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The Evolution of Animal Communication Systems: Questions of Function Examined Through Simulation University of Southampton Research Repository ePrints Soton Copyright © and Moral Rights for this thesis are retained by the author and/or other copyright owners. A copy can be downloaded for personal non-commercial research or study, without prior permission or charge. This thesis cannot be reproduced or quoted extensively from without first obtaining permission in writing from the copyright holder/s. The content must not be changed in any way or sold commercially in any format or medium without the formal permission of the copyright holders. When referring to this work, full bibliographic details including the author, title, awarding institution and date of the thesis must be given e.g. AUTHOR (year of submission) "Full thesis title", University of Southampton, name of the University School or Department, PhD Thesis, pagination http://eprints.soton.ac.uk The Evolution of Animal Communication Systems: Questions of Function Examined through Simulation Jason Noble Submitted for the degree of D. Phil. University of Sussex November, 1998 Declaration I hereby declare that this thesis has not been submitted, either in the same or different form, to this or any other university for a degree. Signature: Acknowledgements Seth Bullock and Ezequiel Di Paolo have probably had the greatest immediate influence on the thesis, and so deserve to be acknowledged first: thanks, guys. I remain grateful to Dave Cliff for taking me on as a student in the first place, back when I was full of wild-eyed ALife enthusiasm. Dave’s devotion as a supervisor is rivalled only by his skill as a rock-climber. He is also very good company—cheers, Dave. Thanks also to Mike Scaife, who kindly agreed to take up the slack when Dave moved on. Mike has been a level-headed and pragmatic presence during the frantic final year of my D.Phil. The research community in the School of Cognitive and Computing Sciences at Sussex has been extremely supportive, and has gone some way towards restoring my faith in academia. There are too many people to mention individually, but some names stand out. A big thank you to Jo Brook, Julian Budd, Ron Chrisley, Stephen Eglen, Inman Harvey, Phil Jones, Adrian Thompson and Mike Wheeler, plus all the members of the ALERGIC, E-Intentionality, CogPhi and EvoComm discussion groups. I would also like to thank Linda Thompson and Magdalena Portmann, without whom nothing in COGS would work. But it’s not over yet. There’s a large group of people, some of them at Sussex and some not, without whom I couldn’t have done it. Their contributions to the thesis range from philosophy to beer to just being patient with me (sometimes all three at once), and I want to thank them all: Argiri Arfani, Guillaume Barreau, Tom “the Cincinnatti Kid” Beament, Hilan Bensusan, Vicki Carruthers, Sirem Erkek (my en tatlı bebek), Berkan Eskikaya, Sevan Ficici, Ronald Lemmen, Mar Muriana-Lopez,´ Sean Noble, Manuel de Pinedo Garc´ıa (“Market Diner, before!”),´ Jorge Ram´ırez- Uresti, Oli Sharpe, Tim Taylor, and Richard Watson. Many people at SAB 96, EvoLang II, ALife VI, and SAB 98 were also helpful. Thanks. Thank you to John Maynard Smith and Peter Caryl for the advice and encouragement, and thanks to Bruce MacLennan for being such a good sport about chapter 5. For their financial support I would like to thank the Association of Commonwealth Universities and the British Council, who were foolish enough to take me on as a Commonwealth Scholar. At this point I should probably also thank Mum and Dad, who gave a lot more than just cash. Finally, my undying gratitude goes to various Australian record companies who, through inaction, encouraged me to do a D.Phil. in the first place. Preface Parts of the thesis are based on work that has been previously presented or published. Chapter 4 borrows heavily from a paper presented at the Fourth European Conference on Artificial Life (No- ble, 1997). Chapter 5 is based on a paper published in the Proceedings of the Fourth Conference on the Simulation of Adaptive Behavior (Noble & Cliff, 1996)—Dave Cliff was a co-author of this paper but all of the work actually presented in the chapter is my own. Chapter 7 is based on a paper published in the Proceedings of the Sixth Conference on Artificial Life (Noble, 1998b) and presented at the Second International Conference on the Evolution of Language (Noble, 1998a); an extended version of the work is currently in press in the journal Adaptive Behavior. Chapter 8 is based on a paper published in the Proceedings of the Fifth Conference on the Simulation of Adaptive Behavior (Noble, 1998c). The Evolution of Animal Communication Systems: Questions of Function Examined through Simulation Jason Noble Summary Simulated evolution is used as a tool for investigating the selective pressures that have influenced the design of animal signalling systems. The biological literature on communication is first re- viewed: central concepts such as the handicap principle and the view of signalling as manipulation are discussed. The equation of “biological function” with “adaptive value” is then defended, along with a workable definition of communication. Evolutionary simulation models are advocated as a way of testing the coherence of a given theory. Contra some ALife enthusiasts, simulations are not alternate worlds worthy of independent study; in fact they fit naturally into a Quinean picture of scientific knowledge as a web of modifiable propositions. Existing simulation work on the evo- lution of communication is reviewed: much of it consists of simple proofs of concept that fail to make connections with existing theory. A particular model (MacLennan & Burghardt, 1994) of the evolution of referential communication in a co-operative context is replicated and critiqued in detail. Evolutionary simulations are then presented that cover a range of ecological scenarios; the first is a general model of food- and alarm-calling. In such situations signallers and receivers can have common or conflicting interests; the model allows us to test the idea that a conflict of interests will lead to an arms race of ever more costly signals, whereas common interests will result in signals that are as cheap as possible. The second model is concerned with communication during aggressive interactions. Many animals use signals to settle contests, thus avoiding the costs associated with fighting. Conventional game-theoretic results suggest that the signalling of aggression or of strength will not be evolutionarily stable unless it is physically unfakeable, but some recent models imply that cost-free, arbitrary signals can be reliable indicators of both intent and ability. The simulation, which features continuous-time perception of the opponent’s strategy, is an attempt to settle the question. The third model deals with sexual signalling, i.e., elaborate displays that are designed to persuade members of the opposite sex to mate. The results clarify the question of whether such displays are the pointless result of runaway sexual selection, or whether they function as honest and costly indicators of genetic quality. The models predict the evolution of reliable communication in a surprisingly narrow range of circumstances; a serious gap remains between these predictions and the ethological data. Future directions for simulation work are discussed. Submitted for the degree of D. Phil. University of Sussex November, 1998 Contents 1 Introduction 1 1.1 Ways of explaining animal behaviour . 1 1.2 Theoretical models and functional explanations of animal communication . 3 1.3 The use of computer simulations . 4 1.4 Problems in defining communication . 5 1.5 Human language . 6 1.6 Overview . 7 1.6.1 Outline of the thesis . 7 1.6.2 Original contributions . 9 2 The biological literature 10 2.1 Darwin on communication . 10 2.1.1 Signalling of emotional state . 11 2.1.2 Sexual advertisement signalling . 11 2.2 The ethological view . 12 2.2.1 Derived activities . 12 2.2.2 Ritualization . 14 2.2.3 Ethology and group-selection thinking . 15 2.3 The rise of behavioural ecology and sociobiology . 15 2.3.1 Group selection or selfish genes? . 16 2.3.2 Animals as maximizers and the application of game theory . 17 2.3.3 A new perspective on signalling . 19 2.3.4 Reinterpretation of the ethological data . 21 2.4 Zahavi’s handicap principle . 22 2.4.1 Paradoxical logic of the theory . 22 2.4.2 Controversial status and variant interpretations . 23 2.4.3 Variable costs and benefits for signallers . 25 2.5 Dawkins and Krebs on communication . 26 2.5.1 Information or manipulation? . 26 2.5.2 Two kinds of signal co-evolution . 28 2.6 Receiver psychology . 29 2.6.1 Sensory biases . 31 2.6.2 Information requirements of receivers . 32 2.7 Communication and sexual selection . 32 2.7.1 The phenotypic gambit . 34 Contents 7 3 Conceptual issues in the study of communication 36 3.1 The concept of function in biology . 36 3.2 Communication and related concepts . 39 3.2.1 Some problem cases . 39 3.2.2 Attempts to define animal communication in terms of behaviour . 41 3.2.3 The role of information . 43 3.2.4 Intentional communication? . 45 3.2.5 Millikan’s alternative classification scheme . 47 3.3 A functional definition of communication . 50 4 Artificial life as a method for studying evolution 54 4.1 Some matters of terminology . 55 4.1.1 Artificial life . 55 4.1.2 Models and simulations . 56 4.2 This thing called science . 57 4.3 Artificial life as science . 57 4.3.1 Does artificial life qualify? . 57 4.3.2 The concept of emergence . 59 4.4 Artificial life and the return of the analytic-synthetic distinction .
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