Cover Page: C Half-title Page: i Title Page: iii Copyright Page: iv Dedication Page: v Contents Page: vii Figures Page: ix Acknowledgments Page: xi Introduction Page: 1 The nature of this book Page: 4 A pluralistic, multi-component perspective Page: 4 Plan of the book Page: 7 Section 1 The lay of the land: an overview of disciplines and data relevant to language evolution Page: 11 1 Language from a biological perspective Page: 13 1.1 A biological approach to the “hardest problem in science” Page: 15 1.2 A comparative, pluralistic approach Page: 17 1.3 The faculty of language: broad and narrow senses Page: 20 1.4 Debates and distinctions in language evolution: an overview Page: 24 1.4.1 Communication and language Page: 24 1.4.2 Genes and environment: nature via nurture Page: 27 1.4.3 Innateness and learning: language as an instinct to learn Page: 30 1.4.4 I-language and E-language: cultural and biological evolution of language Page: 32 2 Evolution: consensus and controversy Page: 35 2.1 Introduction Page: 35 2.2 Evolution: the beginnings Page: 35 2.2.1 Natural selection Page: 37 2.3 Categories of selection: sexual, kin, and group selection Page: 39 2.3.1 Sexual selection Page: 39 2.3.2 Inclusive fitness and kin selection Page: 41 2.3.3 “Group selection” – a highly ambiguous term Page: 42 2.4 The comparative method: the biologist’s time machine Page: 44 2.5 Controversies and resolutions in contemporary evolutionary theory Page: 46 2.5.1 Mutation, saltation, and the modern synthesis Page: 47 2.5.2 Resolution: evolutionarily stable strategies Page: 50 2.5.3 Punctuated equilibrium and sudden evolutionary change Page: 51 2.5.4 Macromutations and gradualism Page: 53 2.5.5 Resolution: evo-devo and deep homology: genetic conservation down the ages Page: 55 2.5.6 Selection and constraints: limits on adaptation and natural selection Page: 57 2.5.7 Shifts in function: adaptation, preadaptation, and exaptation Page: 63 2.6 The evolution of behavior: constraints of the “four whys” Page: 66 2.6.1 Explaining behavior: Tinbergen’s “four whys” Page: 68 2.6.2 The role of behavior in evolution Page: 70 2.7 Summary Page: 71 3 Language Page: 73 3.1 Sensitive periods for language acquisition Page: 73 3.2 Understanding linguists: an interdisciplinary dilemma Page: 76 3.3 Modern linguistics and the interface with biology Page: 77 3.3.1 Western linguistics: description not prescription Page: 78 3.3.2 Generative linguistics: mental, formal, and biological Page: 79 3.3.3 Biolinguistics: exploring the biological basis for language Page: 80 Encapsulation Page: 81 Innateness Page: 83 The poverty of the stimulus Page: 84 3.3.4 The biological basis for language: terminology and “Universal Grammar” Page: 87 3.3.5 Historical linguistics revisited: glossogeny and natural selection Page: 88 Components of language: a survey Page: 93 3.4 Phonology Page: 93 3.4.1 Phonology: a generative system Page: 94 3.4.2 Blurry borders: phonetics, phonology, and syntax Page: 95 3.4.3 Signals and the structure of phoneme inventories Page: 97 3.4.4 Sequence memory, hierarchy, and the particulate principle Page: 99 3.5 Syntax Page: 102 3.5.1 Introduction: the challenge and complexity of syntax Page: 102 3.5.2 What is syntax? Page: 103 3.5.3 Many flavors of modern syntax Page: 105 3.5.4 The autonomy of syntax: formalism and functionalism Page: 106 3.5.5 Computability and the theory of computation Page: 107 Formal language theory Page: 108 3.5.6 Formal language theory and music Page: 110 The formal characterizations of musical “syntax” Page: 114 3.5.7 Syntax summary: what needed to evolve Page: 115 3.6 An appetizer: four hypotheses about the evolution of syntax Page: 116 3.7 Semantics Page: 119 3.7.1 The study of meaning in language Page: 119 3.7.2 Formal semantics and propositional meaning Page: 120 3.7.3 Mentalist semantics and the semiotic triangle Page: 122 3.7.4 Child language acquisition: the acquisition of word meanings Page: 125 3.7.5 Constraints on guesses about word meaning Page: 127 3.8 Pragmatics Page: 129 3.8.1 Pragmatics: context is everything Page: 129 3.8.2 Pragmatic inference: context and interpretation Page: 131 3.8.3 Inferential models of communication Page: 132 3.8.4 Symmetry of signaler and receiver: shedding a misleading intuition Page: 134 3.8.5 The evolution of inference: conceptual components Page: 135 3.8.6 Biological components of the theory of mind Page: 136 3.8.7 Autism and “mindblindness” Page: 139 3.8.8 Mitteilungsbedürfnis: the human need to share meaning Page: 140 3.9 Chapter summary: multiple components of language Page: 140 4 Animal cognition and communication Page: 143 4.1 Animal cognition: exorcising Skinner’s ghost Page: 144 4.2 Overview of animal cognition and communication Page: 147 4.3 The study of animal cognition Page: 148 4.4 Animal cognitive capabilities: the basic toolkit Page: 149 4.4.1 Categorization and learning Page: 150 4.4.2 Memory Page: 150 4.4.3 Time and planning Page: 150 4.4.4 Inference and reasoning Page: 151 4.4.5 Number Page: 152 4.4.6 Cross-modal matching Page: 152 4.4.7 Serial order Page: 152 4.5 Specialized forms of intelligence: physical and social intelligence Page: 153 4.5.1 Animal tool use and “physical intelligence” Page: 153 4.5.2 Animal interactions and “social intelligence” Page: 156 4.5.3 Dogs and gaze following: a simple trick? Page: 159 4.5.4 Avian social intelligence Page: 160 4.6 Social learning, culture, and traditions: “animal culture” Page: 161 4.6.1 Vocal traditions Page: 161 4.6.2 Non-vocal traditions Page: 162 4.7 Inter-species communication: animals’ latent abilities to use language-like systems Page: 164 4.7.1 Ape “language” studies Page: 166 4.7.2 Communication between humans and other vertebrates Page: 168 4.7.3 Are constraints on word learning adaptations “for” language? Page: 170 4.8 Animal cognition: conclusions Page: 171 4.9 Animal communication Page: 173 4.9.1 Continuity and discontinuity: a false dichotomy Page: 175 4.9.2 Signals: a key distinction between innate and learned signals Page: 176 4.9.3 Emotional expression and “reflexive” communication in animals Page: 179 4.10 Structure: phonological and syntactic phenomena in animal communication Page: 181 4.10.1 Non-random ordering Page: 182 4.10.2 Phonological syntax and animal “song” Page: 183 4.10.3 Meaningful syntax Page: 184 4.11 Semantics and the meaning of animal signals: reference and intentionality Page: 186 4.11.1 Pragmatic inference in animal communication Page: 186 4.11.2 Functionally referential signals Page: 187 4.11.3 Interpreting functional referentiality Page: 189 4.11.4 Pragmatic signalers: are animals intentionally informative? Page: 191 4.12 The evolution of “honest” communication: a fundamental problem Page: 194 4.12.1 How can “honest” signals evolve? Page: 195 4.12.2 Other routes to honesty: shared interests and communication among kin Page: 198 4.12.3 Kin-selected communication systems Page: 199 4.13 Chapter summary Page: 201 Section 2 Meet the ancestors Page: 203 5 Meet the ancestors Page: 205 5.1 From a single cell to Miocene primates Page: 205 5.2 In the beginning: the first cells and the genetic code Page: 208 5.3 Eukaryotes: the origins of cellular biology Page: 211 5.4 Early metazoans: epigenesis, the Urbilaterian, and the developmental toolkit Page: 213 5.5 Getting a head (and jaws): the first fish and the vertebrate nervous system Page: 215 5.6 Onto the land: proto-tetrapods Page: 220 5.7 Finding a voice: early tetrapods and vocal communication Page: 222 5.8 In the shadow of dinosaurs: amniotes and early mammals Page: 224 5.9 The End-Cretaceous extinction begins the age of mammals Page: 227 5.10 Early primates: sociality, color vision, and larger brains Page: 228 5.11 Early apes and the last common ancestor Page: 230 5.12 Chapter summary: from the first cell to the last common ancestor Page: 232 6 The LCA: our last common ancestor with chimpanzees Page: 234 6.1 Reconstructing the LCA Page: 234 6.1.1 Communication Page: 236 6.1.2 Sociality Page: 237 6.1.3 Tool use, hunting, and medicine Page: 238 6.1.4 Violence Page: 239 6.2 The ape’s impasse: the hominoid mother’s dilemma Page: 241 6.3 Male parental care Page: 243 6.4 Evolving paternal care and monogamy Page: 245 6.5 Implications for language evolution: Why us and not others? Page: 247 6.6 Summary Page: 249 7 Hominid paleontology and archaeology Page: 250 7.1 What the fossils tell us Page: 250 7.2 Paleospecies: naming fossil hominids Page: 251 7.3 A broad overview: major stages in human evolution since the LCA Page: 255 7.4 The earliest hominids Page: 257 7.5 Australopithecines: bipedal apes Page: 259 7.6 The Oldowan Industry and the genus Homo Page: 263 7.7 A major transition in human evolution: Homo erectus Page: 265 7.8 Neanderthals: our large-brained sister species Page: 268 7.9 The common ancestor of Neanderthals and AMHS Page: 270 7.10 Anatomically modern Homo sapiens: Out of Africa Page: 273 7.11 AMHS and the Upper Paleolithic “Revolution” Page: 275 7.12 The evolution of human brain size Page: 278 7.12.1 Absolute brain size Page: 279 7.12.2 Relative brain size Page: 280 7.12.3 Encephalization quotient (EQ) Page: 281 What does brain size tell us? Page: 283 7.13 Reorganization of neural connectivity Page: 285 7.13.1 Fossil endocasts Page: 286 7.14 The brain as an expensive tissue Page: 288 7.15 Integrating the strands: brain size and brain structure in human evolution Page: 290 7.16 Summary: from the LCA to modern Homo sapiens Page: 292 Section 3 The evolution of speech Page: 295 8 The evolution of the human vocal tract Page: 297 8.1 Speech is not language, but is important nonetheless Page: 297 8.2 Vertebrate vocal production: basic bioacoustics Page: 299 8.2.1 The pulmonary airstream Page: 300 8.2.2 The voice source Page: 301 8.2.3 The vocal tract filter Page: 303 8.2.4 Independence of source and filter in vocal production Page: 306 8.3 The reconfigured human vocal tract Page: 307 8.3.1 People are strange Page: 307 8.3.2 The role of the descended larynx in speech Page: 310 8.3.3 Application to fossil hominids Page: 312 8.4 The comparative data I: mammal vocal production Page: 315 8.4.1 Dynamic reconfiguration of the mammalian vocal tract Page: 315 8.4.2 Permanently descended larynges in nonhuman mammals Page: 318 8.4.3 The function of the descended larynx: size exaggeration Page: 321 8.5 Comparative data II: Is speech perception special? Page: 324 8.5.1 Frequency sensitivity Page: 324 8.5.2 Categorical perception Page: 325 8.5.3 Other potentially special aspects of speech perception Page: 326 8.6 Implications of the comparative data Page: 327 8.7 Reconstructing the vocal abilities of extinct hominids Page: 329 8.7.1 The vocal tract skeleton Page: 329 8.7.2 Other proposed fossil cues to vocal anatomy Page: 332 8.7.3 Proposed neurally based cues to vocal control Page: 333 8.7.4 Summary Page: 336 9 The evolution of vocal control: the neural basis for spoken language Page: 338 9.1 Neural control over speech: the central evolutionary event Page: 338 9.2 Evolving learned vocalizations: phylogeny and function Page: 338 9.2.1 Vocal imitation and song Page: 340 9.2.2 Function and phylogeny of complex vocal imitation Page: 341 9.3 Ontogeny of complex vocal imitation Page: 343 9.3.1 Sensitive periods Page: 343 9.3.2 Babbling and vocal imitation Page: 345 9.4 Neural mechanisms underlying complex vocal imitation Page: 346 9.4.1 Shared mechanisms Page: 347 The brainstem chassis Page: 347 9.4.2 The midbrain control region Page: 348 9.4.3 Cortical control regions Page: 349 9.4.4 Vocal control in comparative perspective Page: 352 9.4.5 Implications of the novel circuitry in humans Page: 355 9.5 The molecular genetic basis of complex vocal motor control Page: 356 9.6 FOXP2 and complex vocal motor control Page: 358 9.7 Summary: the vocal tract and its neural control Page: 362 10 Models of the evolution of speech and phonology Page: 364 10.1 Evolving speech Page: 364 Four models of speech evolution Page: 365 10.2 Lieberman’s model: beyond Broca’s area Page: 365 10.3 MacNeilage’s frame/content model of vocal evolution Page: 366 10.3.1 Synthesis Page: 370 10.4 Deacon’s “leveraged takeover” model: Speech as spandrel? Page: 371 10.5 Carstairs-McCarthy: from speech to syllables to syntax Page: 372 10.6 Bridges from speech to phonology Page: 374 10.6.1 Motor constraints on phonological structure Page: 374 10.6.2 Perceptual constraints and phonological structure Page: 376 10.6.3 Vocal imitation, glossogeny, and dialect formation Page: 377 10.7 Computer models of phonological change: simulating glossogeny Page: 380 10.7.1 Modeling the development of phoneme inventories Page: 381 10.7.2 Commentary: explanation in computer simulations Page: 383 Section 4 Evaluating phylogenetic models of language evolution Page: 387 11 Historical overview: Western theories of language origin before Darwin Page: 389 11.1 In the beginning: the first words Page: 390 11.2 The onomatopoetic theory of word origins Page: 391 11.3 The expressive or interjectionist theory Page: 392 11.4 Alternative origins in sociality or song Page: 393 11.5 Max Müller’s attack on evolution and language origin theories Page: 394 11.6 Charles Darwin’s theory of language evolution Page: 397 11.7 Protolanguage in theories of language evolution Page: 399 12 Lexical protolanguage Page: 401 12.1 Introduction Page: 401 12.2 The discontinuity between animal communication and language Page: 402 12.3 “Living fossils” of protolanguage: contemporary windows onto protolanguage Page: 403 12.4 Catastrophic syntax? Page: 407 12.5 Jackendoff’s model: protolanguage plus incremental evolution of syntax Page: 410 12.6 The selective pressures underlying lexical protolanguage Page: 413 12.7 The evolution of cooperative communication: solving a central problem Page: 414 12.8 Dunbar: grooming, “free-riders,” and gossip Page: 417 12.9 Deacon: meat and monogamy; symbolism and group cohesion Page: 420 12.10 Fitch: the origin of information sharing via kin communication Page: 424 12.10.1 Stage 1: kin selection for information exchange Page: 425 12.10.2 Stage 2: reciprocal altruism – no evolution needed Page: 427 12.11 Whence syntax? Page: 429 13 Signs before speech: gestural protolanguage theories Page: 433 13.1 Introduction: From hand to mouth? Page: 433 13.2 Gesture and speech Page: 434 13.3 Signed language Page: 437 13.4 Gestural theories of language origin: a brief history Page: 438 13.5 Gordon Hewes: father of modern gestural protolanguage theories Page: 440 13.6 Arguments against gestural protolanguage Page: 442 13.7 Arbitrariness, indexing, and duality of patterning as key advantages of speech Page: 446 13.8 The neuroscience of gesture: laterality and mirror neurons Page: 448 13.8.1 Cerebral lateralization as evidence for gestural protolanguage Page: 449 13.8.2 Cross-modal cognition Page: 451 13.9 Cross-modal cognition and mirror neurons: Arbib and Rizzolatti’s model Page: 452 13.10 Critiques of the mirror system hypothesis Page: 455 13.11 Arbib’s move “beyond the mirror”: the extended mirror system hypothesis (EMSH) Page: 457 13.12 Critiques of Arbib’s extended hypothesis Page: 461 13.13 Summary: taking stock of gestural protolanguage Page: 464 14 Musical protolanguage Page: 466 14.1 Introduction: phonology remains puzzling Page: 466 14.2 Charles Darwin’s theory revisited: “musical protolanguage” Page: 470 14.3 Prosodic protolanguage: a contemporary update Page: 474 14.4 Prosodic protolanguage and modern music Page: 478 14.5 Adding meaning to prosodic protolanguage: Jespersen’s model and the origins of meaning Page: 481 14.6 Analyzing holistic protolanguage Page: 484 14.7 Modern versions of musical protolanguage theory Page: 485 14.7.1 Mithen’s “Hmmmm” model Page: 486 14.7.2 Steven Brown’s “musilanguage” model Page: 487 14.7.3 Group selection Page: 489 14.7.4 Sexual selection Page: 490 14.7.5 Kin selection Page: 492 14.8 Critiques of musical hypotheses Page: 494 14.9 Holistic protolanguage today: Alison Wray’s model of holistic protolanguage Page: 496 14.10 Critiques of Wray’s holistic protolanguage Page: 498 14.11 Simon Kirby’s simulations of holistic/analytic transitions Page: 501 14.12 Synthesis and prospects Page: 503 15 Conclusions and prospects Page: 508 Glossary Page: 513 Appendix: list of species names Page: 519 References Page: 521 Author index Page: 605 Subject index Page: 607 Species index Page: 611