The oxford handbook of language evolution maggie tallerman

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LANGUAGE EVOLUTION

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

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# Editorial matter and organization Maggie Tallerman and Kathleen R Gibson 2012

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1 3 5 7 9 10 8 6 4 2

Preface and Acknowledgements xi

Kathleen R Gibson and Maggie Tallerman

3 Language or protolanguage? A review of the ape language

10 Evolution of communication and language: insights from

parrots and songbirds 109Irene M Pepperberg

11 Are other animals as smart as great apes? Do others provide

better models for the evolution of speech or language? 120Kathleen R Gibson

PA RT I I T H E B I O LO G Y O F

L A N G UAG E E VO 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 133Kathleen R Gibson and Maggie Tallerman

13 Innateness and human language: a biological perspective 143

W Tecumseh Fitch

14 Evolutionary biological foundations of the origin of

language: the co-evolution of language and brain 157Szabolcs Sza´mado´ and Eo¨rs Szathma´ry

15 Genetic influences on language evolution: an evaluation

Karl C Diller and Rebecca L Cann

16 Not the neocortex alone: other brain structures also

contribute to speech and language 176Kathleen R Gibson

17 The mimetic origins of language 180Merlin Donald

18 Evolution of behavioural and brain asymmetries in primates 184William D Hopkins and Jacques Vauclair

19 Towards an evolutionary biology of language through

comparative neuroanatomy 198Wendy K Wilkins

20 Mirror systems: evolving imitation and the bridge from

praxis to language 207Michael A Arbib

21 Cognitive prerequisites for the evolution of indirect speech 216Frederick L Coolidge and Thomas Wynn

22 The anatomical and physiological basis of human speech

production: adaptations and exaptations 224Ann MacLarnon

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 E VO LV E ?

23 Introduction to Part III: The prehistory of language: When

and why did language evolve? 239Kathleen R Gibson and Maggie Tallerman

24 Molecular perspectives on human evolution 250Rebecca L Cann

25 The fossil record: evidence for speech in early hominins 258Bernard A Wood and Amy L Bauernfeind

26 The genus Homo and the origins of ‘humanness’ 273Alan Mann

27 The Palaeolithic record 282Thomas Wynn

28 Musicality and language 296Steven Mithen

29 Linguistic implications of the earliest personal ornaments 299Francesco d’Errico and Marian Vanhaeren

30 Inferring modern language from ancient objects 303Rudolf Botha

31 Natural selection-itis 313David Lightfoot

32 The role of hominin mothers and infants in prelinguistic

Dean Falk

33 Infant-directed speech and language evolution 322Bart de Boer

34 Displays of vocal and verbal complexity: a fitness account

of language, situated in development 328John L Locke

35 Tool-dependent foraging strategies and the origin of language 340Kathleen R Gibson

36 Gossip and the social origins of language 343Robin I M Dunbar

37 Social conditions for the evolutionary emergence of language 346Chris Knight and Camilla Power

PA RT I V L AU N C H I N G L A N G UAG E :

T H E D E V E LO P M E N T O F A

L I N G U I S T I C S P E C I E S

38 Introduction to Part IV: Launching language:

the development of a linguistic species 353Maggie Tallerman and Kathleen R Gibson

39 The role of evolution in shaping the human language faculty 361Stephen R Anderson

40 The origins of meaning 370James R Hurford

41 The origins of language in manual gestures 382Michael C Corballis

42 From sensorimotor categories and pantomime to grounded

symbols and propositions 387Stevan Harnad

43 The symbol concept 393Terrence W Deacon

44 Words came first: adaptations for word-learning 406Robbins Burling

45 The emergence of phonetic form 417Michael Studdert-Kennedy

46 The evolution of phonology 423Peter F MacNeilage

47 The evolution of morphology 435Andrew Carstairs-McCarthy

48 What is syntax? 442Maggie Tallerman

49 The origins of syntactic language 456Derek Bickerton

50 The evolutionary relevance of more and less complex

forms of language 469Andrew Carstairs-McCarthy

51 Protolanguage 479Maggie Tallerman

52 The emergence of language, from a biolinguistic point of view 492Cedric Boeckx

PA RT V L A N G UAG E C H A N G E , C R E AT I O N ,

A N D T R A N S M I S S I O N I N M O D E R N H U M A N S

53 Introduction to Part V: Language change, creation,

and transmission in modern humans 505Maggie Tallerman and Kathleen R Gibson

54 Grammaticalization theory as a tool for

reconstructing language evolution 512Bernd Heine and Tania Kuteva

55 Domain-general processes as the basis for grammar 528Joan Bybee

56 Pidgins, creoles, and the creation of language 537Paul T Roberge

57 What modern-day gesture can tell us about language evolution 545Susan Goldin-Meadow

58 Monogenesis or polygenesis: a single ancestral language

for all humanity? 558Johanna Nichols

59 Prehistoric population contact and language change 573Brigitte Pakendorf

60 Why formal models are useful for evolutionary linguists 581Kenny Smith

61 Language is an adaptive system: the role of

cultural evolution in the origins of structure 589Simon Kirby

62 Robotics and embodied agent modelling of the

evolution of language 605Angelo Cangelosi

63 Self-organization and language evolution 612Bart de Boer

64 Statistical learning and language acquisition 621Katharine Graf Estes

65 A solution to the logical problem of language evolution:

language as an adaptation to the human brain 626Nick Chater and Morten H Christiansen

This volume represents a snapshot of current knowledge and theories in the field oflanguage evolution, and is written from the widely differing perspectives of thenumerous academic disciplines that contribute to this fascinating and tantalizingfield of study The editors invited many dozens of the most eminent academics tocontribute; a number, sadly, felt unable to take the time to write for a generalaudience, but 60 scientists at the top of their game did accept the invitation Tothese colleagues, we extend a heartfelt thanks for all your efforts, including the timespent drafting and re-drafting chapters to our exacting standards It is your life’swork that has helped to make this volume a success, and we are grateful to you forsharing this research with the general reader

What has resulted from these endeavours is a collection which, we hope, will be astandard work of reference for years to come Language evolution is, however, adiscipline that is moving extremely quickly Even as we write, many new discoveriesand developments are being reported and published Of course, some of theseapparent breakthroughs will prove ephemeral or illusory, but others will doubtlesschange the field We have tried to avoid endorsing too soon any results which arenot yet proven, but at the same time we have attempted to be—and to encourageour contributors to be—as up to date as possible

The target audience for this volume comprises scholars and students of all thedisciplines that contribute pieces of the language evolution puzzle: linguists,biologists, archaeologists, ethologists, neuroscientists, anthropologists, psycholo-gists, geneticists, palaeontologists, and more We have encouraged contributors towrite with this general audience in mind as much as possible, and we hope that theresults will be widely accessible to readers of all backgrounds

We would like to extend a special massive thanks to Laura Bailey, a PhD student atNewcastle University, England, for her painstaking work in compiling and checkingthe list of references and finalizing the subject index We probably drove her almostinsane, but she completed her task on time and without complaint We gratefullyacknowledge the support of the Faculty of Humanities and Social Sciences at New-castle, both in funding Laura’s work and in allowing Maggie Tallerman leave fromteaching for one semester in order to complete this volume We also thank JohnDavey at Oxford University Press for his help and support over the long timespan ofthis project Finally, we’d like to express our gratitude to our partners, S J and John,

for their patience and forbearance in the face of our continuing need to work when

we should have been taking holidays with them

Maggie Tallerman, Newcastle, EnglandKathleen R Gibson, Houston, Texas

January 2011

8.1 Part of the repertoires of two neighbouring song sparrows 98

13.1 Multiple suggested interpretations of ‘innateness’ 150

18.1 Mean handedness indices for chimpanzees, bonobos, and baboons 186

18.2 Orofacial asymmetries in a chimpanzee 188

18.3 Three-dimensional magnetic resonance imaging reconstruction of a

18.4 Mean asymmetry index measured from a sample of chimpanzee

magnetic resonance imaging scans 191

25.1 Hominin grades—speciose taxonomy 259

27.1 Basic knapping procedure or gesture 283

27.2 Mode 1 tools from the site of Peninj 284

27.3 A 1.4-million-year-old handaxe from Tanzania 286

27.5 Painted images of horses, cattle, and deer from Lascaux, ca 17 kya 294

30.1 Structure of the compound inference about the language used by

the inhabitants of Blombos Cave 305

30.2 Structure of non-compound inference about language evolution 312

43.1 C S Peirce’s categorical scheme for a taxonomy of sign forms 397

54.1 Basis of reconstruction 517

54.2 Six hypothesized layers of grammatical development 519

61.1 The three complex adaptive systems that give rise to language

and their interactions 591

61.2 Example grammars from a run of Brighton’s simulation 598–9

22.1 Summary of the evolution of human speech production features

and potential speech capabilities 234

25.1 Species included in a splitting taxonomy 260

58.1 Expected retentions at various rates with 5000-year increments 564

58.2 Survival rates of morphological ergativity in nouns 565

58.3 Retention rates for inclusive/exclusive distinction 567

58.4 Survival rates for grammatical gender 568

Stephen R Anderson is the Dorothy R Diebold Professor of Linguistics, ogy and Cognitive Science at Yale University His interests in Linguistics cover all ofthe major subfields, although his work in recent years has focused on the theory ofmorphology Among his publications, the 2004 book Doctor Dolittle’s delusion(Yale University Press) discusses the gulf between human language and the com-munication systems of other species

Psychol-Michael A Arbib was born in England, grew up in Australia, and is now sity Professor, Fletcher Jones Professor of Computer Science, and Professor ofNeuroscience (etc.), at the University of Southern California His first book wasBrains, machines and mathematics His 39th book, co-edited with Derek Bickerton,

Univer-is The emergence of protolanguage: HolophrasUniver-is vs compositionality (Benjamins, 2010).Amy L Bauernfeind is a doctoral student in Hominid Paleobiology at the GeorgeWashington University, where she studies comparative primate neurobiology Herresearch interests include variation in neural structure, neurodevelopment, andmetabolism in the primate brain

Derek Bickerton (PhD Cambridge) is Professor Emeritus of Linguistics at theUniversity of Hawaii He is best known for his work on creole languages, includingthe controversial Bioprogram Hypothesis, and on the evolution of language Hispublications in the latter field include Language and species (University of ChicagoPress, 1990), Adam’s tongue (Hill & Wang, 2009), and Biological foundations andorigins of syntax (MIT Press, 2009, edited with Eo¨rs Szathma´ry)

Cedric Boeckx is Research Professor at the Catalan Institute for Advanced Studies(ICREA), and a member of the Center for Theoretical Linguistics at the UniversitatAuto`noma de Barcelona His publications include Islands and chains (Benjamins,

2003), Linguistic Minimalism (OUP, 2006), Bare syntax (OUP, 2008), and Language

in cognition (Wiley-Blackwell, 2009) He is founding co-editor of the Open Accessjournal Biolinguistics, and the founding editor of OUP’s Oxford Studies in Biolin-guistics monograph series

Rudolf Botha is Emeritus Professor of General Linguistics at the University ofStellenbosch and Honorary Professor of Linguistics at Utrecht University Hisresearch includes work on the evolution of language, morphological theory and

word formation, and the conceptual foundations of linguistic theories He is theauthor of twelve books, including Unravelling the evolution of language (Elsevier,

2003) He was the organizer of the Cradle of Language Conference held in 2006 inStellenbosch, South Africa

Robbins Burling has been concerned for a half-century with the linguistics andethnology of north-eastern India and Bangladesh He became interested in theevolution of the capacity for language when colleagues in archaeology asked ‘Whendid language begin?’ and he could only reply ‘I dunno’ He is the author of Thetalking ape: How language evolved (OUP, 2005)

Joan Bybee was at the University at Buffalo from 1973–1989 and is now guished Professor Emerita at the University of New Mexico Her book The evolu-tion of grammar (University of Chicago Press, 1994, with Perkins and Pagliuca)documents how unrelated languages create grammar using similar mechanisms.Works arguing that domain-general principles give rise to linguistic structure arePhonology and language use (CUP, 2001), Frequency of use and the organization oflanguage (OUP, 2007), and Language, usage and cognition (CUP, 2010)

Distin-Angelo Cangelosi is Director of the Centre for Robotics and Neural Systems at theUniversity of Plymouth His PhD is from the University of Genoa, and he has beenvisiting scholar at the University of California, San Diego and the University ofSouthampton He edited Simulating the evolution of language (Springer, 2002), and

is currently co-editor-in-chief of the journal Interaction Studies His primary work

is on language learning and evolution in robots and embodied agents

Rebecca L Cann is a Professor of Genetics at the University of Hawaii, Manoa.Her interests include human evolutionary genetics and the molecular conservationgenetics of endangered species She is interested in the shared properties of modernendangered species and the early stages of human evolution These include smallpopulation sizes, gender differences in behaviour, infectious disease risks, andgeographical isolation

Andrew Carstairs-McCarthy is a former Professor of Linguistics at the University

of Canterbury, Christchurch, New Zealand His main research interests have beenmorphology (Allomorphy in inflexion, Croom Helm, 1987) and the evolution oflanguage (The origins of complex language, OUP, 1999) These two interests con-verge in his most recent book, The evolution of morphology (OUP, 2010)

Nick Chater is Professor of Behavioural Science at Warwick Business School Hisresearch focuses on fundamental principles of cognition, and especially onreasoning, decision making, and language His has co-edited or co-written sixbooks, and has written over 200 papers He is a Fellow of the Cognitive ScienceSociety and Associate Editor of the journal Psychological Science

Dorothy L Cheney graduated from Wellesley College and received her PhD fromthe University of Cambridge, where her advisor was Robert A Hinde She was apost-doctoral fellow at Rockefeller University, working with Peter Marler Togetherwith Robert Seyfarth, she is the author of How monkeys see the world (University ofChicago Press, 1990) and Baboon metaphysics (Chicago, 2007) She is currentlyProfessor of Biology at the University of Pennsylvania.

Morten H Christiansen is Professor in the Department of Psychology and Director of the Cognitive Science Program at Cornell University, as well as ExternalProfessor at the Santa Fe Institute His research focuses on the interaction ofbiological and environmental constraints in the processing, acquisition, and evo-lution of language He is the author of more than 125 scientific papers and hasedited volumes on connectionist psycholinguistics, language evolution, andlanguage universals

Co-Frederick L Coolidge is a Professor in the Department of Psychology at theUniversity of Colorado at Colorado Springs He has a PhD in psychology andcompleted a postdoctoral fellowship in clinical neuropsychology His research interestsinclude behavioural genetics, cognitive archaeology, and psychological assessment Hehas received three Fulbright Fellowships to India and has published five books Hehas published articles in the Journal of Human Evolution, Cambridge ArchaeologicalJournal, PaleoAnthropology, Journal of Anthropological Research, and elsewhere.Michael C Corballis, a New Zealander, completed his PhD from McGill Univer-sity, and taught in the Department of Psychology there before being appointed tothe University of Auckland in 1978, where he is currently Emeritus Professor ofPsychology He has published widely on various aspects of cognitive neuroscience,including laterality, imagery, and language He is the author of From hand to mouth(Princeton University Press, 2002), and his new book The recursive mind (Prince-ton) will appear in 2012

Terrence W Deacon is Professor and Departmental Chair in Anthropology at theUniversity of California, Berkeley His research combines human evolutionarybiology and neuroscience, with the aim of investigating the evolution of humancognition It extends from laboratory-based cellular-molecular neurobiology to thestudy of semiotic processes underlying animal and human communication, espe-cially language Many of these interests are explored in his 1997 book, The symbolicspecies: The coevolution of language and the brain (W W Norton)

Bart de Boer did his PhD with Luc Steels at the Vrije Universiteit Brussel on thetopic of self-organization in vowel systems An adaptation of this thesis waspublished as The origins of vowel systems (OUP, 2001) He has also worked withPatricia Kuhl at the University of Washington on the evolutionary implications of

infant-directed speech Currently he works at the Universiteit van Amsterdam onthe biological evolution of the vocal tract and on cultural evolution of speech.Francesco d’Errico is a CNRS Director of Research attached to the University ofBordeaux, France Focusing on the origin and evolution of symbolic behavioursand complex bone technologies, his research has questioned the dominant para-digm of a sudden European origin of modern cultures and identified the presence

of older symbolic traditions in Africa, Europe, and the Near East He has publishedseveral books and more than 200 papers, mostly in international journals.Frans B M de Waal is a biologist with a PhD from the University of Utrecht, inthe Netherlands He is C H Candler Professor of Psychology at Emory University,and Director of the Living Links Center at the Yerkes National Primate ResearchCenter, both in Atlanta, GA He is the author of numerous books, including Theage of empathy (Random House, 2009)

Karl C Diller is researching the evolutionary genetics of language in the Cannlaboratory (genetics) at the John A Burns School of Medicine, University ofHawaii His PhD in linguistics is from Harvard University He was formerlyProfessor of Linguistics at the University of New Hampshire, where a specialinterest of his was the neurolinguistic foundations of second language acquisition.Merlin Donald is Professor Emeritus, Queens University, Ontario; HonoraryProfessor, University of Aarhus, Denmark; and former Chair of Cognitive Science,Case Western Reserve University, Ohio He is a Fellow of the Canadian Psychologi-cal Association and the Royal Society of Canada, and the author of Origins of themodern mind: Three stages in the evolution of culture and cognition (Harvard, 1991),and A mind so rare: The evolution of human consciousness (Norton, 2001)

Robin I M Dunbar is Professor of Evolutionary Anthropology at the University

of Oxford, a Fellow of the British Academy, and co-director of the British emy’s Centenary Research Project His principal research interests focus on theevolution of sociality in mammals, with particular reference to ungulates, pri-mates, and humans

Acad-Dean Falk is a palaeoanthropologist who specializes in brain evolution She is aSenior Scholar at the School for Advanced Research in Santa Fe, New Mexico and

at the Department of Anthropology, Florida State University Her books includeFinding our tongues: Mothers, infants and the origins of language (Basic Books,

2009), and Bones to pick: How two controversial discoveries changed perceptions ofhuman evolution (University of California Press, 2011)

W Tecumseh Fitch is Professor of Cognitive Biology at the University of Vienna.His main interests are in bio-acoustics and the evolution of cognition, particularlythe evolution of human speech, language, and music, all studied from a broad

comparative perspective He is author of The evolution of language (CUP, 2010) Heconducts experimental research on vocalization and cognition in humans and avariety of vertebrates, including chimpanzees, seals, deer, dogs, alligators, and parrots.Kathleen R Gibson is Professor Emerita of Neurobiology and Anatomy, Univer-sity of Texas Medical School, Houston and Professor Emerita in the Department ofOrthodontics, U.T School of Dentistry She has co-edited six books on thedevelopment and evolution of brains and cognition including ‘Language’ andintelligence in monkeys and apes (CUP, 1990), Tools, language and cognition inhuman evolution (CUP, 1993), Modelling the early human mind (McDonald Insti-tute for Archaeological Research, 1996), and Evolutionary anatomy of the primatecerebral cortex (CUP, 2001) She is currently co-editor with Jim Hurford of theOxford Series, Studies in the Evolution of Language.

Susan Goldin-Meadow is the Beardsley Ruml Distinguished Service Professor atthe University of Chicago, where she has taught for over 30 years Her researchinterests focus on gesture—the home-made gestures children create when notexposed to language, and the gestures we all produce when we talk She is thefounding editor of the journal Language Learning and Development, and is the currentpresident of the International Society for Gesture Studies

Katharine Graf Estes’ research investigates the processes underlying early languageacquisition and the connection between early phonological and lexical representa-tions She received her PhD in developmental psychology from the University ofWisconsin-Madison in 2007 She is currently an Assistant Professor at the Univer-sity of California, Davis She has received funding from the National Institutes ofHealth and the National Science Foundation

Stevan Harnad, External Member of the Hungarian Academy of Science, is author

of or contributor to over 300 publications, including Origins and evolution oflanguage and speech, Lateralization in the nervous system, Peer commentary onpeer review: A case study in scientific quality control, Categorical perception: Thegroundwork of cognition, The selection of behavior: The operant behaviorism of

B F Skinner, Scholarly journals at the crossroads: A subversive proposal for electronicpublishing, and Cognition distributed: How cognitive technology extends our minds.Bernd Heine is Emeritus Professor at the Institute of African Studies (Institut fu¨rAfrikanistik), University of Cologne His 33 books include Possession: Cognitivesources, forces, and grammaticalization (CUP, 1997); Auxiliaries: Cognitive forces andgrammaticalization (OUP, 1993); Cognitive foundations of grammar (OUP, 1997);with Derek Nurse, African languages: An introduction (CUP, 2000); A linguisticgeography of Africa (CUP, 2008); with Tania Kuteva, World lexicon of grammatica-lization (CUP, 2002); Language contact and grammatical change (CUP, 2005); The

changing languages of Europe (OUP, 2006); and The genesis of grammar (OUP,

2007)

William D Hopkins is Professor of Psychology and Neuroscience at Agnes ScottCollege He also holds the appointment of Research Scientist within the Division ofDevelopmental and Cognitive Neuroscience at the Yerkes National Primate ResearchCenter of Emory University His research focuses on the evolution of language andspeech, and on the evolution of manual skills, such as grasping, bimanual coordina-tion and tool use, as they relate to the emergence of population-level behavioural andbrain asymmetries in non-human primates

Jim Hurford is Emeritus Professor of Linguistics at the University of Edinburgh

He has an interest in reconciling traditions within and outwith linguistics whichhave tended to conflict He works in a framework in which representation ofgrammars in individual minds interacts with properties of language used incommunities, emphasizing the interaction of evolution, learning, and communi-cation He constructed early computer simulations of language evolution, and hisbooks include The origins of meaning (OUP, 2007) and The origins of grammar(OUP, 2011)

Vincent M Janik is a Reader in Biology in the School of Biology, University of

St Andrews, UK Previously, he was a postdoctoral research fellow at the WoodsHole Oceanographic Institution, USA and a Royal Society University ResearchFellow at the University of St Andrews His research concentrates on vocal com-munication in marine mammals and the evolution of complexity in animalcommunication and cognition

Simon Kirby is Professor of Language Evolution at the University of Edinburghand co-founder of the Language Evolution and Computation Research Unit Hehas pioneered the application of computational and mathematical modellingtechniques to traditional issues in language acquisition, change, and evolution

In particular, he has developed an experimental and computational approach tocultural evolution called Iterated Learning, which treats language as a complexadaptive system operating on multiple interacting timescales

Chris Knight was for many years Professor of Anthropology at the University ofEast London He is best known for his 1991 book, Blood relations: Menstruation andthe origins of culture He helped initiate the Evolution of Language (EVOLANG)series of international conferences and has published widely on the evolutionaryemergence of language and symbolic culture

Tania Kuteva is Professor of English Linguistics at the Heinrich-Heine-University

of Du¨sseldorf, Germany Apart from a number of articles in international journals,she has also published two books with Cambridge University Press and three withOxford University Press, on grammaticalization, language contact, grammatical

typology, and language evolution, four of which are co-authored with BerndHeine She is on the editorial board of the journal Studies in Language.

David Lightfoot writes on syntactic theory, language acquisition, and historicalchange He argues that internal language change is contingent, taking place inbursts, and that this entails construing language acquisition as ‘cue-based’ He haspublished eleven books, most recently How new languages emerge (CUP, 2006) He

is a fellow of the American Association for the Advancement of Science and of theLinguistic Society of America His main appointments have been at McGill,Utrecht, Maryland, and Georgetown He served as Assistant Director of theNational Science Foundation, returning to Georgetown to direct programmes inCommunication, Culture & Technology and Interdisciplinary Cognitive Science.John L Locke is Professor of Linguistics, Lehman College, City University of NewYork He completed his PhD at Ohio University and took postdoctoral training atYale University and at Oxford His most recent books are Eavesdropping: Anintimate history (OUP, 2010) and Duels and duets: Why men and women talk sodifferently (CUP, 2011)

Ann MacLarnon is Director of the Centre for Research in Evolutionary pology at Roehampton University She has worked on a wide variety of areas inprimatology and palaeoanthropology, with an emphasis on comparative ap-proaches Research topics include reproductive life histories and physiology, stressendocrinology and behaviour, and aspects of comparative morphology includingthe brain and spinal cord Work on this last area led to the unexpected discoverythat humans evolved increased breathing control for speech

Anthro-Peter F MacNeilage has written over 120 papers and one monograph (The origin ofspeech, OUP, 2008) on the nature and evolution of complex action systems Hisconceptual contributions include the Frame/Content theory of the evolution ofspeech and the Postural Origins theory of the evolution of primate handedness He

is a Fellow of the Acoustical Society of America and the American Association forthe Advancement of Science

Alan Mann (PhD University of California, Berkeley) is Professor of Anthropology

at Princeton University and at the University of Pennsylvania (Emeritus) Hisprimary interest is the fossil evidence for human evolution and in particular theorigins of humanness His current research focuses on the evolution of the Nean-derthals and their relationships to modern peoples He is also co-director of theexcavation of a Middle Palaeolithic Neanderthal site in the Charente Department

of southwest France

Steven Mithen is a Pro-Vice-Chancellor at the University of Reading His researchinterests concern early prehistoric communities and the evolution of humanintelligence, language, and music His authored and edited books include After

the ice (Weidenfeld & Nicolson, 2003), The singing Neanderthals (Weidenfeld &Nicolson, 2005), The early prehistory of Wadi Faynan (Oxbow Books, 2007), To theislands (Two Ravens Press, 2010), and Water, life and civilisation (CUP, 2011) Hewas elected a Fellow of the British Academy in 2003.

Johanna Nichols is Professor Emerita in the Department of Slavic Languages andLiteratures and Affiliate Professor Emerita in the Department of Linguistics,University of California, Berkeley She works on Slavic and other languages of thewestern steppe periphery, languages of the Caucasus, typology, and languagespreads She is the author of Linguistic diversity in space and time (University ofChicago Press, 1992), Ingush-English dictionary (RoutledgeCurzon, 2004), andIngush grammar (University of California Press, 2011)

Brigitte Pakendorf obtained a PhD degree in molecular anthropology in 2001 and

a PhD degree in linguistics in 2007 She is currently leader of a Max PlanckResearch Group at the MPI for Evolutionary Anthropology in Leipzig, Germany.Her research focus is on the interdisciplinary investigation of prehistoric popula-tion and language contact

Irene M Pepperberg (S.B, MIT, 1969; PhD, Harvard, 1976) is a Research Associateand Lecturer (Harvard) and an Adjunct Associate Professor (Brandeis) She was avisiting Assistant Professor (Northwestern), tenured Associate Professor (Univer-sity of Arizona), and visiting Associate Professor (MIT’s Media Lab, Ecole NormaleSuperieure) She has received John Simon Guggenheim, Selby, and RadcliffeFellowships, is a Fellow of AAAS and other professional societies, has publishedover 50 peer-reviewed papers, 60 reviews, and two books

Amy S Pollick received her PhD in neuroscience and animal behaviour fromEmory University, where she conducted research on a variety of communicativebehaviours in non-human primates She was Director of Government Relations atthe Association for Psychological Science in Washington, DC, and is now anadjunct professor at Gallaudet University

Camilla Power completed her PhD under Leslie Aiello at the University ofLondon She is currently Senior Lecturer in Anthropology at the University of EastLondon, specializing in Darwinian models for the origins of ritual and religion, andAfrican hunter-gatherer gender ritual, having worked in the field with women of theHadzabe in Tanzania

Paul T Roberge is Professor of Germanic Languages and Joint Professor ofLinguistics at the University of North Carolina at Chapel Hill He is also Extraor-dinary Professor of General Linguistics at the University of Stellenbosch Hisresearch interests are historical Germanic linguistics, sociohistorical linguistics,pidgins and creoles, and Afrikaans He teaches language evolution at UNC-CH

in collaboration with his colleague, Elliott Moreton, and has written on the windowpotential of pidgins.

Robert M Seyfarth graduated from Harvard University and received his PhDfrom the University of Cambridge, where his advisor was Robert A Hinde Hewas a post-doctoral fellow at Rockefeller University, working with Peter Marler.Together with Dorothy Cheney, he is the author of How monkeys see the world(University of Chicago Press, 1990) and Baboon metaphysics (Chicago, 2007) He iscurrently Professor of Psychology at the University of Pennsylvania

Peter Slater is Emeritus Professor of Natural History at the University of

St Andrews Much of his work over the past 30 years has been on sound nication in birds and mammals and he is co-author, with Clive Catchpole, of thebook Bird song: Biological themes and variations (CUP, 2nd edition 2008)

commu-Katie Slocombe obtained a BSc in psychology from the University of Nottingham,and completed her PhD in chimpanzee communication at the University of

St Andrews under the supervision of Klaus Zuberbu¨hler As a BBSRC post-doctoralresearch fellow she continued to investigate vocal behaviour in wild and captivechimpanzees She became a lecturer in the department of Psychology, University ofYork, in 2007, where her research into chimpanzee communication and cognitioncontinues

Kenny Smith is a lecturer in the School of Philosophy, Psychology and LanguageSciences, University of Edinburgh, with interests in the evolution of communica-tion, human language, and the human capacity for language He uses a mix ofmodelling and experimental techniques to address these questions

Michael Studdert-Kennedy holds a BA in classics from the University of bridge and a PhD in experimental psychology from Columbia University He is aformer President of Haskins Laboratories in New Haven His research interestsinclude speech perception, hemispheric specialization for speech, and the develop-ment/evolution of speech He is the author of many papers and the editor or co-editor of six books

Cam-Szabolcs Sza´mado´ received his PhD in ecology and theoretical biology at theEo¨tvo¨s Lora´nd University (Hungary) in 2004 His main research topic is the cost

of honesty in animal communication, which he investigates by means of gametheoretical modelling He works with Eo¨rs Szathma´ry on questions surroundingthe evolution of human language

Eo¨rs Szathma´ry is the co-author with the late John Maynard Smith of The majortransitions in evolution (Freeman, 1995) and The origins of life (OUP, 1999) Hismain interests include evolution in the origin of life and Darwinian approaches to

brain dynamics He is at Collegium Budapest and the Parmenides Foundation(Munich).

Maggie Tallerman has spent her professional life in northeast England, at Durhamthen Newcastle University, where she is currently Professor of Linguistics Heredited and authored books include Language origins: Perspectives on evolution(OUP, 2005), Understanding syntax (Hodder/OUP, third edition 2011), and Thesyntax of Welsh (co-authored with Borsley and Willis; CUP, 2007) She startedworking on evolutionary linguistics in case a guy on a train asked her wherelanguage came from, though some think her real work is on Welsh

Marian Vanhaeren is a Researcher at the French CNRS, specializing in the study ofancient personal ornaments which she investigates using technological and tapho-nomical analyses, reference collections, microscopy, GIS, and statistics Her researchfocuses on the oldest traces of symbolic thinking in the European and AfricanPalaeolithic as well as on ethnocultural diversity, exchange networks, and socialinequalities among Upper Palaeolithic populations She has co-authored more than

30articles

Jacques Vauclair is a Professor of Developmental and Comparative Psychology atthe University of Provence (Aix-en-Provence, France) and a senior member of theInstitut Universitaire de France He is director of the Research Center in thePsychology of Cognition, Language and Emotion in Aix-en-Provence His field

of interest concerns the comparative study of the lateralization processes in objectmanipulations and communicative gestures in human infants and in non-humanprimates

Wendy K Wilkins is currently the Executive Vice President and Provost at NewMexico State University Her PhD in linguistics is from UCLA She has taughtlinguistics at UMass Amherst, the University of Washington, Arizona State Uni-versity, Michigan State University, and at several institutions in Mexico City With

a primary research background in syntactic theory, most recently she has worked

on the evolutionary neurobiology of language and comparative linguistic andmusical cognition

Bernard A Wood is the University Professor of Human Origins at the GeorgeWashington University and Adjunct Senior Scientist at the National Museum ofNatural History, the Smithsonian Institution His research centres on increasingour understanding of human evolutionary history by developing and improvingthe ways we analyse the hominin fossil record

Thomas Wynn is Professor of Anthropology at the University of Colorado, ado Springs, where he has taught since 1977 He has published extensively inpalaeolithic archaeology, with a particular emphasis on cognitive evolution Hisbooks include The evolution of spatial competence (University of Illinois Press, 1989)

Color-and The rise of Homo sapiens: The evolution of modern thinking (with F Coolidge,Wiley-Blackwell, 2009).

Klaus Zuberbu¨hler is a Professor of Psychology at the University of St Andrews

He holds an undergraduate degree in zoology and anthropology from the sity of Zurich and a doctoral degree in psychology from the University of Penn-sylvania He conducts research on the primate roots of human language, focusing

Univer-on the cognitive underpinnings of primate communicatiUniver-on His research has beenpublished in major journals, including Nature, PNAS, and Current Biology

as touching with legs, trunks, or feelers Communication via chemical signals iswidespread; for example, moths use pheromones to attract conspecifics in the dark.

Electric organ discharge (for instance, in electric fish such as mormyrids) alsooccurs, but is much rarer Numerous social insect species have highly sophisticatedcommunication systems, such as the ‘dance’ used by honey bees, where complexmovements produced by a returning forager indicate the distance and direction ofpollen and nectar resources Animal communication systems are thus immenselyvaried in form.

Some animals communicate not only with conspecifics, but also understand atleast some of the signals produced by other species; for example, Diana monkeyslearn the message conveyed by the alarm calls of neighbouring Campbell’s monkeys(Zuberbu¨hler, Chapter 5) Moreover, symbiotic relationships sometimes producecommunication across unrelated species The honey guide bird (Indicator indica-tor) leads honey badgers to bees’ nests by making a sound that attracts the badger,which then breaks into the nest, allowing both animals to reap the rewards.Mammals employ extensive vocal communication with conspecifics, often inaddition to using visual display, chemical messages, and tactile communication.Humans are no exception here, and our ancient primate communication systemencompasses many non-verbal signals, including laughter, crying, smiles, frowns,cries of pain, and postures of aggression and appeasement (Burling, Chapter 44).With a very small amount of cultural diversity, these signals are human universals.But all animal communication systems, including our own non-verbal signalling,share a (negatively-defined) property: these systems cannot combine signals toproduce new meanings In fact, they generally do not combine signals at all Forlanguage, though, this property is fundamental The combinatorial principle,exploited at different levels of organization, is a crucial, distinctive attribute oflanguage The overarching property, shared by no other animal system, is the open-ended productivity of language: humans combine signals to produce an infinite set

of distinct meanings, and can convey to conspecifics any topic that can be thought

of, including absent, hypothetical, and fictitious events and entities

In every meaningful sense, language is an autapomorphy, i.e a derived traitfound only in our lineage, and not shared with other branches of our monophyleticgroup (say, the group of primates, or the group of great apes) We also have nodefinitive evidence that any species other than Homo sapiens ever had language.However, it must be noted straightaway that ‘language’ is not a monolithic entity,but rather a complex bundle of traits that must have evolved over a significant timeframe, some features doubtless appearing in species that preceded our own.Moreover, language crucially draws on aspects of cognition that are long estab-lished in the primate lineage, such as memory: the language faculty as a wholecomprises more than just the uniquely linguistic features We do not know, though,

if and how language has itself shaped properties such as memory (or vice versa), sothe role of extra-linguistic factors is hard to evaluate

We anticipate that both animal communication and animal cognition will shedlight on the evolution of language, but in exactly what ways is hotly debated

Biologists do not expect evolution to throw up a radically new complex systemwith no evolutionary precursors, so it is good scientific practice to look for relevantprimitive features in closely-related species, i.e phylogenetically ancient featuresthat may have preceded language or been prerequisites for language; see Hopkinsand Vauclair, Chapter 18; Wilkins, Chapter 19 Another established methodology is

to search for examples of convergent evolution: functionally or formally similarfeatures appearing in species that do not share a recent common ancestry Unfor-tunately, for language such features are not easily detected For specific traits, thereare indeed both analogues (unrelated but superficially similar features) and homo-logues (features with a shared common ancestry) in other animal systems Theseinclude such common features as vocalization and cultural transmission But itseems clear that language as a system has no ‘simpler’ analogues or homologues inother animals In fact, language is exceptional in almost all aspects

Language obtains its unique expressive power by exploiting a few distinct formalprinciples that operate over numerous subsystems and at different levels of orga-nization These tools have little or no parallel in the animal kingdom First, andperhaps most critically, language combines elements at all levels Starting withsound systems (MacNeilage, Chapter 46), each language combines elements from

an individual set of digitized sounds known as phonemes: discrete, contrastivesound segments In turn, phonemes are often considered to be combinations of adiscrete set of phonological features, such as [þ/ voice] and [þ/ sonorant](though see MacNeilage, Chapter 46, for discussion) Phonemes combine inlanguage-specific permutations to form syllables, and syllables combine to formmorphemes and words; sign languages have equivalent manual systems (Goldin-Meadow, Chapter 57) Words are combined, both in morphology, to form com-pounds (greenhouse, dog-house, icehouse, outhouse), and in syntax, to form headedphrases

Second, elements are ordered in predictable ways Each language has its own set

of phonotactic constraints, governing which sound sequences are permissible andwhich are not; English, for instance, allows word-initial sequences of /pl/ (play) and/kl/ (clay) but not */tl/ Morphemes are ordered in fixed ways (unþafraid, butfearþless; handþfulþs, not *handsful); see Carstairs-McCarthy, Chapter 47, onmorphology Words are also sequenced predictably, for instance by having ausual order of heads and complements across phrasal categories, though languagesare not always dogmatic about this

Third, language exploits hierarchical structure at several levels Syllables arehierarchically structured, with a nucleus and coda combining to form a rhyme(say, vocalic nucleus /i/ plus coda /Nk/ forming ink), and then an onset, say dr or sl,combining optionally with the rhyme, forming drink or slink Hierarchical struc-ture also operates at the level of morpheme combinations: given the morphemes

unþfriendþly we get the structure [un[friendly]] rather than *[[unfriend]ly],whereas with untidily (unþtidyþly) the structure is [[untidy]ly] Semantics

exploits hierarchical structure, for instance by partitioning meanings into differentlevels of specificity: spaniel, setter, retriever are types of dog, dogs are a type ofmammal, mammals a type of vertebrate and so on Syntax exploits hierarchicalstructure by combining words into phrases, and phrases into larger phrases andclauses (Tallerman, Chapter 48) It is also widely argued that texts in discourse (e.g.conversations or stories) are hierarchically structured In addition to these organi-zational principles, mappings occur between all linguistic levels, including mostbroadly between sound and meaning This, then, is the formal basis of language.Even at the lowest levels of organization, there are strikingly few parallels inanimal communication systems In human phonological systems, a relativelysmall, closed set of meaningless elements (sound segments and their visual equiva-lents in sign languages) combine to produce meaningful elements Though birdsong is sometimes described as having ‘phonological syntax’ (Marler 1977), theterm is rather misleading In bird song, discrete acoustic units also exist, and arecombined and sequenced in rule-governed ways, but there is no compositionality;whatever the sequences of notes or motifs, the message never changes and no newinformation is produced (Hilliard and White 2009; Slater, Chapter 8) There is noproductivity in the combinations Bee dances display a limited compositionality(Kirby, Chapter 61) but again, no productivity As we move up the levels oflinguistic organization, we find fewer parallels still in animal systems Hierarchicalstructure exists in some bird song and some whale song (Janik, Chapter 9), but it isalways limited (Hurford 2011) There is no recursion (self-embedding) and nosemantic compositionality.

A priori, we might expect that the natural communication systems of our closestliving relatives, the great apes, would be nearest to language—perhaps rather likelanguage, but with a smaller vocabulary and a simpler grammar But this isabsolutely not the case Even at the most fundamental level, that of sound produc-tion, we have a different morphology of the supralaryngeal vocal tract from that ofchimpanzees, with humans showing clear specializations for speech production(MacLarnon, Chapter 22) Moreover, humans have evolved far greater neurologicalcontrol over their vocalizations than other primates Although much is stillunknown about the subtleties of communication systems in other primates, it isclear that there is really nothing analogous to human sound systems, lexicon,semantics, or grammar Some call combinations do seem to occur in wild chim-panzees (Slocombe, Chapter 7) but as yet there is no evidence that these acquire acompositional meaning; see Tallerman, Chapters 48 and 51

Certain animal systems have something that at first glance seems to resemble aprimitive vocabulary For instance, among other calls, some monkey species have asmall set of distinct alarm calls, each produced in response to a different predator(such as the well-known vervet monkey calls, ‘eagle’, ‘leopard’, and ‘snake’; Seyfarthand Cheney, Chapter 4) These have attracted much interest in the languageevolution literature, doubtless because of the relatively close relationship between

humans and monkeys It should be noted, though, that domestic chickens alsohave distinct alarm calls for different predators, a system as sophisticated as those

of monkeys, and, additionally, have referential food calls; moreover, prairie dogs(which are rodents) employ perhaps the most highly sophisticated systems ofanimal alarm calls (Gibson, Chapter 11) Are alarm calls or food calls parallel towords? They share one property—arbitrariness—with human vocabulary: they arenon-iconic, meaning that they do not sound like the entities they represent; seeDeacon, Chapter 43 Alarm calls are often described as having functional reference;that is, they are prompted by external events (such as the appearance of a leopard)rather than merely conveying an animal’s internal state, such as fear or aggression

We might then assume that the leopard alarm in some way ‘means’ leopard Butthis is not necessarily so—the leopard call may alternatively be associated in thehearer’s mind with the leopard-specific escape route, climbing a tree; see Hurford,Chapter 40; Tallerman, Chapter 51, for discussion

Alarm calls differ from words in all other respects; see Tallerman, Chapter 48.They are not formed from different permutations of a discrete set of sounds, butrather, are holistic Both the calls themselves and the broad contexts that provokethem are innate Conversely, words, both forms and meanings, are learned byhuman infants, and crucially, new words are learned by each speaker throughoutlife Monkey alarm calls are primarily used when the particular predator is present,and sometimes to deceive conspecifics into thinking a predator is around Even inthe latter case, calls are wholly situation-specific; calls cannot be used predicatively

to discuss a predator, past, present, or hypothetical Alarm calls are indexical,meaning that they have a causal link to what they represent—normally, thepresence of the predator induces the appropriate alarm call Alarm calls thus alsolack the property of displacement that is crucial in language: words are not tied to

or produced in a precise context, but can be used whenever the concept theyrepresent floats into our minds

Words are thus true symbols, whereas animal calls, even if functionally tial, are not; symbolic reference, which must be acquired by learning, is explored indetail by Deacon, Chapter 43, and by Harnad, Chapter 42 Critically, word mean-ings are established between a community of speakers and agreed by convention.Part of what this entails is that the meaning of a word can change very quickly,providing other members of the language community adopt the new meaning(think of net, web, or drive) Alarm calls, in contrast, have a fixed meaning andessentially form a closed set The total repertoire of calls in any animal species istiny, numbering no more than a few dozen distinct calls, whereas the vocabulary ofall human languages numbers tens of thousands of items (Tallerman, Chapter 48).The gulf between the small set of essentially innate and relatively inflexible callsfound in all other primate species and the massive, open-ended, and learnedvocabularies of human beings reveals one of the major innovations in languageevolution that must be accounted for This is no mere matter of degree—in

referen-vocabulary, something altogether different has evolved from anything seen in thenatural communication systems of other animals (Burling, Chapter 44).

The evolution of a massive, learned vocabulary store (Tallerman 2009) is just one

of the unique aspects of language Only in language do we find the extensivecategorization that, for instance, divides the lexicon into discrete categories such

as noun, verb, adjective, each category with its own distinctive behaviour Thecategories themselves are unlikely to be innate, since they differ from language tolanguage, but the ability to categorize in this way, and on the basis of little data,appears to be uniquely human Evidence of children’s abilities in generalizing overcategories has been well known at least since Berko (1958) In the wug test, childrenare told that a mythical creature in a picture is a wug; when asked what two suchcreatures are called, they have no difficulty in replying /wVgz/, thus using thecorrect plural form of the noun

Only language conveys propositional meanings (‘idea units’; loosely, what wecall sentences, such as The bird flew past the window), and of course these areunlimited in scope Only language exhibits all the paraphernalia of syntax (Taller-man, Chapter 48), including headed phrases, recursion, long-distance dependen-cies, and expressions such as each other or themselves that can only be interpretedusing other expressions (Kim and Mel hurt each other) Only language displays theproperty of duality of patterning (see Tallerman, Chapter 51), with combinations

on two levels of organization: meaningless units (phonemes) are combined intomeaningful morphemes/words, and words are combined into phrases Otherhighly distinctive properties arise on each level of linguistic organization; eventhe speech signal itself displays significant adaptations both in production andperception (see Pinker and Jackendoff 2005 for an overview)

Given the limited nature of the evidence obtainable from studying animalcommunication systems, how do researchers hope to break into the evolutionarypuzzle that is language?

1.2 WH AT C O U N T S A S E V I D E N C E I N

L A N G UAG E EVO LU T I O N?

The previous section introduced the major novelties of the language faculty, whichincludes notable discontinuities with animal communication systems This givesrise to a fundamental dilemma in the field of language evolution Language seems

to display many features with no precursors, yet general evolutionary principlessuggest that a complex trait like language, which is not under the control of anysingle gene or related group of genes, must have evolved in large part from simpler

precursors Frustratingly, we have no direct evidence for any aspect of languageevolution, and no uncontroversial indirect evidence Moreover, what is consideredpossible evidence differs from discipline to discipline, as we now discuss.

The comparative method is an obvious place to start; see chapters in Part I, alsoFitch (2010a) There are two ways in which this method can be employed The firstinvolves comparing similar traits within a clade For humans, the set of primates as

a whole or the smaller set of great apes would be most relevant; for instance, tooluse by other great apes is an established trait, so it seems likely that the lastcommon ancestor of all great apes, including humans, was able to use simpletools This trait is thus a homologue, involving a shared common ancestry (SeeWood and Bauernfeind, Chapter 25, for discussion of likely features of the lastcommon ancestor between panins, i.e chimpanzees/bonobos, and hominins, i.e.creatures that are on the human line of descent, though not necessarily directancestors to Homo sapiens.) Relevant here too are the natural communicationsystems of closely-related species, and also their latent language-related abilities,such as the ability to learn arbitrary symbols under human instruction (Gibson,Chapter 3) The alternative way of employing the comparative method involvescomparing the convergent evolution of similar traits across a number of unrelatedlineages For instance, bipedal locomotion in humans, kangaroos, and birds is notdue to common ancestry, so is an analogous trait across the three lineages.Analogues are useful because they may have evolved in different lineages undercomparable selection pressures, such as a similar habitat, diet, or predation pattern.The problem, as noted above, is that homologues and analogues to essentialproperties of language are not easily established in animal systems, and no otherspecies has a language faculty, so the comparative method is difficult to applystraightforwardly For discussion in this volume, see especially Arbib (20); de Waaland Pollick (6); Gibson (3 and 11); Hopkins and Vauclair (18); Hurford (40);Pepperberg (10); Slocombe (7); Tallerman (48); Zuberbu¨hler (5)

The discipline of palaeoanthropology examines the fossil record, and from skullendocasts may uncover anatomical evidence of brain structure of potential rele-vance to language, including brain size, external cortical reorganization, and hemi-spheric asymmetries (Wilkins, Chapter 19) Unfortunately, we cannot study paststages of brain evolution in any depth, since endocasts provide no evidence ofinternal brain structure Similarly, with the exception of an occasional hyoid bone,

we have no fossilized remains of the vocal tract (see MacLarnon, Chapter 22; Woodand Bauernfeind, Chapter 25) However, even if we had clear evidence of theemergence of modern vocal tract structure, we would not necessarily know how

to interpret it A broadly modern structure, for instance, could initially haveevolved as a spandrel, that is, as a by-product, perhaps of bipedalism or changes

in dental function (MacLarnon, Chapter 22) In that case, speech capabilities couldstill have been lacking if neural adaptations had not yet occurred Even if we werecertain that a modern vocal tract provided full speech capabilities, this would not

necessarily imply the presence of a full language faculty, since this encompasses farmore than speech.

Recently, molecular biology has provided another possible source of physicalevidence: the use of genetic material from hominin fossils, and the tracking ofchanges in DNA in hominin populations (Cann, Chapter 24; Pakendorf, Chapter

59) Again, however, these methods are fraught with difficulties and controversies.For instance, which class of DNA (mitochondrial, Y-chromosome, or autosomal)should be considered the most reliable? This is as yet a young field, and newdiscoveries and constant developments in technology should provide more answers

in future decades

Another line of enquiry looks not at hominin fossil remains themselves, but atthe artefacts left by our ancestors Archaeologists have argued that inferences can bemade about the development of symbolic communication and linguistic complex-ity by looking at tools and other implements, or personal ornaments such as beads,thus assuming some link between linguistic skills and cognitive sophistication, asevidenced in the material record Moreover, if a certain level of cultural complexity

is attested both in known societies and in prehistoric societies, it seems reasonable

to assume that a similar level of complexity occurs in cognition too In this volume,the chapters by Boeckx (52); Botha (30); d’Errico and Vanhaeren (29); Donald (17);Mann (26); Mithen (28); and Wynn (27) discuss the relevance of the archaeologicalrecord and the difficulties inherent in interpreting it; see also Cann, Chapter 24.There are many possible drawbacks to using technological advances to infer thepresence of the language faculty, not least because crucial artefacts made ofdegradable materials may be absent from the record: for instance, early homininsmay have utilized plant materials, including bark, leaves, wood, grass, and reeds,and animal soft parts, including hides, furs, and feathers We only have to think ofthe exponential increase in the complexity of our own artefacts between 1850 and

2010to realize that there is no simple chain of inference between sophistication inthe archaeological record and the presence of language (see also Botha, Chapter

30) Moreover, new archaeological findings readily overturn previous conclusions.For example, it was once thought that art, beads, and some forms of stone-flakingappeared only in Upper Palaeolithic times, i.e after about 35,000 BP We now knowthat beads, other putative forms of symbolism, and advanced flaking techniqueslong predate the Upper Palaeolithic (Brown et al 2009; d’Errico and Vanhaeren,Chapter 29; McBrearty and Brooks 2000) And until quite recently, tools compris-ing more than one component—such as harpoons or bows and arrows—werethought to have originated only within the last 20,000 years (Coolidge and Wynn

2009b; Wynn, Chapter 27) but a recent find suggests that arrows were beingproduced as much as 64 kya (thousand years ago) (Lombard and Phillipson

2010) Thus, the archaeological record requires us to frequently re-evaluate dence and arguments

evi-Given the problems with the physical record, both of hominin fossils andassociated artefacts, many have turned to other potential sources of evidence forlanguage evolution Jackendoff (2010: 65) suggests that reverse engineering pro-vides the most productive methodology: ‘We attempt to infer the nature ofuniversal grammar and the language acquisition device from the structure of themodern language capacity’ Thus, one form of evidence comes from the study ofnormal language and its acquisition; in this volume, see in particular the contribu-tions by Bickerton (49); Burling (44); de Boer (33); Falk (32); Goldin-Meadow (57);Graf Estes (64); Locke (34); MacNeilage (46); Studdert-Kennedy (45); Tallerman(48 and 51) However, the danger of confusing ontogenetic and phylogeneticprocesses must always be guarded against here There is no reason to think thatany specific evidence concerning the origins of language can be gained fromstudying the acquisition of modern languages Moreover, infants learning languagetoday have a full language faculty, which clearly is not the case for the earliesthominins.

Linguists often suggest that evidence can be obtained by extrapolating fromother modern contexts; for instance, from observable ‘language genesis’ in adultsand children, including the formation of pidgins and creoles (in this volume, seeBickerton, Chapter 49; Carstairs-McCarthy, Chapter 47; Roberge, Chapter 56),homesign, and emergent sign languages (Goldin-Meadow, Chapter 57) A verystrong line of linguistic research (and one of the few areas widely considered toprovide good evidence by practitioners of disparate linguistic theories) involves thestudy of grammaticalization It is widely argued that putative prehistoric stages oflanguage can be reconstructed by studying known linguistic trajectories ofchange—specifically, the ways in which grammatical elements are formed fromlexical elements In this volume, see especially the contributions of Bybee (55);Carstairs-McCarthy (47); and Heine and Kuteva (54) The importance of gramma-ticalization is also emphasized by Bickerton, Chapter 49; Corballis, Chapter 41; andChater and Christiansen, Chapter 65

There are also, however, applications of reverse engineering in spheres otherthan the narrowly linguistic In the field of cognition, Coolidge and Wynn (Chapter

21) investigate the evolution of modern thinking, specifically the emergence ofindirect speech acts There are also applications of reverse engineering in evolu-tionary biology; in this volume, see in particular Sza´mado´ and Szathma´ry, Chapter

14, who discuss the co-evolution of language and the brain Since there must besome relationship between genes and language, it has sometimes been assumedthat there are readily identifiable genes ‘for’ language, or aspects of language (forexample, FOXP2 was often carelessly reported in the popular press as the ‘languagegene’) Diller and Cann, Chapter 15, evaluate the evidence concerning geneticcorrelates for language, and conclude that language is highly unlikely to beassociated with any single genetic mutation; see also Cann, Chapter 24

Two related, relatively new applications of reverse engineering have becomemajor fields of investigation in language evolution: computational and mathemat-ical modelling, and robotics, including embodied agent models Formal modelsallow the predictions of theories of language evolution to be tested empirically, bybuilding in the assumptions to be tested and seeing if they indeed result from themodel: if the model produces these results, the assumptions are borne out In thisvolume, see especially Cangelosi (62); Kirby (61); and Smith (60); also de Boer (63);MacNeilage (46); Studdert-Kennedy (45) As both Kirby and Smith discuss, resultsand predictions obtained from the formal models can further be tested on humansubjects in the laboratory.

In sum, though there will inevitably be much speculation in a field of this nature,

we believe that serious advances have been made in the past few decades in terms ofbuilding an evidence-based discipline In the next section we consider in moredetail the properties of language as a biological system

1.3 LA N G UAG E E VO LU T I O N A N D B I O LO G Y

We start by examining the uniqueness of language in biological terms, in son with other animal communication systems Language is a complex amalgam oflifelong learning (nonetheless including a critical period) and innateness; see Fitch,Chapter 13 Most researchers agree that both aspects are crucial to language, butmany controversies arise over where the line should be drawn (see the followingsection) The aspects uncontroversially considered to be learned are, of course,vocabularies and idiosyncratic lexical properties of distinct languages, transmittedfrom generation to generation (a trait known as traditional transmission) Vocab-ulary is added beyond the critical period for language acquisition, a feature withfew clear analogues in other animal communication systems

compari-Simple communication systems which combine vocal learning and innatenessare found in some animals (notably, songbirds), but the contributions made byeach aspect are easier to tease apart, since experiments can be performed whichwould be impossible with human subjects Here we see a marked contrast with thecommunication systems of non-human primates, in which learning plays a mini-mal part—it is more a case of fine-tuning the acoustic properties of calls, and oflearning the specific contexts in which it is appropriate to use each call Vocallearning does play a vital role in the communication systems of some non-primates, however, especially in bird species (Slater, Chapter 8), in a number ofmarine mammals (whales, dolphins, sea lions etc.; Janik, Chapter 9), and in somebats Among vocal learning birds, there are certain parallels to language learning:

song learning can involve a sensitive period, outside of which the learned systemwill either be incomplete or abnormal; song is traditionally transmitted (i.e.learned from adult models); and a stage analogous to babbling (known as sub-song) occurs However, Gibson (Chapter 11) points out that language learning isactually very different from the learning of bird song, which often has a sensitiveperiod followed by a long ‘quiet’ period, with song emerging only in adulthood; seeHurford (2011) Moreover, young birds raised without an appropriate adult model(e.g solely female instead of male relatives) will nonetheless sing, even though thesong is not adult-like This shows that there is an innate stratum, some basis for thesong which is not entirely learned.

In the case of language, the child undoubtedly brings crucial cognitive tions to the learning process, yet without linguistic input, full language does notdevelop Despite the necessity for learning, the language faculty (whatever itcontains) provides such a powerful drive that in the absence of normal linguisticinput, something language-like can emerge A clear example comes from the deafchildren of hearing parents: from the non-linguistic gestures that the parents make

contribu-to communicate with the child, a structured system—known as ops spontaneously in the child’s communication (see Goldin-Meadow, Chapter 57).This is not language, but has indisputable linguistic properties And tellingly,when speakers of different homesign systems get together in a naturalistic setting,

homesign—devel-a shhomesign—devel-ared system with more linguistic properties soon emerges, homesign—devel-as in the well-knowncase of Nicaraguan Sign Language, also discussed by Goldin-Meadow Over thecourse of a couple of ‘generations’ of schoolchildren, this sign system developedinto full language It is also well documented throughout the world that whencontact occurs between groups with no shared language, restricted linguisticsystems develop, known as pidgins (Roberge, Chapter 56), and these may in duecourse become full languages, learned natively by children Given such evidence, it isdifficult to conclude that language has no genetic component We will assume, then,that there have been significant adaptations in our species with respect to a languagefaculty

Strikingly, though, and unlike animal communication systems, language differsradically in its superficial form in distinct geographical locations—we have mutu-ally unintelligible ‘languages’ in different regions, rather than distinct ‘dialects’, as isthe case in some bird song and whale song systems The superficial diversity oflanguage systems has no discernible consequences for language learning; infantsseem equally capable of learning any ambient language (or indeed, learning half adozen or more languages in their environment), and take around the same amount

of time to get to the same stages, whatever language they are learning This factalone suggests the presence of an innate predisposition for language learning.Another biologically distinctive property of language concerns its function (seealso below) If indeed language has biological ‘function’ at all, it is difficult todiscern what the primary function might be, or might have been while language

was evolving The function of animal communication systems, on the other hand,typically revolves around reproduction, including mate attraction, pair bonding,and defence of territory Even learned animal systems thus have a very limitedmessage There is a biological imperative for songbirds to learn their songs: birdsthat don’t learn the species-appropriate song are less likely to be chosen as mates,and song quality is perceived by potential mates as a fitness indicator (Slater,Chapter 8) Song is, thus, an honest signal (Zahavi and Zahavi 1997), in a waythat language is not Song is ‘costly’, as it takes time to produce, and so reveals thequality of the singer: only a bird that is already in good condition can afford tospend time singing rather than feeding; song may also draw the attention ofpredators Conversely, producing language requires virtually no calorific expendi-ture above and beyond that needed for overall brain growth and maintenance; itdoesn’t take up valuable time that could be used to forage and it doesn’t requirethat the speaker be in good condition One of the questions surrounding theorigins of language is therefore how a ‘cheap’ communication system of this naturemight have arisen; for discussion in this volume, see especially Donald, Chapter 17;Dunbar, Chapter 36; Falk, Chapter 32; Knight and Power, Chapter 37.

It is also notable that language involves developments on three distinct butinteracting timescales (Kirby, Chapter 61; Carstairs-McCarthy, Chapter 47; Sza´-mado´ and Szathma´ry, Chapter 14) The first of these is biological evolution:whatever is in the language faculty (and its precursors in early hominins) must

be genetically transmitted under selective pressures—the transition in homininsfrom no language to language is a biological fact, so must conform to knownbiological processes Language in its earlier forms can therefore be assumed to havebeen adaptive, i.e to have conferred fitness benefits on its users At the very least,whatever neurological, physical, or other changes accompanied an evolving lan-guage faculty had to have no negative impact on selection

Biological change thus encompasses all the phylogenetic changes in homininswhich are prerequisites for language, including the evolution of crucial abilities notshared with other primates, or at best, only minimally developed in non-humanprimates For the speech modality, these prerequisites include full vocal control(the ability both to vocalize and to suppress vocalization at will), vocal imitation,and vocal learning; see MacLarnon, Chapter 22; MacNeilage, Chapter 46 A certainamount of vocal flexibility is in fact attested in modern primates (see Slocombe,Chapter 7; Zuberbu¨hler, Chapter 5), taking the form of acoustic modification ofcalls It is now also known that some primates can both vocalize volitionally andsuppress vocalization under certain circumstances Slocombe also reports on someevidence showing that both vocal imitation and learning occur, for instance indifferent ‘dialects’ of vocalizations in chimpanzees; if there are novel vocalizations,they must be learned and must spread via imitation But even before those traitsemerged, our ancestors must have developed the ability to understand that con-specifics are communicating deliberately; to infer the mental states of other

individuals (see Gibson, Chapter 3; Hurford, Chapter 40; Knight and Power,Chapter 37); and to engage cooperatively in all sorts of tasks, which eventuallyincluded discourse (Tomasello 2008) These traits are all expressed to some extent

in modern apes, so probably existed in the last common ancestor of apes andhumans Language placed a premium on these abilities For a protolanguage toemerge, hominins needed to develop expanded abilities in such domains, leadingultimately to the ability to learn, store, and retrieve a vast intersecting network ofarbitrary symbols (words; Deacon, Chapter 43), and the crucial property ofdisplacement (the ability to refer to entities remote in time or space; Hurford,Chapter 40); see also Tallerman (2009)

Using these conventional symbols relies in turn on the capacity to imitate,rehearse, and refine the practical skills required (Burling, Chapter 44; Corballis,Chapter 41; Donald, Chapter 17) The use of vocabulary also relies on a sharedconceptual system, which probably developed directly from primate cognition(Hurford, Chapter 40), and requires ‘the ability to associate gestures or vocaliza-tions with concepts’ (Burling, Chapter 44; see also Corballis, Chapter 41): this must

be one of the critical first steps in language evolution For full language, more isneeded—the major development being the compositional syntactic abilities whichare the main impetus in generative grammar for assuming an innate languagecapacity; see Bickerton, Chapter 49; Tallerman, Chapter 48, for an outline ofsyntactic processes

The second timescale involves cultural transmission: individual languages aretransmitted across generations, and within populations of speakers This has led toproposals that languages themselves adapt to become more learnable (Christiansenand Chater 2008; Chater and Christiansen, Chapter 65) Many developments onthis timescale are known from attested language change, in particular the processesknown as grammaticalization, whereby lexical items evolve into functional items(auxiliaries, complementizers, demonstratives, determiners, and so on) Mostlinguists assume that similar processes were operative in the evolution of thefull language faculty, so that the earliest protolanguages—simpler precursors tolanguage—may well have distinguished no categories other than protonounsand protoverbs (Hurford 2003a; Heine and Kuteva 2007, Chapter 54; Tallerman,Chapter 51)

Cultural transmission involves not only vertical transmission, between parentsand children, but also horizontal transmission of various kinds, both within andacross communities This includes transmission between speakers of differentlanguages, in cases of language contact: see Pakendorf, Chapter 59 Such contactcan lead to interesting mismatches between the genetic and linguistic heritage in apopulation, as Pakendorf outlines Since population contact is likely to have beenextensive throughout our evolution, language contact between linguistic groupshas very likely contributed much to language evolution itself (Nichols, Chapter 58)