Syntactic rules in avian vocal sequences as a window into the evolution of compositionality

Understanding the origins and evolution of language remains a deep challenge, because its complexity and expressive power are unparalleled in the animal world. One of the key features of language is that the meaning of an expression is determined both by the meanings of its constituent parts and the syntactic rules used to combine them; known as the principle of compositionality. Although compositionality has been considered unique to language, recent field studies suggest that compositionality may have also evolved in vocal combinations in nonhuman animals. Here, we discuss how compositionality can be explored in animal communication systems and review recent evidence that birds use an ordering rule to generate compositional expressions composed of meaningful calls. Also, we suggest that birdsongs, particularly when incorporating calls, may represent unrecognized examples of compositionality in animal communication. Finally, we outline future research directions to uncover the development, neural mechanisms and evolution of compositionality.

[1]  Selvino R. de Kort,et al.  Blackcaps (Sylvia atricapilla) increase the whistle part of their song in response to simulated territorial intrusion , 2015 .

[2]  Thierry Aubin,et al.  Does true syntax or simple auditory object support the role of skylark song dialect? , 2013, Animal Behaviour.

[3]  Klaus Zuberbühler,et al.  Conceptual Semantics in a Nonhuman Primate , 1999 .

[4]  Andrea M.-K. Bierema,et al.  On the Meaning of Alarm Calls: A Review of Functional Reference in Avian Alarm Calling , 2013 .

[5]  W. Fitch.  The Evolution of Language , 2010 .

[6]  Timothy J. O'Donnell,et al.  Evolutionary Linguistics: A New Look at an Old Landscape , 2007 .

[7]  Dominique A. Potvin,et al.  Birds orient their heads appropriately in response to functionally referential alarm calls of heterospecifics , 2018, Animal Behaviour.

[8]  Sabrina Engesser,et al.  Meaningful call combinations and compositional processing in the southern pied babbler , 2016, Proceedings of the National Academy of Sciences.

[9]  Douglas G. Smith,et al.  An Investigation of Song-Based Species Recognition in the Red-Winged Blackbird (Agelaius Phoeniceus) , 1980 .

[10]  Johan J Bolhuis,et al.  Meaningful syntactic structure in songbird vocalizations? , 2018, PLoS biology.

[11]  David Wheatcroft,et al.  Call combinations in birds and the evolution of compositional syntax , 2018, PLoS biology.

[12]  David Wheatcroft,et al.  Wild Birds Use anOrdering Rule to DecodeNovel Call Sequences Graphical Abstract Highlights , 2017 .

[13]  Marisa Hoeschele,et al.  Neural Correlates of Threat Perception: Neural Equivalence of Conspecific and Heterospecific Mobbing Calls Is Learned , 2011, PloS one.

[14]  Stephen Pruett-Jones,et al.  Danger may enhance communication: predator calls alert females to male displays , 2010 .

[15]  David Wheatcroft,et al.  Species replacement reduces community participation in avian antipredator groups , 2016 .

[16]  Toshitaka N. Suzuki.  Communication about predator type by a bird using discrete, graded and combinatorial variation in alarm calls , 2014, Animal Behaviour.

[17]  J. C. Bremond,et al.  Recherches sur la sémantique et les éléments vecteurs d’information dans les signaux acoustiques du rouge-gorge (Erithacus Rubecula L.) , 1968, La Terre et La Vie, Revue d'Histoire naturelle.

[18]  Toshitaka N. Suzuki.  Referential mobbing calls elicit different predator-searching behaviours in Japanese great tits , 2012, Animal Behaviour.

[19]  Robert D. Magrath,et al.  Calling in the Face of Danger: Predation Risk and Acoustic Communication by Parent Birds and Their Offspring , 2010 .

[20]  H. Gelter,et al.  Song differences between the Pied Flycatcher Ficedula hypoleuca, the Collared Flycatcher F. albicollis, and their hybrids , 1987 .

[21]  Koji Fujita,et al.  Recursive Merge and Human Language Evolution , 2014 .

[22]  Bernard Lohr,et al.  Call learning in black-capped chickadees (Parus atricapillus) : The role of experience in the development of 'chick-a-dee' calls , 2010 .

[23]  J. Hailman,et al.  The ‘chick-a-dee’ calls of Parus atricapillus: A recombinant system of animal communication compared with written English , 1985 .

[24]  Michael Griesser,et al.  Referential Calls Signal Predator Behavior in a Group-Living Bird Species , 2008, Current Biology.

[25]  R. Zann,et al.  Structure, sequence and evolution of song elements in wild Australian Zebra finches , 1993 .

[26]  Klaus Zuberbühler,et al.  Language evolution: Semantic combinations in primate calls , 2006, Nature.

[27]  M. Ryan,et al.  What do animal signals mean? , 2009, Animal Behaviour.

[28]  L. Garamszegi,et al.  Flycatcher song in allopatry and sympatry – convergence, divergence and reinforcement , 2004, Journal of evolutionary biology.

[29]  Toshitaka N. Suzuki.  Assessment of predation risk through referential communication in incubating birds , 2015, Scientific Reports.

[30]  Toshitaka N. Suzuki.  Calling at a Food Source: Context-Dependent Variation in Note Composition of Combinatorial Calls in Willow Tits , 2012 .

[31]  April M. Becker,et al.  MOBBING CALLS OF BLACK-CAPPED CHICKADEES: EFFECTS OF URGENCY ON CALL PRODUCTION , 2002 .

[32]  R. Berwick,et al.  Songs to syntax: the linguistics of birdsong , 2011, Trends in Cognitive Sciences.

[33]  F. J. Pelletier.  The Principle of Semantic Compositionality , 1994 .

[34]  Torben Dabelsteen,et al.  Song Features Essential for Species Discrimination and Behaviour Assessment By Male Blackbirds (Turdus Merula) , 1992 .

[35]  Simon W. Townsend,et al.  Functionally Referential Communication in Mammals: The Past, Present and the Future , 2013 .

[36]  A. Kozhevnikov,et al.  Temperature Manipulation in Songbird Brain Implicates the Premotor Nucleus HVC in Birdsong Syntax , 2015, The Journal of Neuroscience.

[37]  P. Marler,et al.  Monkey responses to three different alarm calls: evidence of predator classification and semantic communication. , 1980, Science.

[38]  Lise Menn,et al.  Methods for studying language production , 1999 .

[39]  Toshitaka N Suzuki,et al.  From bird calls to human language: exploring the evolutionary drivers of compositional syntax , 2018, Current Opinion in Behavioral Sciences.

[40]  Richard Bertram,et al.  Independent Premotor Encoding of the Sequence and Structure of Birdsong in Avian Cortex , 2014, The Journal of Neuroscience.

[41]  Klaus Zuberbühler,et al.  Food-associated calling sequences in bonobos , 2009, Animal Behaviour.

[42]  K. Okanoya.  The Bengalese Finch: A Window on the Behavioral Neurobiology of Birdsong Syntax , 2004, Annals of the New York Academy of Sciences.

[43]  Toshitaka N. Suzuki.  Long-Distance Calling by the Willow Tit, Poecile montanus, Facilitates Formation of Mixed-Species Foraging Flocks , 2012 .

[44]  David Wheatcroft,et al.  Reproductive interference via display signals: the challenge of multiple receivers , 2015, Population Ecology.

[45]  Timothy Q. Gentner,et al.  Recursive syntactic pattern learning by songbirds , 2006, Nature.

[46]  M. Manser,et al.  The acoustic structure of suricates' alarm calls varies with predator type and the level of response urgency , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[47]  K. Zuberbühler,et al.  Campbell's monkeys concatenate vocalizations into context-specific call sequences , 2009, Proceedings of the National Academy of Sciences.

[48]  Noam Chomsky,et al.  वाक्यविन्यास का सैद्धान्तिक पक्ष = Aspects of the theory of syntax , 1965 .

[49]  A. Qvarnström,et al.  Song similarity predicts hybridization in flycatchers , 2006, Journal of evolutionary biology.

[50]  Raoul A. Mulder,et al.  A Novel Context for Bird Song: Predator Calls Prompt Male Singing in the Kleptogamous Superb Fairy‐wren, Malurus cyaneus , 2010 .

[51]  Noam Chomsky,et al.  Minimalist inquiries : the framework , 1998 .

[52]  R. Seyfarth,et al.  The assessment by vervet monkeys of their own and another species' alarm calls , 1990, Animal Behaviour.

[53]  Toshitaka N. Suzuki.  Parental alarm calls warn nestlings about different predatory threats , 2011, Current Biology.

[54]  L. Wallin,et al.  Integration of a call into the song of the Collared Flycatcher: adaptive compensation for booadcast efficiency? , 1987 .

[55]  David Wheatcroft,et al.  Repetition rate of calls used in multiple contexts communicates presence of predators to nestlings and adult birds , 2015, Animal Behaviour.

[56]  K. Zuberbühler.  Referential Signaling in Non-Human Primates: Cognitive Precursors and Limitations for the Evolution of Language , 2003 .

[57]  Erick Greene,et al.  Allometry of Alarm Calls: Black-Capped Chickadees Encode Information About Predator Size , 2005, Science.

[58]  Toshitaka N Suzuki,et al.  Experimental evidence for compositional syntax in bird calls , 2016, Nature Communications.

[59]  M. C. Baker,et al.  VOCAL ONTOGENY OF NESTLING AND FLEDGLING BLACK-CAPPED CHICKADEES POECILE ATRICAPILLA IN NATURAL POPULATIONS , 2003 .

[60]  Klaus Zuberbühler,et al.  Causal knowledge of predators' behaviour in wild Diana monkeys , 2000, Animal Behaviour.

[61]  K. Riebel,et al.  Are good ornaments bad armaments? Male chaffinch perception of songs with varying flourish length , 2003, Animal Behaviour.

[62]  Todd M. Freeberg,et al.  Chick-a-dee call variation in Carolina chickadees and recruiting flockmates to food , 2009 .

[63]  Noam Chomsky,et al.  The faculty of language: what is it, who has it, and how did it evolve? , 2002 .

[64]  K A Shiovitz,et al.  The process of species-specific song recognition by the indigo bunting, Passerina cyanea, and its relationship to the organization of avian acoustical behavior. , 1975, Behaviour.

[65]  Toshitaka N. Suzuki.  Semantic communication in birds: evidence from field research over the past two decades , 2016, Ecological Research.

[66]  E. Balaban,et al.  Bird song syntax: learned intraspecific variation is meaningful. , 1988, Proceedings of the National Academy of Sciences of the United States of America.

[67]  Gábor Győri.  Animal communication and human language , 1995 .

[68]  Nobuyuki Kutsukake,et al.  Foraging intention affects whether willow tits call to attract members of mixed-species flocks , 2017, Royal Society Open Science.

[69]  Willem H. Zuidema,et al.  Simple rules can explain discrimination of putative recursive syntactic structures by a songbird species , 2009, Proceedings of the National Academy of Sciences.

[70]  N. Davies,et al.  Learning fine-tunes a specific response of nestlings to the parental alarm calls of their own species , 2004, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[71]  K. Zuberbühler,et al.  Meaningful call combinations in a non-human primate , 2008, Current Biology.

[72]  P. Slater,et al.  Bird Song: Biological Themes and Variations , 1995 .

[73]  Jeffrey Podos,et al.  The evolution of bird song: male and female response to song innovation in swamp sparrows , 2001, Animal Behaviour.

[74]  Todd M Freeberg,et al.  Receivers respond differently to chick-a-dee calls varying in note composition in Carolina chickadees, Poecile carolinensis , 2002, Animal Behaviour.

[75]  B. Partee,et al.  Mathematical Methods in Linguistics , 1990 .

[76]  Toshitaka N. Suzuki.  Alarm calls evoke a visual search image of a predator in birds , 2018, Proceedings of the National Academy of Sciences.

[77]  Jeffrey R. Lucas,et al.  Chick-a-dee call syntax, social context, and season affect vocal responses of Carolina chickadees (Poecile carolinensis) , 2004, Behavioral Ecology and Sociobiology.

[78]  Svein Haftorn.  Ontogeny of the vocal repertoire in the Willow Tit Parus montanus , 1993 .

[79]  Marc F. Schmidt,et al.  At the interface of the auditory and vocal motor systems: NIf and its role in vocal processing, production and learning , 2013, Journal of Physiology-Paris.

[80]  M. Manser,et al.  The information that receivers extract from alarm calls in suricates , 2001, Proceedings of the Royal Society of London. Series B: Biological Sciences.

[81]  Todd M. Freeberg,et al.  Complexity in the Chick-a-Dee Call of Carolina Chickadees (Poecile Carolinensis): Associations of Context and Signaler Behavior to Call Structure , 2008 .

[82]  Stephen T. Emlen,et al.  An Experimental Analysis of the Parameters of Bird Song Eliciting Species Recognition , 1972 .

[83]  Andrew N. Radford,et al.  The development of alarm call behaviour in mammals and birds , 2009, Animal Behaviour.

[84]  Robert W. Ficken,et al.  VOCAL REPERTOIRE OF THE BLACK-CAPPED CHICKADEE , 1978 .

[85]  Carel ten Cate,et al.  Zebra Finch Song Phonology and Syntactical Structure across Populations and Continents—A Computational Comparison , 2016, Front. Psychol..

[86]  Jack P. Hailman,et al.  Combinatorial animal communication with computable syntax: Chick-a-dee calling qualifies as ‘Language’ by structural linguistics , 1986, Animal Behaviour.

[87]  Torben Dabelsteen,et al.  Coding in the song of the wren: importance of rhythmicity, syntax and element structure , 2000, Animal Behaviour.