, 57f, 65f, 77, 82, 88, 97, 117, 166f, 170, 308f, 358, 389, 403, 407, 438, 486 Premack, David, 135, 159, 164, 166, 168 Reby, David, 196, 309, 319, 321f, 328 Seyfarth, Robert, 117f, 134f, 148, 157, 160, 180, 185–193, 237, 324, 403, 416, 425, 431, 452, 457, 461f Steels, Luc, 122, 383, 430, 505 Studdert-Kennedy, Michael, 102, 374, 467, 499 Tallerman, Maggie, 179, 485, 496, 498–502 Tinbergen, Niko, 27, 31, 68ff, 80, 194, 338 Tomasello, Michael, 31, 86, 88, 116f, 140, 144, 146, 158f, 162f, 192, 232, 234, 430f, 435, 440ff, 454f, 457, 464f, 495, 505 Trivers, Robert L., 42, 199, 415f, 491 Wallace, Alfred R., 39f, 149, 397 Whiten, Andrew, 156, 162f, 238, 240f Wrangham, Richard W., 188, 234, 238ff, 267f Wray, Alison, 431, 467, 480, 486, 496504 Zuberbăuhler, Klaus, 148, 165, 185, 188ff, 237 Zuidema, Willem H., 51, 102, 381f Subject index Acheulean industry, 256, 266f action syntax, 431 agents, 51, 381, 382, 383, 501f alarm calls, 24, 74, 135, 165, 180, 185, 187ff, 199, 236, 425, 469, 471, 482 alliteration, 184, 377 allometry, 282 amniotes, 224, 225 analogous, 36, 46, 57, 65, 99, 229, 262, 306, 310, 381, 409, 488 animal cognition, 17, 81, 122f, 134, 143ff, 149, 159, 171f, 281, 396, 402 anticipatory cognition, 151, 172 argument from the poverty of the stimulus, 85 artificial language, 381, 404, 440 audience effects, 180 audiogram, 324, 325 Aurignacian, 275f automata, 111, 179f babbling, 28, 69, 84, 169, 297, 345, 346, 361, 372, 375f, 383, 399, 443, 460, 464, 471 basal ganglia, 83, 118, 349, 358ff, 365f, 369f, 458 base pairs, 49, 102, 210, 356f bases, 49, 50, 66, 208 basicranium, 329, 330f basihyoid bone, 300, 312, 323, 331, 334 beats, 435, 475, 479 bilaterians, 214, 215 biological, 74f, 77–81, 87–94, 98, 110, 115f, 121ff, 136 bipedalism, 251, 255, 257, 259ff, 279, 332, 373, 482, 493, 495 bonobo, 236, 240, 264 brainstem chassis, 347, 349 Broca’s area, 287f, 350, 351, 359, 365, 369, 453f, 458 cache, 151, 160 call usage learning, 338 categorical perception, 98, 99, 146, 325ff, 475 central nervous system, 62, 215, 218, 328 cerebral laterality, 449 chimpanzee, 13ff, 27, 73, 77, 81, 107, 135, 138, 154, 158, 163f, 166, 173, 188, 207, 233, 234–244, 258, 260, 262, 265, 272, 275, 279, 292, 301, 308f, 320, 323, 330, 332ff, 352, 356, 359, 364, 366, 372, 422, 429f, 443, 458, 493 Chordata, 215 chunking, 100, 141 cis-regulatory elements, 357 clades, 45, 46, 59, 147, 187, 252, 268, 272, 285, 363, 426f, 491, 503 co-evolution, 91, 92 concealed ovulation, 246, 493 co-speech gestures, 435, 436 coarticulation, 374, 375 code model, 131, 133 codon, 49, 50, 210 cognitivist, 86, 87, 124 color vision, 21, 45f, 206, 228, 229, 230f combines, 115, 116, 138 common ground, 133 comparative method, 18, 44, 45f, 70, 149, 206, 213, 217, 234, 249, 275, 334, 471, 483 complex gill bars, 217 complex vocal imitation, 338, 340–362, 382 constraints, 35, 51, 57–68, 79, 86, 88, 95ff, 100, 106, 110, 116, 117f, 125–129, 141, 149, 170f, 174, 185, 195f, 218, 220, 232, 326, 342, 344, 354, 367, 372–385, 411, 418, 420, 429ff, 467f context-free grammar, 110, 113, 114 context-driven inference, 130, 141 continuity, 47, 149, 175, 267, 355, 396, 402, 457, 461f, 497f control of vocalization, 180, 302, 348, 360 convergent evolution, 46, 56, 93, 147, 184, 341, 470, 495, 503 cooperation, 134, 140, 159, 248, 410, 414, 415ff, 489f cortical control systems, 347 corvids, 136, 147, 160, 429 creoles, 379, 406f, 461 critical periods in early visual development, 344 cross-modal cognition, 451, 452f cue, 96, 114, 143, 178, 193, 195, 303, 305, 322, 329, 335 608 Subject index cultural transmission, 19, 89, 92f, 239, 377f, 380, 383, 469, 504 cumulative cultural change, 162, 378 cytoskeleton, 212 deceptive alarm calls, 180 declarative pointing, 441 deictic, 435 describe, 78, 81, 119f deuterostomes, 214 dialects, 34, 161, 188, 345, 378, 437, 471 discontinuous, 48, 149, 175f discrete infinity, 104, 116 displacement, 18f, 469 duality of patterning, 19, 94, 442, 446f, 467, 469, 476 echopraxia, 456 efficacy costs, 197 emblematic, 435 encapsulation, 81, 88, 107 encephalization quotient (EQ), 281 episodic memory, 145, 146, 172, 239 ethologists, 68, 69f, 80, 172, 184, 194, 340, 377 eukaryotes, 210ff evo-devo, 55, 205, 213, 218, 222 evolutionarily stable strategy, 51, 415 exaptation, 24, 63, 64, 65, 117, 179, 205, 215ff, 305, 328 executive, 82 Expensive Tissue Hypothesis, 288f eye-direction detector, 137, 138 facial shortening, 332 finite state grammar, 114, 182, 185 fixation, 60, 207, 273, 291, 360, 430, 504, 506 food calls, 180, 187f, 193, 237, 422 formal language theory, 107, 108–115, 182 formal semantics, 120, 121f, 141 formal, 79, 80, 105ff, 120ff, 182 formalist, 106, 107 formant frequencies, 196, 299, 304ff, 310f, 319, 321ff, 326, 370 FOXP2, 54, 56f, 358, 359–363 frame/content theory, 366 frugivory, 229 functional, 69, 83, 86ff, 105, 139f, 150, 187, 189, 194, 205, 215, 266, 269, 279, 286ff, 290, 331, 334, 353, 356, 358, 360, 418, 437, 459, 471, 477, 498 functionalist, 86, 87, 106 fundamental frequency, 299, 302f, 305 gaze detection, 158 gaze following, 136, 158, 159, 172 gaze prohibition, 159 gene duplication, 54, 205, 215, 219, 229 generated, 79, 84, 94f, 99f, 111f gestural protolanguage, 433, 434–465, 470, 495, 498, 507 gestures, 25, 75, 97, 99, 166, 178, 192, 368, 393, 433, 435ff, 439ff, 447f, 454, 457, 464, 468, 472 gibbons, 176, 235f, 246, 332, 340f, 421 gossip, 198, 410, 417, 418, 419, 423ff, 428, 443 gradualism, 35, 46, 52f great apes, 43, 81, 157, 163, 166, 192, 205, 231, 234ff, 238, 242, 246f, 249, 261, 291, 308f, 323, 334, 427, 434, 440f, 464, 479 Gricean maxims, 133ff, 141 grooming, 193, 238, 368, 410, 417, 418, 419, 425, 441, 486 handedness, 449, 451 handicap, 195, 196, 197f haplotype, 274 Hauser Kaspar, 73ff, 343, 354, 405 hemispheric asymmetry, 287, 440 hierarchical structure, 99, 100, 101, 104, 116, 121, 138, 183f hierarchical, 97, 99, 101, 104, 114, 116, 121, 138, 141, 147, 153, 173, 183f, 187, 367, 373f, 431, 468 holistic protolanguage, 484, 486, 496, 497–504 holistic, 182, 402, 410, 442, 467, 476, 480, 484, 485ff, 496–505 homeobox genes, 53 homologous, 45, 46, 56, 219, 222f, 225, 330 “honest” signals, 195 honeybee dance “language”, 200 hyoid apparatus, 329, 330, 331 hyoid bulla, 333, 334 hypoglossal canal, 330, 333 iconicity, 437, 438f, 446ff imprinting, 344 index/indexical, 195, 280f inheritance, 37, 47f, 50, 71, 77, 91, 163, 473 innate call system, 176, 371 innate constraints, 89, 171, 430 innate human call system, 177 intentionality, 91, 136f, 189, 186, 190, 192, 441, 454 intentional stance, 135, 136 intonation, 96, 480 “invisible hand” models, 91, 383 Subject index jawed vertebrates, 218, 330 jawless fish, 215, 217, 218 KE family, 358, 359, 361f Kebara hyoid, 333 kin communication, 42, 198, 199ff, 424–429, 487, 494 kin selection, 41, 42ff, 198, 416, 424ff, 459, 489, 492f, 503f laryngeal air sacs, 308, 333 larynx, 64, 217, 222, 223, 224f, 267, 297, 300ff, 307–337, 338, 347f, 351, 353ff, 362, 364, 368, 375, 445, 459f lateral cortical system, 350 lateral line system, 223 LCA: last common ancestor, 46, 63, 154, 164ff, 168, 173f, 185, 201, 206f, 230ff, 234–249, 250, 255, 258, 262, 291ff, 323, 366, 370, 400, 405, 410, 418, 420, 434, 440f, 452f, 457f, 461, 466, 469, 488, 493, 495, 503, 505f Levallois technique, 257, 269, 272 lunate sulcus 257, 269, 272 lungs, 36, 63, 220, 221, 223f, 299ff, 307, 354 Machiavellian, 156, 197, 428 mammalian auditory ossicles, 225 manipulation, 135, 140, 194, 497 MCPH gene family, 291 mechanism, 21ff, 46, 64, 67, 69, 75, 77, 79, 81ff, 87, 93, 98ff, 105, 118, 130, 136ff, 190, 208, 217, 222, 301, 313, 338, 344, 346, 383, 430ff, 452, 455, 475, 490 medial cortical system, 349 memes, 42, 89 mental, 79, 89, 101, 103, 122ff, 130, 136f, 138, 144f, 148f, 151, 172f, 190ff, 266, 284, 291, 358, 398, 411, 436, 455, 470ff metaphorical, 435 midbrain control center, 347, 349f middle ear, 63, 217, 223, 225, 324, 445 mildy context-sensitive, 111 mitochondria, 211, 273f mitochondrial DNA, 211, 273, 274 Mitteilungsbedăurfnis, 130, 140f mobbing, 188, 190 morphemes, 19, 94, 103, 104, 395, 404, 406, 467 multi-modal animacy detector, 137, 138 multiregional hypothesis, 273 musilanguage model, 487, 489f mutual exclusivity, 129, 130, 170, 411 MYH16, 263, 356, 357 myoelastic-aerodynamic theory, 301 nativist, 86, 191 natural selection, 21, 37, 38, 39, 41ff, 45, 47f, 50f, 57f, 60, 62f, 65f, 88f, 91, 101, 117, 200f, 248, 284, 291, 342, 371, 384f, 397, 501f Neanderthals, 205, 251f, 256, 268–271, 277, 279, 312f, 317, 333ff, 360, 362, 365, 411, 479, 486 neocortex, 206, 218, 224f, 226, 227, 285f, 350, 352, 360, 409 neural crest, 216, 217 Nicaraguan Sign Language, 379, 406 offspring survival, 232, 241 Oldowan Industry, 463 ontogeny/ontogenetic, 18, 69, 94, 285f, 377, 413, 429, 439, 441 optimization, 21, 50, 51, 58, 381 Out of Africa hypothesis, 257, 266, 273, 274, 279, 403 pantomime, 146, 433, 435, 439, 441f, 448, 458, 464f, 466 parity, 97, 134, 141, 247, 324, 453, 456f particulate principle of self-diversifying systems, 102 perceptual magnet effect, 327 periaqueductal gray (PAG), 348f petalia, 24, 288 phonation, 301, 302, 355, 363, 367, 370f, 375 phrase, 23, 73, 80f, 94, 96, 99f, 103ff, 114f, 118ff, 124, 128, 373, 377, 379, 404, 406, 416, 475, 480, 484, 488, 500, 503 phrases, 14, 80f, 94, 99f, 103, 104, 114f, 118ff, 121, 124, 128, 169, 183, 344, 402, 412, 435, 468, 475f, 480, 484f, 488, 496f, 500, 502f phylogenetic tree, 45, 274 phylogeny/phylogenetic, 34, 70, 76, 94, 117, 160, 235, 252, 274, 336, 366, 377, 387, 429, 433, 440, 453, 459, 480 pidgins, 379, 406f, 433 point mutation, 50 population-level change in allele frequencies, 49 pragmatic inference engine, 141, 194 preadaptations, 63, 64, 171, 475 prescribe/prescriptivism, 78 principle of contrast, 129 productivity/openness, 19, 469 prokaryotes, 210, 211f, 273 propositional meaning, 114, 120, 121, 469, 475f, 503 prosodic protolanguage, 474–505 protolanguage, 176, 266, 293, 343, 392, 395–400, 401–432, 433–465, 466–507 609 610 Subject index protomusic, 477, 478, 479f, 488 protostomes, 214 pseudowords, 95, 373, 467 punishment, 198, 245, 416f, 428 pushdown automaton, 113 reciprocal altruism, 416, 417, 419, 427f recursive structures, 100 reference, 105, 121, 123, 124f, 150, 166, 171, 186, 198, 200f, 282f, 298, 373, 403, 466f regulatory genes, 30, 53ff, 357 repeatability, 480 repetition, 115, 183, 369, 480 residuals, 281 rhyme, 116, 184, 377, 411 saltationists, 48, 53 scale, 110, 479 second-order intentionality, 136, 138, 192 segments, 95, 97, 366f, 369, 475, 499, 502 self-embedding, 100, 104, 116, 118 semantics, 18, 20f, 76, 82, 85, 87f, 93ff, 103, 105ff, 114, 116, 118ff, 141, 175, 182, 186, 291, 373, 404, 419, 426, 435, 437, 441, 463, 467 sensitive period, 73, 74, 75, 343ff sequencing, 100, 141, 273, 356, 358, 379, 503 serial homology, 216, 219 serialization, 77, 117 sexual dimorphism, 249, 250, 259, 261f, 266f, 303, 493 sexual selection, 39, 40, 41f, 196, 201, 248, 342, 343, 399, 472, 474, 481, 490f shared attention mechanism, 137, 138 signals, 18f, 25, 30, 61, 94, 97, 100f, 107, 110, 114, 116, 118f, 124f, 130ff, 134, 141, 146, 148, 165, 173–201, 212, 223, 314, 339f, 346, 353f, 363, 373, 377, 416, 429, 431f, 467f, 472, 474f, 494ff, 501, 503f signed languages, 25f, 97, 297, 433, 435, 437ff, 442, 463, 495 silent substitutions, 210, 359 sociality, 44, 159, 228, 231, 237, 240, 248, 290, 293, 424, 505 sound symbolism, 437, 447, 448 source-filter theory of vocal production, 306 spandrel, 24, 64, 65, 117, 371 speech error data, 367 speech, 13ff, 19, 21, 54, 56f, 76f, 78, 81f, 84, 94–101, 117, 121, 133, 141, 146, 148, 166–171, 174ff, 183, 188, 195f, 216ff, 223f, 237, 249, 287, 292, 297–336, 338–363, 364–384, 391f, 395f, 400, 409, 413, 433–465, 467, 471–505 stability of hierarchical structure, 101 strategic costs, 197 stress-timed, 376 structure dependence, 104, 108 supplementary motor area, 349, 369, 458 symbol grounding problem, 122, 123 symbolic, 64, 108f, 165, 272, 277, 293, 410, 420, 421, 422, 424, 436, 464 synonymous, 50, 108 syntactic, 18, 65, 75, 77, 80, 85ff, 94, 96, 100, 103ff, 110, 116ff, 128f, 148, 169, 181, 183, 185, 313, 366, 379, 401, 406ff, 412, 430f, 444, 467, 475, 477f, 484, 498, 501, 503 syntax, 18ff, 58, 73, 76ff, 79ff, 85ff, 93ff, 100, 102–119, 121, 126, 141, 155, 166ff, 181ff, 287, 291, 298, 351, 365, 372ff, 377, 383, 399f, 401–413, 422, 429ff, 433, 435, 437, 454, 466f, 474ff, 481, 484, 495ff, 500ff taxonomic assumption, 128 template learning, 85 tetrapods, 63, 206, 220, 221ff, 299 the intentional stanc, 136 the principle of contrast, 129 theory of mind, 19, 130, 131, 135–141, 158, 160f, 172, 194, 201 Theory of Mind Mechanism, 137, 138 third inferior frontal convolution, 287 tools, 14, 145, 153, 154, 155, 168, 173, 178, 238, 240, 248, 250, 253, 255ff, 260, 262–267, 270, 272, 276, 279, 292, 396, 398, 426 transcription, 50, 53f, 56, 214, 357f, 360f transitions, 112, 221, 312, 338, 484, 501 turbulence, 301 Turing machine, 108ff vertebrates, 21, 31, 54, 56, 59, 62f, 67, 137f, 147f, 150f, 166, 168f, 171, 175, 181, 186f, 199, 206, 214f, 216, 217ff, 222, 224, 232, 247f, 278, 281f, 288, 299, 301, 306, 321f, 324, 327, 330, 340, 347ff, 353, 368, 370, 405, 426, 430f, 481, 495 Viki (chimpanzee), 13, 15, 27, 73, 163, 352, 372 vocal folds, 299f, 301, 302f vocal imitation, 18, 57, 65, 301, 338, 339–362, 366, 377, 382, 385, 391, 401, 470 vocal tract, 28, 63, 97, 217, 270, 297–336, 338, 347, 352, 354, 361f, 364f, 367f, 373, 375, 403, 439, 471, 475, 479, 498 vocal tract articulators, 305 vocal tract normalization, 314, 315, 319, 326 voice timbre, 299, 305 whole object assumption, 127, 128f word spurt, 129 Species index chimpanzee, 13ff, 27, 81, 136, 154, 158, 163ff, 173, 188, 207, 233, 234, 236–244, 258, 260, 262, 265, 272, 275, 279, 292, 301, 308, 309, 320, 323, 330, 332ff, 352, 356, 359, 364, 366, 372, 422, 429f, 443, 458 280, 301, 309, 311, 315, 316, 317ff, 332f, 339, 344, 364, 411, 471, 500 harbour seal, 14, 340, 346 red deer, 196f, 272, 319, 320, 321f dogs, 24, 45, 81, 123f, 128f, 134, 137, 147, 158f, 164ff, 170, 173, 176f, 179, 189, 191f, 207, ... [250] section the evolution of speech The evolution of the human vocal tract [297] The evolution of vocal control: the neural basis for spoken language [338] 10 Models of the evolution of speech... The sensual eye is just like the palm of the hand The palm has not the means of covering the whole of the beast – From Rumi’s Tales from the Masnavi (translated from Persian by A J Arberry) Language, ... constrain, theories of language evolution Furthermore, once these constraints are taken seriously, many aspects of contemporary evolutionary scenarios (“evolutionarios”) of language evolution