Biosemiotics Volume 16 Series editors Jesper Hoffmeyer, Professor emeritus in biochemistry, University of Copenhagen Kalevi Kull, Professor in biosemiotics, University of Tartu Alexei Sharov, National Institute of Aging, Baltimore Aims and Scope of the Series Combining research approaches from biology, semiotics, philosophy and linguistics, the ield of biosemiotics studies semiotic processes as they occur in and among living systems. This has important implications and applications for issues ranging from natural selection to animal behaviour and human psychology, leaving biosemiotics at the cutting edge of the research on the fundamentals of life. The Springer book series Biosemiotics draws together contributions from leading scholars in international biosemiotics, producing an unparalleled series that will appeal to all those interested in the origins and evolution of life, including molecular and evolutionary biologists, ecologists, anthropologists, psychologists, philosophers and historians of science, linguists, semioticians and researchers in artiicial life, information theory and communication technology. More information about this series at http://www.springer.com/series/7710 Timo Maran Mimicry and Meaning: Structure and Semiotics of Biological Mimicry Timo Maran Department of Semiotics University of Tartu Tartu, Estonia ISSN 1875-4651 ISSN 1875-466X (electronic) Biosemiotics ISBN 978-3-319-50315-8 ISBN 978-3-319-50317-2 (eBook) DOI 10.1007/978-3-319-50317-2 Library of Congress Control Number: 2016961294 © Springer International Publishing AG 2017 This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of the material is concerned, speciically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a speciic statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, express or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional afiliations. Printed on acid-free paper This Springer imprint is published by Springer Nature The registered company is Springer International Publishing AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland Preface This book is a result of 20 years of my interest in reading and thinking about the bizarre phenomenon of mimicry. This journey has brought me from the standard scientiic world view of animal ecology to a small but friendly community of semiotics, in other words, from treating mimicry as a defensive adaptation to recognising it as a peculiar sign structure in an open semiotic space of ecological relations. The topic of mimicry made it possible for me to work together with Kalevi Kull, to whom I owe a lot for his academic guidance. This scholarly cooperation resulted in the defence of a doctoral thesis on the semiotics of mimicry in 2005 (published also as Maran 2008a). Since then, I have focused on several related topics and, in the process, have developed semiotic tools to analyse mimicry, discerning speciic sign structures active in mimicry and locating the phenomenon within the broader ecological framework. The present book elaborates and reorganises texts of my earlier research papers published on mimicry and also includes materials from several conference presentations and manuscripts that have not been previously published. Earlier publications have been thoroughly reviewed and updated. The focus of this book is on mimicry as a sign process. The reader will not ind lengthy discussions on the evolutionary dynamics of mimicry and genetic causes of speciic mimicry cases that often dominate biological literature. Evolutionary perspectives on mimicry are discussed with great precision by other authors in other publications (for overviews, see, e.g., Brower 1988; Ruxton et al. 2004; Wickler 1968; Cott 1957). My emphasis here is mostly on mimicry as a formal communication structure, in other words, on sign processes that take part in mimicry and their possible ecological effects. The study of mimicry has a long history that has resulted in hundreds of books on the topic, but it has also brought along some stereotypical understandings and concepts. By using biosemiotics as paradigmatic framework, my attempt is to treat mimicry in a way that could provide some new perspectives. As for any object of science, the choice of concepts to be used and questions to be asked partly constrains and inluences the results that will be gained. The present book contains eight main chapters that are accompanied with three short excursions. In these interludes, I discuss historical and philosophical aspects of mimicry and open up some premises that have led me to approach mimicry in the v vi Preface way that it is presented in this book. I start the book with an overview of the types and typologies of the mimicry, paying special attention to how the understanding of the tripartite mimicry structure was formed. Thereafter, I discuss semiotics of mimicry from different viewpoints: disciplinary, structural and communicative. I pay special attention to the relationship between mimicry and iconic signs. In the second part of the book, the attention shifts to the dynamics of sign processes going on in mimicry and what the roles and choices of the different participants are. I also consider the position of the human observer in regard to mimicry and propose semiotic tools for modelling mimicry. In the inal chapters, I discuss some semiotic insights pertaining to the evolution of mimicry and make propositions to position mimicry within broader interspeciic communication networks of the ecosystems. For the possibility of writing this book, I owe a debt of gratitude to Kalevi Kull, Peeter Torop, Riin Magnus, Silvi Salupere, Elin Sütiste and other colleagues at the Department of Semiotics, University of Tartu, for creating a good academic environment, for collegial support and for thinking along with me. Interest in mimicry has put me in contact with the Prague Biosemiotic School, whose works have been of great relevance for me. I would especially like to emphasise the long-lasting cooperation with Karel Kleisner, as well as consultations with philosopher Stanislav Komárek. In biosemiotics, my major guides and beacons have been Thomas A. Sebeok, Jesper Hoffmeyer and Almo Farina. I also want to pay reverence to authors whom I have not met in person, but whose written words have been a great source of inspiration for my thinking about mimicry: Henry W. Bates, Edward Poulton, Jakob von Uexküll, Wolfgang Wickler, Malcolm Edmunds, John Maynard Smith, Adolf Portmann, Richard I. Vane-Wright and Roger Caillois, among others. In this book, the material from the following papers has been used in a thoroughly reviewed and updated form: Maran, T. (2015). Scaffolding and mimicry: A semiotic view of the evolutionary dynamics of mimicry systems. Biosemiotics (Springer), 8(2), 211–222; Maran, T. (2012). Are ecological codes archetypal structures? In T. Maran, K. Lindström, R. Magnus, M. Tønnessen (eds.). Semiotics in the wild. Essays in honour of Kalevi Kull on the occasion of his 60th birthday. (pp. 147– 156). Tartu: University of Tartu Press; Maran, T. (2011). Becoming a sign: The mimic’s activity in biological mimicry. Biosemiotics (Springer), 4(2), 243–257; Maran, T. (2010). Semiotic modeling of mimicry with reference to brood parasitism. Sign Systems Studies (University of Tartu Press), 38(1/4), 349–377; Maran, T. and Kleisner, K. (2010). Towards an evolutionary biosemiotics: Semiotic selection and semiotic co-option. Biosemiotics (Springer), 3(2), 189–200; Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica (De Gruyter Mouton), 167(1/4), 223–248; Maran, T. (2007). Mimicry. In P. Bouissac, A. Lewis (Eds.). Semiotics Encyclopedia Online. E.J. Pratt Library, Victoria University (http://www. semioticon.com/seo/); Maran, T. (2003). Mimesis as a phenomenon of semiotic communication. Sign Systems Studies (University of Tartu Press), 31.1, 191–215. I thank the respective copyright owners for their courtesy for using these materials. Writing of this book would not have been possible without the all-encompassing support of my family. I am also grateful to Jamie L. Kruis for her help in language editing and to the editors in Springer for their technical assistance. I express my Preface vii gratitude to the Institute of Philosophy and Semiotics at the University of Tartu, Estonia, Norwegian-Estonian Research Cooperation Programme (grant EMP151), Estonian Research Council (grant 2–44) and European Regional Development Fund (Centre of Excellence in Cultural Theory) for their inancial support that made it possible to do the research on which the present book is based. Tartu, Estonia Timo Maran Contents 1 Biosemiotics of Mimicry: Introductory Notes ...................................... 1.1 On a Biosemiotic Approach ........................................................... 1.2 Deining Biological Mimicry ......................................................... 1 4 7 2 First Excursion: The History of the Mimicry Concept ....................... 11 3 The Structure of Mimicry ...................................................................... 3.1 Mimicry Types ............................................................................... 3.2 Mimicry in Relation to Other Adaptations..................................... 3.3 Typologies of Mimicry................................................................... 3.4 Mimicry Systems—Wolfgang Wickler’s Account ......................... 3.5 Critical Discussion of the Triadic Mimicry Model ........................ 15 16 19 23 28 30 4 Semiotics of Mimicry .............................................................................. 4.1 Semiotic Interpretations of Mimicry.............................................. 4.2 Mimicry as a Communicative Interaction ...................................... 4.3 Mimicry as a Sign System ............................................................. 4.4 The Umwelten of the Receiver and the Human Observer ............. 35 36 38 42 49 5 Iconicity and Mimicry ............................................................................ 5.1 If Mimic Is a Sign Then What Does It Stands For? ....................... 5.2 Peirce’s Second Trichotomy and Animal Communication ............ 5.3 Peircean Categories and the Three Basic Mimicry Types ............. 55 55 58 61 6 Second Excursion: Importance of the Object ...................................... 65 7 Different Perspectives in Mimicry System ............................................ 7.1 Mimic’s Activity and Intentionality ............................................... 7.2 Resembling the Environment and Becoming a Sign ..................... 7.3 The Receiver’s Perspective and Ambivalent Signs ........................ 69 71 74 78 8 Modelling Mimicry ................................................................................. 8.1 Toolbox for Modelling Mimicry .................................................... 8.2 Applying Semiotic Modelling to Brood Parasitism ....................... 8.3 Towards Comparative Modelling ................................................... 83 84 88 96 ix x 9 Contents Mimicry and Semiotic Evolution ........................................................... 9.1 Semiotic Selection: Deinition and Examples................................ 9.2 Mimicry and Semiotic Scaffolding ................................................ 9.3 Evolution of Mimicry in the Bio-semiosphere .............................. 101 104 109 115 10 Third Excursion: An Epistemology of the Uncertain .......................... 119 11 From Abstract Mimicry to Ecological Codes ....................................... 11.1 Abstract Mimicry: When the Meaning Comes First ...................... 11.2 Connecting Umwelten, Sharing Codes .......................................... 11.3 Ecological Codes and Archetypal Structures ................................. 123 124 128 132 12 Conclusions .............................................................................................. 137 References ........................................................................................................ 139 Index ................................................................................................................. 157 

Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Mimicry and Meaning: Structure and Semiotics of Biological Mimicry Timo Maran Department of Semiotics Institute of Philosophy and Semiotics University of Tartu Jakobi 2-311 Tartu 51014 Estonia timo.maran@ut.ee http://kodu.ut.ee/~timo_m 1 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. PREFACE This book is a result of twenty years of my interest in reading and thinking about the bizarre phenomenon of mimicry. This journey brought me from the standard scientific worldview of animal ecology to a small but friendly community of semiotics; in other words, from treating mimicry as a defensive adaptation to recognising it as a peculiar sign structure in an open semiotic space of ecological relations. The topic of mimicry made it possible for me to work together with Kalevi Kull, to whom I owe a lot for his academic guidance. This scholarly cooperation resulted in the defence of a doctoral thesis on the semiotics of mimicry in 2005 (published as Maran 2008a). Since then, I have focused on several related topics and in the process, have developed semiotic tools to analyse mimicry, discerning specific sign structures active in mimicry and locating the phenomenon within the broader ecological framework. The present book elaborates and reorganises texts of my earlier research papers published on mimicry, and also includes materials from several conference presentations and manuscripts that have not been previously published. Earlier publications are thoroughly reviewed and updated. The focus of this book is on mimicry as a sign process. The reader will not find lengthy discussions on the evolutionary dynamics of mimicry and genetic causes of specific mimicry cases that often dominate biological literature. Evolutionary perspectives on mimicry are discussed with great precision by other authors in other publications (for overviews, see e.g. Brower 1988; Ruxton et al. 2004; Wickler 1968; Cott 1957). My emphasis here is mostly on mimicry as a formal communication structure; in other words, on sign processes that take part in mimicry and their possible ecological effects. The study of mimicry has a long history that has resulted in hundreds of books on the topic, but it has also brought along some stereotypical understandings and concepts. By using biosemiotics as paradigmatic framework, my attempt is to treat mimicry in a way that could provide some new perspectives. As for any object of science, the choice of concepts to be used and questions to be asked partly constrains and influences the results that will be gained. The present book contains eight main chapters that are accompanied with three short excursions. In these interludes, I discuss historical and philosophical aspects of mimicry and open up some premises that have led me to approach mimicry in the way that it is presented in this book. I start the book with an overview of the types and 2 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. typologies of the mimicry, paying special attention to how the understanding of the tripartite mimicry structure was formed. Thereafter I discuss semiotics of mimicry from different viewpoints: disciplinary, structural and communicative. I pay special attention to the relationship between mimicry and iconic signs. In the second part of the book, the attention shifts to the dynamics of sign processes going on in mimicry and what the roles and choices of the different participants are. I also consider the position of the human observer in regard to mimicry and propose semiotic tools for modelling mimicry. In the final chapters, I discuss some semiotic insights toward the evolution of mimicry and make propositions to position mimicry within broader interspecific communication networks of the ecosystems. For the possibility of writing this book, I owe a debt of gratitude to Kalevi Kull, Peeter Torop, Riin Magnus, Silvi Salupere, Elin Sütiste and other colleagues at the Department of Semiotics, University of Tartu for creating a good academic environment, for collegial support and for thinking along with me. Interest in mimicry has put me in contact with the Prague Biosemiotic School, whose works have been of great relevance for me. I would especially like to emphasize the long-lasting cooperation with Karel Kleisner, as well as consultations with a philosopher, Stanislav Komárek. In biosemiotics, my major guides and beacons have been Thomas A. Sebeok, Jesper Hoffmeyer and Almo Farina. I also want to pay reverence to authors whom I have not met in person, but whose written words have been a great source of inspiration on my thinking about mimicry: Henry W. Bates, Edward Poulton, Jakob von Uexküll, Wolfgang Wickler, Malcolm Edmunds, John Maynard Smith, Adolf Portmann, Richard I. Vane-Wright, and Roger Callois, among others. In this book, the material from the following papers have been used in a thoroughly reviewed and updated form: Maran, T. (2015). Scaffolding and mimicry: A semiotic view of the evolutionary dynamics of mimicry systems. Biosemiotics (Springer), 8(2), 211–222; Maran, T. (2012). Are ecological codes archetypal structures? In: T. Maran, K. Lindström, R. Magnus, M. Toennessen (eds.). Semiotics in the wild. Essays in honour of Kalevi Kull on the occasion of his 60th birthday. (pp. 147–156). Tartu: Tartu University Press; Maran, T. (2011). Becoming a sign: The mimic's activity in biological mimicry. Biosemiotics (Springer), 4(2), 243–257; Maran, T. (2010). Semiotic modeling of mimicry with reference to brood parasitism. Sign Systems Studies (University of Tartu Press), 38(1/4), 349–377; Maran, T. and Kleisner, K. (2010). Towards an evolutionary biosemiotics: Semiotic selection and 3 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. semiotic co-option. Biosemiotics (Springer), 3(2), 189–200; Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica (DeGruyter Mouton), 167(1/4), 223–248; Maran, T. (2007). Mimicry. In: P. Bouissac, A. Lewis (Eds.). Semiotics Encyclopedia Online. E.J. Pratt Library, Victoria University (http://www.semioticon.com/seo/); Maran, T. (2003). Mimesis as a phenomenon of semiotic communication. Sign Systems Studies (University of Tartu Press), 31.1, 191– 215. I thank the respective copyright owners for their courtesy for using these materials. I am grateful to Jamie L. Kruis for her help in language editing and to the editors in Springer for their technical support. I also express my gratitude to the Institute of Philosophy and Semiotics at the University of Tartu, Estonia; NorwegianEstonian Research Cooperation Programme (grant EMP151); Estonian Research Council (grant 2-44); and to European Regional Development Fund (Centre of Excellence in Cultural Theory) for their financial support that made it possible to do the research on which the present book is based. 4 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. TABLE OF CONTENTS PREFACE ...................................................................................................................... 2 TABLE OF CONTENTS ............................................................................................... 5 1 BIOSEMIOTICS OF MIMICRY: INTRODUCTORY NOTES ................................ 6 1.1 On a biosemiotic approach...................................................................................9 1.2 Defining biological mimicry ..............................................................................14 2 FIRST EXCURSION: THE HISTORY OF THE MIMICRY CONCEPT .............. 19 3 THE STRUCTURE OF MIMICRY ......................................................................... 23 3.1 Mimicry types ....................................................................................................24 3.2 Mimicry in relations to other adaptations ..........................................................27 3.3 Typologies of mimicry .......................................................................................33 3.4 Mimicry systems—Wolfgang Wickler’s account ..............................................39 3.5 Critical discussion of the triadic mimicry model ...............................................42 4 SEMIOTICS OF MIMICRY .................................................................................... 48 4.1 Semiotic interpretations of mimicry ..................................................................50 4.2 Mimicry as a communicative interaction ...........................................................52 4.3 Mimicry as a sign system ...................................................................................56 4.4 The Umwelten of the receiver and the human observer ....................................64 5 ICONICITY AND MIMICRY ................................................................................. 70 5.1 If mimic is a sign then what does it stands for? .................................................70 5.2 Peirce’s second trichotomy and animal communication ...................................74 5.3 Peircean categories and the three basic mimicry types ......................................77 6 SECOND EXCURSION: IMPORTANCE OF THE OBJECT ................................ 82 7 DIFFERENT PERSPECTIVES IN MIMICRY SYSTEM ....................................... 86 7.1 Mimic’s activity and intentionality ....................................................................88 7.2 Resembling the environment and becoming a sign ...........................................92 7.3 The receiver’s perspective and ambivalent signs...............................................98 8 MODELLING MIMICRY ...................................................................................... 104 8.1 Toolbox for modelling mimicry.......................................................................105 8.2 Applying semiotic modelling to brood parasitism ...........................................110 8.3 Towards the comparative modelling ................................................................120 9 MIMICRY AND SEMIOTIC EVOLUTION ......................................................... 125 9.1 Semiotic selection: Definition and examples ...................................................129 9.2 Mimicry and semiotic scaffolding ...................................................................136 9.3 Evolution of mimicry in the bio-semiosphere .................................................143 10 THIRD EXCURSION: AN EPISTEMOLOGY OF THE UNCERTAIN ............ 147 11 FROM ABSTRACT MIMICRY TO ECOLOGICAL CODES ........................... 152 11.1 Abstract mimicry: When the meaning comes first ........................................154 11.2 Connecting Umwelten, sharing codes............................................................159 11.3 Ecological codes and archetypal structures ...................................................164 12 CONCLUSIONS................................................................................................... 170 REFERENCES ..........................................................................................................173 INDEX .......................................................................................................................197 5 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 1 BIOSEMIOTICS OF MIMICRY: INTRODUCTORY NOTES On a rainy October day five years ago, I messed up my mushroom roast. We had spent a long day hiking in thick north-east Estonian forests near Lake Peipus, gathering different Boletus species from this chilly and foggy mushroom heaven. Later, back home, after spending hours cleaning and preparing mushrooms, the smell of freshly roasted mushrooms floated in the air and my mind was prepared for the dinner. The first morsel brought me, however, painfully back to reality as the roast had a distinctively bitter taste that overshadowed all other flavours and spices. In the forest we had probably mistakenly picked a bitter bolete Tylopilus felleus among young porcini Boletus edulis. One of such specimens is usually enough to make you throw away your dish. I was not a victim of, well, mimicry, but of my limited ability to distinguish similar species that had different properties or applicability. The same dilemma is faced by many species who act as receivers in mimicry, as they too need to distinguish between organisms that are edible or inedible, harmless or dangerous, species-mates or predators and so on. Even the bitter taste of Tylopilus felleus is supposedly part of chemical defence system that mushrooms have against some fungivorous insects (Hackman and Meinander 1979: 53; Spiteller 2015). This personal story characterises well the dominant themes in my approach to mimicry. First, mimicry, as I understand it, is a semiotic phenomenon—it includes a particular organism that has a problem in making correct interpretations in regard to the objects in its environment, within the limits of its perceptual sphere, based on the sign system it is using, and taking into account its competencies and earlier experiences. Second, mimicry is an ecological phenomena in the sense that it includes many different species of the given ecosystem. Not only does it create communicative and ecological connections between species that take part in that particular mimicry interaction (as the receiver, carrier of mimetic signals and object of imitation), but mimicry is often open to other species that can occasionally encounter and become deceived by the confusing resemblance. There appears to be some sort of mimetic landscape or mimicry potentiality in the ecosystem. There are many resemblances and relations in nature that have not yet formed distinct mimicry systems, but have potential to do so in the future under the right circumstances. Third, mimicry is not considered in this book as a pure object of biological science, but more as a hybrid object in the sense of Bruno Latour (1993) that connects spheres of biology and 6 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. culture. People in everyday life often have a strong relation to the mimeticity of nature. This could emerge as problems making taxonomical distinctions between similar species of berries, birds, and insects, or as folk narratives about cuckoo children or were-wolves. A facet of the same topic is expressed in human relations with companion species who are selected and shaped by humans according to some image or preference (for example, the Komondor and many other sheep dog breeds resembling sheep), or in a situation where valerian or catnip is offered to a cat. The natural scientific understanding, too, often relies upon and works with conceptual distinctions and models that have a cultural basis and are rooted in the long history of the concepts of mimesis and imitation in philosophy and aesthetics. At the same time I acknowledge the complexity of mimicry as a topic in biology. In many specific mimicry cases and topics such as developmental biology of mimicry, mimicry rings of the Heliconius butterflies, complex relations between Batesian and Müllerian mimicry, mimicry in brood parasitism, and others, the research literature dates back several decades and these subjects take full effort and many years to master. Having not worked as a field biologist for a long time now, I may be somewhat superficial in discussing these specific mimicry cases. At the same time, what I hope to offer a biologist is a supplemental viewpoint toward mimicry studies from the perspective of another discipline. A different perspective brings along different concepts and questions asked, allowing even the topic of mimicry, the history of which dates back about 150 years, to be seen afresh. If the reader happens to be a true-minded natural scientist, then I hope he or she can bear with me. I use concepts that come from semiotics, linguistics and other humanities disciplines, and apply explorative thinking probably more freely than is custom in natural sciences. My concern is not always about what mimicry is in terms of facts, but what it could be, if we would be willing to shift our frame of reference a little. A few questions that could be relevant to biology, and that I aim to cover in this book, are the following: What are relations of ecological and communicative processes in mimicry? What are organisms’ prerequisites and possibilities to take part in and act on mimicry? How can sign processes constrain or influence the development of mimicry? How can human perceptional bias and interpreting activity relate to mimicry, and what possible influence does this have on mimicry theory? For semiotics and especially for biosemiotics, I hope to offer a thorough treatment of a topic that truly has a semiotic nature. It needs to emphasized that 7 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. communication, as an exchange of information and messages between organisms, appears to be the core of mimicry indeed. Biosemiotics could benefit from having more object-focused research than is customary, which could consequently lead to novel developments of the biosemiotics theory. Studying mimicry appears to open up many theoretical questions that could be relevant for the general field of biosemiotics. Some of these discussed in this book include: What are the relationships between signs and their (dynamic) objects in biosemiosic processes? What are the limits or borders of a sign, both in regard to the number of different interpreters as well as the scope of possible interpretations? What is the relationship between mimicry and iconicity? What is the role of abstract or symbolic sign complexes in ecological semiotic systems? In a biosemiotics framework that deals with semiotic processes in intraorganismic, interorganismic and ecological levels, my research focuses mostly on semiotic and communicative relations between organisms and the role of semiosis in ecosystems. My primary concern is semiotic ecology of mimicry—how different species can become intertwined by mimicry in an ecosystem, and what the role and effect of mimicry as a semiotic process could be on this broader ecological-semiotic realm. Much of what I write about mimicry could be generalizable to other types of semiotic-ecological relations, which Jesper Hoffmeyer (2008: 189) has denoted as semethic interactions in the ecosystem. In my understanding, the level of ecological relations is the most natural level for studying semiotic processes in nature, and yet much work needs to be done in this area in biosemiotics. The main issue from a biosemiotic perspective would be how ecological relations and processes at the ecosystem level translate into qualitative forms that can be perceived by organisms in their subjective and local presence. My other area of interest within this book lies in the semiotic dynamics of mimicry. On the one hand, I try to observe how this diverse phenomenon of nature is translated into a scientific concept, i.e. how it is specified and defined and how, at a certain point, it obtains an identity of its own. The concept is further applied to describe biological processes and to make typological distinctions, and some problems emerge in this process. The major source of problems in mimicry studies appear to be incompatibility between the diversity of the biological world and human attempts to describe this diversity through a unified theory and by applying clear conceptual models. In this book, a different methodological approach is taken as my 8 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. aim is not to establish a strong unified theoretical core. Rather, I make use of works of various authors – from Thomas Sebeok to Roman Jakobson and from Jakob von Uexküll to Wolfgang Wickler – and this loosely organised set of ideas provides me a modelling devise suitable for bringing forth and analysing different aspects of mimicry as a complex phenomenon. On the other hand, I focus on the position of the organism in the mimetic interaction: what its semiotic activities and behavioural dilemmas are and how this contributes to mimicry resemblance. It is foremost the activity of the living organisms—their memory and abilities to distinguish and make mistakes—that influence what features are promoted in the evolution of mimicry resemblance and what will diminish in time. Thus my interest in mimicry has a twofold focus—humans as cultural sign users and animals as semiotic beings, and how mimicry as a semiotic phenomenon emerges when their different activities are being juxtaposed. Thus my approach corresponds to what has been recently called bi-constructivist or multiconstructivist ethology (Lestel et al. 2014; Jaroš 2016). Dominique Lestel and colleagues (2014: 128) describe bi-constructivism: “ethology as the science of the human interpretation of animal interpretation” that takes “axiomatic the subjectivity of animals and the situational emplacement of their human observers as living beings themselves.” I think that as a practical means of research, biosemiotics allows to understand this dynamic quite well as it provides a framework and methodological tools to also take into account the semiotic activity of other animals. It is not just humans that create the description of the reality, but other animals also. Therefore, it would be important to attribute the position of the subject to other animals. Humans are not the only observers and researchers of mimicry. Other species, within the limits of their perceptual capacities and cognitive distinctions, also try to make sense of what is what in mimicry. 1.1 On a biosemiotic approach To understand what the possible benefits could be to approach mimicry from a biosemiotic viewpoint, we would first need to take a short look at biosemiotics. Biosemiotics, or semiotic biology, is a paradigm in which the general aim is to study semiotic processes in a non-human world. If semiotics is historically considered as a field of humanities that studies language and other language-based sign systems, then 9 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. with the emergence of biosemiotics, prelinguistic semiotic processes are taken into focus. Biosemiotics studies qualitative semiotic processes that are considered to exist in a variety of forms down to the simplest living organisms and to the lowest levels of biological organization. The major semiotic process that biosemiotics works with is a sign process or semiosis, understood as a mediated relation where something stands for or represents something else. Through mediation, semiosis reveals information that would otherwise remain unknown while the same time, hides other properties of the entities that form the sign relation (that would be accessible without sign relation). Taking a simple example from the human society, wearing a uniform reveals the profession of the policeman but hides his/her personality. In addition to semiosis, biosemiotics also studies more specific types of semiotic processes such as communication, naming, categorisation, deception and others. In a biosemiotic understanding, sign processes presume the participation of the living organism— interpreter—for whom the sign would have meaning by taking into account its physiology, ecology, motivation and experience. Emphasizing the role of sign processes and interpretation in nature makes it possible to restore the subjectness or agency of living organisms, who in turn are considered to influence larger ecological and evolutionary processes. Biosemiotics as a paradigm emerged from the comparative semiotic studies of animal communication carried out by Hungarian-American semiotician Thomas A. Sebeok in the 1960s (called zoosemiotics at that time, Sebeok 1972, 1990b). Later reconstructions of the history of the field trace biosemiotics back to German romantic biology, primarily to Umwelt theory of Jakob von Uexküll (1982), and to the semiotics of the American pragmatist philosopher Charles S. Peirce. The description of structural resemblances between the genetic code and human language presented in the 1960s, among others by Roman Jakobson (1971), teacher of Sebeok, made an important contribution to the emergence of biosemiotics. The word biosemiotics itself was first used by Friedrich S. Rothschild in 1962 (see Kull 1999b), and it began to be used more widely in the early 1990s. Biosemiotics is also characterized by a diversity of approaches; better-known interpretations include the Copenhagen-Tartu group (Kull et al. 2009; Hoffmeyer 2008), Prague biohermeneutics (Markoš et al. 2009; Markoš and Faltýnek 2011), code biology (Barbieri 2008, 2010, 2012), and the 10 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. cybernetic approach (e.g. Pattee 2008; Brier 2013). 1 Nowadays biosemiotics can be considered as part of a larger shift in biology towards organism-centred approaches: evolutionary developmental biology, systems biology, epigenetic studies, extended Darwinian synthesis and others (Gilbert 2016; Gilbert and Epel 2015; Lindholm 2015; Noble 2006; West-Eberhard 2003). Early efforts to organize the field of biosemiotic studies largely followed taxonomical logic: in addition to zoosemiotics, the semiotic study of plants— phytosemiotics—was proposed (Krampen 1981; Kull 2000). Alternative classifications follow one of two systematic approaches, the first being the hierarchical logic of biological processes, distinguishing between endosemiotics (the study of semiotic processes inside the organism), zoosemiotics (the study of semiotic processes between organisms) and ecosemiotics or ecosystem semiotics (the study of semiotic aspects of ecological processes, e.g. Nielsen 2007). The other approach is based on the mechanisms of sign processes, distinguishing between the study of vegetative (based on analogical iconic relations), animal (based on physical linkage and indexical relations), and cultural semiosis (based on conventional symbolic relations) (Kull 2009). In a similar vein, Sharov and Vehkavaara (2015) have later introduced a distinction between protosemiosis (when direct associations are made between signs and actions) and eusemiosis (when associations are mediated by a signobject relation). In regard to the semiotics of mimicry, an ecosemiotic approach is relevant, defined as “the study of sign processes which relate organisms to their natural environment” (Nöth 2001:71) or as the semiotic discipline investigating “human relationships to nature which have a semiosic (sign-mediated) basis” (Kull 1998: 351). More recently, we have specified ecosemiotics to be “a branch of semiotics that studies sign processes as responsible for ecological phenomena (Maran and Kull 2014: 41). It may also be said that ecosemiotics is concerned with the semiotic processes that relate to or address the broader context of living biological processes. In all these levels and types, describing the properties of sign systems, semiosis and semiotic regulation is in the foreground. A most general philosophical principle of biosemiotics is the interconnectedness of sign processing (semiosis) and life: some biosemiotics authors 1 For overviews of the biosemiotics paradigm, see Favareau (2009); Kull (1999a, 2004); Barbieri (2009). 11 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. regard sign activity as an important property of living organisms, and some see semiosis as the very condition of life (Sebeok 2001). In its understanding of sign processes, biosemiotics mostly proceeds from the semiotics of Charles S. Peirce, for whom a sign is “something which stands to somebody for something in some respect or capacity” (CP 2.228). Such an understanding differs in important respect from the other major semiotic tradition, the semiology of Swiss linguist Ferdinand de Saussure: different from the Saussurean two-part model of the sign, the Peircean sign is tripartite (including the ‘object’) and does not rely on the existence of language— which make Peircean semiotics suitable for describing sign processes outside the human species. On the other hand, in contrast to the concept of signals in biology, signs are not fixed in biosemiotics: anything can be a sign for an organism, if interpreted as meaningful. Another major semiotic process, communication, adds the concepts of sender, repertoire and code that connect the parties of communication and make mutual understanding possible. This allows us to inquire about both the general properties of biological communication systems as well as the special position of human language therein. For instance: is the combination of analogical and digital codes necessary for the development of complex biological systems, as argued by Hoffmeyer and Emmeche as early as 1991? What are the similarities and differences between the codes that cells use for interpreting DNA and the codes of human language? Are there special rules for communication by which the members of the different species interact? One of central theoretical foundations of biosemiotics and especially of zoosemiotics (the semiotic study of animals) is Jakob von Uexküll’s (1926, 1982, 1992) Umwelt theory that gives biosemiotics a subject-centred perspective. Umwelt theory describes the organism’s alignment with its environment, i.e. what is shaped by perceptual and cognitive capacities of the particular species and organized by meanings that an animal can attach to living and non-living entities in its environment. Thus Umwelt theory describes interconnections between the organism’s bodily constitution (physiology), particular environment (ecology) and meaning making (semiotics). An important principle for biosemiotics is to consider semiotic and biological processes as they appear to the organism and to treat biological communities as the sum of interconnecting Umwelten. The existence of Umwelten in 12 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. animals can also be taken as a basis for ethical programs (Beever 2012; Tønnessen 2010, 2011). As a methodological approach, biosemiotics uses a combination of wellstructured concepts, typological distinctions and research questions with a practical application to a research situation. A good example of such an approach is Thomas A. Sebeok’s (1990b: 111–112) zoosemiotic research framework that consists of six questions: 1. How does an animal that acts as a sender formulate and code a message? 2. How are messages transferred, through what channel and under what circumstances? 3. How does an animal that acts as a receiver in the communicative situation decode and interpret the message? 4. What is the possible repertoire of a specific species? 5. What are the properties of the code as used by a specific species? 6. What is the meaning of messages and, relatedly, what role does contextual information play in interpretation? Sebeok’s approach is very systematic by being based on the classical model of communication introduced by Shannon and Weaver (and as derived from the works of Roman Jakobson). Sebeok addresses different stages and aspects of animal communication and asking such questions could provide a systemic overview of the communication abilities of the given species. Such a systemic approach also makes it possible to compare communication between different species using descriptions that depart from a single conceptual framework. Kull et al. 2008 have further proposed a list of prospective research questions in biosemiotics, including: How does the world in which any individual organism finds itself appear to that organism and what are the methods that allow studying of such subjective worlds (Umwelten)? What are the general biological functions that are made possible through the phenomenon of semiosis? What are the major modes of biosemiosis? How can anything that initially does not have a function obtain a function? These questions clearly differ from the type of research questions asked in mainstream evolutionary biology, which also leads to different research hypotheses and results. To bring some examples of practical research where biosemiotics has been applied: there is an ongoing search for minimal biological entities with semiotic competence (e.g. “autocell” model in Deacon 2006, see also Deacon 2012). Additionally, biosemiotic concepts have been fruitfully applied in landscape ecology to study the engagement of different species with the environment, including their use of resources, interaction and conflicts with one another and with human influence (Farina 2008; Farina, Belgrano 2006). Biosemiotics has also had many ramifications 13 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. in literary and cultural studies (Coletta 1993, 1999; Maran 2014a; Siewers 2011; Tüür 2009; Wheeler 2008). 1.2 Defining biological mimicry 2 The common everyday understanding of a concept or phenomenon can often be found in dictionaries or textbooks. For heuristic reasons, I will start by introducing mimicry with these general sources and thereafter, point out limits and inconsistencies of everyday definitions, and will dig deeper into logics of the mimicry. Dictionaries and reference books most often explain mimicry by resemblance; sometimes other clarifying criteria such as concealment, protection or mistaken identity are added. Mimicry can be defined, for instance, as: “the close external resemblance of an animal or plant (or part of one) to another animal, plant, or inanimate object” (NODE 2001: 1175); “the resemblance, through natural selection, of one organism to another or to a natural object, as a natural aid in concealment” (AHD 1981: 834); “the resemblance shown by one animal species, esp. an insect, to another, which protects it from predators” (Collins 1994: 993); “the superficial resemblance that an organism may show to some other animate or inanimate structure, and which serves as a means of concealment” (Trident Webster’s 1995: 635). As a biological phenomenon, mimicry includes many dimensions and phenomena that everyday definitions and understandings do not usually emphasize. For example, common descriptions of mimicry are often limited to the resemblance of colours and forms. Many cases of mimicry are indeed visual. Mimicry studies were launched with the descriptions of visual resemblance between butterfly species (Bates 1862). But mimicry is not limited to visual perception. It can take place in auditory, chemical, tactile or any other channel and frequency that animals use for communication. Related to that, besides being a similarity of body structures, mimicry can also be dynamic and include resemblances of gestures, movements and action patterns. Another widespread understanding is that mimicry is predominantly a defence behaviour against predators. Although most mimicry cases are indeed based on predatory relations, mimicry exists in other ecological relations and functions, such 2 This chapter is partially based on Maran, T. (2007). Mimicry. In: Bouissac, Paul; Lewis, Ann (Eds.). Semiotics Encyclopedia Online. E.J. Pratt Library, Victoria University. http://www.semioticon.com/seo/. Used with permissions. 14 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. as symbiosis, parasitism or competition. Many mimicry relations also make use of social relations between different individuals of the same species, including sexual relations between mates, fostering behaviour between parents and offspring or between different individuals in a social species (e.g. in social insects). The third quite common misunderstanding about mimicry is that the object of imitation should inevitably be a specific species or object. In addition to close resemblances with specific biological species (as is common in mimicry between butterflies), more abstract features such as mammalian eyes (eye-spots in many caterpillar and fish) or specific movements (the worm-like movements of an outgrowth of anglerfish) are imitated. Most research in mimicry has been done in different biological paradigms. This has also shaped the hypotheses raised and questions asked in mimicry studies. Biologists have paid much attention to the evolutionary aspects of the phenomenon— the influence of mimicry resemblance to the participants in terms of evolutionary success and selection advantage. Evolutionary views have also influenced the definitions used and concepts chosen to explain mimicry. For instance, the British entomologist Richard I. Vane-Wright defines mimicry as follows: “Mimicry occurs when an organism or group of organisms (the mimic) simulates signal properties of a second living organism (the model), such that the mimic is able to take some advantage of the regular response of a sensitive signal-receiver (the operator) towards the model, through mistaken identity of the mimic for the model” (Vane-Wright 1976: 50). The three participants of this definition—mimic, model and operator or (signal) receiver—form the obligatory part of most contemporary theoretical accounts of mimicry. To emphasize the relatedness of the participants in mimicry, some authors (e.g. Wickler 1965, 1968: 239–242) use the notion of mimicry system to indicate “an ecological setup that includes two or more protagonists, performing three roles” (Pasteur 1982: 169). Different authors and directions of study tend to focus on some particular participant and emphasize its position. For instance, in the works that rely on NeoDarwinism and Modern Synthesis, more emphasis is put on the cost and benefit of the participating species, fitness and manipulation. Delbert Wiens, a specialist of plant mimicry, emphasizes the evolutionary approach by introducing the Neo-Darwinian concept of fitness to the mimicry definition: “I define mimicry as the process whereby the sensory systems of one animal (operator) are unable to discriminate consistently a 15 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. second organism or parts thereof (mimic) from either another organism or the physical environment (the models), thereby increasing the fitness of the mimic” (Wiens 1978: 367). Andrew Starrett develops this view further by introducing gained fitness as the bases for conceptual unity of mimicry, crypsis, social imitation and other types of resemblances (Starrett 1993). Some radical Neo-Darwinian approaches explain mimicry in terms of manipulation or exploitation: “natural selection has selected for a resemblance between members of one population A and a second population B, thereby allowing one population to exploit the other” (Hauser 1998: 271). Systematic mimicry studies were launched together with Darwinian evolution theory in the 1860’s, although single examples of mimicry were known much earlier. It was foremost thanks to the two biologists, Henry W. Bates and Alfred R. Wallace, working in tropical areas, that mimicry became a topic to be studied in the biological science (Bates 1862; Wallace 1871). It was especially Henry William Bates, who in the Amazonian basin observed a detailed similarity between the butterfly species of two different families of the Heliconidae and the Pieridae that might have resulted from the coevolution of palatable and unpalatable species under variation and natural selection. Bates interpreted mimicry as a supporting argument for the Darwinian evolutionary view of nature. Since that time, mimicry studies have been intertwined with the development of evolution theory, and interest toward mimicry has followed the ups and downs of Darwinian thought (see Kimler 1983). Besides being a theoretical issue, mimicry has been an important object for empirical studies in modern biology. The main directions in mimicry studies include: dynamics of mimic and model populations in various selection situations and environmental conditions studied both in nature and in computer modelling; behaviour of predators and other signal receivers with regard to mimics and models, receivers’ abilities to discriminate and learn differences between mimics and models; and variability of mimicry features of species, including genetic and geographical variability of mimics and models (see Brower 1988; Ruxton at al. 2004). Although most biological studies focus on the evolution of mimicry and limit themselves to a physical similarity between organisms, there are also approaches that emphasize communicative aspects. For instance, a German zoologist Wolfgang Wickler provides a rather semiotically oriented description in the entry of the Encyclopædia Britannica: “Mimicry is a biological phenomenon characterized by the 16 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. superficial resemblance of two or more organisms that are not closely related taxonomically. This resemblance confers an advantage—such as protection from predation—upon one or both organisms through some form of ‘information flow’ that passes between the organisms and the animate agent of selection” (Wickler 1998: 144). Richard Vane-Wright also emphasizes the roles of the communicative process and interpreting organism in mimicry: “[…] the information flow concerning the resemblance between the model and the mimic mediated via the signal-receiver […] is the agent which brings about the mimetic resemblance, or at least maintains or improves it” (Vane-Wright 1976: 28–30). In emphasising communicative and semiotic aspects even more, we can specify mimicry from a semiotic viewpoint to be not a resemblance of one organism to another but rather a resemblance of its messages (features or signals) to the messages originating from another being (that usually belongs to a different species), to some feature of the environment, or to generalizations of either of those. Such resemblance should be recurrent and confusing to the third participant in a communicative relationship, the result of which should be relevant to the organism that emits mimetic messages. Unlike most biological definitions of mimicry, the specification given here is not based on the conceptual grounds of the evolutionary process. Rather, it emphasizes that mimicry is a communicative and behavioural situation—the choice that has many implications for practical analysis as we see later in this book. There are alternative explanations for mimicry resemblances that do not rely on natural selection, fitness or other Darwinian concepts (for an overview of the history of Non-Darwinian mimicry studies, see Komárek 2003). An old tradition rising from German biological philosophy explains mimicry as a coincidence due to the limited number of structural combinations in living organisms (Eimer 1897). Another quite common approach explains mimicry resemblances as the result of the influence of the physical conditions of species that live in similar environments (e.g., Stephenson 1946). Evolutionary developmental biology seems to have given some empirical support to the understanding that mimicry resemblances may also have nonheritable causes (such as the influence of environmental conditions on the early ontogenetic development of butterfly wings, French 1997; Naisbit et al. 2003; Nijhout 1984). Finally, there have been opinions that the functionality of mimicry resemblances is simply human over-interpretation (Heikertinger 1954). 17 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Based on the short overview given here, two conclusions can be made: mimicry is a biological topic with a long and rich history, and there is a variety of different approaches to mimicry studies. This point is important for a semiotic understanding of mimicry for paradigmatic reasons. As an argument supporting evolutionary theory, mimicry has historically been a ground for many contested disputes. Here I would like to approach mimicry without relying on this historical and theoretical baggage as something self-evident. The way forward is to scrutinize mimicry critically as a scientific concept rooted into practices and theories of biology and at the same time, consider it as biological phenomenon that derives from the semiotic and ecological relations of different living organisms. Both mimicry as a concept and mimicry in its own rights have a certain ontological reality, reality that in both cases is constrained and contextualized by local environments of scientific discourse and living nature. As a concept and as a phenomenon, mimicry is changing in time and is dependent on the activities of the partakers, which also brings along the changing dynamics between these two poles. The diversity of different mimicry theories and types indeed appear to support a more heterogenic style of reasoning. 18 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 2 FIRST EXCURSION: THE HISTORY OF THE MIMICRY CONCEPT 3 In order to study mimicry in biology or semiotics, it would be useful to have an overview of the depth of meaning and different ramifications of the concept. Historically, mimicry as a concept derives from far outside biology and belongs to the concept family of mimesis, mime, imitation, etc. It is important to note here that mimicry as a peculiar biological phenomenon also has a connection to the human experience of misperceiving environmental objects or animals and to our abilities of imitating sounds and gestures of other organisms. Mimicry is related to the general ways in which we use our perceptual and cognitive capacities to make the environment meaningful and how these capacities can be directed, tricked or manipulated. Natural phenomena with several partially contradicting properties has probably been attractive to the human observer since prehistoric times. American anthropologist of religion Stewart E. Guthrie has described the existence of objects that are hard to classify as a cause of animistic belief systems (Guthrie 1993). An ability to perceive something as something else appears to have a strong linkage with illusion, belief, magic, imitation and imagination. Mimicry also has a relation to artistic creativity—mimesis—which is also supported by the etymology and history of the mimicry concept. The notion of mimesis (mimēsis) has roots in Greek Antiquity drawing originally from human imitation of natural sounds. In the earliest written works of Ancient Greece that contain the notion of mimesis, it has been used in diverse contexts to indicate the particular characteristics of the object or the phenomenon. For instance, in the extant fragment of Aeschylus’s tragedy Edonians, the sound of musical instruments has been described as mimetic, resembling the voices of roaring bulls (Halliwell 2002: 17). In the time of Plato and Aristotle, ‘mimesis’ emerges at the centre of various philosophical debates concerning metaphysics, moral issues, arts and human nature, etc., which has ensured the idea a place at the heart of Western thought for centuries, being especially relevant for the performing arts (Gebauer, Wulf 1995). The works of Antique authorities later became a common source to refer to when 3 This chapter is partially based on following sources: Maran, T. (2003). Mimesis as a phenomenon of semiotic communication. Sign Systems Studies (University of Tartu Press), 31.1, 191–215; Maran, T. (2007). Mimicry. In P. Bouissac, A. Lewis (Eds.). Semiotics Encyclopedia Online. E.J. Pratt Library, Victoria University. http://www.semioticon.com/seo/. Used with permissions. 19 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. using the notion of mimesis, and also today Plato’s “Cratylus”, “Republic”, “Sophist” and “Laws” or Aristotle’s “Poetics” and “Rhetoric” have quite often been taken as the point of departure in historical overviews and even in conceptual analyses. Aristotle especially uses the notion of mimesis as an intrinsic component of the artistic process in poetry, fine art, music and dance (Aristoteles 1997: 1.1447al3-28). To understand the concept of ‘mimesis’ and its different interpretations, it is important to emphasise the historical link between ‘mimesis’ and actual performative and artistic activities. ‘Mimesis’ has never been a pure theoretical category. For instance, philosopher Günter Gebauer and anthropologist Christoph Wulf (1995) in their monograph “Mimesis: Culture—Art—Society” stress the link between mimesis and practical embodied knowledge. The roots of mimesis lie in the oral tradition and as such it is the essence of mimesis to be dynamic and to include body-related motions, rhythms, gestures and sounds (Gebauer and Wulf 1995: 316). They emphasise that mimesis originates from practice, and therefore it is in the nature of mimesis to overcome any theoretical restrictions and structural frameworks. The decreasing of that dynamism and the coalescence of the notion of mimesis in Western thought is primarily connected with the advancement of literary culture and the related connection between mimesis and copying or reproduction. In regard to the contemporary use of the notion and also in relation to biological mimicry, the tension between the creative and static aspect appears to be a significant issue. Historian of aesthetics Stephen Halliwell regards the period when ‘mimesis’ was translated into Latin and ‘imitatio’ was chosen as an equivalent to be the decisive turning point in the history of the concept. Later on, in the Middle Ages and Renaissance, ‘imitation’ and its parallels in other languages were used to indicate the concept. Halliwell argues that translation changed the nature of the concept considerably, reducing it for centuries to mere imitation with negative connotations. He writes: No greater obstacle now stands in the way of a sophisticated understanding of all the varieties of mimeticism, both ancient and modern, than the negative associations that tend to colour the still regrettably standard translation of mimesis as “imitation”, or its equivalent in any modern language […]. Although it cannot be denied that the greater part of the history of mimeticism has been conducted in Latinized form (i.e., through the vocabulary of imitatio, 20 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. imitari, and their derivatives and equivalents), it is now hazardous to use “imitation” and its relatives as the standard label for the family of concepts. (Halliwell 2002: 13). Parallel to its use in biology, the concept of mimicry has also been adapted in the humanities, partly through its use in biology and partly from the ancient concept of mimesis directly. For instance, in postcolonial discourse, anthropologist Graham Huggan has described mimicry as an aggressive and disruptive imitation, which has the purpose of interfering, ridiculing or subordinating the subject under imitation (Huggan 1997: 94–95). Another well-known postcolonial writer Homi Bhaba uses the notion of mimicry to describe the situation of postcolonial culture “as almost the same, but not quite” and characterizes it by the designations “ambivalence”, “ironic compromise”, and “incompleteness” (Bhaba 1994: 122–127). In human psychology, the concept of mimicry is often used to indicate a deceptive or unconscious imitation between humans, especially with reference to facial gestures and body movements (e.g., Gambetta 2005; van Baaren et al. 2004; Zepf et al. 1998). It appears that the conceptual family of mimesis, mimicry, imitation and others has lost much of its historical meaning as an active and creative process and has become more of an equivalent to a mechanical coping and reproduction, a systematic device of creating resemblance. Such an approach is perhaps most evident in the field of memetics, which has attempted to describe culture as being composed of fixed multiplying units (Blackmore 1999; cf. Deacon 1999; Bouissac 2000, 2001). In the memetic view, the artistic and also interpretational aspect of mimesis is reduced in the extreme, and what is left is a pure function of copying. We can see, however, that such use of the concept is not supported by the general history of the notion and also, that there are alternative connections between the biological and cultural realm that place their emphasis on the creative aspect of mimesis. For example, there are different approaches of biomimicry as technical engineering (Benyus 2002) that is inspired by forms and processes of nature. The long and rich history of the mimicry concept may bond together notions that at first glance have different meanings and are spread between various fields of study. The field is rich with antagonistic standpoints and traditions, and there are also loans, rediscoveries and metaphoric usages that could transcend millennia. For instance, French zoologist Georges Pasteur, when classifying different mimicry types, 21 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. distinguishes Aristotelian mimicry by referring to a passage from Historia Animalium, in which Aristotle describes how a brooding bird may pretend to be wounded if it encounters a dangerous creature near its nesting place (Pasteur 1982: 190). Another such terminological crossing is lending the concept of ‘mimesis’ from aesthetics to biology. For denoting the imitation of the forms of nature, ‘mimesis’ was first used, to my knowledge, by German biologist Franz Heikertinger (1925), and later became a common practice. On such occasions, intellectual traditions with different origins meet to produce new layers of meaning. 22 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 3 THE STRUCTURE OF MIMICRY A property that makes mimicry both fascinating and difficult to comprehend is its diversity. Mimetic resemblances can occur as colours and forms in the visual medium, as imitations of hissing, buzzing and other sounds, or as similarities of chemical components in pheromones. Mimicry can be based on different ecological relations (predation, parasitism, symbiosis, competition) and the number and composition of involved species can vary to a great extent. Mimicry can take place inside organisms at the cellular level, a phenomenon that is known as molecular mimicry. In some cases, mimicry requires the coordinated behaviour of several individuals to create or enhance the mimetic effect (e.g. myrmecomorphic jumping spiders aggregating to emphasise their resemblance with ants, Nelson and Jackson 2009). Indeed, there seems to be little in common between the cases of fully behavioural mimicry, such as the mimic octopus Thaumoctopus mimicus, in which the cephalopod uses its tentacles to create sporadic imitations of flatfish, sea snakes, lionfish and other sea creatures, and the stillness of the perfect resemblance between the eggs of the common cuckoo Cuculus canorus and those of reed warblers, pipits, redstarts and other hosts species, whose nests the cuckoo uses to lay its eggs. The huge diversity of mimicry resemblances has probably led mimicry theory to develop, on the one hand, towards typological and classificatory reasoning and on the other hand, towards structural thinking and formalisation of mimicry as a notion. The challenges of the diversity at the object level have been met with cataloguing and organising various mimicry cases and distilling the conceptual core of mimicry that would make it possible to bridge these different cases. A typological approach is necessary for having a comparative approach to different mimicry cases and, in fact, for having a broader comprehension of mimicry as a phenomenon at all. At the same time, as American biologist Adrian Wenner has reminded us in his account of animal communication (Wenner 1969), typological descriptions are never neutral or innocent, as they start with a logical basis and criteria that determine the limits of typology (what is counted as mimicry) and possibilities for comparison (what features are used as a basis of comparison). Typological approaches to mimicry have a story to tell about what features are accepted as meaningful characteristics in nature and what are left below the threshold of significance. Making categories is always a semiotic procedure of selecting, picking up and leaving out. 23 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 3.1 Mimicry types 4 There exist many old and well-studied types of mimicry such as Batesian, Müllerian and aggressive mimicry that are commonly accepted by the scientific community. In addition, one can find many novel mimicry types that an individual researcher or school has proposed and used occasionally. Opinions about the delineation of the mimicry concept and its relations with neighbouring phenomena also differ. Most authors exclude, for instance, camouflage coloration from mimicry, but the position of biological mimesis, understood as a resemblance with an element of the non-animate environment (stone, twig, leaf, etc.), is not so clear-cut. Some authors consider this as a subtype of mimicry whereas others regard it as a separate phenomenon (Endler 1981; Pasteur 1982: 171–173). The list that is presented hereafter is not exhaustive of all mimicry types, but is meant to give an overview of the diversity of mimicry types and concepts used for naming these phenomena. Historically, the oldest and best-known mimicry type is Batesian mimicry, named after the British entomologist Henry Walter Bates, who in his fieldwork in the Amazon River described the resemblance of many butterfly species from the families Papilionidae, Pieridae, Lycaenidae a.o. to Ithomia and Methona butterflies (family Nymphalidae) (Bates 1862). Batesian mimicry is the resemblance of an edible and harmless species to some poisonous or otherwise non-edible species that signals its unsuitability to possible predators by aposematic colouring or by other conspicuous signs. Besides tropical butterflies, Batesian mimicry is well documented not only on other insects such as hoverflies (Howarth and Edmunds 2000; Waldbauer 1988) and ants (Ito et al. 2004), but also on snakes (Wüster et al. 2004), fish (Randall 2005: 301– 310), and even on plants (Augner and Bernays 1998). A contemporary of Bates, German entomologist Fritz Müller (1878) provided a different explanation of deceptive resemblances. He showed mathematically that it is advantageous for several unpalatable species to share the same aposematic coloration because the predator then learns more quickly which, in turn, strengthens the effect of this specific colour pattern. This phenomenon has become known as 4 This chapter is partially based on Maran, T. (2007). Mimicry. In: P. Bouissac, A. Lewis (Eds.), Semiotics Encyclopedia Online. E.J. Pratt Library, Victoria University. http://www.semioticon.com/seo/. Used with permissions. 24 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Müllerian mimicry. Some authors argue that this is not actually mimicry and have suggested the term Müllerian convergence, because it is not possible to indicate which species is the mimic, which is the model and thus who resembles whom (Wickler 1965; Pasteur 1982: 193–194). Also, species with an edibility spectrum from palatable to highly noxious may share a similar appearance. Such groups are known as Batesian-Müllerian spectrum, quasi-Batesian mimicry or arithmetic mimicry (Huheey 1976; Mallet and Joron 1999; Someren and Jackson 1959). The edibility spectrum can also take place inside one species, e.g. in the case where butterfly larvae eat on plants that have different toxicity levels. Such mimicry type has been called auto mimicry (Ruxton 2004: 176–182). In aggressive mimicry (also known as Peckhamian mimicry) the mimic does not belong to prey animals but is the predator. Aggressive mimicry helps a predator organism approach its prey or lure the prey to approach the predator. Aggressive mimicry became known by the works of American entomologist Elizabeth G. Peckham, who described a similarity of the spider Peckhamia picata to ants as an adaptation for hunting (Peckham 1889). Such adaptations are common in many carnivorous animals of different groups: spiders (Pekár and Křál 2002), fireflies (Lloyd 1986), some fish (Even and Geoffrey 2004) and carnivorous plants (Moran 1996). The object of aggressive mimicry can be: 1) receiver, who is also a prey animal (red wolf fish Erythrinus erythrinus mimicking its prey killifish Rivulus agilae (Brosset 1997); 2) common food object of the receiver (flatfish Asterorhombus fijiensis, who moves a special membrane in the front of its mouth which is attractive to smaller fish on which the flatfish preys upon (Amaoka et al. 1994); 3) symbiont of the receiver as a false cleaner fish Aspidontus taeniatus, that resembles cleaner wrasse Labroides dimidiatus, which has a symbiotic relationship with many larger fish species by searching and eating their ectoparasites (Wickler 1968: 157–176). The term reproductive mimicry covers various examples in which resemblance aids the mimic in reproduction. Reproductive mimicry is common in many orchids, for instance in fly orchids Ophrys (Schiestl 2005: 257–258). Blossoms of these plants resemble insects in form and smell (Ayasse, Schiestl et al. 2003). They deceive male insects into copulating with the blossoms and the pollen becomes transferred during this pseudo-copulation. Reproductive mimicry also appears in brood parasitism, such as in the resemblance of cuckoo eggs to the eggs of their passerine host species (Aviles and Møller 2004; Payne 1977: 8–10). Intraspecific mimicry, understood as a 25 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. mimicry system occurring within one species, often has a reproductive function as well. A good example of intraspecific mimicry is the resemblance of the anal fin pattern of the male African mouth-brooding fish Haplochromis burtoni with the eggs of the same species. The resemblance has an important role for the reproductive behaviour of the species, as the female fish has a specific fostering behaviour of hatching eggs in the mouth. While spawning, the female fish tries to gather along with the eggs the pattern of the male fin into its mouth, and thus catches milt for fertilization (Wickler 1968: 222–227). Within intraspecific mimicry, some authors distinguish sexual mimicry as a specific resemblance between members of different sexes that has functionality in intraspecific communication (e.g. genital masculinization in female spotted hyenas, Muller and Wrangham 2002). A similar concept is self-mimicry, where a part of an animal body resembles another part of it. Rainey and Grether (2007) introduce the concept of competitive mimicry in which the mimic, due to its resemblance, gains greater access to a defended resource than closely related non-mimics that also compete for the resource. The model in competitive mimicry can belong to the same or different species as mimic and the model can be the competitor or other related party (e.g. predator). Besides these categories, there are many mimicry types that some researchers have used occasionally or that are introduced to denote a specific concept or hypothesis. Behavioural mimicry describes the resemblance of the dynamic behaviour of mimic to model (McIver and Stonedahl 1993); instances of behavioral mimicry are locomotor mimicry and escape mimicry (Srygley 1999). Satyric or imperfect mimicry is a partial or approximate resemblance or combination of similar and dissimilar features (Howse and Allen 1994). In satyric mimicry (e.g. in hoverflies and Saturniid moths), mimetic signs are presented in an unexpected context, or different types of mimetic signs are presented together that lead to an ambiguous interpretation space (Howse 2013). This mimicry type is believed to operate by causing confusion in signal receivers. A similar concept is polymorphic mimicry, in which case the single species exhibits different functional resemblances with different models (Joron 2005). Fully behavioural mimicry also includes vocal mimicry, a concept used primarily to denote the ability of many birds to imitate songs of other species and environmental sounds (Goodale and Kotagama 2006; Wickler 2013). In historical mimicry or aide memoire mimicry, the mimic and model may have temporally different population dynamics and the mimic benefits from the receiver’s earlier experience or memory 26 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. (Rothschild 1984). Mertensian or Emsleyan mimicry is the name for the mimicry complex in which deadly poisonous coral snakes Micrurus imitate moderately poisonous but more numerous species. Such a reverse mimicry system is believed to exist because the learning process is quicker in cases where the receiver remains alive after the encounter (Wickler 1968: 111–121; Mertens 1966). A very ambivalent category is floral mimicry or plant mimicry, which unites mimicry cases in which the mimic is a plant (Roy and Widmer 1999; Dafni 1984; Lev-Yadun 2014). Plants actually take part in many of the above-mentioned mimicry types, such as Batesian, aggressive or reproductive mimicry, and there are also mimicry types that are specific to plants (see Wiens 1978: 369, 371). In dispersal mimicry, the propagules of a plant resemble fruits or other food sources to signal receivers that are usually birds. In weed mimicry, weeds (false flax Camelina sativa, common wild oat Avena fatua) or secondary crops resemble the crops (flax, wheat) with which they grow (e.g. Wickler 1968: 40–45). In weed mimicry, humans and, in modern times, agricultural machinery serve as signal receivers. A close concept to mimicry is mimesis, in which the mimic resembles, with its body form and patterns, the physical or living element of the environment (stones, twigs, fungi, plant leaves, Heikertinger 1925). Georges Pasteur has distinguished two principal types of mimesis, cryptic mimesis in which the mimic resembles a common element of the environment that does not have any specific meaning for the receiver, and phaneric mimesis (or masquerade) in which case the model is well discernible but unpleasant to the receiver (e.g. bird faces, insect carcases, Pasteur 1982: 183). A related concept is plant-part mimicry, where an organism (usually an insect) resembles a stick, leave, bark or other floral object that is common in the environment and that is not sought after by the predators (Robinson 1981: 16). Mimicry and mimesis are also often related with other protective adaptations such as countershading, disruptive coloration, false heads and the like. 3.2 Mimicry in relations to other adaptations In all of its diversity, mimicry does not exist in a vacuum, but is surrounded by other natural phenomena. Mimicry is often combined or correlated with other phenomena (convergence, polymorphism), or there may exist a functional relation between them—for instance, escape behaviours may be used along with fixed mimicry 27 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. resemblances. Many authors have proposed criteria that would allow us to distinguish mimicry from other similar or connected adaptations. Such attempts often lead to typological accounts that contextualise mimicry among other protective or visual adaptations of animals. For the sake of clarity, we can call these typological approaches meta-typologies, as distinguished from the typologies of mimicry analysed in the next subchapter that aim to organise relations between different mimicry types. One of the earliest authors who treated mimicry and other defective adaptations in a complex and systematic manner was British zoologist and evolutionary scholar, Edward B. Poulton. In 1890, Poulton published a monograph „The Colours of Animals. Their Meaning and Use, Especially Considered in the Case of Insects”, where he proposes a classification of animal colourations that proceed from the question of whether the colouring makes an animal similar to another organism or environmental object (apatetic colours), or whether its aim is to signify and express some property of the organism itself (sematic colours, see table 3.1). The group of apatetic colours is further divided into concealing colours (cryptic colours in Poulton’s termininology) and deceptive colours (pseudosematic colours). Under concealing or cryptic colours, he distinguishes between defensive (procryptic) and aggressive (anticryptic) resemblances to environmental colours. As opposed to concealing colours, deceptive or pseudosematic colour similarities are those in which warning colours and other distinctive colourations are mimicked. Pseudosematic colours are further divided into defensive (pseudaposematic) and aggressive (pseudepisematic) types. The name of the first class refers to deceptive similarity to warning colouration and the name of the second class to deceptive similarity to some species-specific properties. At the first logical level that is related to the properties of the organism itself (sematic colours), Poulton further distinguishes warning colouration (aposematic colours) and features that are used primarily for communication among members of the same species (episematic colours or recognition marks). In addition to these types, Poulton brings out a separate class of colours that are used in courtship rituals and other forms of intraspecific communication (epigamic colours), and are therefore the objects of a different type of selection mechanism—Darwinian sexual selection. Table 3.1. Classification of animal coloration (Poulton 1890: 338). 28 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. I Apatetic colours A Cryptic colours II Sematic colours B Pseudosematic colours 1 Procryptic colours 1 Pseudaposematic colours 1 Aposematic colours 2 Anticryptic colours 2 Episematic colours 2 Pseudepisematic colours III. Epigamic colours Poulton’s classification is remarkable in its clear structure and systematic use of concepts derived from Greek roots. From the many original concepts that Poulton introduces, only cryptic and aposematic (warning) colouration are still commonly used today. 5 Poulton’s approach especially focuses on living organisms and their appearance, which is different from many later typologies with a more theoretical flavour. Poulton’s observation of the special relationships between defensive mimicry, warning colouration and aggressive mimicry on the one hand, and speciesspecific properties on the other hand is also very accurate. In most cases of aggressive mimicry, mimics indeed imitate the colour patterns and body forms of their models in a species-specific way (myrmecomorphism by several spider species serves as a good example). Some ideas from Edward Poulton’s typology were later elaborated by Czech philosopher and historian of science, Stanislav Komárek, a leading specialist in the history of mimicry studies and the author of the outstanding book “Mimicry, Aposematism and Related Phenomena. Mimetism in Nature and the History of its Study” (Komárek 2003). He combines three properties—conspicuousness, edibility and resemblance. Using these criteria, he distinguishes four basic categories of semantic or meaningful coloration (Komárek 2003: 71): 1) aposematic colouring – (semantic, inedible, dissimilar to other species); 2) Müllerian mimicry – (semantic, inedible, similar to other species); 3) Batesian mimicry – (semantic, edible, similar to other species); 4) pseudaposematic colouring – (semantic, edible, dissimilar to other species). 5 The typology was in active use in first half of the 20th century, being discussed in length, for instance, in Carpender and Ford (1933). 29 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Differently from Poulton, who listed Batesian mimicry under pseudaposematic colouring, Komárek uses the concept of pseudaposematism to emphasise the role of uncommon characteristics and nonspecific resemblances, such as vivid speciesspecific spots and patterns on the butterfly wings that may confuse the potential predator (Komárek 2003: 71). This understanding of pseudaposematism principally treats it as an open category—if other types are constrained by a similarity to another species or by the necessity of being conspicuous and easily perceivable, then pseudaposematism is accidental but meaningful. Pseudaposematism broadens the reach of semantic colours in nature beyond the rigid and theory based types and opens it up to the spontaneusness of natural forms and patterns. Komárek’s work is grounded on the German biological tradition and has an affinity to Adolf Portmann’s philosophical biology, who emphasised the role of sematic but non-functional appearances in animals as forms of “self-representation” (Portmann 1964: 66–67; cf. Kleisner 2008). Both typologies of Poulton and Komárek find their bases in the relationship between the mimic and model species involved in the resemblance. Another possibility would be, however, to ground the typology in the relationship between the mimic and the receiver. In that vein, British zoologist Malcolm Edmunds presents in his monograph, “Defence in Animals. A Survey of Anti-Predatory Defences”, a distinction between primary and secondary adaptations (Edmunds 1974). He describes primary adaptations to be defensive adaptations that operate “regardless of whether or not there is a predator in the vicinity” (Edmunds 1974: 1). This group includes mostly species-specific and inborn adaptations like anachoresis (physically hiding by burrowing in the ground or hiding under other surfaces), different concealing adaptations (cryptic colouring, countershading, cryptic mimesis, aposematic adaptations, also Müllerian mimicry) and Batesian mimicry. Secondary adaptations, on the other hand, are processes and behaviours that are launched during an encounter with the predator and for these organisms’ behaviour, knowledge and decisions play a decisive role. As secondary adaptations, Edmunds distinguishes between withdrawal (to the burrow or under the shell), escape (evasive maneuvering, flashing colourful patterns during the escape), deimatic behaviour (deimatic postures, eyespots and false heads that are actively demonstrated by the organism), distraction of the attack (faking injuries, eyespots in butterflies and body parts that can be constricted), counter defensive behaviours and group behaviours (defensive associations between the 30 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. members of the same or different species). Malcolm Edmunds’ own long-time research topic was deimatic displays and escape behaviour, especially in praying mantis species (Edmunds 1972, 1976). Edmunds’ overview emphasizes the role of the underlying ecological relations between predatory and prey species as well as complex relations between bodily properties and behavioural displays of the species involved. In the early 1980’s, a heated discussion took place in the Biological Journal of the Linnean Society over the criteria to delimit mimicry and to distinguish between mimicry and concealing colour adaptations (mostly cryptic colouration and cryptic mimesis). The discussion started with a paper by British entomologist Richard VaneWright that proposed a criteria of mimicry to be: that the mimic imitates the signal in which the receiver has interest in and that the receiver makes a mistake in determining the mimic’s identity (Vane-Wright 1980). What was in principle at stake here, at least from a biosemiotics perspective, is the question of the receiver’s interpretational activity and frame of reference (a search image) and its relevance to the definition of the mimicry concept. The discussion, in which J. L. Cloudsley-Thompson (1981) and Michael H. Robinson (1981) also took part, was concluded and synthesised in a paper by eminent evolutionary biologist John A. Endler (1981), in which he formulated the criteria for distinguishing logically basic mimicry types and other adaptive colour patterns. Endler based this typology on three criteria: 1) whether the model is a living organism or an environmental background; 2) whether the receiver distinguishes between several (groups of) species; and 3) whether the receiver’s ability to distinguish mimics and models is dependent on the environmental background. These criteria allowed John Endler to describe six types of colour adaptations as follows: 1. Crypsis that is defined as: „a colour pattern of S is cryptic if it resembles a random sample of the background (P) perceived by R at the time and age and in the microhabitat where S is most often sought after by R.” (Endler 1981: 28). 2. Masquerade is a detailed imitation of stones, leaves, twigs, etc. The concept of masquerade includes plant-part mimicry (Robinson 1981: 16) as well as phaneric mimesis (in which the model is a clearly discernible object of the environment (Pasteur 1982: 183). 3. Batesianism covers Batesian mimicry but also reproductive and dispersal mimicry, and some cases of aggressive mimicry. The distinctive property of Batesianism, according to Endler, is that the model has specific disgusting or 31 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. attractive properties; that making the distinction between model and mimic is not dependent on the environmental background; and that the mimic has an effect on the model’s population dynamics or evolution. 4. Müllerianism is resemblance between several aposematically coloured species—it is not dependent on the environmental background but has an effect on population dynamics and evolution of the involved species. In addition to Müllerian mimicry, Endler also includes Mertensian mimicry in this category, which refers to the mimicry of coral snakes in Central America, in which more poisonous species are supposedly imitating less poisonous ones (Mertens 1966). 5. Polymorphism is understood as the existence of different morphs of the same species, where every morph resembles its specific environmental background. Having different morphs makes it more difficult for the predator to learn how to catch the particular prey item (that is, to develop a cognitive search image) and the ecological pressure of predation is divided between the different morphs based on their concealment and relative abundance. 6. Convergence, understood by Endler as the situation in which species in the same environmental background resemble one-another because they share the same receiver (e.g. predator), who, through communicative interactions and selective behaviour, shapes their evolution in a direction in which they become more similar to one-another. Endler’s typological account is based on clear, logical criteria that successfully organise relations between crypsis, mimicry, polymorphism and other types. In this typology, mimicry proper would belong to the Batesianism, whereas all other categories would describe adjunct phenomena. At the same time, all six types of adaptations that Endler describes have a semiotic character, as they take part in communicative relations and are influenced by these. All distinguished types can be filled with the real examples from nature (which is not true in many other typological attempts). From a critical side, Endler’s typology, as many other typological descriptions, cannot avoid contradiction between rigid typological schematisation and the diversity of actual mimicry phenomena in nature. This results in dividing one and the same phenomenon between several types. For instance, Endler’s typology divides aggressive mimicry into three categories: cryptic aggressive mimicry would belong to first type, detailed aggressive mimicry to the second type and aggressive mimicry, where the model is a specific animal species, to the third type (Endler 1981: 28–29). 32 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. The meta-typologies described in this subchapter depart from different logical bases (Poulton’s signifying and resembling, Edmund’s primary and secondary properties, and Endler’s background dependence), and therefore it is difficult to combine or unify these. 6 From a semiotic perspective, we may claim that communicative adaptations are complex research objects with characteristics and features that may also partly contradict each other. Perhaps it would be more fruitful to consider meta-typologies as heuristic devises that underline the specific structural properties and relations of mimicry and other connected phenomena. Thus, metatypologies by Poulton and Endler emphasise the dependence between mimicry and aposematic colours or environmental colours. This will underline the connection between mimicry phenomena and the ecological relations between organisms. Edmund’s typology on the other hand emphasises distinction between static and behavioural defensive adaptations and by doing this, points out the complex relations between ontogenetic and phylogenetic processes in mimicry. Komárek’s fourfold typology opens mimicry up to include novel and occasional colours and forms. Metatypologies also provide us with criteria of exclusion; that is, they frame mimicry by describing secondary phenomena that are not considered to belong under mimicry. 3.3 Typologies of mimicry To organise the diversity of different mimicry cases, researchers have proposed various typologies to organise the landscape of mimetic phenomena. As with all metatypologies, this is an essential question about the criteria used to categorise different mimicry cases. 7 The bases of typology can be selected differently, which will also alter affinity and distance between mimicry types. Many typologies take what could be called an ethological-ecological route and pay attention to questions of what function mimicry has for the mimic, what the ecological relationship is between species upon which mimicry resemblance is built, and what the properties are of imitated signals. Another approach that is more inclined towards Darwinian biology and evolutionary thinking focuses on questions like: what the evolutionary effects of the mimicry system on different participants (mimics, models and receivers) are and 6 Broad overviews of different visual adaptations also include Oren Hasson’s unified typology of the signals (Hasson 1997) and Andrew Starrett’s typology of the adaptive resemblances (Starrett 1993). 7 In addition, many loosely organized lists of different mimicry types have been published that do not seem to follow any typological criteria (e.g. Dafni 1984; Wickler 1968). 33 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. how each and every one of these is connected by cost-benefit-relations. Many approaches use two logical layers for building a typological account. In such approaches, the secondary criteria is often a question of what species take part in the case of mimicry and how the positions of mimic, model and receiver are filled by specimens of the different species. This issue of the combination of species again has a direct connection to evolutionary thinking as it relates to differences in evolutionary dynamics. For instance, competition in and between species has different dynamics that guide treating intraspecific and multispecies mimicries as separate from oneanother. A good example of a functionally focused mimicry typology is Delbert Wiens’ systematic description of plant mimicry. Wiens departs from the question of what the role of mimicry is for the mimic, or, to be more specific, which biological functions of the plant the mimicry helps to fulfill. The starting point for the analysis are different ecological relations: antagonistic relations like predation, where plants and animals can both fulfill the positions of predator and prey, and mutualistic, or reciprocally beneficial relations (e.g. pollination). Wiens also pays attention to the traditional mimicry types (cryptic mimesis, Batesian mimicry) and distinguishes four basic functions in plant life where mimicry becomes useful: acquiring food, defence from predators, reproduction through spreading pollen and distributing seeds (Wiens 1978: 373). German biologists Helge Zabka and Günter Tembrock (1986) also take the function of mimicy as one criterion of their typology in their paper, “Mimicry and crypsis—a behavioural approach to classification”. They distinguish between: 1) imitating signals that have a defensive function for the organism (Batesian and Müllerian mimicry and cryptic mimesis); 2) imitation that enables animals to feed (aggressive mimicry and aggressive mimesis); and 3) imitation that optimises reproduction. Zabka and Tembrock stress that in the last category, especially, the differences between plant and animal mimicry systems are radically large, as problems in reproduction for sessile and free-ranging organisms are principally different (Zabka and Tembrock 1986: 159). The second criterion used by Zabka and Tembrock is the type of mimicked signals based on the receiver’s perception. The distinction is made between imitating the signals that are relevant for the receiver (characteristics of prey, eyes of the predator, etc.) and signals that imitate nonrelevant environmental features (environmental background, foliage, void, etc.). 34 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. When describing reproductory mimicry, Zabka and Tembrock make an additional distinction between communicative and non-communicative releasers as objects of imitation. Communicative releasers are understood to be signals that have evolved with a communicative function whereas informational releasers (cues) are passive features that the receiver perceives and can be used to shape behaviour (Zabka, Tembrock 1986: 167). Zabka’s and Tembrock’s paper comes close to biosemiotics in their attention to the mimic’s aims in mimicry as well as to the functionality and communication in mimicry (e.g. by distinguishing between the relevant and nonrelevant environment for the animal receiver). 8 In the beginning of this sub-chapter, the second principal class of mimicry typologies was described as proceeding from the species combination and cost-benefit relations between species. Here the most thorough typology is proposed by British entomologist Richard I. Vane-Wright in his 1976 paper „A unified classification of mimetic resemblances” (Vane-Wright 1976). Vane-Wright proceeds from the works of German zoologist Wolfgang Wickler, who proposed the concept of the tripartite mimicry system (that will be discussed in detail in the next sub-chapter). In VaneWright’s typology, mimic, model and receiver form a static basis of the mimicry structure. Changing variables of the mimicry are: 1) the effect of the model on the receiver (positive or negative); 2) the effect of the mimic on the receiver (positive or negative); 3) the effect of the mimic on the model through the behaviour and activities of the receiver (positive or negative). These three variables allow for Vane-Wright to distinguish eight different mimicry categories that he denotes by using the following nomenclature: 1) synergic (the effect of mimic on model is positive); 2) antergic (the effect of mimic on model is negative). Both of these categories can be divided further to four subtypes: 1) warning (the effect of both model and mimic on the receiver is negative); 2) inviting (the effect of both model and mimic on the receiver is positive); 3) aggressive (the effect of the mimic on the receiver is negative, but the effect of the model on the receiver is positive); 4) protective (the effect of mimic on the receiver is positive, but the effect of the model on the receiver is negative). Developed typology allows Vane-Wright to discuss, for instance, antergic protective mimicry (which includes what is usually described as Batesian mimicry) and synergic aggressive 8 Günter Tembrock’s recent studies were explicitly related to biosemiotics. For instance, he has proposed an overview of various types of semiosis in an animal world (Tembrock 1997). 35 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. mimicry (e.g. the mimetic behaviour of the angler or monkfish Lophius piscatorius, that uses its long worm-like appendix to lure the smaller fish on which it feeds upon). This is an example of synergic mimicry, as the adaptation of the mimic also turns out to be beneficial to the model by decreasing the number of natural enemies of the marine worms (model). In addition to the distinction of effects on the mimic, model and receiver described above, Vane-Wright introduces the distribution of different species in the mimicry structure as a second level logical criterion of his typology. This allows him to distinguish between disjunct mimicry (all three participants—the mimic, the model and the receiver belong to different species), conjunct mimicry or intraspecies mimicry and three types of bipolar mimicries. In bipolar mimicry types, the model and mimic can belong to the same species (S1+S2), the mimic and receiver can belong to the same species (S2+R), or the model and receiver (S1+R) can belong to the same species. The latter case occurs, for instance, in many myrmecomorphic insects that live in ant nests, resemble ants and communicate with ants, and who thus act as both the models and the signal receivers (McIver and Stonedahl 1993; Kleisner and Markoš 2005). Vane-Wright further combines two bases into a single typology, and provides a general table in which 40 possible mimicry types are presented as number-letter combinations. For instance, the mimicry situation where eggs laid in the nests of the host species by the common cuckoo are similar to the eggs of the host belongs in Vane-Wright’s typology to the category VIIB, which denotes antergic aggressive S1+R-type two-species mimicry. For many of the types that Vane-Wright distinguishes, it is not possible to find real life examples in nature. Although Richard I. Vane-Wright’s typology is very thorough and gives a good overview of the diversity of mimicry cases, it has not found much use in the following research literature. In practical mimicry studies, traditional categories such as Batesian, Mertesian or Peckhamian mimicry still appear to be handier compared to the Vane-Wright’s systematic and very complex nomenclature. Vane-Wright’s typology has also been criticised as being too artificial and projecting logical distinctions onto natural phenomena beyond necessity. Zabka and Tembrock point out that using species as a basic concept of the mimicry typology limits its use to the species level and does not allow for describing mimicry systems in which it is not possible to specify participants down to the species (Zabka and Tembrock 1986: 172). The identity of the mimic, the model and the receiver may be fixed at the species 36 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. level, but inasmuch as mimicry is based on semiotic relations, this is not a necessary condition as we will see later in the discussion on abstract mimicry. Having a predominant focus on the species level appears to be another heritage of evolutionary biology, as it is the level of the species (or population) that are considered to be the carriers of adaptations and thus are primary objects of the evolutionary processes. The next author that should be discussed in relation to typologies of mimicry is French zoologist Georges Pasteur (1982). He takes Wolfgang Wickler’s systematic treatment of mimicry and Vane-Wright’s complex typology as points of departure, and connects these to practical studies. The first basis of Pasteur’s typology is the question of whether the receiver (Pasteur uses the concept of “dupe” in this position) has any interest in the model or not, which in principle comes close to Zabka’s and Tembrock’s distinction between relevant and non-relevant environment (Pasteur 1982: 182–184). In the case of camouflage and cryptic mimesis, the model is indifferent to the receiver and based on this, Pasteur describes these phenomena as adjunct to mimicry. In situations where the model has some relevance to the receiver, it can be either attractive or disgusting. With some smaller exceptions, this distinction corresponds to the difference between protective (model is disgusting to the receiver) and reproductive mimicry (model is attractive to the receiver). Pasteur obtains the second layer of typology from the works of VaneWright—the distinction between disjunct, conjunct and bipolar mimicry systems based on the question of how different species fill the roles of the mimic, model and receiver. The third basis of Pasteur’s typology is the function of the mimicry relation in regard to ecological relations between the mimic and model. Here he distinguishes seven different categories: aggressive, aggressive/reproductive, reproductive, reproductive/mutualistic, mutualistic, commensialist, and protective. Combining these three different criteria allows Pasteur to create a very complex mimicry typology. Differently from Vane-Wright, however, Pasteur does not aim to map all possible configurations, but uses partly classical, partly originally construed mimicry types that he names based on the discoverer or literary source (for instance Browerian mimicry, Dodsonian mimicry, Vavilovian mimicry, etc). Pasteur describes altogether 18 mimicry types and illustrates these using real-life examples from the research literature. His author-based nomenclature follows the historical tradition of naming mimicry types, and has therefore been used quite a lot in later research literature (e.g. Barrows 2011; McElroy 2014). 37 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. When comparing different attempts to typologise mimicry, four basic criteria or questions can be brought out that are used as bases to establish the typology: 1) What is the nature of the signal or feature being imitated (does the receiver have interest in this or not, is it communicative or not; is it background, warning colouration or feature specific to the given species)? 2) What is the function of the mimicry to the mimic (does the mimicry help the mimic in protecting itself, catching prey or reproducing, is the mimicry based on commensialism, mutualism or other types of ecological relations)? 3) What is the evolutionary or ecological effect on the participants (what are the cost/benefit relations between mimic, model and receiver or the effect of the mimicry on every participant)? 4) Who are the participants of the mimicry systems (what are the possibilities for representatives of different species to become combined in the mimicry system (also by distinguishing intraspecies mimicry)? These different criteria can be combined into a more precise, basic question motivating the mimicry typologies as follows: who mimics what for whom, on what cause and with what effect? Proposed typologies well characterise various mimicry types through their relations and differences, but at the same time all typologies reviewed in this subchapter have their problematic aspects. The problem can appear as an inconsistency of the typology, that compels one and the same mimicry type to be divided between different categories, and also as a situation where a specific mimicry case may resist classification or fall outside of the limits of typology. Furthermore, in many species, different mimicry relations are expressed simultaneously and in combination. I discuss these different logical problems of mimicry typologies in detail after introducing the tripartiate mimicry model proposed by German biologist Wolfgang Wickler in the next subchapter. Problems applying mimicry typologies indicate that mimicry in nature is a complex phenomenon that does not submit well to rigid classificatory schemas. However, I think there is even more to this phenomenon. I suggest that contradicting typological attempts is not an accidental feature of mimicry, but tells us something essential about what mimicry is. Namely, at the ontological level, mimicry is not a solid class of entities with a common origin that developed from the same initial conditions, or follows the same biological laws. Rather, mimicry is a secondary phenomenon that can spontaneously emerge in very different natural conditions, given that the participants (mimic, model and receiver) start interacting with one-another through deceptive communication. There appears to 38 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. be a great deal of spontaneity in many mimicry cases. Think for the moment about the acts of laying eggs in the nest of another species or trying to copulate with a plant instead of a member of one’s own species. Such cases appear to expel rationality of the biological or evolutionary processes and are rather based on occasional mistakes or on the creative interpretation on behalf of the animal subjects. Therefore, specific mimicry cases are, in fact, very independent from one-another while at the same time dependent on the particular communicative setup and context. What follows is that we should not take mimicry as a combination of physical forms and categorisation arising from this, but we should rather focus on what the semiotic possibilities are for the deceptive resemblance to emerge and function. The semiotic approach is not able to solve all the problems of the structural and typological descriptions of mimicry, but it better articulates causes for these problems and shows some novel ways in which a structural approach can be used more productively for modelling mimicry systems. 3.4 Mimicry systems—Wolfgang Wickler’s account Typologies of mimicry have developed in parallel with the emergence of understanding mimicry as a formal system. Such a systemic approach to mimicry becomes widespread in research literature starting from 1960’s. The advancement of the systemic approach to mimicry could be related to the general tendency of that time to use mathematical and cybernetic approaches for analysing biological phenomena, as well as to the impact of theoretical ecology that modelled relations between species in ecological communities. Under the systemic approach to mimicry, I include authors and works that clearly distinguish and pay attention to three participants of mimicry— the mimic, the model and the receiver—and treat these as an interconnected mimicry system. The systemic approach also tends to see relations between the mimic, model and receiver as central factors in the evolution and dynamics of mimicry. The systemic approach thereby purifies the mimicry system and brings the unity of the mimic-model-receiver setup to the foreground in comparison to other evolutionary and ecological processes. A central figure in developing the systemic approach to mimicry is German zoologist Wolfgang Wickler with his much-cited paper “Mimicry and the evolution of animal communication”, published in the journal Nature in 1965. Wickler was a colleague of Konrad Lorenz in the Max Planck Institute for Comparative Ethology 39 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. and a true member of the classical ethology school. In his paper, Wickler raises a problem that earlier accounts of mimicry were mostly derived from observations of specific mimicry cases, and that commonly used theoretical concepts like protective or aggressive mimicry did not cover the full diversity of mimicry systems in nature (Wickler 1965: 519). As a solution, he proposes describing mimicry as a formal structure of three participants in which their interrelations could be expressed by mathematical symbols based on the evolutionary effect of the participants to oneanother (whether it is positive or negative). Wickler describes mimicry through the roles of the mimic, model and receiver as follows: A signal is emitted by two different signal-senders (S1, S2) which have at least one signal receiver (R) in common that reacts similarly to both of them. One of the senders is called a model, the other a mimic, […] if it is profitable (+) for the receiver to give the reaction to one of the senders, but unprofitable (–) to react in the same way towards the other. That means, that if the signals from the two senders could be distinguished by the receiver, individual experience and/or selection would favour different reactions. (Wickler 1965: 519). Wickler calls such a communicative situation the model-mimic-receiver system or later just the mimicry system. The way in which Wickler formalises mimicry allows for quick and clear representation of different triadic relations between species. For instance, a situation where it is profitable for the receiver (R) to give a certain behavioural response to the signal (+) sent by one sender (S1), but harmful (-) to respond in the same way to a similar signal sent by a second receiver (S2) can be expressed as a formula: S1 +R– +S2. In such a situation, the first sender (S1) should be called a model and second sender (S2) a mimic. In Wickler’s schematisation, when mimicry is evolutionarily beneficial to the mimic, it is expressed by a + sign before S2. For the model S1, the evolutionary effect of mimicry relation depends on the behaviour of the receiver: it can be beneficial + (if the model has warning coloration and the typical reaction of the receiver is to avoid the model),– damaging (in the case of aggressive mimicry, where the model is the typical prey of the receiver and where the mimic imitates the prey with an aim to catch the receiver) or neutral (where the model is an object or feature of the non-living environment). 40 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Wickler’s schematisation was meant to be quite universal and to allow for describing triadic relations between species beyond mimicry. For instance, he discusses De Ruiter’s experiment, where insectivorous birds were let into an enclosure where caterpillars with camouflage colouring were hiding in the bush branches. After finding some moth larvae, birds started to snap everything that resembled larvae (both real caterpillars and branches). Wickler formalizes such a misleading situation with the formula S1– +R– –S2, where S1 signifies the mimic caterpillars and S2, the models (that is, bush branches and vegetation). For birds (R), it is beneficial to catch larvae (S1) but damaging to eat branches (S2), whereas pecking is detrimental to both moth larvae and branches, expressed by the minus signs. Wickler a uses similar approach to describe Müllerian mimicry (S1+ +R+ + S2) and concludes similarly to many other authors, that as Müllerian mimicry is beneficial to all participants, it should not be taken as a case of mimicry, but rather as an example of convergence. Wickler’s systemic approach that focuses on the three participants of mimicry and their interrelations makes it possible to describe and compare mimicry cases that take place in many different ecological relations and communicative media. Although centred on evolutionary cost-benefit relations, the tripartite approach also provides a good ground for describing biosemiotic and communicative processes and their effects on mimicry, which will be developed in the following chapters. For a semiotics of mimicry, Wickler’s observation of the receiver’s role in a mimicry system is especially relevant. According to him, it is the receiver’s long-lasting learning activity that determines the dynamics of the mimicry system (Wickler 1965: 519). In certain cases, as for instance in Mertensian mimicry, the type of receiver’s leaning process (whether its reaction is innate or acquired during ontogenesis through experience) could even be essential in determining which participant in mimicry acts as a model and which one as a receiver (Wickler 1968: 241). In 1968, Wolfgang Wickler published a monograph „Mimicry in Plants and Animals”. The book included the popular overview of mimicry types with a thoroughgoing discussion of examples in the framework of Darwinian evolutionary theory. The final part of the book presents, however, Wickler’s own understanding of the mimicry system as a specific configuration of mimic, model and receiver. The fluent scientific-popular style and good set of examples made Wickler’s tripartite model of the mimicry quite broadly known. His approach was later elaborated by 41 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. British entomologist Richard I. Vane-Wright, who emphasised in his papers (VaneWright 1976, 1980, 1981) the importance of informational processes in mimicry. In regard to the receiver’s behaviour, Vane-Wright distinguished two aspects: receiving information and reacting to it, i.e. sensorial and motoric functions. There appears to be an implicit parallel to Jakob von Uexküll’s (1982) functional cycle that contains two sign-dependent connections: perception and effect that both connect an organism with the environmental object. Vane-Wright (1976: 30) further calls organisms, whose reaction depends on perceived information and who act based on learned knowledge or innate instincts, to be “sensitive receivers”. In an indirect way, the structural approach to mimicry appears to have some affinity to the thinking of old Austrian-German biological tradition. Wickler’s approach provides the concept of mimicry a certain autonomy and self-sufficiency. Mimicry cases can be modelled as separate phenomena on their own, distinct from the surrounding ecological and biological conditions. This turns out to be both a strength and a weakness of the tripartite approach. On the positive side, this makes formal description not only applicable to biological phenomena, but also to imitations in various human and animal communicative situations and sign systems on similar grounds. Formalising mimicry systems has promoted mathematical modelling of mimicry (Huheey 1964, 1976, 1988; Holmgren and Enquist 1999; Speed 1999) and using game theory in the study of mimicry systems (Bacharach and Gambetta 2001; Augner and Bernays 1998). In practical biological fieldwork, Wickler’s systemic approach has been applied, for instance, by Greene and McDiarmid (1981), Wong and Schiestl (2002), Grim (2005) and others. 3.5 Critical discussion of the triadic mimicry model 9 The tripartite model of mimicry is an effective tool for formal and comparative description, but it also appears to have problematic aspects that may distort our understanding of mimicry. The tripartite model presumes the existence of strong 9 This subchapter is partially based on Maran, T. (2010). Semiotic modeling of mimicry with reference to brood parasitism. Sign Systems Studies (University of Tartu Press), 38(1/4), 349–377. Used with permissions. 42 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. relations between three participants that are mostly concretized as species. 10 This presumption may, but does not necessarily, correspond to an actual situation in nature. It is more frequent that one of the three positions of the triad is filled by several species. We can consider here the case of a protective mimicry system, where a broad-bordered bee hawk-moth Hemaris fuciformis resembles a tree bumblebee Bombus hypnorum for a European pied flycatcher Ficedula hypoleuca. In this mimicry system, the hawk-moth can be considered the mimic, the bumblebee the model and the flycatcher the receiver. But the resemblance between the broadbordered bee hawk-moth and the tree bumblebees can be confusing to shrikes Lanius sp. also, and to several other insectivorous bird groups beside flycatchers. In a similar light, the position of mimics and models can be filled with more than one species. Furthermore, the number of species that can participate in a mimicry system is often not limited in principle; as, in addition to the dominant participating species, there may be occasional participants, for instance omnivorous birds, for whom the moths form a small part of the diet and for whom confusing a hawk-moth with a bumblebee is a rare event. In some other mimicry systems, the involved species cannot be clearly divided between mimics and models and the participating species rather form a fuzzy set of resemblances, called the Müllerian–Batesian mimicry complex or arithmetic mimicry. In arithmetic mimicry, the difference in edibility or dangerousness of the involved species is not clearly established, or it may change between individuals or during their life course. Openness of a mimicry system to occasional participants is not the only problem with the triadic description of mimicry. There are also many mimicry complexes in which more then three participants are entangled in a structural way with clear roles and functions. A good example is large parasitic hoverflies from the genera Volucella and cuckoo bumblebees Psithyrus, who both have appearances similar to various bumblebee species (Plowright and Owen 1980). These parasitic insects can be often found in the vicinity of bumblebee nests. The genus Volucella includes more then forty species of hoverflies, many of which are commensals or parasites of the bumblebees. Female Volucella try to enter into bumblebee nests and 10 This does not necessarily mean that there are three species involved in the mimicry system. Quite often two species fill the three roles: for instance the model and the receiver can belong to the same species, as is the usual case in aggressive mimicry. Common mimicry typologies also acknowledge such possibility and include the category of bipartite mimicry systems. 43 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. lay their eggs there, and their larvae are later feed upon nest debris or bumblebee larvae. Cuckoo bumblebees Psithyrus are closely related taxonomically to bumblebees (forming a subgenus of Bombus), but they do not live collectively in nests like other bumblebees. Instead, they are parasites of collective bumblebee species (see Figure 3.1). Female cuckoo bumblebees seek and try to enter bumblebee nests and if successful, they lay eggs there. The aggressiveness of host workers are suppressed by pheromones that appear to be species-specific, and the parasitic strategies of cuckoo bumblebees include a number of different methods (oophagy, larval ejections, preventing worker oviposition, Fisher 1988; Küpper, Schwammberger 1995; Honk et al. 1981; Martin et al. 2010). The resemblance of Volucella and Psithyrus to the bumblebee hosts has a double function: it helps parasites to enter the bumblebee nests and also grants protection against birds that may have unpleasant experiences with real bumblebees. The tripartite mimicry model has difficulty accommodating such a mimicry system. In the mimicry typologies of Vane-Wright and Pasteur that proceed from the Wickler’s tripartite model, the cuckoo bumblebee would belong to different types depending on whether it is encountered by a bumblebee (class VIIB in VaneWright’s classification, Kirbyian mimicry in Pasteur’s classification) or by an insectivorous bird (class VIA in Vane-Wright’s classification, Batesian mimicry in Pasteur’s classification). [Enter figure 3.1 here] Figure 3.1. Bumblebee species (models) and corresponding cuckoo bumblebees (mimics). Upper row, from left: Red-tailed bumblebee Bombus lapidarius, whitetailed bumblebee Bombus lucorum, garden bumblebee Bombus hortorum. Lower row, from left: Cuckoo bumblebees Psithyrus rupestris, Psithyrus bohemicus, Psithyrus campestris (From the collection of The Zoological Museum of the University of Tartu, photo by courtesy of author). A third source of difficulty with the tripartite model of mimicry is the possibility that one and the same individual can be simultaneously be involved in more than one mimetic resemblance; a combination that can in some cases have structural importance for the mimicry system. As early as 1890, Edward Poulton described in detail the protective adaptations of the puss moth’s caterpillar Cerura vinula that 44 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. combines camouflage coloration, warning patterns and specific warning posture (Poulton 1890: 269–278). Another example is an aggressive mimicry system of the monkfish Lophius piscatorius that combines a cryptic resemblance of its body surface (mimic) to the seafloor rich in algae and other plants (model), and the resemblance of its foremost fin ray (mimic) to a worm (model). The first type of resemblance serves to make the monkfish hard to notice and the second type helps to lure and catch smaller fish (receiver). Both resemblances support each other and are active during the same communicative interaction between the monkfish and its prey species. Such instance of mimicry cannot be easily accommodated by the classical mimicry triad. A fourth source of problems is the under-determinacy of the model in this approach. German theoretical biologists Zabka and Tembrock have argued that in many cases, the model cannot in principle be reduced to a single species, as for instance in the case where decaying meat is mimicked by carrion-flowers to attract flies, which are looking for carcasses to lay their eggs (Zabka and Tembrock 1986: 172). One of such plants is Titan arum Amorphophallus titanum that produces a three meter high inflorescence with a spathe coloured dark red inside and a distinctive smell that resembles rotten meat. Titan arum is pollinated mostly by scarabids, carrion beetles, rove beetles and other destruents (Jürgens and Shuttleworth 2015: 370). The same type of phenomena is described by Georges Pasteur as semi-abstract or abstract homotypy (Pasteur 1982: 191) and by myself as abstract mimicry (Maran 2007: 239– 243), examples of which cover a wide array of phenomena from eyespots to deimatic displays. In the case of abstract mimicry, the object of imitation appears to be some abstract meaning complex in nature and its physical expression, for instance dangerous situations, are marked by a sudden change of affairs. In the forthcoming discussions, we shall see that abstract mimicry is not an exceptional situation but rather such abstractness is a property of many mimicry systems that indicates the semiotic nature of the resemblance (see Chap. 11.1 “Abstract mimicry: when the meaning comes first” for detailed discussion). When trying to systematize these examples, it appears that the systemic approach to mimicry has problems in: 1) defining the set of elements (species) that belong to the (mimicry) system as a whole; 2) determining the location of elements with regard to predefined classes of mimics, models, and receivers; 3) presence of classificatory error, i.e. the same element can belong to more than one class in the system; 4) possibility for the different mimicry systems to become merged. A partial 45 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. solution to this complex classificatory problem would be to reconsider the mimicry triad as a logical and conceptual relationship between three entities and not to take this as a necessarily ecological relationship between three species. In such an approach, we would take the mimicry triad to be a heuristic device or a modelling tool that would help us to map the species relations in the deceptive resemblances. In some cases, the involvement of actual species may correspond to the roles in the mimicry triad, but this is not, by any means, an inevitable condition. Another possibility would be to consider the mimicry system as having a double layered structure, consisting of a layer of ecological relations between species and a layer of semiotic relations of sign. Such a semiotic approach is helpful in regard to the under-determinedness of the model, but it cannot solve problems rooted in the ecological level in terms of the openness of the mimicry system to occasionally participating species. When considering mimicry as a double-layered structure, species are indeed the actual biological entities that are involved in different ecological relations such as predation, competition, parasitism and others, and their number and evolutionary characteristics can also change because of a particular relation. The second, semiotic layer would, however, explicate a specific relation of resemblance, in which case we may ask “What resembles what to whom in what respect?” On this level we are dealing with specific qualities and their similarity in the eyes of a particular beholder. A Danish biosemiotician Jesper Hoffmeyer has described such semiotic layer accompanying ecological relations as semethic interactions (Hoffmeyer 2008a: 189): “Whenever a regular behavior or habit of an individual or species is interpreted as a sign by some other individuals (conspecific or alter-specific) and is reacted upon through the release of yet other regular behaviors or habits, we have a case of semethic interaction.” (Hoffmeyer 2008b:15). The two layers—ecological relations and sign relations—can combine with each other in different ways, and one goal of the ecologically oriented biosemiotics could be describing possible configurations in such a system. In addition to the rigidness of the triadic system of mimicry, a radical differentiation that is often assumed to exist between mimicry and other adaptations can also become an obstacle for studying resemblances in nature. In research literature, arguments can be often found for regarding one or another resemblance in nature as mimicry. Such arguments often presuppose a distinct category of mimicry with clear boundaries. However, when we take into account the many spontaneous 46 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. similarities between species, the involvement of facultative species in open mimicry systems, and the fact that the recognition of mimics and models by the signal-receiver is probability based and not an absolute, then the clear demarcation line between what is mimicry and what is not is not so easy to draw. Instead, we should perhaps talk about a mimetic landscape in nature or about the capability of natural forms to create confusion. In such an approach, the concept of mimicry would become reserved for the most well-formed examples of deceptive resemblances while keeping in mind the probabilistic and fuzzy nature of the phenomenon. 47 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 4 SEMIOTICS OF MIMICRY Biological mimicry is often described as a deceptive resemblance of some physical traits between representatives of different species. In such cases, attention predominantly stays at the physiological level and focuses on the evolution of mimetic features. In mimicry studies, much less consideration is given to the ways in which such resemblances are achieved, expressed and managed by specific individuals in specific behavioural encounters. In this chapter, I analyse the semiotic and communicational aspects of the mimicry system: what the semiotic features of mimicry are, how deceptive communication takes place, what mimicry is as a sign structure, and what the common interpretations of mimicry have been in the field of semiotics. As hinted in the previous chapter, mimicry appears to have two-level structure. On a biological or ecological level, mimicry is constituted by groups of organisms that form evolutionary units and are connected by ecological relations (predation, symbiosis, competition, etc.). On the semiotic level, mimicry is a specific resemblance between messages or signs transferred in communication. In a semiotic or communicative perspective, the three participants of mimicry become two senders (the mimic and the model) and a receiver in their communicative interactions. According to this view, resemblance does not occur between mimics and models as organisms, but rather between messages as these are perceived by the third participant of the mimicry system—the receiver in its Umwelt. The mimetic message (a cue or a signal) is similar to some message of another organism, some feature of the environment, or a generalization of either of those. Based on this reasoning, the essential research question in mimicry from a semiotic perspective becomes: “What resembles what to whom in what respect?” The apparent similarity of this question to the Peircean definition of sign as “something which stands to somebody for something in some respect or capacity” (CP 2.228) is not a coincidence, but points to the deep semiotic nature of mimicry. Introducing the duality of semiotic and ecological levels into the mimicry system brings along major shifts in understanding mimicry. From a semiotic perspective, the resemblance between the message of the mimic and the message of the model is not just a construction of human scientific thought, but is rather an actual confusing situation as it is perceived by the receiver in its subjective world or 48 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Umwelt. This means that in order to studying mimicry, we need to take into account the specifics of the receiver’s Umwelt as well as its difference from the Umwelt of the human observer, including human symbol-based modelling and cultural narratives in regard to mimicry studies. The semiotic approach emphasizes the activity of particular individuals in a mimicry situation, as it is their behaviour, their decision and appearance in the world that creates the deceptive resemblance. Here we may rely on the old Baldwinian notion that no trait or signal can participate in the evolutionary process if it is not developed to its organic form and used by a particular individual (Baldwin 1896: 443–444). Depending on the animal group and species, an individual can bring messages into existence in the following ways: 1) influencing their development (choice of environment, organism’s activity in metabolism); 2) learning how to communicate (learning to use one’s bodily resources and acquiring species-specific repertoire of signs); 3) choosing specific locations and partners for communication (recognising species-mates, members of other species and the right situations for communication); 4) making choices in a particular communicative situation (selecting between signals and behaviours, and organising these into sequences). Paying attention to specific sign relations and mapping these out would enable us to overcome restrictions of the tripartite mimicry model. Such a view would allow sign relations of the mimicry system to be combined in various ways—for instance, the angler-fish in its in communicative interaction with a prey species simultaneously communicates two mimetic messages: the skin pattern that resembles the seafloor and the fin ray that resembles a small wiggling worm. From a semiotic perspective, there is no logical restriction for combining a message of contradictory parts. In human linguistic communication, such a message type is known as an oxymoron (e.g. “honest thief”, “sweet pain”), which can empower novel meanings but also question the normality of the sign system. 49 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 4.1 Semiotic interpretations of mimicry 11 In semiotics, mimicry is seldom explicitly defined, although several authors have used it as an illustrative example or argument. 12 The fact that biological mimicry has been altogether included in the field of semiotics is largely due to the activities of Thomas A. Sebeok. He was the first one to express the opinion that mimicry could be a semiotic phenomenon, and therefore his role in introducing mimicry to the semiotic community remains fundamental. As general editor, Thomas A. Sebeok included in the “Encyclopedic Dictionary of Semiotics” an article on mimicry written by the wellknown British ecological geneticist, Edmund B. Ford. The one and a half page long overview includes notes on polymorphism, sexual mimicry and species combination, and explains the difference between Batesian and Müllerian mimicry, but unfortunately does not develop any explicit semiotic perspective (Ford 1986). In his own writings, Thomas A. Sebeok has given thought to the mimicry phenomenon on several occasions. In the essay “Can animals lie”, Sebeok contrasts mimicry as naturally evolved stratagems with deliberate acts of lying (Sebeok 1990a: 95–96). In his essay “Iconicity”, Sebeok demonstrates iconicity in nature by referring to mimicry alongside other phenomena such as the scent marks of social insects in which the intensity of the emitted pheromone matches the amount of nearby food resources, and the tropic communication between aphids and ants (Sebeok 1989: 115–117). Here we should note that iconic signs (and the property of iconicity) is one of the three basic sign types distinguished by an American semiotician Charles S. Peirce based on the relationship between the sign (representamen) and its object. For icons, this relation is based on similarity; that is, the connection between sign and object is established because the sign evokes the same or similar sensation or reaction in the interpreter as does the object of that sign. Presumably due to the influence of Thomas A. Sebeok’s writings, the connection between mimicry and iconicity has been repeatedly expressed in semiotic literature, and in general semiotic overviews, mimicry is often referred to as an 11 This subchapter is partially based on Maran, T. (2011). Becoming a sign: The mimic's activity in biological mimicry. Biosemiotics (Springer) 4(2), 243–257 and Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica (DeGruyter Mouton), 167(1/4), 223–248. Used with permissions. 12 A short overview of the concept of mimicry has been also published in “S – European Journal for Semiotic Studies” by the mimicry historian Stanislav Komárek (1992). In biosemiotics, mimicry as a specific phenomenon has also been shortly discussed in relation to recognition and species concept (Kull 1992), intentionality in evolutionary processes (Hoffmeyer 1995), and types of information valuation in communication (Sharov 1992). 50 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. example of iconicity in the natural world. Winfried Nöth writes in his “Handbook of Semiotics” about connection between mimicry and iconicity: “visual and olfactory icons occur in the form of mimicry” (Nöth 1990: 124) and “for nondeceptive purposes, iconicity is relatively rare in animal semiosis” (Nöth 1990: 163). In a similar vein, neurobiologist Terrence W. Deacon considers the relationship between the moth and the tree bark it resembles to be iconic (Deacon 1997: 75–76). Eminent entomologist and mimicry scholar Philip E. Howse (2013: 205) further describes resemblance in the design of the Lepidopteran wings, where “eyespots can often be seen as part of the design of the head of a potential predator, which commonly includes representations of teeth, or a beak, such that the image becomes a more convincing portrayal of the predator” as iconic with making reference to semiotic terminology. Aside from the widespread statement that mimicry is an example of iconicity in nature, we will find two alternative interpretations in semiotic research literature. 13 The first alternative explanation is that mimicry does not obtain all necessary characteristics of the iconic sign. The second possible view is that biological mimicry is more complex then a common iconic sign. The first position is taken, for instance, by a Swedish cognitive and visual semiotician Göran Sonesson (2010) who discusses biological mimicry in the framework of the theory of pictoriality and aesthetics. He describes mimicry as an extreme example of the iconic phenomena and claims that “In mimicry […] neither sender nor receiver is in any way involved with anything that resembles a sign. The mimic and the model, and thus the iconicity, only exists for the outside observer.” The criterion that inhibits treating mimicry as an iconic sign appears for Sonnesson to be a strict definition of sign that relies upon differentiation between the signifier and the signified and upon an asymmetrical relationship between these (one part being more directly experienced than the other by an interpreter). Such a position enables the argument that there is no participant in the mimicry system for whom the sign relation would actualise and therefore, there is no reason to indicate that mimicry is iconic. The second position, that mimicry is more complex then a simple iconicity, is developed in a thorough study on the semiotics of mimicry made by a group of 13 My own understanding of this issue is somewhat more complex and is presented in the Chap. 5, “Iconicity and mimicry”. 51 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. authors João Queiroz, Frederik Stjernfelt and Charbel Niño El-Hani (2014; also ElHani et al. 2010; Queiroz et al. 2012). Their point of departure is the Peircean classification of signs and they demonstrate possibilities to use Peirce’s sign typology for the semiotic study of mimicry. They use the mimicry system of the fireflies Photuris with a reference to James E. Lloyd’s lifelong research (1975, 1986) on these magnificent insects as a test case and argue that mimetic signs go far beyond simple iconicity. In their view, mimicry needs to have a propositional character and thereby it corresponds to a separate sign type: the dicisigns: “[the] double reference of the dicisign is the reason why it may claim something (iconic) about something (indexical)—and this is why a proposition may be true or false, depending on whether the iconic quality claimed actually exists in the object referred to. And this is why deceptive signs must necessarily be dicisigns.” (El-Hani et al. 2010: 46–47). For future reference, it is relevant to point out that João Queiroz, Frederik Stjernfelt and Charbel Niño El-Hani attribute a symbolic character to mimicry; that is, they consider at least some mimicry examples to be based on law-like regularities. This statement will be brought up later in the discussion on ecological codes and abstract signs. At present, we could note that mimicry has been a relevant topic for semiotics and is mostly discussed in connection with resemblance between the sign and its object. 4.2 Mimicry as a communicative interaction When analysing mimicry in the semiotic discipline, different viewpoints can be taken and different aspects emphasised. For instance, mimicry could be described as communicative phenomenon or as a specific type of sign structure. The first approach would consider mimicry as an interaction between participating organisms, with relation to the classical transmissional communication model that includes sender, receiver, message, signal, environmental noise (corresponding to the pragmatic dimension of the sign relations 14). The second approach would focus on the specific signs of mimicry, describe the possible types, combinations and series of these (corresponding to syntactic and semantic dimensions). 14 Distinction between syntactic, semantic and pragmatic dimensions of sign comes from the works of eminent American semiotician Charles Morris. Syntactics would be concerned with relations between different signs (sign vehicles), semantics with the relation between the sign and its meanings or objects referred to and pragmatics with the relations between signs and interpretants or participants of communication (Morris 1971a: 21–22). 52 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Mimicry as a communicative interaction appears to follow the general logic of communicative processes, being dependent on channel or medium, communicative code (rules and habits of combining different messages and interpreting those) and participants’ intentions and/or biological functions. A good analytic tool for distinguishing and analysing such aspects would be Roman Jakobson’s model of functions of linguistic communication, where he distinguishes between six components in communication and six corresponding communication functions. Jakobson proposed his communication model in the late 1950's, combining various sources and influences including the works of Karl Bühler, Bronisław Malinowski, and Russian formalists. Basic components of Jakobson’s communication model are: 1) addresser (sender); 2) addressee (receiver); 3) message; 4) context that surrounds the message or what the message refers to; 5) code, which forms the basis of coding and decoding the message; and 6) contact, including both the communication channel, as well as the psychological contact between the sender and the receiver (Jakobson 1981: 21–22). By asking what happens if any of those factors dominates in communication, Jakobson distinguishes the following six functions of communication: 1) emotive or expressive that focuses on the the sender’s (addresser’s) own intention or attitude; 2) conative that orients to the receiver (addressee) and aims to influence it; 3) poetic function, that is directed to the properties of the message, for instance its internal aesthetics; 4) referential function, which focuses on the meaningful relations with the context; 5) metalingual function, in which the communication makes a reference to the communicative code at the meta-level; and 6) phatic function, in which case the aim of communication is to establish or keep the contact between the sender and receiver (Jakobson 1981: 22–27). Jakobson’s functions were originally developed in the studies of human linguistic communication. Later, it has been shown (for instance by Dario Martinelli 2010: 77–80) that they can also be effectively used in the context of animal communication. Although mimicry is a relatively simple communicative phenomenon, Jakobson’s functions also appear to be operational in the context of mimicry. Thus mimicry can predominantly be about giving a false impression about the mimics’ intention, thus having an emotive dominant (for instance behavioural mimicry such as broken wing display in Charadriidae and others), or it can be focused on influencing the receiver’s interpretation and behaviour, thus having a conative dominant (sexual mimicry as that of fly orcid Ophrys sp.). Referential 53 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. function is in the foreground of most cases of Batesian mimicry that abuse the reference between aposematic coloration of the model and its poisonousness or other defences. Metalingual function appears to be in the foreground in, for instance, satyric mimicry and other mimicry cases that combine several contradicting messages to create the mimetic effect. In such cases, it is the code relation between the animal’s appearance and its applicability that is taken advantage of and undermined by the mimic. Poetic function appears to be in the foreground of cases where the message’s inner structure is what makes imitation or deception possible (e.g. in vocal mimicry). Phatic function can be considered dominant in cases in which the mimic manipulates the communicative contact with the receiver. The latter is true in most cases of cryptic mimesis and camouflage coloration. It appears that functionally, mimicry cases are quite diverse as they make use of different facets of communication. This structural diversity can be well analysed based on Jakobson’s typology of different communicative functions. Mimicry as a communicative interaction also relates to other communicational phenomena in nature. Namely, mimicry is often dependent on the predominant communicative interaction that takes place between the model and receiver. Depending on the mimicry case, model–receiver communicative interaction can take place between aposematic insects (models) and insectivorous birds (receiver), between females and males of solitary wasps (as in the case of reproductive mimicry of the fly orchid), as chemical communication among countless ants in the ant colony (exploited by many myrmecophilic insects, e.g. larvae of blues Aloeides dentatis, Maculinea rebeli) and so on. Correspondingly, mimicry as a communication system can be schematized as the interaction or mergence of two communicative interactions—model-receiver and mimic-receiver—, where the mimic is sending a similar message to that of the receiver, thereby intercepting the model-receiver communication. Such a schematisation has been used by Australian military scholars Carlo Kopp and Bruce Mills (2002) in the context of information warfare studies by contextualising biological mimicry among several other possibilities of destructive communication (see Figure 4.1). Interpreting mimicry as a combination of two transmissional communication sequences allows for describing many specific aspects of mimicry (communicative dominant, the role of channel and noise, etc.). [enter figure 4.1 here] 54 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Figure 4.1. Mimicry system integrated with the Shannon-Weaver communication model (modified after Kopp and Mills 2002). At the same time, we should remind ourselves that transmissional models simplify the actual communicative situations in several respects. Important features missing in transmissional communicative models are feedback and the possibility for mutual interaction. In mimicry systems, the receiver’s feedback to the mimic and model allows cyclicity or self-referentiality to enter into mimicry. The receiver’s feedback makes it possible for interpretation to become an agency influencing the resemblance of the mimic and model (this will be discussed further in the Chap. 9.1. “Semiotic selection: definition and examples”. In classical evolutionary accounts, the receiver’s selective feedback is mostly considered to be an epiphenomenon or secondary process in comparison to genetic causes and heritability of mimetic features. Focusing on the communication process, however, can change this classical view by departing from a single communicative situation and considering behavioural, developmental and evolutionary processes as expansions of the communication situation at different levels. From the perspective of communicative interaction, feedback allows the relation between senders and receivers to become dynamic, and in the long run, leads to changes in messages used and the sign system in general (see Figure 4.2). From this perspective, every communicative interaction can be understood as consisting of at least three layers: 1) one-time event between specific individual in a specific time, space and environmental context; 2) generalization of interactions between specific individuals, which accumulate in individual learning and experience; 3) relationship between participating species, with co-evolutionary changes. Respectively, feedback in communication can also take place: 1) in the limits of a singular communicative situation (e.g. flight manoeuvring of a mimic butterfly and an insectivorous bird); 2) in the ontogeny of the participating individuals (e.g. different effect of encountered hoverflies and wasps on the learning of a toad of how to avoid wasp-like insects); 3) in the phylogeny as a fine-tuning of colour resemblance between mimics and models by selection caused by different ecological (e.g. predation) pressures. [enter figure 4.2 here] 55 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Figure 4.2. Different feedback cycles that can have effect upon communication. Distinction between autocommunicative (proprioceptive and exteroceptive) and communicative feedback can be made. The latter divides between instant feedback, ontogenetic and phylogenetic feedback. Among mimicry cases, the relevance of these feedback cycles may differ and furthermore, when considering a single mimicry case, participants may rely on different feedback cycles. For instance, if we consider communication in a mimicry system between small song birds (e.g. a great tit Parus major) and moths with eyespots (e.g. eyed hawk-moth Smerinthus ocellatus), then birds have a greater chance to learn from their individual experiences, whereas moths need to rely more on instant behaviour and evolutionary fixed adaptations. The difference is caused by the different lethality rate, but also by different cognitive capacities and life spans of the participants. Mimicry as a communicative situation appears to have a structural diversity in regard to functional dominants of communication, and to location of the feedback in the system. Also the balance between the sender’s and receiver’s activity can differ. In some examples, mimicry depends more on the receiver’s capacity of perception (e.g. many cases of camouflage) where the activity of the mimic is minimal. In other cases, it is predominantly the mimic’s complex behavioural activity that creates the mimetic effect. Whatever the balance is between the activities of the mimic and receiver in the specific mimicry system as communication, they need to fit well together for the mimicry to be operational. A communicational approach helps to highlight different possibilities for such coupling to emerge. 4.3 Mimicry as a sign system In common understandings, a criterion that determines whether the case under observation is mimicry is a mistake made by the receiver when perceiving deceptively similar messages of the model and the mimic. At a closer look, however, a mistake because of deception appears to be a very coarse description of the sign processes that actually take place in a mimicry system. In this chapter, I shall discuss some detailed possibilities for combinations of signs that make it possible to confuse and deceive in 56 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. mimicry. The topics dealt with in this subchapter correspond to the domains of syntactics (relations between different sign vehicles) and semantics (relations between signs vehicles and their meaning or object referred to) in semiotics. The third domain of semiotics: pragmatics (use of signs in communication) was addressed in the previous subchapter. In general, we can consider a sign system to consist of a set of signs (more correctly, sign vehicles) and rules or regularities (formalised or not) that are used to combine the signs for producing meaningful messages. Messages with a complex semantic and syntactic structure are mostly characteristic of intraspecies communication and especially human linguistic communication, but there are also good examples in mimicry to consider. In treating mimicry as a sign system, I will use a well-known distinction between paradigmatic (associative) and syntagmatic dimensions of utterances. The distinction itself comes from the language semiotics of Ferdinand de Saussure (2011[1916]: 122–125) and is one of the core elements of structural analysis. In a broad sense, the syntagmatic axis describes the sequence of signs—how signs can be ordered to form a series (such as a sentence). The paradigmatic axis describes types or sets—which signs are replaceable with which other signs in a given message. The paradigmatic axis is related to synonyms, cognitive types and associations that the subject is able to make or discern. For a semiotic analysis of mimicry as a sign system, we would first need to distinguish between different sign systems or sets that take part in the mimicry interaction. Based on the discussions of the previous subchapter, there is a sign system/set that is used in communication between the model and the receiver; and the sign system/set that is used for deceptive communication between the mimic and the receiver. To understand the sign structures in mimicry, we should also analyse the relation between these two sign systems as a separate issue. All three topics have their own specifics and peculiarities. For the sake of clarity, I will refer in the following discussion to the model-receiver sign system as sign system 1, and signs exchanged between the mimic and receiver as sign system 2. In addition, there may be a third (3) sign system partially incorporated in mimicry, which involves signs used by the mimic species for its own intraspecies communication needs. In some cases, there is a complex interplay between the mimic’s intraspecific sign system 3 and mimicreceiver sign system 2, as we will see from the examples at the end of this subchapter. 57 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. In the following discussion, the typological diversity of mimicry also needs to be taken into account. It is easier to describe the model-receiver relation as a sign system in cases where the model is a specific animal; that is, when we have a true communicative relationship (that includes sender and receiver). In camouflage, where the relationship is rather based on environmental semiosis or signification (the animate sender is missing), the reference to the model-receiver sign system is also an approximation. In such a case, sign system 1 is rather based on the way in which the receiver categorises and interprets the surrounding environment and how environmental affordances relate to the animal perceptual system, but there is no reason to talk about repertoire or communication code. In camouflage, the sign set of the model is an array of environmental colours and their specific patterns (base coloration of the environment, tonal variations, shades, alternation, rhythms and patterning). On the paradigmatic axis, different cues are easily interchangeable with one another (where exactly the specific colour patch is located in the environmental background does not change the meaning of the environment in most cases). The syntagmatic axis appears to be almost missing (in respect to camouflage, there is no meaningful series in environmental colouring). In other mimicry cases, there can exist very complex signs systems 1 used for communication betwee the model and receiver. For instance, in the symbiotic relationship between cleaner wrasse Labroides dimidiatus and its host species, a full set of bodily and behavioural signs exist that cleaner wrasse presents in a dynamic manner. Cleaner wrasse is adapted to feed upon ecoparasites and dead skin tissue of the larger fish, and many inhabitants of the coral reef (morays, mullets, mantas and others) are willing to use this service. The communication code of the cleaner wrasse includes the specific habitat, delineated body shape and colour pattern of the horizontal blue and pale stripes, as well as specific movements of approaching other fishes in a rhythmic dance pattern (Wickler 1968: 157–176; Stummer et al. 2004). More precisely, the invitation movements of the cleaner wrasse have been observed as following: “tactile dancing involved the cleaner oscillating its posterior body in a dorsal-ventral manner […] within 15 cm of the client and mainly in one place while often contacting the client with its body. Tactile stimulation without dancing involved the cleaner contacting the client’s body with its pelvic fins and often also nibbling on the client’s body.” (Grutter 2004: 1082). The combination of these signs is taken as a key for recognising a cleaner wrasse by host species, and if positively identified, the 58 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. host becomes motionless and allows the wrasse to enter its mouth and gills for searching parasites. The invitation dance of the cleaner wrasse is both syntagmatically and paradigmatically constrained; that is, the wrasse needs to send specific signs in a specific manner for the receiver to be recognised as a symbiont (this symbiotic relationship is also imitated and taken advantage of by a mimic—the false cleanerfish Aspidontus taeniatus). Sign system 1 is deriving from the features of the organism or from the affordances of the environment and is initially related to other properties of these (environmental structures and their diversity, physiology and ecology of the species, e.g. the cleaner wrasse’s delineating body shape as suitable for entering into hosts’ buccal cavities). In sign system 2, used between the mimic and the receiver, the set of signs used in communication has much narrower limits or boundaries. The problem arises from the fact that the mimic as a species in most cases has a different body structure, physiology, ecology and the behaviour than the model. In many cases, the mimic is situated very far from the model taxonomically as well. Yet, the mimic needs to use its bodily and semiotic resources to produce a coherent set of mimetic messages in order to confuse the receiver. Thus, an essential question in analysing sign system 2 is: how are the mimetic signs construed and distinguished from the mimic’s other semiotic resources (e.g. those that are used by the mimic in interactions with its own species mates, sign system 3)? Let us illustrate this topic with the reproductive mimicry of the fly orchid Ophrys insectifera by asking how the plant forms the mimetic message as distinct from its other perceivable features (see figure 4.3). Genus Ophrys (bee orchids) include more than thirty species, with many subspecies and hybrids that have a remarkable reproduction strategy involving bees, wasps, flies and other insects (Schiestl 2005: 258). The blossoms of orchids have a close resemblance to the body shapes and appearance of their pollinators, which is used to induce copulation behaviour in the male insects. The sexual mimicry of the fly orchid also includes chemical signals that are effective from close range and that elicit sexual behaviour in males, who try to copulate with the flower labellum (Ayasse et al. 2003: 225). In Europe, Ophrys species have gone through an intense speciation and divergence process because of their relations with various pollinator species (Breitkopf et al. 2015). The fly orchid Ophrys insectifera is mostly pollinated by digger wasps 59 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Argogorytes, whereas other Ophrys species are specialised for deceiving other species of Hymenoptera (e.g. bees from genera Andrena, Euceral, Melecta, and Anthophora). [enter Figure 4.3 here] Figure 4.3. Sexual mimicry of the fly orchid Ophrys insectifera (photo courtesy of author). In the mimicry system between the female digger wasp (model), male digger wasp (receiver) and fly orchid (mimic), in sign system 1, it is not a problem for a receiver to perceive the additional physiological characteristics of the model—a male digger wasp can perceive very different signs about a female: that it is an insect, has wings and large compound eyes, can fly, etc. These contextual signs either support the message of the model or, at least do not contradict it. On the contrary, in sign system 2, contextual information about the physiology and behaviour of the mimic can severely interfere with the mimetic communication. Therefore, the question we should ask is not how the fly orchid causes the male digger wasp to confuse its flowers with a female digger wasp, but rather how it succeeds in hiding the fact that it is a plant. The fly orchid creates the false identity, for instance, by the placement of mimetic messages (shape, colour and smell) on the tiny area of its surface while the rest of the plant does not emit any messages to the digger wasp’s Umwelt and cryptically dissolves into background vegetation. It appears that there are several means that the mimic species can use to keep the deceptive message (sign system 2) distinct from its overall bodily and semiotic construction. For instance, mimetic may: a) localize mimic signs into a specific area, b) increase the distance or contrast (spatially, temporally or medium-based) between mimetic and non-mimetic signs, c) use symbolic or symbol-like signs that have meaning apart from their background, position or location (such as when imitation of animal eyes stands as a symbol of the animal and the imitation of the full animal becomes not necessary), d) using super-releaser’s (Tinbergen’s term for signals that evoke over-enhanced response, e.g. a gigantic dummy egg that are preferred by gulls in comparison to normal sized eggs) that are based on a model-receiver sign system. In sign system 2, we can also distinguish both syntagmatic and paradigmatic aspects. These dimensions are valid for the organisation of the set of mimetic signs (e.g. the false clearer fish needs to follow the same syntagmatic and paradigmatic 60 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. organisation of signs as the cleaner wrasse for its deception to be successful). The relation between the mimic’s mimetic display and own physiological constitution also has both a syntagmatic and paradigmatic dimension. Mimetic signs are formed and enhanced on a non-mimetic background by procedures following the paradigmatic dimension described above (localization of signs, using symbols and super-releasers), but the mimic also needs to organize its mimetic messages temporally on a syntagmatic axis, for instance to solve issues, or how to enter and exit from the mode of mimetic presentation. Inability to organize the mimetic sign system and make a shift between mimic and non-mimic modes of communication may lower the success of the deception or it may create problems in the mimics’ intraspecific communication (intertwining sign systems 2 and 3). A good example of the relevance of the syntagmatic axis for entering and leaving the mimetic modes are the strategies used by cuckoos and other brood parasites when laying eggs into the nests of their host species. A most critical stage in this process appears to be the presence of the female brood parasite in the vicinity of the host nest. This is the very act of laying the egg where the parasites own species identity and its mimic identity are simultaneously present and where entering and leaving from the mimetic stage takes place. For this reason, in most brood parasites, the act of laying eggs is kept very short (according to Møksnes et al. 2000: 253, 12 seconds for the common cuckoo). Another type of problem may rise when the mimic itself encounters problems in differentiating between its species-specific communication system and its mimic displays, or when, for some reason, the mimic's species-specific communication becomes substituted by its mimetic identity. In such cases, the mimic may lose its ability to use speciesspecific sign systems for intraspecific communication and take over the sign system used by the model organism or develop a combination of those two (as is the case in the communicative relationship between brood parasitic indigobirds Vidua and their hosts—weaver finches Estrildidae, Payne et al. 2000, see Chap. 7.2. “Resembling the environment and becoming a sign” for detailed overview). Following these discussions, we are now ready to consider the relations between sign system 1 (model-receiver) and sign system 2 (mimic-receiver). Let us keep in mind two aspects: that sign system 1 forms a relatively well organized and self-sustained set of communicative entities, based on code or other regularities shared between the sender (model) and the receiver, and that sign system 2, on the contrary, rearranges the mimic’s own bodily structures and expressional capacities to 61 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. produce the deceptive message. Sign system 2 does not have much structural organisation outside of the mimetic display, but is rather an extension of the mimics’ general physiological and semiotic capacities. The constitutional logic of sign systems 1 and 2 are therefore different, and this incompatibility produces the space for possible combinations and ways for the confusion in the receiver to emerge. The deceptive perceptual similarity in mimicry is just one among many possibilities. The space of combinations of how sign systems 1 and 2 relate and what their effect is on the receiver also depends on criteria based on which the receiver categorises different perceptions and objects into distinct types. Here we enter into the topic of the receiver’s psychology and categorical perception that is studied in many different paradigms: psychology (Medin and Barsalou 1987), neurology (e.g. Solan and Ruppin 2001), anthropology (MacLaury 1991) cognitive linguistics (Lakoff and Johnson 1980) and others. In general, there are different mechanisms for forming perceptual categories that can be listed as deriving from: 1) perceivable similarity between objects; 2) common properties between objects under comparison; 3) space of limited properties (size, colour, pattern, etc.), that are used to evaluate objects; 4) prototype or model that is used as a basis of comparison; and 5) context of the perceived object (Blough 2001; Hampton 2001: 20–26). Some of these processes relate to perceptual resemblance, others are based on the cognitive processes of creating models or in integrating contextual information. Nevertheless, it is important to perceive that the receiver’s psychology is not the only cause of mistake or deception in mimicry, but the space of combinations between sign system 1 and sign systems 2 can also be a source for the confusion. A possible case of this is a situation where sign system 2 used by a mimic violates the systemic integrity of sign system 1. Here the mimic produces messages that blur or disrupt the codes or regularities used in sign system 1, which results in the receiver’s interpretation to become unreliable and its behavioural response delayed. As discussed before, this type of mimicry should be connected with the concept of code and metalingual function in Jakobson’s list of communicative functions. An example of such mimicry type is satyric mimicry, where mimetic signs are presented in an unexpected context where different types of mimetic signs are presented together that lead to ambiguous interpretation, or where different mimetic species combine similar appearances with unexpected escape strategies (Howse 2013). What makes satyric mimicry effective is not the perceptual similarity, but the underlying communicative 62 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. code that connects form (appearance) with content (meaning, function), and that becomes unreliable due to the mimic’s action. Another possibility for confusion to emerge in mimicry appears to relate to the relationship between the mimic’s bodily and semiotic capacities and sign system 1. In its aim to display the forged identity, the mimic may not be able to establish full distinction between what it is and what it pretends to be. In such a situation, the receiver may get messages with two different and contradicting meanings—the mimic signals simultaneously its own identity and that of the model. Such a situation may actually be much more common than generally considered in mimicry studies, especially when considering the diversity of possible receivers having different perceptual systems and cognitive capacities. The presence of two contradicting sign systems 1 and 2 causes cognitive confusion in the receiver, as the receiver needs to figure out how to combine contradicting signs and what the meaning is of the full situation. Let me present here a speculative interpretation of eye-spots—the colourful concentric and paired patterns in butterfly wings—to illustrate this aspect. In classical mimicry theory, the eyespots have been mostly interpreted as being deceptive for small insectivorous birds, who are assumed to mistake them for owl eyes and face. Eye-spot display has been found to trigger the anti-predatory response in some bird species (Olofsson et al. 2013; De Bona et al. 2015). From a semiotic approach, it can be hypothesised that the receiver does not always confuse the eye-spot display for the face of an owl, but that two alternative sign sets and interpretations may be active simultaneously. In some cases (e.g. the European Peacock Aglais io) there are vivid eye-spots present in the wing coloration that as a whole is also quite conspicuous (see Figure 4.4). Therefore, it seems accurate to assume that the receiver perceives simultaneously the peacock as a butterfly and its eyespots as standing for the predatory animal. Conveying two contradicting set of messages makes the communicative encounter controversial and its outcome hard to predict—a bird would almost need to make a decision for what to do with a strange butterfly with owl’s eyes in its wings. [enter Figure 4.4 here] Figure 4.4. Eyespots and conspicuous coloration of the European Peacock Aglais io (photo courtesy of author). 63 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. To conclude this subchapter, describing mimicry as just a deception based on perceptual similarity simplifies the space of possibilities that sign systems provide to undermine the communication between the model and receiver. To include the full space of possibilities for our interpretation of mimicry, we should pay separate attention to the sign systems used in model-receiver (1) and mimic-receiver (2) communicative interactions and we should describe as well the different functional aspects that communication may have. Some arguments developed in this chapter will later be put to use in the discussion of semiotic modelling of mimicry. 4.4 The Umwelten of the receiver and the human observer 15 Analysing mimicry from a semiotic viewpoint makes it necessary to take into account the position of the human observer in regard to the mimicry system. In a scientific view, the description of mimicry is ideally considered to be fact-based, neutral and independent from human psychology and biological make-up or cultural context. In actual research we see, however, that human perceptual capacities are relevant and it is the mistaking of the mimic for model by humans that often initiates the forthcoming scientific research. Even Henry Walter Bates’s discovery of mimicry in Heliconid butterflies was launched by his endeavour to determine species correctly and by the observation that some individuals did not fit well with the existing taxonomy. In early mimicry studies of the 19th century, the similarity of mimic and model for the human eye was commonly treated as a criteria for describing the relation as protective coloration. To give an occasional example, H. L. Osborne, in his report on mimicry in marine molluscs to Science, describes camouflage in nudibranch as follows: “Placed upon a mass of Sargassum in an aquarium, the Scyllaea was hard to find, so closely did it imitate the appearance of the leaves […] It seems to me that there can be but little doubt that this creature presents another interesting case of mimicry” (Osborne 1885: 9–10). A tendency to emphasize apparent visual resemblances was also notably present in the writings as well as artistic works of American naturalist and artist Abbott Thayer, who depicted many animals in oil as having camouflage coloration in 15 This subchapter is partially based on Maran, T. (2007). Mimicry. In: P. Bouissac, A. Lewis (Eds.). Semiotics Encyclopedia Online. E.J. Pratt Library, Victoria University. http://www.semioticon.com/seo/. Used with permissions. 64 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. their natural habitat (Thayer 1909; Boynton 1952). He also made an attempt to use his artistic endeavour in the research practice: I made some wooden eggs about the size of a Woodcock’s body, and provided them with wire legs to poise them six inches above the ground. Most of these I colored in imitation of the color-gradation of a grouse or hare; earth-color above, to pure white beneath; while to two others I gave a coat of earth-color all over, above and below; then set the whole like a flock of ‘shore birds,’ on the bare ground in a city lot. I then summoned a naturalist and let him begin at forty or fifty yards to look for them. He saw immediately the two monochrome ones; but although told exactly where to look, failed to find any of the others, until within six or seven yards, and even then only by knowing exactly where to look (Thayer 1896: 318). In more recent studies, humans have been used in the position of receivers to assess the effectiveness of the mimicry systems (Golding et al. 2005; Moksnes and ØSkaft 1995). For instance, Yoanne Golding and her colleagues (Golding et al. 2005) judge the effectiveness of the hoverfly mimicry based on the discrimination skills of university students and schoolchildren. Without a concurrent comparative analysis of the perceptual capacities of humans and receiver, adopting a human to the receiver’s position in the mimicry system may lead to biased results due to the specifics of the human Umwelt and hidden anthropomorphism deriving from this. As living and perceiving beings, humans are limited by their Umwelt as well as by conceptual resources and cultural models. Therefore, the human's position as an observer of the mimicry resemblance cannot be neutral by any means. For a semiotically literate study, the specifics of the human Umwelt should be analysed as part of the research situation. At the level of prelinguistic processes, we should consider in which way the perceptual capacities of the human observer are positioned in relation to the mimic, model and receiver of the of mimicry system. As an observer and biological organism, humans belong to primates; that is, they have well-developed binocular colour vision (400–750 nm), intermediate capacities for auditory perception (20– 20000 Hz) and less then mediocre tactile and chemical perception. Through their natural perceptual organs, humans have relatively good access to visual and audible 65 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. messages. At the same time, the human observer lacks direct access to the messages in the chemical communication channel used by most mammals and by many insect species. We often compensate our limited perception by using special technical equipment: bat detectors to perceive echolocation at the range of 20– 200 kHz, UVcapable cameras to record ultraviolet colour patterns (<380 nm) and many other technical aids. The problem of technically mediated perception is, however, that we are not able to induce the research hypothesis from the direct observations, but need to construe the research problem based on the information that is received indirectly. This means, that in studying communication systems outside our perceptual reach, we first need to have quite a precise understanding of what to look for. Due to this epistemological hindrance, mimicry systems that are located beyond our perceptual range still remain largely unexplored. For apprehending the diversity of perceptual systems and signals in mimicry that remain outside of our perceptual reach, it would be helpful to adopt a biosemiotic perspective. This means acknowledging that qualitative perception and interpretation are not just epiphenomena of the underlying genetic processes, but have an essential importance for organisms to interact with their environment. From the biosemiotic perspective, every living organism is a centre of experience that construes its own surrounding world by using its physiological, cognitive and behavioural resources. Such an understanding stems from the works of Estonian-German biologist Jakob von Uexküll. In his writings, Uexküll claims that all organisms live in their speciesspecific subjective worlds or Umwelten, and that they are interrelated with the environment by functional cycles of perception and action, which are organized and mediated by meanings (Uexküll 1982: 27–33). Animal Umwelt consists only of those properties of the environment that are mediated by the animal sense organs and that are interpreted by the animal as meaningful. Uexküllian perspective is relevant to mimicry studies as it allows us to emphasise the organism’s specific position in mimicry. Namely, the organism’s behaviour and subsequent effects on other participants in mimicry appear to be Umwelt dependent (Maran 2007). In classical ethology, in a similar manner, the concept of search image or searching image (e.g. Pietrewicz and Kamil 1979, the concept in the English version was proposed by Luuk Tinbergen 1960 based on Uexküll’s concept of Suchbild) has been employed as the set of the object’s representative features that the specific animal uses in searching for food, mate or some environmental resource. 66 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Evidently some biological resemblances and mimicry systems are more easily accessible to humans than others. As mentioned before, there is a strong bias in mimicry studies towards visual mimicry, and this inclination seems to follow the human perceptual profile. Given the relative use of perceptual modalities by different animal groups in communication, there is no reason to assume that visual mimicry would be more abundant compared to, for instance, chemical mimicry. On the other hand, limits of human perception conceal several mimicry systems partially or to a full extent that makes their study problematic. For instance UV-patterns of plants and their role in plant-insect interactions became systematically accessible only since the 1950’s, when UV photography become widely accessible and used in research (e.g. Kevan et al. 2001). This created possibilities to study mimicry resemblances visible in UV wavelengths, as for instance deceptive signals of the insectivorous Nepenthes species. Carnivorous pitcher plant Nepenthes rafflesiana appears cryptic to the human eye. In UV range, it appears to have vivid patterning in the upper pitchers that have a considerable effect on catching flying pollinators, as it exploits the colour code used in flower-pollinator communication (Moran 1996). In UV mimicry, the entire mimicry system remains concealed from the human observer as messages sent by both models and mimics are outside of our perceptual reach. There are, however, mimicry cases, in which both the model’s and mimic’s messages are perceivable for humans, but they do not appear similar to us because of the specifics of our perceptual organs and Umwelt. For instance, Swedish ecologist Anders L. Nilsson has described the mimicry system between red helleborine Cephalanthera rubra and bellflowers Campanula where receivers are mostly solitary bees from the genus Chelostoma (Nilsson 1983). For the human observer, red helleborine and bellflowers are clearly discernible: the orchid has red flowers with a violet tone, while bellflowers have deep blue coloration with some violet hue depending on the species. For solitary bees, however, who pollinate bellflowers, the visual perception range is shifted towards higher wavelengths of the spectrum and they do not perceive reddish colours well. Therefore the difference between Helleborine and bellflower flowers remains concealed by the solitary bees and it can be considered as a fully operational mimicry system. For the human observer, noticing this type of mimicry in nature remains problematic and we may assume that many mimicry systems on the borders of our perceptual reach remain undescribed. It is also quite difficult for us to interpret the systemic properties of the environments that are unfamiliar to us, such as for instance 67 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. in underwater communication, where fading light with the increase of depth drastically and systematically changes resemblance between colour patterns (Cheney and Marshall 2009). In addition to the differences between the perceptual organs of the receiver and human observer, the differences in Umwelt structure and sign categories may also make it difficult to find potential mimicry cases. For instance, in the intraspecific mimicry of the ciclid Haplochromis burtoni, the male fish have a group of round spots on the anal fin. This specific pattern has an essential role for the reproduction of the fish. Namely, Haplochromis burtoni are mouth breeders. That is, their females gather roe into their mouth during the spawning, incubate it and carry thereafter fish larvae in mouth for several weeks. The female’s instinct to gather roe may prevent fertilisation, and to solve this problem, a peculiar intraspecific communication system has evolved. The male’s colourful spots on its anal fin direct the female to also grasp some milt with the roe, which makes fertilisation possible. (Wickler 1968: 221–227) The colourful spots on the anal fin of the male Haplochromis burtoni hardly resemble anything to the human observer, but to the female of the same species, to whom the dropping roe is an essential sign during the mating, the similarity is probably quite explicit. Based on the difference between Umwelten, we may easily overlook some mimicry resemblances just because the involved entities do not have any significance in our own Umwelt. In addition to perceptual differences and different Umwelt structures between human observers and receivers, the effect of human cultural interpretations of mimicry should also be taken into account. Some mimicry cases have an age-old history of research dating back to Greek Antiquity, some are related to wide-spread cultural narratives and beliefs, some involve species that are especially significant due to their place in natural history or because of a cuteness factor (so called charismatic species), whereas others are described de novo with very little earlier cultural context involved. I will make an attempt to discern the number of ways in which mimicry may gain relevance for human cultural processes as a narrative or a model of thinking. Mimicry may actualise in human culture as: 1. A typological confusion, where humans encounter similar species (e.g. making error in determining different mushrooms or other species); 2. An ontological problem of breaking borders between animal groups or realms (e.g. image of the skull in death's-head hawkmoth, Acherontia); 3. A similarity principle used in sympathetic magic or healing practices 68 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. (e.g. in ethnopharmacology). 4. An imaginative or real deception where the human may become a victim (e.g. motives of changelings or mermaids). 5. An imaginary or real strategy used by humans to cross borders of cultural or ontological domains (shapeshifters, carnival, role play); 6. In hunting strategies, where the human imitates some signs of the prey animal or an object neutral to it (use of camouflage, decoys and animal calls in hunting); 7. Artefacts and technical solutions that enhance human creativity by resembling biological structures (so called biomimetics). The different ways that mimicry can be relevant and participate in human culture may also have an effect on how we approach different mimicry systems in the biological world. To sum up the discussion, mimicry systems appear to be well describable for the human observer in cases where his/hers and the receiver’s perceptual organs and Umwelt structures are similar to one-another. If the Umwelten of the receiver and human observer are very different, the observer needs to make a more conscious effort to perceive and describe the mimicry resemblance. For describing mimicry systems that are perceptually inaccessible to humans, elaborate conceptual tools and theoretical models could be helpful in compensating for the Umwelt difference. 69 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 5 ICONICITY AND MIMICRY As it was shortly discussed in a subchapter 4.1 “Semiotic interpretations of mimicry”, the connection between mimicry and semiotics was originally established in the works of Thomas A. Sebeok. He was the first who expressed an opinion that mimicry is a semiotic phenomenon, discussed mimicry in several essays and included it as a separate keyword in semiotic handbooks. Also due to Sebeok’s interpretation, mimicry has later been mostly treated in semiotics as an example of iconicity in nature. The connection between mimicry and iconicity has been expressed and discussed, e.g. by Winfried Nöth, Frederik Stjernfelt, John W. Colleta, Göran Sonesson and others. However, taking mimicry as an example of an iconic sign, or a sign based on similarity between the representamen and object, is not as simple of an issue as it might appear at first glance. In the present chapter, the relations between mimicry and iconicity will be analysed by discussing the different types of iconicity in nature and the necessary conditions of the sign. I will rely here predominantly on Peircean semiotics and will later also discuss different mimicry types based on Peirce’s sign typology. 5.1 If mimic is a sign then what does it stands for? 16 In handbooks of semiotics, mimicry is often described as an example of iconic signs in nature (Nöth 1990: 163; Sebeok 1994: 84). It is easy to regard mimicry as an iconic sign since it intuitively seems to us to be a resemblance in the natural world that is meaningful or functional. According to Peirce, an iconic relation is one of the possible connections between the sign and the object it refers to (the other two being indexical and symbolic relations). Therefore, the question of iconicity has to do with the connection between the sign and its referent and not with any other dimension of the sign. An icon is a sign whose “qualities resemble those of that object, and excite analogous sensations in the mind for which it is a likeness” (CP 2.229), and “such a sign whose significance lies in the qualities of its replicas in themselves is an icon, image, analogue, or copy. Its object is whatever that resembles it its interpretant takes 16 This subchapter is partially based on Maran, T. 2011. Becoming a sign: The mimic's activity in biological mimicry. Biosemiotics, 4(2), 243–257. Used with permissions. 70 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. it to be the sign of, and is as sign of that object in proportion as it resembles it.” (MS [R] 7:14/5)17 The condition of resembling qualities between the mimic and model seems to be met in most examples of mimicry. In addition resemblance, a sign also needs to have a representational character—the sign (in a narrow sense, representamen) needs to stand for its object. Thus when assessing the connection between mimicry and iconicity, we should ask—what is the sign in mimicry and for what does it stands for? This question was also asked by Swedish semiotician Göran Sonesson (2010) with a negative conclusion that mimicry does not have a sign structure because the criterion of representativeness is not met. Posing this question, we should also differentiate between the position of the human observer and that of the signal receiver as two different interpreters of the mimicry system, as was done in the previous chapter. For instance, to the human observer, the yellow and black stripes of a hover-fly may resemble the coloration of wasps and thus signify a functional mimicry system. The same is probably not the case for a small insectivorous bird (such as a great tit Parus major) who has had unpleasant experiences with wasps. For the great tit, a hover-fly does not signify or represent (to stand for) a wasp in a strict sense, and therefore the position of the representamen cannot be allocated to the hover-fly and the position of the object to the wasp. In such a mimicry case (that follows the logic of Batesian mimicry), the relationship between the mimic and the model appears to be, on most occasions, about sameness/difference and not about representation. But at the same time, the iconic reference appears to be present in many cases of abstract mimicry (e.g. eyespots and deimatic displays). In cases where the mimic actively creates or adjusts the mimetic display, iconic sign creation should also be considered to be taking place (cf. Chap 7.1). So can a mimetic organism function as a sign and what would it stand for? A possible line of argumentation could follow the thinking of American semiotician Charles Morris. He has emphasised that iconicity is a matter of degree rather than type, so that iconic signs have a great variety from vague resemblance to close similarity between the representamen and its object (Morris 1971b: 273). If this is so, there should be a class of icons in which the representamen’s similarity with its object 17 Original manuscript, Robin Catalogue id. Derived from http://www.commens.org/dictionary/term/icon at 31.01.2016. 71 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. is perfect. Such icons could be named absolute icons, and an example of this would be a mirror image (Eco 1984: 207–211). If there is absolute similarity between the representamen and its object, the relation seems to lose its representational character and turn into sameness. In principle, one could argue that this applies to the mimicry system: that for the signal receiver, mimics have often become absolute icons of models. Another possible direction of argumentation for the sign nature of mimicry would be to take a much wider, perhaps even a pansemiotic perspective. This appears to have been the choice of Thomas A. Sebeok, who takes hold of Peirce’s description of icon as a sign where “there is a mere relation of reason between the sign and the thing signified” (CP 1:372). Departing from this quotation, Sebeok argues that mimicry “must be integrated, in toto, with the far more general and deep theory of iconicity” (Sebeok 1994: 84). However, there is a third possibility to show the sign nature of mimicry, that I believe is more productive and less controversial. To understand this, we need to consider a general zoosemiotic principle of communication in nature. In the majority of animals we find colours, shapes, sounds, smells and other features that are specific to the particular species and express the animal’s species identity for its species mates as well as for symbionts, competitors and representatives of other groups. Thus, the red plumage of a male bullfinch’s (Pyrrhula pyrrhula) chest can be considered as a sign with meaning: here is a male bullfinch (and not a female or young or a greenfinch). Ethologist Peter Marler, who has followed Charles Morris’s typology of signs, has described such signs that can give different types of information (speciesspecific, sexual, individual, motivational, and environmental) about their carrier as designative signals (Marler 1961: 302). Concerning the present discussion, the most relevant from these types would be species-specific designators and also sexual designators, that in the above-mentioned case would express: I belong to the class of male bullfinches. Focusing on the designators in biocommunication would allow us to connect mimetic resemblance with a more general iconicity in nature. In Peircean terminology, individual signs can be described as tokens that stand for their type; that is, they are iconic signs whose primary semiotic function is to indicate their belonging to their class of signs. In addition, they can relate to some other interpretants, such as applicability of this class of objects. For instance, the red plumage of the bull-finch 72 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. may signify a suitable catch for a pine marten (Martes martes). 18 In the words of Charles S. Peirce: “In order that a Type may be used, it has to be embodied in a Token which shall be a sign of the Type, and thereby of the object the Type signifies” (CP 4:537). The token—type relationship is intrinsically iconic. The existence of such sign relation is based on the general condition of semiotic processes that individual signs form groups and that the semiosphere has a discrete structure. One consequence of this argumentation is that in mimicry, not just entire organisms should be described as bonded by sign relations, but rather the specific cues or messages that take part in semiosis. This would support the general idea expressed in the present book that mimicry has a double layered structure: one of ecological relations between species and one of semiotic relations of signs. Thus, the question of whether the mimic can refer iconically to the model (for the receiver) finds a negative answer, whereas the question of whether the signs in the animal body or communication can have an iconic referent warrants more serious consideration. In a similar way, the species-specific types described above do not comprise the entire body and all expressions of an animal individual, but are rather focused on some characteristics that work as species-specific markers. This is similar to the everyday experience of working with field guides, when naturalist does not need to pay attention to the whole appearance of an animal but can concentrate on some speciesspecific features. To sum up, rather than signifying it belonging to its own species or group, a mimetic sign indicates that its carrier belongs to the type of some other species. For a small insectivorous bird, the mimesis of the leaf insect indicates its belonging to a type of tame and monotonous foliage. The threat display of a stick insect, where an insect flashes colourful spots on its wings, indicates that a stick insect has a connection to some indistinct and unexpected danger. The mimetic effect and tension arise from a discrepancy between the type of organism that the mimic essentially is and the type of organism that the mimetic sign that it carries implies it to be. Consequently, the mimic’s activity in becoming a sign is aimed at resembling a certain type, and a mimetic sign is basically a false designator. 18 This secondary relation appears to correspond to what Frederik Stjerfeld has called dicisign structure of animal communication, which makes it possible to „convey information—to rely claims, assert statements true or false” (Stjernfelt 2014: 54). 73 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 5.2 Peirce’s second trichotomy and animal communication To better understand mimicry as a sign phenomenon, it is not enough to focus on the Peircean icon only. We should also consider other possibilities of the icon–index– symbol trichotomy. To avoid a possible misunderstanding, let me state clearly that the second Peircean trichotomy of the sign is considered to describe the sign-object relation exclusively and not any other aspect of the sign. Peirce’s second trichotomy distinguishes possibilities for how the sign can relate to its object: I had observed that the most frequently useful division of signs is by trichotomy into firstly Likenesses, or, as I prefer to say, Icons, which serve to represent their objects only in so far as they resemble them in themselves; secondly, Indices, which represent their objects independently of any resemblance to them, only by virtue of real connections with them, and thirdly Symbols, which represent their objects, independently alike of any resemblance or any real connection, because dispositions or factitious habits of their interpreters insure their being so understood. (EP 2:460-461) From these three sign types, index is considered to be most characteristic to animal communication. As Winfied Nöth writes in his “Handbook of Semiotics”: “it is hard to find signs in animal communication devoid of indexicality. The degree of indexicality in animal semiosis is definitely higher then in human semiosis.” (Nöth 1990: 163). The common examples of indexical signs are, for instance, landmarks used for navigation by migrating birds, claw marks of a bear in a tree trunk corresponding to the height of an animal and stripes and spots in animal coloration that draw attention to significant body areas. Animal tracks are indexes as they refer to the presence of an animal as well as its moving direction and behaviour in the environment (cf. Vladimirova and Mozgovoy 2003). The shape, size and distance of animal footprints, rhythm and orientation of tracks can provide a lot of information about the animal’s speed, motivation and intentions. Based on the type and speed of animal movement, field guides distinguish track patterns, such as a walk, trot (distinguishing side trot and straggle trot), pacing, lopes, gallops and different types of jumps (Liebenberg et al. 2010: 33–50). In the framework of evolutionary biology, indexes are treated as signals that ensure truthful communication due to the motivated 74 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. relationship between the signal form and the content (Maynard Smith and Harper 1995; for discussion, see Maran 2009). The presence of symbolic signs in animal communication is a much-disputed topic. There is a widespread opinion in biosemiotics (e.g. Deacon 1999; Kull 2009) that true symbols and symbolic communication exists only in human culture. There are also alternative views. For instance, João Queiroz and Sidarta Ribeiro analyse alarm calls of vervet monkeys and argue that this is an example of symbolic communication. Vervet monkeys, as it is well known, use the system of different alarm calls to refer to different predatory groups (ground predators, flying predators, snakes, Cheney and Seyfarth 1990). In their article “The biological substrate of icons, indexes and symbols in animal communication”, João Queiroz and Sidarta Ribeiro (2002) describe an experiment with vervets, in which different alarm calls were recorded and played back to monkeys. Animals responded to the experiment by looking around for a referent, and then fleeing to nearby refuges according to the specific type of call played (‘leopard’ calls evoked tree climbing, ‘eagle’ calls evoked bush hiding). Queiroz and Ribeiro conclude that alarm calls hold a previously established relationship to the predators they stand for, and are symbols because they operate in a sign-specific way, even if the an external referent is absent. Thomas A. Sebeok has also stressed that applying symbols to animal communication depends on the specific definition used but by “invoking the principle of arbitrariness, the idea of a conventional link between a signifier and its denotata, Peirce’s ‘imputed character,’ or the notion of an intentional class for the designatum—animals demonstrably employ symbols.” (Sebeok 1990b: 42). In evolutionary biology, signals used in courtship such as the tails of the peacock or lyrebird are considered to be symbolic by John Maynard Smith and David Harper on the basis that “form is unrelated to actual fighting ability” (Maynard Smith and Harper 2003: 59). Several arguments indicate that the presence of symbolic communication in animals should be considered probable, at least, when focusing on the sign relations that are based on the phylogenetic level; that is, sign relations that are inborn and not rooted in psychological processes. To be clear, the icon-index-symbol distinction is considered here as applicable to the relationship between the representamen and the object, and the question is to what extent and in what way the objectrepresementamen connection is motivated or habit based. The discussion of the cognitive capacities of the animal interpreters should, in my understanding, be 75 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. addressed in the framework of Peirce’s semiotics as the third trichotomy (rheme— dicisign—argument). 19 The question of whether animals are able to use conventional signs would translate into the question of whether animals can make use of arguments. Argument, according to Peirce: “is a sign which is understood to represent its object in its character as sign” (EP 2: 292). Thus, argument introduces a distinction between object-level and meta-level semiosis, which, indeed, occurs very rarely in animal communication, if at all. When describing iconic sign relations in nature, attention is turned to similarities. In his essay “Iconicity”, Thomas Sebeok presents an example of a symbiotic relationship between ants and aphids’, which is successful because of the topological similarity between an ant’s head and an aphid’s abdomen (Sebeok 1989: 116). Similarities are employed in communication systems by many animals, for instance in mating rituals that repeat many elements of fostering behaviour between offsprings and adults. Female passerine birds beg for food, by taking a remarkably similar posture to youngsters of the same species. In many ungulates such as antelopes or gazelles, the male can push the female’s abdomen with his head during the mating ritual in a similar way that youngsters do before suckling the milk (Wickler 1974). Diagrammatic or relational icons are described in pheromone signals of social insects and alarm calls of the passerine birds, where the intensity or length of the signal corresponds to the size or relevance of the referred object (e.g. Templeton et al. 2005). In the case of the icon, the sign is evoked just because of the representamen’s qualities. According to Peirce: “An Icon is a sign which refers to the Object that it denotes merely by virtue of characters of its own, and which it possesses, just the same, whether any such Object actually exists or not” (CP 2: 247). This quotation also expresses the paradoxical nature of the icon. As the manifestation of Firstness, pure iconicity should not refer to anything else (cf. Eco 2000: 100). When it does so, something second must be added, and the iconicity recedes to the iconic sign that is often called “hypoicon” (Ransdell 1997[1986]). Sometimes these two aspects are differentiated as iconicity and iconic signs, the latter being mediated by similarity, or iconic ground. Göran Sonesson writes: “Perception of similarities (which is an iconic 19 Distinction of rhema, dicent and argument has been seldom used in analysing animal communication (a positive example being Stjernfelt 2014). 76 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. ground) will give rise to an icon only when it is combined with the sign function” (Sonesson 2012: 84). Peirce has described icons as “originalian sign” (CP 2:92) that is explained as a “mere possibility of something, at its beginnings, nascent” (Braga 2003: 50). The peculiarity of icons to lie at the border or at the beginning of semiosic activity seems relevant for mimicy studies, as mimicry also functions in the perceptual threshold by lingering between perception and non-perception, recognizing and non-recognizing, meaning-making and lack thereof. To make practical use of the Peircean categories for comprehending mimicry, we should not take icon, index and symbol as pre-given types, but rather as relative categories. This would mean that indicating some natural phenomenon to be an icon, index or symbol should be done in the context of this particular sign process. It also means that different sign properties do not need to exclude one-other but that an object can participate simultaneously in sign relations that are iconic, indexical or symbolic. Furthermore, different sign types include each other in such a way that an index may include iconic properties and symbol may include iconic and indexical properties. “A Symbol is a law, or regularity of the indefinite future. […] But a law necessarily governs, or “is embodied in” individuals, and prescribes some of their qualities. Consequently, a constituent of a Symbol may be an Index, and a constituent may be an Icon.” (EP 2:274). The same is true for the index. Peirce wrote: “In so far as the Index is affected by the Object, it necessarily has some Quality in common with the Object, and it is in respect to these that it refers to the Object. It does, therefore, involve a sort of Icon” (EP 2: 291–292). The interrelatedness of Peircean sign types is relevant for a semiotics of mimicry, as it allows us to describe mimicry not just as an example of iconic signs, but also as an index if the mimic is perceived as spatially related to the model, and as symbol if the sign relation embraces some natural communicative convention. Instead of seeing mimicry as belonging to the fixed category of icons, we should rather look for the relational dominant of the specific mimicry system and focus on different ways that mimicry makes iconicity occur in nature. 5.3 Peircean categories and the three basic mimicry types I have argued before that mimicry seems iconic to humans mostly because of our external position as an observer in relation to the mimicry triad, and that makes it 77 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. convenient for usto compare mimics and models and to interpret their relationship as iconic. For the receiver inside the mimicry system, an iconic reference may be present but it does not need to be a necessary condition for the mimicry system to exist. At the same time, this conclusion does not mean that the Peircean sign typology would be useless for organising our understanding of mimicry. Rather, it appears that mimicry cases can be described as having different dominants in regard to the icon–index– symbol triad. I would like to start the following argumentation with a long quotation by Terrence Deacon, who has related iconicity to a specific type of mimicry resemblance—camouflage: Consider camouflage as in the case of natural protective coloration. A moth on a tree whose wings resemble the graininess and color of the bark, though not perfectly, can still escape being eaten by the bird if the bird is and interprets the moth’s wings as just more tree. Admittedly, this is not the way we typically use the term iconic, but I think it illuminates the most basic sense of the concept. [---] Their [moth’s wings] coloration was taken to be an icon because of something that the bird didn’t do. What the bird was doing was actively scanning bark, its brain seeing just more of the same (bark, bark, bark …). What it didn’t do was alter this process (e.g., bark, bark, bark, not-bark, bark…). It applied the same interpretative perceptual process to the moth as it did to the bark. It didn’t distinguish between them, and so confused them with one another. This established the iconic relationship between moth and bark. Iconic reference is the default. (Deacon 1997: 75–76). This passage has received some attention, both positive and rather critical. My aim here is not to enter into dispute over Deacon’s comprehension of iconic reference that has been critically discussed, for instance, by Göran Sonnesson (2010), but rather to emphasize the nature of the example that Deacon gives. He describes the moth lying in the tree bark. The chosen example is significant in two aspects. First, mimic and model are physically together and it is possible to perceive these as one and the same. Second, in camouflage, the decisive point for the receiver lies exactly in perceiving or not perceiving the mimic. Here one can see perception of a moth emerging from nowhere and that allows relating camouflage to iconicity and to other concepts that 78 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. are derived from Peircean firstness. Camouflage is possibly as close to pure iconicity in Peircean terms as it is possible to get. The same is valid to most cryptic coloration, disruptive coloration and cryptic mimesis. These conditions are not, however, applicable to all cases of mimicry, especially when we are dealing with deceptive similarity between representatives of two species. Such one-to-one resemblance is common to classical Batesian mimicry, as for instance similarity of two species of tree frogs, where one is edible and the other is poisonous, e.g. Dendrobates fantasticus (model), Dendrobates imitator (mimic). Both species have conspicuous orange-black patterning and live in the primeval rain forests of Northern Peru. In such a case, the decisive point for the receiver is not in perceiving the mimic—because animals carry vivid coloration and are easy to spot—but rather in recognizing and typifying organisms correctly. This type of mimicry requires comparison, a reference to the second, which is a characteristic feature of indexical semiosis. Depending on the specific mimicry case, the receiver may compare the mimics and models with its generalised image in mind, but there may also be situations when mimics and models are physically together in one place and a one-to-one comparison is possible. The latter is the case, for instance, in egg-mimicry of the common cuckoo and other brood parasites, as well as myrmecomorphic insects inhabiting ant nests. Indexicality appears to be a dominant sign feature in many forms of Batesian mimicry, aggressive mimicry, Mertesian mimicry and phaneric mimesis. In an earlier writings, I have described the distinction between crypsis (cryptic mimesis) and Batesian mimicry from the receiver’s perspective as two different possibilities for how the mimic can find shelter in the receiver’s Umwelt: The difference between mimicry and crypsis becomes even clearer when we consider Uexküll’s concept of Umwelt. When the model of resemblance plays an important part in the receiver's Umwelt, there is mimicry, but when the mimic tries to dislocate its perceivable properties to a world outside of the receiver's Umwelt by resembling some object that does not belong to this receiver's Umwelt, thus making its message meaningless to the signal receiver, there is crypsis. (Maran 2001: 331) 79 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Later, French semiotician Marie Renoue and biologist Pascal Carlier (2006: 121) have considered this distinction to be theoretically inconvenient and to introduce little economical use value. Yet, I am convinced, that for understanding the difference between camouflage and mimicry, the receiver’s Umwelt structures need to be taken into account. Inasmuch as different sign types reflect the specifics of animal Umwelten, the distinction between iconic, indexical, and symbolic mimicry can be taken as a more convenient and formalised manifestation standing for the underlying differences of the receiver’s Umwelt structures. The two mimicry types distinguished above do not, however, cover all possible mimicry cases in nature. There are also many hard-to-classify resemblances that are described as abstract or imperfect mimicry. Good examples of abstract mimicry are eye-marks in the bodies of insects, fishes and amphibians and colourful patches in the bodies of many lizards and insects that are kept hidden in the resting position, but that are flashed during the escape. Characteristic of abstract mimicry is the presence of an abstract model that is not definable down to the species level, or there is an imprecise, ambivalent similarity between the mimic and the model. A unifying aspect of different cases of abstract mimicry appears to be some common or generalised meaning, as for instance big eyes, unexpected movement or warning coloration that all signify something dangerous. Abstract mimicry requires comprehension of this general meaning complex and therefore it seems to have a dominant in symbolicity and thirdness. The issue here is not about typifying or correctly recognising deceptively similar organisms, but rather, to determine if the receiver knows the meaning of the messages and if it relates the specific instance with the general abstract category. To conclude the discussion on mimicry and iconicity—mimicry does not need to fit under iconic signs per se, but at the same time, Peircean categories appear to be good analytical tools to characterise properties of different mimicry cases. Mimicry makes use of the following critical points in the semiotic process: whether the sign is perceived (foreground in iconicity), whether the reference to the second is made (foreground in indexicality) or whether the general meaning of the sign is known (foreground in symbolicity). This corresponds to three possible grounds on which the sign relations in mimicry are built: 1) limited perception of a receiver; 2) bodily similarities (accidental or phylogenetically constrained) between the living forms; 3) conventions in meaning making and communication. It is the third ground of the sign 80 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. processes especially that is independent enough of particular forms of nature that it may connect a broad number of different species, thus producing natural conventions that will be discussed in the final chapter of this book as ecological codes. 81 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 6 SECOND EXCURSION: IMPORTANCE OF THE OBJECT There is an age-old distinction between dyadic and triadic sign models in semiotics. Most of dyadic sign models describe the relationship between the meaning and the carrier of that meaning (e.g. the relationship between content and form or signifier and signified). A classical example of the dyadic sign model is the model of the linguistic unit introduced by a Swiss linguist and semiotician Ferdinand de Saussure. Triadic sign models add a third component to the sign—an external referent in relation to which the sign has its meaning. In the semiotic tradition stemming from the works of Charles S. Peirce, this aspect of the sign is called the “object” and the triadic sign, correspondingly, consists the relationship between the representamen, object and interpretant. The specific relationship between the sign and its object is a controversial topic. One source of problems appears to be the issue of causality or determinacy of the sign process and can be expressed as follows: to what extent is the sign autonomous in regard to its external referent? To explain it a little coarsely, there are two possible extreme answers to this question that lead us, respectively, into two opposite but wide-spread intellectual camps. If there is no autonomy in the sign, then the external referent determines the meaning and interpretation of the sign. Such a position would lead us to positivist biology, classical ethology and animal psychology, where the external stimulus is considered to trigger a behaviour of an organism and communication is described as a series of stimulus-response sequences. To another extreme, the sign is absolutely autonomous in regard to its external referent, which means that there is unlimited freedom for possible interpretations of the sign to occur. In this case, every possible interpretation is a correct one, and the intellectual camps we are approaching are those of cultural relativism and postmodern thinking. Several authors have expressed the view that in addition to these two major intellectual traditions, biosemiotics has a third approach to offer. British semiotician and communication scholar Paul Cobley has described the role of biosemiotics in bridging the general logics of natural sciences and humanities in terms of agency: Biosemiotics introduces agency into nature and, in the process, will re-draw the lines of science and the humanities. The problem with the latter is that 82 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. culture already has agency in bucketloads—indeed, it has more than it knows what to do with. […] The main cultural implication of biosemiotics — as opposed to the most glaring — must be, then, that the dun scientism of mechanistic thought, sociobiology and determinism which has been rejected by culture does not have to be replaced by autonomy, agency, the (free) will and other myths. Mechanism can be replaced by a perspective that is not just culture-friendly (agency in nature) but provides a scientific outlook (continuity of organisms, matter and mind) which can underpin, while qualifying, understanding of culture’s cherished irreducible features, and at the same time also undermining the extravagant individualist, anthropocentric, glotto-centric, claims of rampant culturalism (Cobley 2010: 241). The potential of biosemiotics to surpass the cleavage between natural sciences and humanities that Cobley calls for appears to be at least partly connected with the position of the object and its relation to the sign. In this context, mimicry as a research model has an especially significant position. After all, a central issue in mimicry is how the receiver interprets the relationship between a sign and its object: does it recognize mimics and models correctly and thus create an adequate meaning connection between the sign and the object? Or does it mistake the mimic for the model and establish a meaning connection that is not supported by the properties and applicability of the objects themselves (mimics and models as real organisms)? What appears to lie at the heart of the mimicry system is precisely the question about the relationship between the sign and its object. In mimicry, the interpreter does not appear to have full freedom in interpreting the mimicry situation. Its decisions are constrained by the appearance, properties and applicability of mimics and models as embodied physical organisms. It has the possibility to make a mistake, a possibility that can appear only in cases where the object also has properties that are independent of the sign relations. But at the same time, the receiver’s decision also appears to not be fully determined by the object, as such a situation would exclude any behavioural dynamics and evolutionary development from the mimicry. The mimicry is rather a phenomenon where the sign is partly dependent and partly independent of the object, which introduces a kind of constrained freedom to the system as it is discussed by Jesper Hoffmeyer under the 83 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. concept of “semiotic scaffolding” (this topic will be addressed in Chap. 9.2 “Mimicry and semiotic scaffolding”). A conceptual tool that seems to be suitable for describing this sort of semiautonomous relation is Charles S. Peirce’s distinction between immediate and dynamic object as two aspects of the object of a sign. In Peircean semiotics, the immediate object is a notion denoting the object as it is revealed within a sign and the dynamic object denotes the object outside of the sign. We know the dynamic object by “collateral”, i.e. indirect knowledge (CP 8.314), for instance, “[i]n the example of animal tracks, the immediate object would be the knowledge of an elk as it appears to us by looking at the tracks, and the dynamic object would be the elk as it is, or the elk as the sum of all other experiences of it” (Maran 2010a: 83). Peirce’s approach uniquely allows us to treat two aspects of the object—material object and perception of this—as both existing but still as being connected to each other. At the same time, the correspondence between the dynamic and immediate object is never absolute and complete but rather an approximation, a way forward to. In the words of Peircean scholar Donald Ransdell: [T]he immediate object is the object as it appears at any point in the inquiry or semiotic process. The [dynamic] object, however, is the object as it really is. These must be distinguished, first, because the immediate object may involve some erroneous interpretation and thus be to that extent falsely representative of the object as it really is, and, second, because it may fail to include something that is true of the real object. In other words, the immediate object is simply what we at any time suppose the real object to be. (Ransdell 1977: 169) This special relationship between the dynamic and immediate object also supports Peirce’s pragmaticism, where the sign process is understood as an advancement towards a better correspondence between the sign and its object and correspondingly towards the truth. This can be a goal of scientific inquiry, but as we see from the example of mimicry, it may also be an aim of semiotic process in the natural world. In the context of mimicry, the connection between immediate and dynamic object renders as a connection between the mimic and model as actual biological organisms with their properties and applicability, on the one hand, and mimic and model (or their 84 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. messages) as perceived by the signal receiver, on the other hand. If for Peirce’s pragmaticist view, the final aim of the sign process is to create an ideal correspondence between the dynamic and the immediate object, then in mimicry, it is the receiver who learns through correct and mistaken interpretations about the actual properties of the mimics and models. Thus mimicry becomes a learning device and the receiver becomes someone who aims to acquire better knowledge about the surrounding environment. Such a specific relationship between the sign and object may actually be rather widespread in animal semiosis, but due to two alternative interpretations bonded together, mimicry illustrates this property of living systems in a most excellent manner. 85 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 7 DIFFERENT PERSPECTIVES IN MIMICRY SYSTEM In the first half of this book, I have described the structure of mimicry, its diversity and typologies, given an overview of the semiotics of mimicry and analysed the relationship between mimicry and iconicity. In other words, I have analysed mimicry basically as a static structure or system with a shifting focus on its different aspects. In the present chapter, I will change the perspective and predominantly analyse dynamic issues: how mimicry as a structure operates as approached from the positions of the mimic, the model and the receiver, how they act to fit into the mimicry system, and what the different strategies are that they can use to cope with the other participants in mimicry. When analysing theoretical literature on mimicry, we can see that many works emphasise either one participant of the mimicry system or the relationship between two participants and describe mimicry from this particular viewpoint. Logically it is possible to distinguish between three basic relationships in mimicry: 1. communicative relationship between the model and the receiver; 2. resemblance-based relationship between the model and the mimic; 3. deception-based relationship between the mimic and the receiver. Depending on which relationship is preferred, specific properties of the mimicry system will be put in the foreground. Thus, the relationship between the model and the receiver relates to common forms and models of animal communication. Mimicry is often explicitly related to the properties of the model that provide protection or other benefits to the mimic. An eminent mimicry scholar Hugh B. Cott has noted that: „The theory that the mimicking species are benefited by a superficially deceptive resemblance to a model which is either dangerous or distasteful to their natural enemies, or to one which is not feared or avoided by their prey, rests ultimately upon the validity of the theory of warning coloration—since the animals resembled are themselves typically aposematic.” (Cott 1957: 396). From the position of the communicative encounter between the model and the receiver, the mimic can be seen as an eavesdropper or as a communicative parasite. The mimic disturbs the normal flow of communication between the model and receiver (cf. Endler 1993; Kopp and Mills 2002). The perspective that focuses on the model–mimic relation highlights the issue of their similarity or difference. Such an approach to mimicry reaches back to the 86 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. beginning of mimicry studies. The external similarity was a focal point in the explanations of natural theology (Blaisdell 1982) as well as in early Darwinian mimicry accounts (Bates 1862). In 1862, Henry Walter Bates specifies mimicry to be “resemblances in external appearance, shape, and colours between members of widely distinct families [...] The resemblance is so close, that it is only after long practice that the true can be distinguished from the counterfeit, when on the wing in their native forests” (Bates 1862: 502, 504). From contemporary field research, the relationship between the mimic and model is focused on in treatments that study spatial and temporal dynamics of mimic and model populations. The relationships between mimic and model populations can be analysed by mathematical modelling (nt. Waldbauer 1988; Huheey 1964; Franks and Noble 2004) and by different ecological studies emphasising spatial arrangement of species. The correlated change of the appearance of mimics and models has been described under the concept of mimicry rings that denotes local morphs that include individuals from many different species (Mallet and Gilbert 1995; Mallet and Joron 1999). Especially in tropical rain forests, butterflies form mimicry rings that either inhabit neighbouring microhabitats or even different horizontal vegetation layers. The third possible approach in the study of mimicry is to focus on the relationship between the mimic and the receiver, and often pays attention to strategies and adaptations that mimics use to mislead receivers. This approach is well suitable to behavioural ecology and other Neo-Darwinian paradigms that stress the deceptive component as not just characteristic to mimicry, but to all communicative relations (e.g. in interspecies courtship rituals, communication between adults and offspring, etc., Maynard Smith and Harper 2003: 3–4). Focusing on the relationship between the mimic and receiver could also be a reasonable choice in cases where the mimic’s imitating properties are too abstract for constructing the specific similarity relation with the model (e.g. in abstract mimicry). The deceptive aspect can be in the foreground in behavioural studies as well, as for instance in the experiments of classical ethology on learning capacities of the receiver when encountered with the mimetic organism. Studies by Jane Van Zandt Brower and Lincoln Pierson Brower launched the understanding of resemblance between mimics and models as a behavioural dilemma for the signal receiver (Brower 1960; Brower and Brower 1962). The three perspectives that are distinguished above are definitely not exclusive to one-another. In-depth reviews and monographs often deal with all three 87 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. relationships of the mimicry system. Still, distinguishing between communication, resemblance and deception as three central focuses could be proposed as a useful heuristic devise for organising different understandings of mimicry. 7.1 Mimic’s activity and intentionality20 Distinguishing the three participants in mimicry and their relationships brings along the possibility to analyse mimicry from different perspectives. The position of the mimic and its connection to the model is especially interesting for semiotics, as it relates to the question of deliberate deception and hence intentional semiosis in the animal world. In his essay, “Can animals lie”, Thomas A. Sebeok focuses on the position of the mimic and describes various organisms that carry deceptive characteristics (Sebeok 1990a: 95–96). In his analysis, Sebeok seems to emphasize the position of sender rather than that of receiver. This is in compliance with Sebeok’s later theoretical stance: to describe different types of signs in connection with various modelling strategies, i.e., from the position of the utterer and sign creation than that of the receiver and sign perception. For example, in the book “The Forms of Meaning. Modeling Systems Theory and Semiotic Analysis,” which seems to be one of the most unified syntheses of Sebeok’s views (written together with Marcel Danesi), iconic signs are defined on the basis of the features of sign creation: A sign is said to be iconic when the modelling process employed in its creation involves some form of simulation. Iconic modelling produces singularized forms that display a perceptible resemblance between the signifier and its signified. In other words, an icon is a sign that is made to resemble its referents in some way’ (Sebeok and Danesi 2000: 24). In “Signs: An Introduction to Semiotics”, Sebeok emphasizes the relation between the iconic sign and the concept of ‘mimesis’ (imitation) from the Platonic-Aristotelian discourse (Sebeok 1994: 28; see also Sebeok 1989: 110). From the position of mimic, the process of changing or altering itself (or the surrounding environment) in order to 20 This subchapter is partially based on Maran, T. (2011). Becoming a sign: The mimic's activity in biological mimicry. Biosemiotics (Springer), 4(2), 243–257; and Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica (DeGruyter Mouton), 167(1/4), 223–248. Used with permissions. 88 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. resemble the model can be considered as a creation of iconic resemblance. The connection between mimicry and iconicity is further illustrated by Sebeok’s description of the behaviour of the spider who changes “its surroundings to fit its own image by fabricating a number of dummy copies of itself to misdirect predators away from its body, the live model, to one of several replicas it constructs for that very purpose” (Sebeok 1989: 116). Orb-weaving spider, Cyclosa mulmeinensis, common in Asia, indeed uses remains of its prey and eggsacs to construe objects that, when wrapped into spider silk on the web, resemble the real spider both in size and coloration (Tseng and Tso 2009). As it was observed by Tan and Li 2009, the decoys help to camouflage the spider and lower the predation rate by Hymenopteras and insectivorous birds. In Sebeok’s semiotic treatment, the mimic’s active involvement in creating the deceptive resemblance appears to become a criterion of mimicry. From a biological point of view, emphasizing the mimic’s activity as a representational feature of mimicry is more questionable, as there are also many mimicry cases in nature where the mimic holds quite a modest position. In terms of the mimic’s behavioural activity, the example of self-copying spiders mentioned above is an exceptional situation. To systematize different deceptive behaviours in nature, animal psychologist Robert W. Mitchell has distinguished four levels of deception according to the sender’s freedom to act. On the first level, the sender deceives because it has been designed to do so and cannot do otherwise. On the second level, deception is largely predetermined, but for its expression, the sender needs to come into contact with the receiver and trigger the deceptive demonstration actively. On the third level, the sender is capable of customizing the pre-existing behavioural patterns and repeating the successful deceptions based on experience and learning. On the fourth level of deception, which is characteristic to humans and to some extent anthropoids, the sender takes into consideration the past activities of the receiver and can customize deception depending on the receiver’s response in a particular communicative situation. (Mitchell 1986: 21–27). Many observations have been made of deception in higher mammals such as elephants, chimpanzees or polar foxes, and such cases of deception could even be considered intentional in the sense that they are behavioural novelties, used by specific individuals to settle particular social contentions (Morris 1986; Rüppel 1986; Waal 1986). At the same time, these occurrences, which belong to level three or even four in Mitchell’s classification, are usually not considered as examples of biological 89 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. mimicry. Many classic examples of mimicry rather belong to level two in Mitchell’s classification, as they are characterized by genetic determination and some behavioural activity. However, there also exists a multitude of examples of mimicry that belong entirely to the first category of Mitchell’s classification, because the mimic as an individual does not express any communicative activity to become similar to its model. The involvement of an individual’s behavioural activity in mimicry varies from the absolute minimum in fixed bodily displays to fully dynamic deception. For example, the mimic’s activity is minimal in the expression of most camouflage patterns. The same is also true of many examples of mimicry in plants that resemble either physical objects of the environment, other plants or even animals. Delbert Wiens gives an example of the passiflorous host plants (passion flowers) of the Heliconius butterflies, which have developed modified stipules resembling egg masses of butterflies. As Heliconius butterflies choose vacant plants for laying their eggs to avoid later cannibalism between caterpillars, the plants with false eggs are thus left undamaged by feeding caterpillars (Wiens 1978: 376; Benson et al. 1975: 670–671). In other cases, the mimic’s behavioural activity plays an essential role in bringing the deceptive display into existence. As an example of this, Mark D. Norman and his colleagues describe the mimic octopus (Thaumoctopus mimicus). This species that was discovered only in 1998 and has become a celebrity of animal behaviour research. The mimic octopus uses its flexible body to imitate movements and different body shapes of predators and venomous animals of its marine environment, such as flatfishes, lionfishes and sea-snakes (Norman et al. 2001). Images of different animals are performed mostly by using different positions of arms, e.g. for imitating sea snake posture, “six arms were threaded down a hole and two were raised in opposite directions, banded, curled and undulated” (Norman et al. 2001: 1755). The imitation of the body shape is strengthened by simulation of the movement, for instance, swimming in the pace and rhythm of the fish species. Dynamic mimicry has later been described in other octopus species as well (Krajewski et al. 2009; Hanlon et al. 2010). What is remarkable in this case is that resemblance is created purely through behavioural activities in changing body shape and coloration, as the body of the octopus is flexible and lacking any skeletal structure. In the mimicry display of the Thaumoctopus mimicus, the intentional involvement of the octopus appears indeed to 90 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. be very high (corresponding to at least level three in Mitchell’s scale), as the octopus appears to be able to select the appropriate imitation according to the perceived threat and surrounding environment. Norman and colleagues write: Dynamic mimicry has the unique advantage that it can be employed facultatively, with the octopus adopting a form best suited to the perceived threat at any given time. Evidence for such sophisticated behaviour comes from our observation that on all occasions when sea-snake mimicry was observed it was exclusively a reaction to an attack by territorial damselfishes. Sea-snakes forage by entering burrows, and are predators of damselfishes (Norman et al. 2001: 1757–1758). However, when trying to use individual activity as a general criterion of mimicry, we encounter difficulties in describing the resemblance in camouflage or floral mimicry in terms of iconic sign creation. If the mimic as an individual is similar to the model just by way of genetic determination and does not participate in the creation of resemblance in any active manner, could this similarity be excluded from mimicry and from iconic sign activities? Should it not rather be categorized under natural similarities like, for example, resemblance between thorns and outgrowths of plants and furs of mammals, which do not have any communicative reason but are rather adaptations against forces of physical nature? The similarity of the stipules of Passiflora with Heliconius’ eggs is, however, related to the ecological linkage of those species as well as to communicative contact between them. As a possible way out of this problem, which has also been described in connection with other topics in phytosemiotics, Winfried Nöth has suggested that plant–animal co-evolution could with certain reservations be considered as semiosis of the evolutionary level, where the message is “taken from a large repertoire of morphological structures (colours, forms, etc.)” (Nöth 1990: 167). Jesper Hoffmeyer seems to support this view, arguing that similarities in mimicry may be both nondeceptive on the level of the individual (because the mimic can not send a truthful message) and deceptive on the evolutionary level. Using the resemblance between the Malayan praying mantis Hymenopus bicornis and the pink flowers of Melastoma polyanthum as an example, Hoffmeyer explains: 91 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. I suggest that we include among lies a special category to be termed evolutionary lies, i.e., lies rooted in the kind of intentionality exhibited by lineages in the process of evolving strategies for deceiving individuals from other species. [ . . . ] the single mantis doesn’t lie, but it is nevertheless an integral part of the lying lineage to which it belongs. Seen in the historical setting in which the adaptation took place the “resemblance” between mantis and flower was meant to be a (false) “representation” i.e., it was a lie. (Hoffmeyer 1995). We can conclude that whether or not mimicry is a phenomenon of iconic sign creation depends first on the particular examples of resemblance we observe in the natural world, and second, on our understanding of the mimic as an active party in communication. Instances where the mimic as an individual plays an important role in bringing deceptive resemblance into existence could be understood as examples of iconicity on the level of the organism and its neural activity. Other cases of mimicry could be seen as iconic, presuming that the interpretative and communicative activity is attributed to the species as a whole on the evolutionary level. We can consider mimicry to be a phylogenetic deception that does not need to include the specific cognitive activity of the sender. Here we encounter discussions of final causation or evolutionary intentionality, a topic much discussed in contemporary biosemiotics. 7.2 Resembling the environment and becoming a sign 21 A behaviourally active mimic has different possibilities for expressing its activity. The mimic’s activity can be targeted toward the receiver, or the mimic can obtain deceptive resemblance in the course of active behaviour targeted toward the model (cf. Maran 2008b). Dynamic mimicry of the mimetic octopus described above was a good example of the mimic’s activity directed to the receiver, whereas the mimic’s behaviour is often directed to the model in biological mimesis, cryptic mimesis and camouflage. In these types of deceptive resemblance, the model is the environmental background or its static element, and the relatively persistent contact between the mimic and the model favours the communication. In this subchapter, I will focus on 21 This subchapter is partially based on Maran, T. (2011). Becoming a sign: The mimic’s activity in biological mimicry. Biosemiotics (Springer), 4(2), 243–257. Used with permissions. 92 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. two narrower types of deceptive resemblance—facultative camouflage and frightening displays—with the aim to explicate the mimic’s active role from a semiotic viewpoint. In camouflage and in mimesis, the imitating organism resembles its environmental background or some static element by colour or form. Such resemblance can vary from monochromatic tones of soil or foliage to exact and complex patterns of tree barks, stones, leaves, blossoms, fungi, bird droppings, etc. If resemblance is facultative and not rigidly fixed, the mimic as an individual may have several possibilities to adjust its appearance. There are interesting new findings in evolutionary developmental biology that demonstrate an organism’s ability to select between different developmental paths in its early stages. Some species are able to comprise environmental parameters into their early developmental process in such a way that the course taken will lead to an appearance that is beneficial for the organism in the particular environment. Such adjustable phenotypic plasticity is described, for instance, in butterflies Bicyclus anynana (Satyridae, where placement and size of eyespots in imago is dependent on temperature where the pupae are developing, Brakefield et al. 1998), Helicoverpa armigera (Noctuidae, larvae colouration is dependent on the feeding plant, Yamasaki et al. 2009) and in ground-hoppers Tetrix subulata and Tetrix ceperoi (body colouration appears to be directly influenced by background colouration, Hochkirch et al. 2008). Fully grown individuals may also have several possibilities to regulate their relationship with the environment. There are descriptions of many different groups selecting a resting place or habitat based on the animal’s conformity with the background colours. Although in some cases, selection is induced by indirect mechanisms, e.g. preference for the feeding plant, quite often mimetic individuals actively select the environmental background. Specific correspondences between body coloration and background preferences are found for instance in treefrogs Hyla regilla (Wente and Phillips 2005), grasshoppers Tetrix undulata (Ahnesjö and Forsman 2006), crab spiders Thomisus spectabilis (Heiling et al. 2005) and in many moths (e.g. Catocala sp., Phigalia titea, Sargent 1966, 1969). Other species have adaptations that allow them to change body coloration to become similar to the colours of the surrounding environment. The most exemplary case of such temporal and reversible colour change are chameleons, but also many species of grab spiders, octopods, lizards and flatfishes are known to have the same ability. The third possible 93 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. way for an organism to hide in the environment is to use environmental objects to cover itself. In simpler cases, organisms (for example some rays and flatfish) can use particles of the soil to cover themselves or build a shell (caddis larvae Trichoptera). In these cases, resemblance with the surrounding environment is acquired by passive involvement, just by the random use of materials present in the environment. In other cases, however, organisms search for and use particular objects that would better serve their protection. Such adaptations can be quite complex and take into account ecological relations in specific biotopes as well. For instance, John J. Stachowicz and Mark E. Hay describe the adaptation of the decorator crab Libinia dubia living in waters off the eastern coast of North America that depends largely on the crab’s habitat. In the southern part of the habitat, where the number of natural enemies is higher, decorator crabs plant chemically noxious alga Dictyota menstrualis on their backs. In the northern part of the habitat, decorator crabs gather seaweeds without preferring any noxious species. This behavioural adaptation does not offer chemical protection to decorator crabs in the northern part of the habitat, but at the same time it allows them to blend better into the surrounding environment. Authors suggest that these adaptations may be related to the difference in ecological pressure by predation as well as in foraging strategies of the fishes (Stachowicz, Hay 2000: 65–67). A similar active selection of suitable clothing is described in hermit crab Pagurus bernhardus (Briffa et al. 2008). If the choice is given, crabs prefer gastropod shells for casing that match the colours of the surrounding environment better. However, an active choice is balanced by the threat of predation. When naked, hermit crabs enter the nearest suitable shell, even if it is in great contrast with the substrate, and when cues of predators are present, they remain in the present shell, even if it offers low crypticity. What can we conclude from such examples? By covering themselves with particles of soil, by changing colour, and by planting algae on their bodies, cryptic animals develop a strong connection with their specific perceptual environment or part of it. The creation of such correspondences are actively mediated by iconic signs. Based on this peculiarity, the French philosopher Roger Caillois has even described mimicry as a disorder of spatial perception, or as assimilation into space that is accompanied by a diminished sense of one’s own personality and vitality (Caillois 2003: 101). Using the conceptualisation of Jakob von Uexküll, the surrounding environment becomes both a significant aspect of the mimic’s Umwelt, and also an 94 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. important theme that organises its bodily organisation and life activities. Mimetic organisms are not just limited to themselves—they reach beyond their semiotic selves across their organismic borders (Sebeok 1991c). Through their activity, mimetic organisms obtain a representational character. Such organisms represent the surrounding environment, and it is not an overstatement to say that they develop a sign relation with that environment. The established iconic connection can later be perceived from the different perspectives: from the perspective of the mimetic organism itself, from the perspective of the possible predator or prey animal, or from the perspective of the human observer. Additionally, from these different perspectives, such an organism may become dislocated or inappropriate, when moved out of its natural environment. In fact, facultative camouflage and mimesis provide an advantage here, as they do not predetermine a specific semiotic relationship, but rather offer means and readiness to establish such relationship. The relationship can be recoded and re-established if necessary. What is remarkable in the cases of camouflage and mimesis is that the engagement with the environment is primarily semiotic and only secondarily physical. It is the meaning relation that organises the connection between the animal and the environment, whereas the specific physiological means by which the resemblance is achieved can vary from species to species (chitin cuticule, and skin with different structural and pigmental colors). Also, in cases of fixed mimesis, where the whole animal body is involved in the process of mimesis as it is in leaf insects Phylliidae, physical structures often seem to become subordinated to semiotic relations and meanings. Leaf insects have a flattened body shape with a scattered outline, and their coloration fits well with the vegetation as it includes different shades of green, beige, and brown (see figure 7.1). In many species, the spines that resemble plant veins are clearly visible. The precise mimesis is accompanied by jerky movements that resemble swaying leaves in a gust of wind. Fossil evidence shows that this strange appearance of leaf insects has remained almost the same for more than 45 million years, making this a good example of morphological stasis (Wedmann et al. 2007). With their curvy bodies slowly moving in the foliage, the leaf insects indeed become leaves. This is not merely a metaphoric expression, but rather pointing out that the general appearance of an animal together with its specific cues and signals is the basis for establishing the animal’s identity in the wider ecological community. If the animal’s appearance changes, then the autocommunicative, intra- and interspecific 95 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. communicative relations need to transform. For instance, leaf insects and stick insects are not dependent on sexual reproduction but instead rely mostly on parthenogenesis. Liliana Milani and colleagues comment on the biology of leaf insects and stick insects, stating that “Phasmids show noteworthy abilities to overcome species-specific reproductive isolation mechanisms, including hybridization, polyploidy, parthenogenesis, hybridogenesis and androgenesis.” (Milani et al. 2010: 258) The semiotic niche as the placement of an animal in all possible semiotic and communicative relationships prepares the ground for and influences its ecological niche. [enter figure 7.1 here] Figure 7.1. Mimesis of the leaf insect Phyllium siccifolium (From the collections of Tallinn Zoological Gardens, photo courtesy of the author). In many cases of mimicry, we also witness a discrepancy between semiotic and ecological niches. Kleisner and Markoš have used the concept of seme to express a similar thought. According to their view, a seme is every particular morphology, shape, colour pattern, odour or behaviour according to which the species is immediately recognized. Users of alien semes acquire the image or behaviour of the species that is the subject of imitation (Kleisner and Markoš 2005: 218; cf. Kleisner and Markoš 2009). At the same time, mimetic species need to find ways to overcome difficulties in maintaining their own identity and intraspecific communicational systems, often utilising different communication channels or even the object they imitate for making contact with members of their own species. A most vivid example of such kind is provided by Robert Payne in his studies on indigobirds Vidua that are the brood parasites of the weaver finches Estrildidae (Payne 1977: 12; Payne et al. 2000). As nestlings, indigobirds learn songs from their foster parents and use this later to invite their own mates. Also, female indigobirds have a preference for the song that they heard in their youth. The imitated song thus becomes the uniting feature for indigobirds reared by the same host species, making the later communication possible and ensuring the continuation of the lineage that prefers that particular host species. In the case of indigobirds, the communicational code borrowed by another species becomes the basis of intraspecific identity, and the birds have given up their original means of expression. Furthermore, being based on imprinting, this adaptation is 96 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. flexible. Parasitic indigobirds can switch to another host species, as it has been shown by experiments of Payne and colleagues (2000) when they raised indigobirds in different hosts and demonstrated that their song preference changed accordingly. This extraordinary communication system illustrates the paradoxical nature of a sign. A sign denotes something other than itself, and by doing so, it simultaneously conceals the identity of the sign carrier. When standing for something else, one cannot be just itself anymore. When analysing the position and relations of the mimetic messages to other features of the mimic, we may notice something peculiar. Sometimes the mimetic messages are in stark contrast with the overall organisation of the animal’s body and behaviour. Let us observe here a deimatic behaviour where the activity of the mimic is directed to the receiver. A natural reaction for an animal under attack would be to escape, and the majority of animals do exactly that. However, a common toad Bufo bufo encountered by a grass snake Natrix natrix stays in place, lifts itself up from the ground and emits strange growling sounds while its body becomes bloated from the inhaled air. There is nothing comparable to this threat display in the whole behavioural repertoire of the toad. The toad remains in a trance-like state and performs his display with great vigorousness and energy. The contradictory nature of this situation is obvious. It seems that in some way, the motive of fear in the snake’s Umwelt has entered into the toad’s subjective world and become manifested in its behaviour. Uexküll has described a similar transmission of meanings in the relations between a spider web and a fly: The spider’s web is certainly formed in a ‘fly-like’ manner, because the spider itself is ‘fly-like’. To be ‘fly-like’ means that the body structure of the spider has taken on certain of the fly’s characteristics—not from the specific fly, but rather from the fly’s archetype. To express it more accurately, the spider’s ‘fly-likeness’ comes about its body structure has adopted certain themes from the fly’s melody. (Uexküll 1982: 66). The same kind of adoption of a theme of meaning is present in the toad’s relation with the snake. It appears as if the toad would need to suppress its own identity and urge to flee in order to obtain a new external sign faculty—a sign of threat for the snake. For 97 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. a short time, the toad is not a toad; in its performance, it becomes a much larger and dangerous creature that is capable of scaring the snake away. 7.3 The receiver’s perspective and ambivalent signs 22 An alternative possibility to analyse mimicry as a semiotic phenomenon is to focus on the position of receiver. To the receiver, the mimicry situation may appear very differently from how it appears to the sender. This change is first rooted in a common feature of communication: the emergence of shifts in meanings due to the asymmetry of the processes of formulating and interpreting, coding and decoding. Theatre semiotician Tadeusz Kowzan has described this as different aspects of the sign, which are expressed in the different phases of communication. For example, a sign can be mimetic in its creation and iconic in its interpretation (Kowzan 1992: 71, see also Maran 2003: 205–211). In mimicry, however, the difference of meaning for the sender and the receiver seems to be a more fundamental property. Alexei A. Sharov has explicated mimicry with the term ‘inverse sign,’ where sign has a positive value for the sender (‘transmitter’ in his terminology), but negative for the receiver. Sharov describes female fireflies, which imitate light signals of other species to attract their males in order to eat them as an example of such inverse signs. Sharov specifies that “an inverse sign is always an imitation of some other sign with positive value for the receiver” (Sharov 1992: 365). Similar to many other cases of animal semiosis, the sign relation in mimicry is formed for the receiver from the perceived features or messages of an organism (representamen), the organism as it is physically capable of being interacted with (object), and the meaning connected with the applicability of the organism (interpretant). Connecting applicability with interpretant derives from the semiotics of Charles Morris (1985: 179), which in my understanding is one prospective way to use Peirce’s sign model in animal communication. As the mimic and the model have different potential value or applicability for the receiver, it is in the receiver’s interest to distinguish between them. In general terms, the receiver should recognize organisms and objects correctly based on the perceived cues and messages, and apply appropriate behaviour toward them, while avoiding using the same behaviour toward 22 This subchapter is partially based on Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica (Degruyter Mouton), 167(1/4), 223–248. Used with permissions. 98 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. deceptively similar organisms or objects. The difference between the model and the mimic for the signal receiver may be manifested in the following oppositions: discernible object versus perceptual noise, eatable versus uneatable item, safe versus dangerous organism, cues referring to a general meaning complex versus cues that lack this reference. The existence of objects that are perceptibly similar but have different applicability in the Umwelt of the receiver goes together with different and often diametrically opposite aspirations to react (e.g., catch versus flee). Thus, a common denominator of mimicry seems to be the signal receiver’s effort to make the correct recognition in a situation where perceptibly similar cues, messages, objects or organisms are present. The exact method of how perceptible features are perceived, selected and categorized largely depends on the perceptual capabilities of the receiver, as well as on the specific architecture of the mimicry system. In cognitive psychology, major alternatives are often distinguished as prototype categorization where the receiver compares the perceived object with the generalized mental image, and boundary classification where distinctive features of two classes of objects are focused on. Frederik Stjernfelt argues that boundary categorization “should be expected where there is a crucial behavioral difference in relation to the two categories on each side of the border […] (similar berries, the one being edible, the other poisonous),” and prototype categorical perception should be expected “where the boundaries are fuzzy or simply do not exist (what are the boundaries of phenomena like “danger” […])” (Stjernfelt 2001: 95). In mimicry, both types of attentiveness may actualize depending on the specific mimicry type, perceptional accessibility of the mimics and models, their relative number, consequences of a possible mistake for the receiver, and other factors. For instance, in a case such as the eyed hawkmoth’s warning display, we can expect two equally strong stimuli to be present for a small insectivorous bird. The perceptual cues of the moth correspond to the search image of a suitable prey, whereas false eyes indicate the presence of a possible predator and danger. In flower mimicry, there is a different situation. Studies on bees’ flower constancy indicate the existence of one prevalent search image, which corresponds to the most profitable flower in terms of nectar and pollen in the surrounding environment. If the abundance of the preferred flower decreases, bees begin to ‘make mistakes’ and visit other species, which occasionally leads to the formation of a new search image (e.g., Goulson 2000). For example, in the mimicry system of bellflowers Campanula 99 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. (model) and orchid red helleborine Cephalanthera rubra (mimic), we can expect one prevalent search image to exist for solitary bees (receiver) who use it to search for proper food plants and filter out mimicking orchids as mismatching (Nilsson 1983). Yet another type of situation is present in camouflage colouring or crypsis, where the mimic tries to hide itself in the surrounding environment and/or in the communication noise in the receiver’s Umwelt. We can conclude that the exact combination of signs involved in a mimicry situation varies from case to case. In addition to differences caused by the type of mimicry, mimicry as a sign system and properties of participating species, the physiological status (e.g., hunger versus comfort) of the receiver may also become decisive. Furthermore, he signs perceivable to the receiver may depend on the specific communicational context and vary according to the location where the mimicry situation takes place (for example, whether communication between an insectivorous bird and butterflies takes place on a tree trunk or while flying in the air). In addition to recognition, the active interpretation of the mimic’s messages by the receiver seems to form an important aspect in some mimicry systems. In some cases, signs can be combined to produce new meanings. Keeping that variability in mind, we can, describe biological mimicry as a sign situation for the receiver. Kalevi Kull has studied semiotic aspects of recognition and suggested that the mimic and the model constitute the same sign in mimicry situation. “The organism for whom a certain wasp functions as a sign cannot differentiate between the wasp and its mimic fly. The fly presents the same sign as the wasp, because their patterns are indistinguishable to the interpreter” (Kull 1992: 228). This description corresponds well with such mimicry cases where erroneous recognition is highly prevalent, as it is for example in molecular mimicry where an immune reaction toward the mimic is based on the compatibility of physical structures. In mimicry between organisms, however, some neural activity by the receiver is involved and recognition is therefore probable. An insectivorous bird like the flycatcher may mistake a fly for a wasp and leave it untouched as disgusting, but a different course of events may also take place. The bird may recognize the fly correctly as an eatable object and catch it. These two scenarios should give us reason to distinguish between two perceivably similar but still different signs or sign complexes in the Umwelt of an insectivorous bird—one standing for food and the other for the unpleasant experience of getting stung. 100 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. If the differentiation of mimics from models depends on many contextual factors and therefore reappears in each and every act of communication, then it is not possible to conclude whether there are one or two sign complexes involved in mimicry. Floyd Merrell, describing border situations in semiotic systems, argues that demarcation between sameness and difference is essentially indefinable and ambiguous (Merrell 1999: 460–461). If such an uncertainty is not occasional but rather common to most mimicry situations, then it should be carefully considered whether it forms a principal feature of mimicry and thus also a new type of sign structure. The probability of recognition also seems to be connected with the evolutionary dynamics of mimicry by being a mechanism that allows mimicry to be accommodated in different biological and communicational contexts. In our effort to deal with ambiguous sign complexes, the American semiotician Charles Morris can offer us some guidance. In addition to advancing a theory of semiotics towards psychology and behaviourism, Morris elaborated the complex typology of signs and sign processes. In “Signs, Language, and Behavior” Morris introduces the term ‘sign family,’ defining it as a group of signs which have the same meaning for the interpreter: “A set of similar sign-vehicles which for a given interpreter have the same significata will be called a sign-family” (Morris 1971b: 96). In accordance with his behaviourist stand, Morris unites signs into a sign family on the basis of a similar behavioural reaction released by the interpreter (Morris 1971b: 97). Concerning mimicry, Morris’s concept of sign family can be used to describe mimetic phenomena where similar marks, forms and behaviours are carried out by different organisms. Such a sign family is present, for example, in Müllerian mimicry, where many inedible or dangerous organisms of different descent carry similar colours. However, if the signal receiver’s reactions toward mimics and models are at least partly different, as they are in classical Batesian mimicry, then the presence of several sign families should be expected. Representamens of these different sign families can be similar and even partly overlapping. In connection with the concept of sign family, Charles Morris also points out that a sign may, but not necessarily need to have only one meaning. He contrasts unambiguous and ambiguous signs: “A sign-vehicle is unambiguous when it has only one significatum (that is, belongs to only one sign-family); otherwise ambiguous” (Morris 1971b: 97). The concept of ambiguous signs seems to cover different types of relations between meanings (interpretants). First, there can be situations where 101 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. meanings complement each other (for example, in many polysemic structures of human language). In such cases, meanings can be combined to form more complex interpretation patterns. In nature, for example, river as a biological universal may signify ‘drinking water’ and ‘obstacle in the way’, which may combine and produce new meanings such as demarcation of a living place. Another possibility actualizes when different interpretations or meanings are in opposition and exclude each other because of, for example, contrasting behavioural outcomes. Such situations can also be observed in homonymy, where the same sequence of letters refers to different meanings and the reader needs to make a choice between possible interpretations based on the context. For example, English ‘lie’ can be understood as “rest or place oneself in a reclining or horizontal posture” (from Old English Licgan) or “say or write something with the deliberate intention of deceiving” (from Old English Lēogan) but not both at the same time (Kirkpatrick et al. 1995: 784–785). The second type of ambiguousness is more characteristic of mimicry, at least in cases where the mimic and model are definable as specific species. Therefore it would be more correct to call such a sign combination an ambivalent sign instead of ambiguous sign. An ambivalent sign can be described as a sign structure that fluctuates between one and two signs and whose actual composition and number of signs emerges in the course of interpretation. Different from homonymy where concurrence is largely coincidental, ambivalence in mimicry has structural importance. The perceptual similarity of mimics and models and the opposition in meanings are components of the evolutionary conflict between the mimic and the receiver, and an important feature of the communicative regulation between them. 23 To sum up the receiver’s role in mimicry, it seems that the shifting balance between correct and false interpretations is a characteristic feature of mimicry systems. The dilemma of recognition for the receiver may actualize in perception or in various types of categorization and become manifested on the different levels of consciousness. Supported by communicative feedback and evolutionary regulation, the communicative confusion can evolve through various natural and communicational contexts. The recurring perplexity of situation for the receiver 23 In most cases, representamens in mimicry are exclusive to each-other, but there are also examples where it is possible to combine different interpretants for using the iconic reference with a new communicative function. Such processes will be discussed in more detail in the Chap. 9.2 “Mimicry and semiotic scaffolding”. 102 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. allows us to distinguish a new type of sign formation—ambivalent sign, which is stable in fluctuating between one and two signs. 103 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 8 MODELLING MIMICRY In these pages, I have tried to suggest a view of mimicry that would be systemic and take into account the complexity of mimicry, while at the same time sensitive to the peculiarities of different mimicry cases. Next, I will consider possibilities for developing semiotically inspired research methods that could be applied in the practical analysis of mimicry. In order to do this, semiotic modelling should have a central importance. Semiotic modelling could especially be helpful in a systematic description of mimicry and in formulating research questions for mimicry studies. The concept of modelling is widely used in semiotics, especially in the semiotics of the Tartu–Moscow school (for an overview, see Grzybek 1994; Tondl 2000). Modelling can be defined here as a cognitive process in which a certain phenomenon is presented so that it can be understood, using (conceptual or material) representations which are at least partially based on analogical relations (cf. Lotman 1967: 130–131). An important characteristic of modelling is that a model does not represent the entire set of the properties of its object, but only highlights certain aspects of that object. The question of the type of relationship between the model and the object is meaningful and significant in itself, since the relationship may be specific to a particular species, Umwelt, culture, text, paradigm, etc. The Czech semiotician Ladislav Tondl writes, “[t]he model represents a homomorphic representation, i.e., it is not identical to the original. It means the representation in the sense of the Latin “pars pro toto”, the part instead of the whole” (Tondl 2000: 83). Thus, models are based on analogy and exhibit wide variance in terms of complexity—from the categorising perception of animals and their ability to create associations to humans’ mathematical models and anthropomorphic descriptions of nature. In Tartu–Moscow semiotics, the variety of sign systems used by humans are described as modelling systems, while the Hungarian-American semiotician Thomas A. Sebeok, in contrast, has linked modelling to the Umwelt concept. Sebeok sees the capacity to model structures or phenomena as a quality shared by all animals and distinguishes, in the case of humans, between the levels of zoosemiotic and linguistic modelling (Sebeok 1991b). Language-based modelling is a species-specific feature of humans which in turn gives rise to more complex poetic, ideological, religious, scientific and other models that are described as secondary modelling systems in the semiotics of the Tartu–Moscow school (Lotman 1967: 131). Consequently, modelling 104 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. can be understood in semiotics both as an object level and metalevel process, that is, modelling is both the nature of the semiotic inquiry as well as an important property of the objects being studied. Using semiotic modelling as a tool for analysis in the context of present study would mean establishing a well-forged analogical representation of the object, where the bases or grounds of modelling are acknowledged and actively shaped. Semiotic modelling aims to bring together a number of different viewpoints in a complimentary fashion that would result in a thick yet systemic perception of the object. The critical and systemic attention to the bases of modelling allows the researcher to perceive his/her own position in regard to the object. In the following subchapters, I will first present a semiotic research method of mimicry that consists of five levels or points of description. Thereafter I apply this method for describing a specific mimicry case— the brood parasitism of a common cuckoo—and in the final subchapter, I will make some suggestions for using this method for a comparative study of different mimicry cases. 8.1 Toolbox for modelling mimicry 24 A semiotics-based methodology that I propose consists of a group of various methods to be used in a toolbox-like manner for approaching different types of mimetic resemblances. In building this approach, I have integrated aspects of classical mimicry theory, Jakob von Uexküll’s Umwelt theory, and semiotic and communication analysis into a model of a five-level analysis (the theoretical bases of this toolbox were articulated in the previous chapters). The first level of analysis departs from the classical tripartite Wicklerian model consisting of a mimic, a model and a receiver. Rather than trying to match this to every mimicry system, I would extend it to a general principle and ask what kind of configurations mimicry could take. Attention here is not on the participation of the biological species in the mimicry system but on the semiotic constitution of mimicry: what resembles what to whom in what respect? For instance, mimetic resemblances in monkfish could be expressed as a compound model that includes one mimic and one 24 This subchapter is partially based on Maran, T. (2010). Semiotic modeling of mimicry with reference to brood parasitism. Sign Systems Studies (University of Tartu Press), 38(1/4), 349–377. Used with permissions. 105 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. receiver, but two models: the background of the environment and the movement of a worm-like prey of the fish (see figure 8.1 left). In a similar way, mimics and receivers could also be manifold. The mimetic resemblance of cuckoo bumblebees (subgenus Psithyrus) could in most cases be described as a compound model that includes one mimic and one model (a particular bumblebee species), but two receivers: bumblebee host species (being simultaneously the model and the receiver) and insectivorous birds (see figure 8.1 right). On this level of analysis, it is also possible to distinguish in general terms between mimicry and other types of resemblances. If it is possible to point out all three parties—a mimic, a model and a receiver, given that the receiver is not a human observer—then the resemblance can be regarded a mimicry system. If this condition is not met, then some other type of resemblance, such as similarity due to an evolutionary affinity or convergence, or some resemblance induced by human cultural description, should be suspected. [Enter figure 8.1 here] Figure 8.1. Schematic representation of complex mimicry systems. The unbroken arrow line represents the communicative relation between the model (MO) and the receiver (R) that is exploited by the mimicry system. The dashed arrow line represents the deceptive relation between the mimic (MI) and the model. The dotted line represents the relation of resemblance between the mimic and the model. Left monkfish with two distinct models MO1,2 (a worm and the sea bottom). Right cuckoo bumblebee with two distinct receivers R1,2 (a bumblebee host and an insectivorous bird, a host R1 being at the same time also a model MO). The second level of analysis proceeds from the Umwelt theory of Jakob von Uexküll, and focuses on the corresponding body structures of the mimic, the model and the receiver, or the properties of the original and the imitated signal, as well as the communication channels and behavioural regulation used (Merkwelt and Wirkwelt in Uexküllian terms). On this level of analysis, a more specific description of resemblance can be achieved. The second level of analysis enters the field of qualitative phenomena by describing the specific percepts and activities that are employed in mimicry. Umwelt analysis sensu Uexküll (1982: 52–57) can bring out a correspondence or non-correspondence between the perceptual capabilities of the 106 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. animal receiver and the body structures of the mimic and model. This enables reasoning if the mimic and model are deceptively similar for the receiver, or if they are even perceptible at all. Paying attention to the Wirkwelt and the behaviour of the receiver regarding the mimic and the model allows us to make conclusions about the relevance of the resemblance for the receiver. This is a significant question, as in many cases, the deceptive resemblance between two objects does exist for the receiver, but it is irrelevant just as it is irrelevant for an average human to distinguish between bumblebees and bee hawk-moths that mimic them. Through such argumentation, mimicry becomes grounded in ethological functionality and not in the evolutionary functionality that would be much more difficult to find explicit proof of (see Chap. 9.2 “Mimicry and semiotic evolution” for future discussion). The third level of analysis proceeds explicitly from semiotics as the study of signs and sign systems and focuses on meaningful units that resemble each other in mimicry. This level of analysis addresses the general conditions that make it possible for a confusion to emerge. A starting point for this discussion is an understanding that resemblance in mimicry is not univalent, but that there are different possibilities for resemblance to occur. First, the resemblance can be described as a matter of degree ranging from non-resemblance to absolute similarity. Concepts used in psychological studies of categorical perception, such as boundary perception, common characteristics resemblance, prototype resemblance and others can also be employed in distinguishing between various possibilities for a resemblance to emerge. Mimetic effect can be based on different ways in which the model-receiver and mimic-receiver sign systems interact with one-another. Mimicry can also be described as fixed or adjustable, partial or complete, local or general, individual or collective, embodied or detached, etc. One possibility to distinguish between different mimicry types would follow the categories of firstness, secondness and thirdness as brought out by Peirce’s semiotics (as was discussed in Chap. 5.3). The simplest type of resemblance can be described as a relation between one and many and is common in camouflage, for instance when a moth is lying on tree bark. Here the perception of the moth emerges from nowhere and that allows us to relate camouflage to Peircean firstness. The second type of resemblance is present when we are dealing with deceptive similarity between representatives of two species, where one is eatable and the other poisonous, as is the case in typical Batesian mimicry. This type of mimicry requires comparison, 107 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. or a reference to the second, and therefore seems to relate with Peircean secondness. The third type of mimicry is present, for instance, in the case of eye-marks on the bodies of insects, fish and amphibians, or in the case of colourful patches on the bodies of many lizards and insects. These patches are kept hidden in the normal body position but flashed during escape. It seems that the unifying aspect of such mimicry systems is a common meaning: thus, for instance big eyes, unexpected movement or warning coloration all signify something dangerous for the receiver. Such abstract mimicry requires that the meaning of the message is understood and therefore seems to relate to thirdness. The fourth level of analysis proceeds from a communication analysis of the mimicry system by focusing on the communicative functions and the position of feedback cycles. Communicative functions of mimicry can be mapped according to the distinction of Roman Jakobson between emotive, conative, poetic, referential, metalingual and phatic functions (see Chap. 4.2. “Mimicry as a communicative interaction” for discussion). Distinguishing different communicative functions would allow for bringing forth the dominant of the interaction, that is, the specific aspect in communication that the mimic makes use of in its deceptive signalling. In regard to the feedback in ecological relations, it is possible to distinguish between communicative, ontogenetic and evolutionary levels of feedback regulation. In communicative feedback, resemblance is regulated within a single act of communication. This is a prevalent regulation mechanism in behavioural and adjustable mimicries. Through ontogenetic feedback, an individual’s personal experiences are formed or expressed. In phylogenetic or evolutionary feedback, genotypic adaptations related to this particular communication act develop or manifest. Feedback cycles enable dynamics to enter the act of communication and also enables the sender to change its behavioural and communicative activity with respect to the receiver’s activity. On the metalevel, feedback in communication allows messages to be adjusted and changed, which in the long run enables the development of sign systems. Distinguishing between different types of resemblance regulation allows us to include among mimicry such resemblances that are induced by epigenetic memory, e.g. by animal cultures where the dependence on genetic causation cannot be proved. This level of analysis helps to focus on dynamic, reversible and individual aspects of mimicry resemblance, such as individual song imitations by many birds, for instance the European starling Sturnus vulgaris; and to connect these with mimicry 108 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. studies. Although cognitive mimicry such as imitational bird songs is rather different from mimicry systems with a genetic dominant, there is also a lot of analogy between the two. From a semiotic viewpoint, both bring along the spreading of messages from the repertoire of one species to another, and in both, the dynamics between different sign systems is present. The fifth level of analysis proceeds from the cultural level and focuses on the observer’s perceptual involvement in the mimicry system as well as on the cultural and scientific status of the phenomenon. This level of analysis deals with the human observer in relation to the mimicry system (see also Chap. 4.4. “Umwelten of the receiver and the human observer”). The starting point for this discussion is the understanding that Umwelten of the human observer and of other living beings participating in a mimicry system are likely to be different. Besides deceptive similarities perceptible to the human observer, there may also occur situations when the messages of the model and the mimic are situated with respect to the Umwelt of the human observer in such a manner that from the latter’s viewpoint, they do not seem similar (e.g. the red helleborine Cephalanthera rubra and the bellflower Campanula sp.). Likewise, a whole communication system may be left concealed from the human observer. As Czech biologist Thomas Grim has noted: “similarity for the human eye does not per se support the hypothesis of mimicry. In turn, the absence of similarity for the human eye does not reject the hypothesis of mimicry: mimicry may be cryptically ‘hidden’ in UV-part of the spectrum.” (Grim 2013: 276). The location of a deceptive similarity in relation to the Umwelt of a human observer may bring about a biased choice of the object to be studied and may lead a nature scholar to under- or over-interpret some mimicry cases. Besides the relations between an observer’s perceptual organs and a mimicry resemblance, the cultural-historic component of mimicry also needs attention. There are mimicry systems that have a long history of studies due to the peculiarities of the history of science. For example, mimicry in butterflies is a well-established field partly because of the activity of the nineteenth-century naturalists in collecting insects. A collector’s main interest is to identify items correctly. This desire of the naturalists helped to advance systematics, but at the same time drew attention to the confusions and ambiguities in nature, including mimicry. In addition, descriptions of mimicry resemblances may also include cultural, religious or mythological layers. There are various ways that mimicry resemblances can be actualised in culture. For 109 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. instance, mimicry resemblances can be interpreted as signs of nature with special meaning (so called signatures) that have been believed to have their origins in a supernatural or divine source. The cultural layer, which clearly remains out of the biologists’ scope, can still be of interest in a semiotic approach that interprets resemblances in nature and human culture not as distinct fields, but as a continuous complex phenomenon. 8.2 Applying semiotic modelling to brood parasitism As a practical example of the semiotic analysis of mimicry, I am going to use the methodological tools introduced above to describe brood parasitism as one specific case of deceptive resemblance in nature. More specifically, I will take under observation the semiotic and ecological relations between the old world cuckoos, especially the common cuckoo Cuculus canorus, and its frequent host species, the Passerine birds. This example has been thoroughly studied by contemporary evolutionary biology and is therefore suitable for testing the methodology. I will focus primarily on the resemblance of eggs, though I will shortly discuss other resemblances related to brood parasitism as well. The analysis is based on various sources of biological literature, but my argumentation and point of view remain semiotic. For the interpreting subjects—birds—the primary question in regard to brood parasitism is related to recognizing eggs. The image of the egg as such can be considered to be a ‘biological universal’ that is meaningful to various species. Most animal taxa have some evolutionary experience with eggs, which have been around on our planet for about 500 million years if we start counting from chordates, and even more if we include insects. When we look for the meaning of eggs, the most immediate answer would be ‘reproduction’—eggs stand for the offspring. This goes along with the observation that there are no abstract eggs, eggs always belong to somebody, and in many species eggs are considered worth keeping and defending. On the other hand, many predatory and omnivorous mammals, birds and reptiles view eggs as a possible food source, and some species such as egg-eating snakes (Elachistodon, Dasypeltis) specialize in them. Additionally, egg shells can be used as an indexical sign for detecting the presence of a bird nest, and in order to avoid being 110 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. revealed, some bird species (e.g. the common starling Sturnus vulgaris) carry shell pieces away from their nests. For brood parasites, eggs signify an opportunity to reproduce: cuckoos and other brood parasites tend to lay eggs in nests where some clutch is already present, as empty nests can easily be abandoned by hosts. Brood parasitism is a very widespread biological phenomenon. It is estimated that approximately 1% of all bird species use some sort of brood parasitism, including nearly half of the 130 species of cuckoos, some cowbirds, indigobirds, whydahs, two genera of finches and some ducks (Payne 1977: 1). Most brood parasites are specialized in a specific host species or set of these. The common cuckoo has more than 100 potential hosts, the most usual of these including reed warblers Acrocephalus, leaf warblers Phylloscopus and warblers Sylvia, robin Erithacus, redstarts Phoenicurus, and wagtails Motacilla (Honza et al. 2001: 344) (see Figure 8.2). In some populations of host species, brood parasitism may affect up to 65% of all nests (Moskát and Honza 2002). [Enter figure 8.2 here] Figure 8.2. An egg of a common cuckoo in the clutch of a common redstart Phoenicurus phoenicurus (collection of the Natural History Museum of the University of Tartu, photo courtesy of the author). If using the tripartite model of mimicry in the studies of brood parasitism, some difficulties seem to occur. Case-studies in brood parasitism encounter the ecology and biology of parasites and their hosts as a whole set of adaptations that include many types of similarities with different causes. In practical studies, a question arises — should all similarities be included under the concept of mimicry or should they rather be differentiated. A Czech animal ecologist Tomáš Grim (2005: 70–72) notes that the similarity between the eggs of a cuckoo and its hosts can have several sources: besides mimetic evolution also phylogenetic closeness, or environmental similarities that could influence the coloration of both host and parasite eggs in the same way, or predation pressure by similar visually orientated predators. Even competing female cuckoos can become an evolutionary factor influencing the appearance of eggs when they remove from the host’s nest an 111 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. egg that is least similar to the rest of the clutch — which can be an egg laid by a previous cuckoo (ibid., 71–72). A second argument supporting the claim that egg mimicry cannot be easily separated from the overall ecology of the species is the dependence of the accuracy of recognizing one’s own eggs and imitations on the living habits of the host species. The accuracy is lower in the case of hole-nesting species (e.g. redstarts Phoenicurus phoenicurus) than it is in the case of open-nesting species. This can be explained by lower ecological pressure of brood parasites in regard to hole-nesting species: it is difficult for an adult cuckoo to lay eggs into holes, and it is also difficult for a host parent to throw a cuckoo egg out of the nest, as well as for a cuckoo chick to throw out host eggs (Avilés et al. 2005: 609). It seems that the complexity of brood parasitism exceeds far beyond the simple triadic structure of the mimicry system and issue of similarity and recognition between the brood parasite and host species. In the following discussion, I am going to take the egg mimicry of cuckoos as a test case for the abovementioned research methodology, which can be summarized in the form of five questions: 1) What is the formal structure of the mimicry system? 2) What are the perceptual and effectual correspondences between the participants of mimicry? 3) What are the characteristics of resemblances? 4) How is the mimicry system regulated in ontogenetic and evolutionary processes? 5) How is the mimicry system related to human cultural processes? 1. What is the formal structure of the mimicry system? Compared to the ecological variety of brood parasitism, its descriptions in mimicry studies remain rather sketchy and superficial. In general overviews and typologies of mimicry, attention is usually focused on resemblance and deception, species combination and cost-benefit relations of the participants. Brood parasitism belongs to bilateral mimicry systems, meaning that two biological species are involved. French zoologist Georges Pasteur (1982: 188) describes the cuckoo’s egg mimicry under Kirbyan mimicry (following W. Kirby who noticed this phenomenon in syrphid flies who parasite in the nests of bumblebees). Pasteur writes that this kind of mimicry belongs to the aggressive/reproductive type in which the model and the receiver belong to one species and the mimic belongs to another species. In his thorough mimicry typology, a British entomologist Richard Vane-Wright (1976: 42) describes the common cuckoo 112 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. as belonging to the anthergic aggressive S1+R-type mimicry. In simpler terms, this means that in addition to Pasteur’s criteria of two species where the model (the host’s eggs) and the receiver (the host adult) belong to the same species, Vane-Wright also takes evolutionary influences and interests into account. The mimic’s (cuckoo’s) influence on the receiver is negative and the interests of the mimic and the model do not coincide. From a biological perspective, in most cases of egg mimicry, two participants (with evolutionary memory) are indeed involved in performing the three roles in the mimicry system. At the same time, the mimicry system can involve an open set of host species as receivers and their eggs as models. There are several host-specific egg patterns in the common cuckoo and specific resemblances are genetically determined and transmitted in the maternal lineage. Researchers have described 11 (Honza et al. 2001: 344), 16 in some estimations (Avilés, Møller 2004: 57), of such lineages or socalled “gents”. Eggs with specific patterns are, however, not always laid in the nests of the corresponding hosts, so the number of occasional host species may be much larger. This introduces a restrictive condition: if we take the common cuckoo as a species to be the starting point of the description, we should consider egg mimicry as an open system, where the species of neither models nor receivers (nor mimics on the level of gents) are determined. If we preferred the strict tripartite model of mimicry, the level of description should follow the relations of one specific lineage to its fixed host species. When we make a distinction between the biological and the semiotic layers in the egg mimicry of the common cuckoo, several resemblances become observable. The very resemblance between the eggs of the brood parasite and those of the host species (secondary iconic resemblance) seems to be located between two additional resemblances: the primary and tertiary iconic resemblance. The primary ground of a deceptive resemblance between the eggs of cuckoos and hosts is a phylogenetic resemblance of the egg shapes of birds of corresponding size (this also relates to corresponding behavioural adaptations, such as the readiness of a species that does not have any evolutionary experience with brood parasitism to treat objects of various shapes in their nests as their eggs, as shown by studies in classical ethology (Tinbergen 1951: 45)). The tertiary level of iconic resemblances is the relation of camouflage coloration between egg coloration and the surrounding environment to avoid predation. This may influence the coloration of both host and parasite eggs. In 113 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. addition, there exist resemblances at several other developmental stages, such as correspondences between the behaviour of the cuckoo chicks and the chicks of a particular host species (e.g. Kilner et al. 1999), or similarities in the appearance of the adult cuckoo and hawk species (Paine 1977: 8; Davis and Welbergen 2008). It becomes clear that on the level of semiotic relations, many additional resemblances and connections can be examined. 2) What are the perceptual and effectual correspondences between the participants of mimicry? Although from a biological viewpoint, it may be correct to include the model and the receiver in the same category as in the classification above, from the semiotic and perceptual viewpoints it seems simplifying. Parents of the host species do not need to differentiate between themselves and the other, instead they need to recognize distinct objects—eggs—and decide which eggs are theirs and which are not. We can consider this type of mimicry to be a detached mimicry, where the imitating and imitated objects are distinct from the bodies of the participating animals. Here the second research question, which focuses on specific matching and functional cycles between the species, can provide a more elaborate picture. The main communication channel used to determine the identity of the eggs is mostly the visual channel, which may also include patterns that remain outside the human visual sphere in the UV range. Some studies show that tactile perception of an egg can be relevant as well, as birds appear to differentiate eggs based on their material and touch sensations (Kemal and Rothstein 1988). Besides perceptual sensations, the indexical place-specificity is an important criteria—many species regard all egg-like things in their nest as their eggs, but if an egg falls out of the nest, the birds will not treat it as an egg anymore. The spatial criteria appears to be taken as self-evident in many biological studies, but from a semiotic viewpoint it certainly requires attention. It may be that the spatial arrangement of the eggs have an effect on their interpretation. In some cases birds, are also believed to estimate the number of their eggs and use this estimation as a basis for distinction (Lyon 2003). The difference in communication channels and criteria used to determine the origin of the eggs demonstrates that egg mimicry cannot be considered in terms of a simple resemblance or similarity, but with respect to complicated processes used by birds for making the distinction. The reaction of the parents of the host species towards a suspected false egg can provide information about the importance of deceptive resemblance for the birds. The decision concerning the authenticity of eggs often leads to a harsh behavioural 114 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. reaction: depending on the species, this may involve removing a false egg from the nest, punctuating it or abandoning the nest (Moksnes et al. 1991; Soler et al. 2002). In relation to this, it is also remarkable that birds abandon their nests much more easily if they have eggs, but much more seldom once the chicks have been hatched. A classical evolutionary explanation emphasizes the differences by using an abstract measure of fitness: in the second case, the bird cannot manage to have a second brood and this decreases its fitness significantly. From the viewpoint of Uexküllian semiotics, the difference in number and intensity of possible modalities used to communicate with eggs or chicks can itself can provide an answer. With regard to eggs, parents can use the visual and tactile channels for communication. With regard to chicks, the use of the visual and tactile channels becomes much more abundant, the auditory channel becomes operational, and there appears to be dynamic dialogical communication, thus there are more possibilities for the relationship to develop. Using different communication channels and modes would introduce greater redundancy of communication, which would probably improve the phatic function of communication. That would make breaking off the communicative contact and abandoning the nest much more difficult for the parents. 3) What are the characteristics of resemblances? When we focus on specific characteristics of the resemblance between the eggs, it can be considered as a prototype resemblance or common characteristic resemblance; that is, birds are making the distinction based on some specific characteristics or the generalization of them in the model. In the three-fold categorization given above between the firstnessbased, secondness-based and thirdness-based resemblance, the egg mimicry will be close to the second option. It is a Batesian-type of mimicry, in which the resemblance requires comparison and actual reference to a second. In the case of brood parasitism, we have a rare case in nature where the objects being compared are actually physically together, lying side by side in the nest. Another peculiarity of the eggmimicry is the completeness of mimicry (as opposed to partial mimicry), as the resemblance covers most if not all aspects of a complete singular object: shape, size, weight, colour pattern, surface structure, etc. In most cases of mimicry, the scope of resemblance is much more narrow, for instance being limited to the appearance of insects’ cuticle or the shape of the plant blossoms, as we have seen in previous discussions. In Chap. 4.3 “Mimicry as a sign system” it was argued that for a mimic, it is particularly problematic to enter and exit from the mimetic performance. Also in 115 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. the egg-mimicry, it is the presence of the adult cuckoo in the vicinity of the host nest that breaks the completeness of the mimetic resemblance. To minimise this effect, the act of laying eggs is kept very short and cuckoos use resemblance to the predatory birds to mask their presence in the host nest. In the case of Passiformes, the resemblance between mimic and model eggs is not static but an ecological relationship with cuckoos, and the threat of brood parasitism has influenced their egg recognition mechanisms towards becoming much more accurate. In addition to simple resemblance between mimic and model eggs, there are a lot of additional factors that influence the functionality of the egg mimicry. In general, host birds can distinguish their eggs’ overall appearance, size (Marchetti 2000), colour (Moskat et al. 2008) and pattern (Polačiková et al. 2010), including markings in ultraviolet (Honza et al. 2007). The tertiary iconic relations with the environment can influence the recognition of the brood parasites’ eggs, and individual learning and memory also have an important role in recognition. The dependence of egg coloration on environmental conditions in some species (Eurasian reed-warbler Acrocephalus scirpaceus) makes the recognition process much more complex, as the cuckoo—host egg matching depends on the amount of rainfall in a given spring (Avilés et al. 2007). Some species (for example, Hume’s leaf-warbler Phylloscopus humei) tend to throw out eggs that are bigger than eggs in the clutch on average (Marchetti 2000). Recognition also includes the possibility of a mistake— it has been observed that some birds tend to reject their own eggs if they are “unusual” compared to the rest of the clutch. In addition to the comparison of eggs side by side, egg mimicry seems to include a prototype generation and memorizing of specific types of eggs. In some species like the great reed warbler Acrocephalus arundinaceus and chaffinch Fringilla coelebs, the recognition rate is dependent on the age and experience of the bird (Stokke et al. 2004). The host needs to learn from its first nesting what its own eggs look like (Admundsen et al. 2002: 367). 4. How is the function of communication and how is mimicry regulated in ontogenetic and evolutionary processes? Foregrounding the communicative function in mimicry enables us to bring out the specific aspect of communication that allows the mimic to be successful in its deceptive display. From the six communicative functions that Roman Jakobson distinguishes, it seems to be predominantly the poetic function that is abused in the case of egg mimicry. I make this claim based on the specific qualitative properties that make a female bird accept the egg as part of its brood or 116 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. alternatively to reject it. There is no apparent usage of other communicative functions: no reference is made to an external object or contextual meaning; a cuckoo egg does not express any behavioural activity (which excludes expressive function), no attempt is made to alter the behaviour of the female compared to the normal fostering behaviour, and is are there no manipulation with the communication code or phatic contact. The presence of a poetic dominant could perhaps be an argument to pay more systematic attention to the specific aesthetic qualities that make birds become attached with their nest, eggs and brood. Asking about the regulation of egg mimicry in ontogenetic and evolutionary processes enables us to describe the mimicry system in terms of dynamics and fixation. In the case of brood parasitism, the communicative feedback, understood as feedback within a single act of communication, plays a minimal role as eggs are passive and do not participate in communication with adult birds. The relationship between the eggs and the adult birds rather follows the logic of unidirectional communication or signification (sensu Nöth 2001: 72), so that the adult birds are the active subjects that recognize eggs, attribute meaning to them, and act selectively according to this attributed meaning. The quickest feedback cycle in brood parasitism takes place on the level of the clutch, and depending on the host species, there are between a few and tenfold communication-feedback cycles throughout the life of an individual. Ontogenetic learning and feedback has some role in recognition, as birds may improve their egg-recognition skills on the basis of previous experience. At the same time, there is not much individual variety known to exist in the behaviour of hosts or parasites in this ecological relation, nor is the individual experience known to pass from generation to generation by cultural learning. It seems that egg mimicry is to a large degree (compared to many other mimicry systems) genetically induced and controlled. This is also supported by the existence of various genetically induced gents of the cuckoo that differ by their egg appearances and preferred host species. Such relatively fixed mimicry systems allow fixed bodily adaptations and counteradaptations to develop, as for instance in some species like blackcaps Sylvia atricapilla, egg variation in the clutch seems to have decreased because of brood parasitism (Honza et al. 2004). 5. How is the mimicry system related to human cultural processes? Regarding human cultural involvement and influences, egg mimicry turns out to be an especially rich and interesting case. The phenomenon itself is known long before the rise of modern 117 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. science. For instance Aristotle in his “Historia Animalium” gives a long description and different explanations of cuckoo’s nesting behaviour: The cuckoo, as has been said elsewhere, makes no nest, but deposits its eggs in an alien nest, generally in the nest of the ring-dove, or on the ground in the nest of the hypolais or lark, or on a tree in the nest of the green linnet. It lays only one egg and does not hatch it itself, but the mother-bird in whose nest it has deposited it hatches and rears it; and, as they say, this mother bird, when the young cuckoo has grown big, thrusts her own brood out of the nest and lets them perish; others say that this mother-bird kills her own brood and gives them to the alien to devour, despising her own young owing to the beauty of the cuckoo. (Aristotle 2002, Dd6r). It seems that brood parasitism of the cuckoo has even turned into a cultural model for describing certain parasitic relations, as there are cuckoo bumblebees Psithyrus, cuckoo finches Anomalospiza and cuckoo ants Leptothorax in the zoological nomenclature. There is also a larger cultural mythological background for interpreting brood parasitism, since many European cultures believe in the existence of changelings, or human children swapped by an elf, a troll or some other supernatural creature. The same theme of changelings is much used in contemporary fiction in different variations (e.g. John Wyndham’s “The Midwich Cuckoos”, 1957). It appears that the topic of an alien offspring is a strong cultural narrative and that brood parasitism is also the primary characteristic that people associate with cuckoos. Scientific studies of the egg mimicry of cuckoos can also be shaped (although mostly unconsciously) by this strong cultural narrative (see e.g. Schulze-Hagen et al. 2009; Smith 1999). Although the egg mimicry of cuckoos is scientifically a rather well-established case, a possible methodological error can arise from substituting the position of the receiver with that of the human observer and proceeding from human perceptual and behavioural possibilities. Concerning studies in brood parasitism, this becomes obvious for instance in the experiments with artificial eggs that are produced according to the human perceptual system and understanding of similarity and difference: “The resemblance was so good that, by visual inspection alone, an observer could not distinguish between the artificial eggs and the genuine cuckoo eggs from the same area” (Moksnes and Røskaft 1989: 27). Artificial eggs could also 118 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. be described as “similar in size and mass to real cuckoo eggs, made of hard plastic…the mimetic egg type painted to resemble eggs laid by the blackcap” (Honza et al. 2004: 176). This issue becomes methodologically crucial, as there are differences between human and bird visual systems. The principles of colour distinction in birds are different from mammals (birds having tetrachromatic vision), with many bird groups also being sensitive to ultraviolet light. Some parasitic eggs that appear nonmimetic in visible light are highly similar to host eggs in UV-light (Grim 2005: 75; Polaciková et al. 2007; Honza et al. 2007). In some studies, even human test persons are used to test the similarity or difference of eggs in a clutch and the results are used to argue for the resistance of some species to brood parasitism (Honza et al. 2004: 177). From a semiotic viewpoint, such studies can be interpreted critically and assumed to provide biased results since the position of the receiver (song bird) is at least partly replaced by that of the human receiver. David C. Lahti (2015: 530–531) smartly denotes such uncritical use of artificial objects in behaviour studies as Umwelt gamble: “when we perform experiments that assume continuity between our own perceptual and cognitive infrastructure and that of an animal—for instance, using artificial stimuli—we engage in what we might call an umwelt gamble. We bet that something we think would be perceived in a certain way will indeed be perceived in that way” (Lahti 2015: 530–531). The five-stage semiotic analysis of the common cuckoo’s brood parasitism demonstrates the complexity of the mimics’ and receivers’ relations as well as the several layers of resemblance compared to the simple schematization of the tripartite mimicry system. The method allows for bringing out different aspects of the mimicry system in an organised manner. It shows the specifics of egg mimicry as a detached and complete mimicry resemblance corresponding to Peircean secondness. The validity of the mimicry system could be supported by the strong behavioural reactions of the adult birds (receivers) toward the cuckoo eggs, although at the same time, the human cultural narrative and perceptual involvement can also influence the description. The dynamics and development of egg mimicry takes place mostly at the level of genetic information and phylogenetic feedback, although there is also some individual learning involved. To fully articulate the specific features of egg mimicry, a comparative analysis should be made in other mimicry systems as well, which will be carried out in the next subchapter. 119 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 8.3 Towards the comparative modelling In following subchapter, I will illustrate the proposed research method by analysing two well-known mimicry examples in comparison—brood parasitism of the common cuckoo Cuculus canorus on various passerine host species and Lepidopteran eyespots (e.g. eyed hawk-moth Smerinthus ocellatus). Egg mimicry is supported by additional resemblances, as there are correspondences between the behaviour of the cuckoo chicks and the chicks of a particular host species (for example Kilner et al. 1999) and similarities in the appearance of the adult cuckoo and hawk species (Payne 1977: 8). Eyespots are common feature in animal visual communication systems, found in almost all kinds of ecosystems. This sign type is present in the bodies of organisms living nowadays, as well as in species that were fossilized long ago (Labandeira et al. 2016). In butterflies and moths, eyespots are most common and best studied in Nymphalidae, but exist widely also in other families (e.g. Saturniidae, Sphingidae) as well as in fish, amphibians and reptiles. The common understanding of the function of the eyespots is the “intimidation” hypothesis, according to which the function of the eyes is to deter predators and prevent an attack. The usual reasoning here is that eyespots deter predators because they represent an imitation of vertebrate eyes: this is the “eye mimicry” hypothesis (Blest 1958, see discussion in Kleisner, Maran 2014). Eyespots have yet other functions such as playing a signalling role in sexual selection. In the butterfly Bicyclus anynana (Nymphalidae), for instance, the dorsal wing pattern is supposed to partake in female mate choice, whereas the ventral pattern serves a camouflaging and/or predator-deflecting role (Brakefield and Reitsma 1991; Stevens 2005). The two common mimicry systems—egg mimicry of the common cuckoo and Lepidopteran eyespots mimicry—could be compared using the five-staged research method proposed above. Since a longer discussion of the egg-mimicry of the common cuckoo was given in the previous subchapter, I will present here a comparison between the two mimicry cases in a concise form (see table 8.1, figure 8.3). Table 8.1. Comparative analysis of mimicry in brood parasitism (Cuculus canorus) and Lepidopteran eyespots. Brood parasitism 120 Lepidoptera eyespots Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 1. The formal structure of the mimicry system (see figure 8.3) Bilateral mimicry system Tripartite mimicry system with (model and the receiver protective function. belong to the same species) with the reproductive function. 2. Perceptual and effectual correspondences between the participants Detached mimicry (imitating and imitated objects are distinct from the bodies of the participating animals) operating mainly through the visual channel. Mimicry has high relevance for the receiver and it has developed specific behavioural reactions (punctuating the egg, abandoning the nest). Operating mainly through the visual channel. Mimicry system takes advantage of the dynamic and quick communication between the predators and prey. Receiver's reactions differ radically in regard to whether the sender is recognised as model or mimic. 3. Characteristics of resemblances Prototype resemblance or common characteristic resemblance. Objects compared are physically together and mimicry is complete (resemblance covers most aspects of a single object). Prototype resemblance or common characteristic resemblance with some inclination towards the abstract mimicry (model is diverse or unspecified). In some cases different sign sets (modelreceiver, mimic-receiver) can be combined. Objects compared are spatially distinct from eachother and comparison must rely on the memory of the receiver. 4. Function and regulation in communication Poetic dominant in communication. Communication is mostly genetically induced and controlled with some regulation via ontogenetic learning and feedback. Referential dominant in communication. Communication is mostly genetically induced and controlled with some regulation via ontogenetic learning and feedback. 121 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 5. Relation to human cultural processes Mostly accessible in human Umwelt. Larger cultural mythological background is present for interpreting brood parasitism; common cuckoo acts as a cultural model for describing other parasitic relations (cuckoo bumblebees, cuckoo finches) Accessible in human Umwelt. Eyes are conspicuous structures (semantic organs, cf. Kleisner 2015) for humans and for many other species. At the same time, there are no well-known general cultural narratives in regard to eyespots. [Enter figure 8.3 here] Figure 8.3. Graphic representation of the analysed mimicry systems. Left. Common cuckoo’s (MI) mimicry system cuckoos’ host species (MO1) and hawks (MO2) act as models. The unbroken arrow lines represent the communicative relation between the model (MO) and the receiver (R) that is exploited by the mimicry system (circular arrow represents autocommunication in the host species). The dashed arrow line represents the deceptive relation between the mimic (MI) and the receiver. The dotted line represents the relation of resemblance between the mimic and the model. Right. Mimicry system of the Lepidopteran eyespots, where a butterfly species acts as mimic (MI), taking advantage of the communicative relation between owls or small carnivorous mammals (MO), and small insectivorous birds (R). Note that from the perspective of the mimic, both model and receiver are not specific species, but rather abstract groups (and therefore blurred in the graph). Discussion. There are both considerable similarities and differences between the mimicry in brood parasitism and Lepidopteran eyespots. Both mimicry systems operate on visual communication (2, 5) and are thus well are accessible to the human Umwelt. This proliferates human interest towards these mimicry systems, but at the same time lays ground for over-interpretation (this should especially be taken into account in the case of the brood parasitism that is also a common cultural narrative). Easy visual access to the mimicry may be a cause for disregarding similarities in other communication modalities (UV, tactile characteristics), that is especially relevant for 122 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. the egg-mimicry system. The regulation in both mimicry systems are similar (3, 4), relying primarily on genetic processes at the population/species scale with some input from individual learning and recognition. In the case of brood parasitism (1, 2), the combination of the species involved is more restricted and the participant’s specific behavioural reactions and adaptations are well established (compared to eye-spots). This difference is supported by the different dominant in regard to communicative functions (4): poetic in brood parasitism and referential in the mimicry of eyespots. From a semiotic perspective, the following differences can be brought out between the two mimicry systems (2, 3): egg mimicry in brood parasitism is detached (imitating and imitated objects are distinct from the bodies of the participating animals), comparable (objects compared are physically together), and complete (resemblance covers most aspects of a single object). This creates good evolutionary and cognitive preconditions for fine-tuning the mimetic resemblance, recognition mechanisms and behavioural responses on which many examples are known (for example, throwing out eggs that are bigger than eggs in the clutch on average, Marchetti 2000). Lepidoptera eyespots mimicry, on the other hand, is an open mimicry system (1, 2, 3), that takes advantage of the quick communication between winged organisms and relies on receivers’ recognition capacity, memory and perception of the image of the eye. Eyespots mimicry relies upon the broader semiotic convention in an ecosystem about the metonymic relationship between the image of an eye and an animal that carries this eye. Metonymic relation means in this instance apart standing for a whole; therefore, there needs to be a general understanding in an ecosystem that eyes are signs standing for an animal that is active and potentially dangerous. Eyespot mimicry is much less dependent on the specific receiver species (as there are many of these involved), and rather floats in between the resemblances and meanings shared by the inhabitants of the biological community (see the Chap 11 “From abstract mimicry to ecological code"). The eyespot mimicry system could potentially be considered an open mimicry system (different from brood parasitism) that can easily incorporate new species and images. The analysis confirms that both mimicry systems (1) are well-established and complex in the natural world. It further demonstrates that although both resemblances operate in the visual communication system and have basically the same group— small song birds—functioning as a receiver, the ecological and semiotic characteristics of the mimicry systems under observation are very different. 123 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Comparative analysis of these two mimicry cases hopefully demonstrates the possibilities of semiotic modelling as a research method. It is a tool for getting a quick overview of different facets of the observed mimicry cases. Semiotic modelling surpasses the limits of an evolutionary understanding of mimicry and allows mimicry to be related and analysed in comparison with various other behavioural imitations. It helps further to determine the role of human cultural interpretation in mimicry and gives the researcher a self-critical position for the description. By having a set of different questions as a fixed basis, it also allows comparing mimicry cases that may otherwise be quite difficult to relate to one-another. 124 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 9 MIMICRY AND SEMIOTIC EVOLUTION Mimicry has historically been used as a main argument for supporting Darwin’s theory of evolution. This was the original emphases of Henry W. Bates when he wrote: “I believe the case offers a most beautiful proof of the truth of the theory of natural selection.” (Bates 1862: 513). The chronicler of evolutionary biology, Ernst Mayr, has written that Darwin’s “On the Origin of Species” was easily attackable as it was mostly based on deductive reasoning and included little proof except the analogy between artificial selection and natural selection. Bates’s discovery that allowed for demonstrating the mechanisms of natural selection in nature turned out to be a good argument against such criticism (Mayr 1982: 522–523). In later discussions, evolutionary adaptations, fitness landscapes and other conceptual tools of NeoDarwinism were used for interpreting mimicry: e.g. for determining the positions of the mimic and model in mimicry systems (in Mertensian mimicry, Wickler 1968: 111–121), as criteria in mimicry typologies (Starrett 1993), or for theorizing about the specific mimicry cases. There have also been some alternative attempts to describe mimicry (e.g. Theodor Eimer’s (1897) orthogenesis, Franz Heikertinger’s (1954) nonadaptationist view, Stanislav Komárek’s (2003) historical interpretation), but these have been met with critical objections or have been marginalized in an academic debate. In this chapter, I will discuss the connection between mimicry and semiotic evolution, whereas the general focus of this book still lies in the structural aspects of mimicry and horizontal communicative processes therein. A history of mimicry studies demonstrates that mimicry has not been a most loyal proponent of the Neo-Darwinian understanding of nature. Explaining why and how the specific resemblance could be beneficial for its bearer, or how the mimicry types could have been evolved, often appear to include quite complex logical gymnastics (as evident, e.g. in Mertensian mimicry of coral snakes, cf. Komárek 2003: 114–115 and mimicry complexes of the Heliconius butterflies, cf. Mallet and Gilbert 1995). These difficulties are probably related to the variety and diversity of mimicry in nature and the problematics of organizing different mimicry cases around a single and unified theoretical core of natural selection. When more complex and exceptional mimicry cases are found, more ad-hoc explanations are needed to connect these to the core of the Neo-Darwinian understanding of evolution: differentiated survival of genes and fitness leading to new evolutionary adaptations. 125 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Evolutionary explanations of mimicry are often used as a basis for defining mimicry in general and for finding proof of the existence of the specific mimicry systems. The strongest possible proof for mimicry in an evolutionary perspective would be a demonstration that the change in some evolutionary agent (animate or inanimate) has caused a specific mimicry adaptation to emerge or proliferate. An example of this kind of strong proof are studies on the abundance of dark-winged moths in Great Britain during the rise of the coal-based economy in the nineteenth century (so called industrial melanism), where the darkening of the visible environment correlated with the amount of melanismic moths (e.g. Berry 1990), although even this case has been an object of dispute. Such strong proofs are more than rare, especially in the face of common claims about mimicry as an evolutionary adaptation par excellence. More often the existence of mimicry is argued for by indirect proofs, such as the correlation between the specifics of the receiver’s sensory perception and the resemblance between the mimic and the model; correspondences of the living areas of mimics and models; indications of predatory pressure, such as the relative abundance of the mimic and the model in the diet of a predatory receiver; the location of injury in the mimic’s body regarding mimetic features (such as birds’ beak marks on butterfly wings); behavioural responses of the receiver to specific appearances of models and mimics (warning coloration); the receiver’s ability to learn from unpleasant experiences with models; or experiments with the manipulation of mimetic features, e.g. painting mimetic features onto a non-mimetic organism or covering up mimetic features, etc. These various arguments are indirect in the sense that although they demonstrate different types of correlations between mimics and models or illustrate how an organism benefits from mimetic features, they cannot prove that mimetic features have developed because of the specific relations between the models, the receivers and the mimics. At the farthest end of this scale of validity are cases where human senses are used to determine mimetic relations—if a mimic and a model seem similar to us, a mimicry resemblance is judged to exist. Thus, the problem with the perception of mimicry in mainstream biology has two sides. First, mimicry is often seen as a distinct phenomenon that has definite borders and secondly, it is supposed to be provable as such. However, many classic examples of mimicry in nature, as well as arguments used to explain these, do not correspond to this wide-spread scientific conviction. In comparison, a biosemiotic approach appears to be more dynamic as it does not need to rely on the abstracted 126 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. selection process but sees semiosis as initially local and contextual. Every semiotic process—perception, recognition, communication—that involves the participants of mimicry can also have effect on mimicry as an evolutionary system. On the other hand, as a local process, semiosis includes specific characteristics of the mimic, the model and the receiver and incorporates these into mimicry as an evolutionary system. By focusing on semiosis and sign relations, the semiotic approach is particular, as opposed to abstract, general treatments. In the biosemiotic perspective, the borders of mimicry are necessarily blurry, as they are based on the sign activities and interpretations of the mimicry participants. What counts as a mimetic feature or mimetic resemblance is a part of the sign as it participates in semiosis and becomes manifested through this process. One could talk about efficiency or fitness in regard to mimicry, or define mimicry based on such a property, but these measurements stay outside of the sign as a combination of representamen, object and interpretant, and can, at best, indicate something about the result or outcome of the semiosis. The properties of the sign or message (colour, pattern, perceivable sound) are perceivable in the Umwelt of the interpreter, whereas the effectiveness of the mimicry is the measure given by the viewpoint of the external observer. There can exist correlation between the formal features, interpretation and outcomes of the sign, but this relation is never deterministic in regard to semiosis. Although different interpretations made by the receiver may have different effects on the participants of mimicry, the effectiveness of communication or evolutionary fitness can hardly be measures that the receiver could take into account during the interpretation. Often it just lacks access to this information or does not have cognitive capacities to make the necessary generalisations. What appears to be needed to improve the general understanding of mimicry is focusing on the specific interactions between senders and receivers. Such a view appears to be supported by some eminent mimicry scholars as well, for instance by British biologist Tim Guilford, who emphasises the predator psychology as a factor that could very much influence the properties of prey animals, including the aposematic colouring and mimicry. For instance, Guilford considers the searching image used by a predator to be an entity that influences the characteristics of the mimic. If the predator focuses on the common and more characteristic features of the species, then this may lead to the development of polymorphism as a counter adaptation (Guilford 1992: 378–384). He also describes the repetitious patterns of the 127 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. warning coloration as probably caused by the perceptual psychology of the predator. Patterns of yellow and black, red and black and other similar colour combinations are easily perceivable from very different perspectives and also in environments rich in perceptual noise, for instance in cases when an insect with aposematic coloration is partly covered with foliage (Guilford and Dawkins 1993: 432). The receiver interprets cues and messages as it finds these in its Umwelt: it sees, smells, hears and senses specific colours, patterns, sounds, rhythms, smells and tastes, the dynamics and change as well as resemblance and difference between them. Interpretation may have effects on the messages emitted by senders through feedback and action, but this effect is mediated by way of how and what the receiver perceives as discrete sign complexes in its Umwelt. The receiver’s perception and interpretation is under-determinate, as the searching image it uses does not have a one-to-one correspondence to the physiological properties of the animal that is sought after. Also, animals’ responses to the environment are seldom absolutely stereotypic. This is because the organism in its particular being faces the variability of other organisms, environmental conditions and specific communicative situations. Therefore any communicative encounter (e.g. mimicry) also needs to include some interpretive effort on behalf of the participants. What we have in animal semiosis thus appears to be a delicate balance between the repetition of the earlier habits and communication patterns on the one hand, and ambiguity leading to novel phenomena on the other hand. Recognition as a conformity with the earlier form or model makes it possible for the sender’s and the receiver’s semiotic activities to match to one-another, whereas interpretation of cues and messages allows for accommodating novel situations and development of new communication codes. In this view, evolution and semiosis become abridged, which is another way to express Thomas A. Sebeok’s maxim of the interrelatedness of life and semiosis. As communication scholars Allen Ivey and James Hurst explained it decades ago: „Communication develops through constant process adaptation. Each communication depends on past communications in which the participants have been involved. While strict and precise definitions may be made of communications, no one can predict the future consequences of new communicative acts with absolute certainty.” (Ivey and Hurst 1971: 207). In the following subchapters, I will focus on two themes in which semiotic processes take a lead role. First, I explain the role of the receiver’s semiotic activity 128 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. and Umwelt structures in shaping the properties of the sender (e.g. a mimic) in a process that I have called semiotic selection. The discussions on semiotic selection are formed in dialogue with my good colleague Karel Kleisner from Charles University in Prague. Thereafter I make a connection between mimicry studies and Jesper Hoffmeyer’s concept of semiotic scaffolding and emphasise the temporal structure of mimicry—how earlier stages of mimicry constrain the development of later stages. In both topics—semiotic selection and semiotic scaffolding—the balance between fixed and habitual on the one hand and novel and unexpected on the other hand is an implicit organizing principle. In both cases, it is the activity of an animal organism as a subject that shapes the evolutionary changes in mimicry. 9.1 Semiotic selection: Definition and examples 25 In this subchapter, an attempt is made to organise evolutionary approaches in semiotics while simultaneously proposing some novel terminology. In doing so, I am well aware that semiotic processes that have evolutionary results are diverse and generally not predetermined. Therefore, the following should be considered not as an attempt to build a rigid typology, but rather as a proposition for epistemological models that appear to have some counterpart in natural processes and that could help us analyse these processes better. My main argument is that semiosic processes are crucial for evolutionary change, and I will focus more specifically on mimicry, domestication, and Darwin’s treatment of sexual selection as examples in order to reveal their common underlying theoretical basis. I argue that these phenomena are critically dependent on the peculiarities of the Umwelt of a particular animal interpreter (selector). In all three of these phenomena, an individual gains some cues and signals from another living organism, interprets these, and acts selectively in relation to the other organism. In the case of repeated interaction during the life of the same individuals or over generations, such relation can produce a directed evolutionary influence. The term semiotic selection (sensu stricto selection because of semiosis) has therefore been proposed to embrace all phenomena where subjectspecific interpretation comes into play. The effect of semiotic selection can be 25 This subchapter is partially based on Maran, T, Kleisner, K. (2010). Towards an evolutionary biosemiotics: Semiotic selection and semiotic co-option. Biosemiotics (Springer), 3(2), 189–200. I want to thank a good colleague Karel Kleisner from Charles University in Prague for a long-lasting and fruitful cooperation. Used with permissions. 129 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. suspected, for instance, in cases where there is a specific one-to-one correspondence between the perceptible characteristics of one animal and the perceptual organs of the other. Semiotic selection operates through perception, interpretation and feedback, being thus an evolutionary derivative of Uexküll’s Funktionskreis (Uexküll 1982: 31). Proceeding form Uexküll’s “Bedeutungslehre”, Kalevi Kull (1998) has emphasized the role of human linguistic and interpretational activity in changing our environment through actions. Kull lists several processes that are an inseparable part of human cognitive activity, the effects of which are transposed back into nature through voluntary or involuntary activities. Such processes include: 1) recognition and control, 2) decontextualisation, 3) operation and remodelling (forming), 4) opposition and reduction, 5) understanding and devaluation; 6) selfing and valuation (Kull 1998: 352–354). I extend this principle of semiosically charged change to all living organisms that have an Umwelt. Many of the processes that Kull distinguishes also appear to be present in other animals besides humans: recognition, subsequent categorization and control appear to be quite general properties of animals. There is also an equivalent for opposition and reduction in animals, if recalling Darwin’s principle of antithetic expressions of emotions (Darwin 1872a: 50–66). In biosemiotics, much attention is given to the concept of organic selection, more commonly known as the Baldwin effect after its founder, James Mark Baldwin (1896). Organic selection describes the effect of an organism’s activities directed at itself and its environment and the consequent evolutionary effects. The basic mechanism of organic selection lies in the organism’s choice made in the larger space of possibilities that later limit possibilities of natural selection to operate, as the selection can only operate upon those features that are expressed or used. The choosing of a novel food source by an organism with its subsequent later ecological and evolutionary consequences is a typical example of organic selection. A good explication of this process was provided by Markus Lindhom (2015) in his reinterpretation of the classical case of radial adaptation of Darwin’s finches in the Galapagos Islands. He demonstrates that it has been finches’ learning and behavioural choices in regard to habitat and food that have later lead to the emergence of the specific adaptations and speciation. There is, however, a crucial difference between the concepts of semiotic selection and organic selection; in semiotic selection, it is not the animal itself, but another organism (as an evolutionary unit) that is the object of the selective process. 130 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. In semiotic selection, there may be no evolutionary effect on the selector (the organism making the selection) itself. This difference can be explicated by focusing on parallels between evolution and communication (as pointed out by Ivey and Hurst 1971). It is possible to characterize both evolution and communication by distinguishing two general processes that can be described as outward and inward directed. Different forms, variability in expressions and messages, are created in outward processes (e.g. uttering a message); while in inward processes, the same things are acted upon: they are selected, interpreted and responded to. In communication, outward process corresponds to formulating and sending messages and inward process to receiving and interpreting them; in evolution, mutations and creative behaviour are examples of outward processes, whereas selection thereof is an example of an inward process. If assuming that both processes can involve a semiotically active subject, we can organize different forms of selection based on the subject's activity or non-activity in the outward and inward process (see Table 9.1). Selection is considered here as a general term to denote a two-phase process, in which alternatives are created and acted upon, thus producing change in a biological system. Table 9.1. Four types of selection characterized by the predominant localization of the activity of the semiotic subject. Activity of the semiotic in outward process in inward process Natural selection not active not active Organic selection active not active Semiotic selection not active active Communication as active active subject selection In organic selection, the activity of the semiotic subject seems to develop, calibrate and apply the organism’s capabilities according to a specific situation and 131 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. environmental context. In the words of Baldwin: “we may […] apply the phrase, “Organic Selection,” to the organism’s behaviour in acquiring new modes or modifications of adaptive function with its influence of structure” (Baldwin 1896: 444). In semiotic selection, the semiotic subject does not have any direct influence on the development of another organism’s structure; it can only interpret and choose between the forms that are already laid out before it. Other types of selection that can logically be distinguished in this framework include natural selection (sorting), as it is considered a process in which the activity of the semiotic subject is kept minimal both in the creation and application of biological forms. The influence of climate on the properties of mammalian fur may serve as a good example. Under communicative selection, I describe communicative and cultural interactions in animals (including man), where the semiotic subject is involved in both creating messages and forms and acting upon them. In this type of interaction, the level of semiotic freedom and innovativeness is the highest and often results in changes and novelties that spread horizontally among the members of a population. In mimicry, resemblance induced by an individual leads to interpretation or misinterpretation by another individual that in turn leads to some communicative or behavioural feedback (for instance eating an insect or pollinating a flower) that may well have evolutionary consequences. Such a sequence of actions in which the participation of an organism as an individual in a semiotic process leads to evolutionary change can be called semiotic selection (Maran 2005: 171). Mimicry is just one, albeit remarkable example of such phenomena. Evolutionary processes by semiotic selection are not directly concerned with the physical existence of organisms, but are mediated by communicative or semiotic processes. In semiotic selection, organisms participate as perceivable and responding organisms with their cues and signals in the environment. Thus semiotic selection is essentially qualitative, based on qualias, and not quantitative, although its results are often also quantitative in nature and can be described as such. To exemplify the diversity of the processes of semiotic selection, three examples that belong respectively to mimicry, domestication, and sexual selection will be used. From the numerous examples of mimicry, I focus on a well-studied group of myrmecophilic insects in the butterfly family Lycaenidae (for general discussion, see Fiedler 2006). This group contains approximately 6000 known species, of which 75% have some form of association with ants (Pierce et al. 2002: 132 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 734). Many Lycaenids possess a chemical mimicry of the ant’s pheromones or other chemical substances. For instance, Akino and colleagues show that larvae of the lycaenid Maculinea rebeli (Mountain Alcon Blue, synonym Phengaris rebeli) produce hydrocarbon compounds sufficiently similar to that of Myrmica schencki ant larvae to induce M. schencki workers to carry M. rebeli larvae into their nests (Akino et al. 1999). Many Lycaenid butterflies lay their eggs near the ant nests to ensure that worker ants will pick them up. Lycaenid species also have many other non-specific adaptations serving their encounters with ants, for example Pore cupola organs that secrete substances to pacify ants and a cuticle that can be up to 20 times thicker than that of larvae from other Lepidopteran families (Pierce et al. 2002: 737–738). Such adaptations probably allow Lycaenids to survive ant attacks. More specific communication abilities and defence mechanisms in Lycaenids can develop with the help of semiotic selection by ants. The differential behavioral reactions of both ants (Jordano and Thomas 1992) and Lycaenids (Axén 2000), as depending on the encountered other, are exemplary of the discussion of semiotic selection; in several cases, there is a specific correspondence between the chemical compounds of ants and those of Lycaenid species (Schlick-Steiner et al. 2004). Sexual selection can be addressed by focusing on song-birds such as the European starling Sturnus vulgaris, whose song is known by its variety, complexity, individuality and imitativeness. It should be noted, however, that characteristics, functions and ecology of bird songs vary to a significant degree between different species, and the following discussion on the European starling cannot be applied uncritically to other species. The songs of the European starling are well studied, and many authors find in them an essential role in sexual communication (e.g. Eens et al. 1991: 221). Females of the European starling have been found to prefer males who sing in longer average song bouts and have larger repertoire sizes (Gentner, Hulse 2000; Eens et al. 1991: 231). It is very significant that such preference remains even in cases where males with elaborate songs do not occupy the most preferred nest sites (Mountjoy and Lemon 1996). When talking about sexual selection, it is important to note that the concept is used here in a Darwinian, but not Neo-Darwinian sense. Darwin regarded the sense of beauty in animals as a general by-product of their nervous system, but did not consider aesthetic expression to be an indicator of an animal’s reproductive potential. Darwin thought that the appearance of vivid coloration and melodious songs in birds could be results of female preference and 133 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. selection. In his words: “If man can in a short time give beauty and an elegant carriage to his bantams, according to his standard of beauty, I can see no good reason to doubt that female birds, by selecting, during thousands of generations, the most melodious or beautiful males, according to their standard of beauty, might produce a marked effect.” (Darwin 1872b: 70). The third example concerns domestication, and here, the focus is on the Dachshund, a dog breed of German origin that has been seen in illustrations dating back to the 15th century. Dachshunds were developed specifically to pick up the scent of, chase, and drive badgers and other burrow-dwelling animals out from their cover. This rather specific purpose is expressed in the Dachshunds' low and stretched body, their good sense of smell that is comparable to that of other hounds and terriers, their long ears that prevent mud and dust from getting into the auditory canal, as well as their psychological readiness to enter burrows and fight animals in such cramped spaces. The short legs of Dachshunds are actually due to developmental defects known as chondrodysplasia or chondrodystrophy (Parker 2009). This genetically induced abnormality leads to the premature hardening of the outer layer of bone and thus prevents the limbs from growing to full size. Importantly, this state has probably been caused by artificial selection, generated by the visual preferences of humans, i.e. it corresponds to the criteria of semiotic selection. These three examples differ from each other in several respects (see table 9.2). First, the group of organisms that function as selectors are biologically quite different—in the case of mimicry, Hymenopterans, in sexual selection, Passeriformes, and in domestication, Primates. Corresponding to this, the communicative channels where the selection takes place also differ from each other: in the case of mimicry, the predominant channel is chemical, in sexual selection, auditory and in domestication, visual/tactile. One could also note the difference in ecological relations: the mimicry of ants is based on parasitism, sexual selection belongs to intraspecific relations and the breeding of Dachshunds seems to be related to symbiosis. There is also a relevant difference between the bases of the selection: in the examples above, mimicry is based on similarity/categorization, sexual selection is expectedly based on aesthetic faculty, and for domestication, functionality is a dominant motive. 134 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Table 9.2. Characteristic features of three examples of semiotic selection: mimicry in Lycaenids, sexual selection in European starling; and domestication of the Dachshund. Mimicry in Lycaenids Sexual selection in Domestication European starling of Dachshund Selector Hymenopterans Passeriformes Primates Channel Chemical Auditory Visual Relationship Parasitism Intraspecific relation Symbiosis / parasitism Basis of selection Categorization / similarity Aesthetic faculty Functionality / profitability We may conclude that semiotic selection can occur in very different types of relations, and that a specific analysis of animals’ Umwelten and bodily forms is necessary for studying such relations. In all of these examples, the specific matching occurs between the forms or messages of the sender and the perceptual system of the receiver. As a general concept, Hoffmeyer has used the phrase “semethic interaction” to denote such semiotic relations that lead to the forming of new regular behaviours or habits (Hoffmeyer 2008a: 189). In all given examples of semiotic selection, some image of the selector’s Umwelt is imprinted onto the perceptible features of another organism. In the case of Lycaenids, it is the brood’s smell in the ant’s perceptual world, in the case of the European starling, it is a certain aesthetic preference within the female’s perceptual world, and in the case of the Dachshund, it is the human understanding of the dog’s characteristics suitable for badger hunting. Thus, semiotic selection creates a connection between the animal’s inner perceptual sphere and physical forms in nature, and influences the latter to change in accordance with the former. It also creates iconic linkages between different species, their physical features and perceptual preferences, which can be an important factor for organizing and stabilizing ecological systems. 135 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 9.2 Mimicry and semiotic scaffolding In the concept of semiotic selection, the effect of a receiver on the properties of the mimic due to the receiver’s Umwelt structure and subsequent action was described, but there is also another essentially semiotic process active in mimicry evolution. Namely, how an organism in a mimicry system “selects itself”; that is, what the effects are of its sign activity, interpretation and actions for its subsequent developmental and evolutionary possibilities. I will discuss the developmental and evolutionary dynamics of mimicry by applying the concept of “semiotic scaffolding” that was introduced by Danish biosemiotician Jesper Hoffmeyer. Semiotic scaffolding can be seen here as an elaboration of James Baldwin’s principle of “organic selection” (cf. Chap. 9.1), with an emphasis on the constraints and causation in the temporal dimension. The main argument of this chapter is that although biological mimicry is an evolutionary phenomenon, it does not develop because of abstract selection pressures, but mostly through semiosis, or meaningful relations between organisms (that often include or are mediated by the environment). The evolution of a mimicry system is shaped by both the actual physical conditions (e.g. participants, environmental conditions, and communication channels) and potentials for new semiotic linkages to occur. To say that these relations are meaningful is to say that in mimicry relations, different organisms are bound together through perception, recognition, communication and action and they act in these relations as subjects, interpreting each-others’ perceivable appearances in a species-specific manner while also including their individual ontogenetic experience. The evolution of mimicry can thus be envisioned as a compound process of myriads of interactions between individual organisms, each equipped with their semiotic competences located in the context of different semiotic affordances (Gibson 1986: 127; Maran 2014b). Each act of perception, communication and subsequent action have an effect, although minute, on the mimicry system, thus creating and shaping conditions for future stages of the system. If we consider mimicry not as a static structure, but as a dynamical phenomenon changing through the involvement of living organisms, the question arises of how the earlier stages of mimicry relate and influence the subsequent stages. 136 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. This can be explicated with the help of the concept of semiotic scaffolding (Hoffmeyer 2007, 2014a, 2014b), which can be understood in a wider sense as a succession of stages of evolution, development or sign process in which the previous stages form the conditions for the subsequent stages. In semiotic scaffolding, a subsequent stage of the semiotic process presupposes and is based on the existence and configuration of the previous stage, or in other words, “new scaffoldings have been built on the top of those already operative” (Hoffmeyer 2014b: 107). Jesper Hoffmeyer (2007: 156) has explained this further in a figurative but precise way: “The emergence of new scaffolding devices (unknowingly) function like stepping stones in a river, leading evolutionary processes forward one step at a time and—in average—farther away from the bank at each step.” The criterion that helps us distinguish scaffolding from other similar phenomena (e.g. exaptation, semiotic cooption, Maran and Kleisner 2010) could be a question of whether an earlier developmental or evolutionary step is critically important for the living system to reach the present situation, and whether in this process the system remodels and uses the conditions established by the previous stage. If the system would be unable to reach the present stage without making use of certain conditions of development or evolution actualized in the previous stage, the phenomenon could be considered as scaffolding. When analysing scaffolding from a logical point of view, there appear to be several aspects or sub-types of this phenomena. The basic distinctions lie between ontogeny and phylogeny (cf. Williams et al. 2009), and between physiology and cognition. Of these four types, scaffolding in the ontogeny of an individual, either in prenatal or postnatal development, appears to be most studied. 26 In mimicry systems, we can find a good example of scaffolding in the ontogenetic development of eyespots of butterflies. The evolution and genetics of butterfly wings is long celebrated by researchers as a complex case of interaction between gene-based evolution and development process (Nijhout 1986, 1994; French 1997; Nijhout et al. 2003). Acknowledging the complexity of the topic, I will give here only a cursory overview of the sequence of the developmental stages in the ontogeny of the eyespots on butterfly wings. Also, to keep the discussion brief, I will overlook the diversity of 26 A related concept is developmental scaffolding (cf. Giorgi and Bruni 2015), but I prefer here ontogenetic scaffolding as the former concept has quite diverse meanings in developmental biology, education and developmental psychology. 137 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. different butterfly families (including specifics of poly-morphism and poly-phenism). Veron French (1997) has described the general logic of development of eyespots on butterfly wings (in buckeye Precis coenia) as a multilevel process that begins with the localization of eyespots in a developing pupal wing, specified by the expression of the regulatory gene Distal-less (dll) at the central signalling region (the involvement of specific transcription factors and genes differ depending on the species, cf. Monteiro et al. 2013). The signalling region thereafter produces a focal signal(s) (e.g. a concentration gradient, the different levels of which specify concentric colour rings) that spread across the epidermis (French 1997: 524). As a third step, the exact colours and proportions of the eyespot are determined by the responses of the scale-forming cells in the wing epidermis, whereas the different regions of the epidermis respond differently to the focal signal. We can see that every stage of the development of the eyespot acts as a scaffold for the characteristics of the following step to emerge. Stepwise formation of the eye-spot depends on local developmental conditions as well as on general environmental conditions such as temperature and moisture (Brakefield and French 1999; Otaki 2008). Such regulation through scaffolding produces the variety and diversity of spots on butterfly wings both within and between species. Many possible examples of scaffolding in mimicry will relate to phylogenetic and evolutionary processes. From an almost infinite number of possible examples, we can use the complex communication system between weaver finches Estrildidae and their brood parasites, the indigobirds Vidua (Payne 1977; Payne et al. 2000). I have previously described this amazing communication system (Maran 2011: 250–251, cf. Chap. 7.2): A vivid example of this kind is provided by Robert Payne in his studies on indigobirds Vidua that are the brood parasites of the weaver finches Estrildidae (Payne 1977, p.12; Payne et al. 2000). As nestlings, indigobirds learn songs from their foster parents and use these later to invite their own mates. In addition, female indigobirds have a preference for the song that they heard in their youth. The imitated song thus becomes the uniting feature for indigobirds reared by the same host, so ensuring the continuation of the lineage that prefers that particular host species. In the case of indigobirds the communicational code borrowed from another species becomes a basis for 138 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. intraspecific identity and they have given up their original means of expression. In the context of the present argument on semiotic scaffolding, we can interpret this example as a series of scaffolding steps: 1. Certain ecological conditions of the weaver finches (e.g. effectiveness in occupying nesting sites and skills building elaborate woven nests) has opened up new evolutionary possibilities for brood parasitism by other species. 2. Becoming brood parasites, indigobirds have faced limited possibilities for intraspecific communication between sexes, and parasitic ecological strategy has also laid down the specific conditions for the nestlings of the indigobirds to be in contact with the foster parents and their vocalisations. 3. Acquiring the song as well as the song recognition pattern of the host has turned out to be an effective new means for finding species mates in brood parasites. This series of steps has led to the evolution of a very peculiar and species-specific strategy for intraspecific communication and recognition. Another typological distinction in the scaffolding process can be made between physiological and cognitive processes of scaffolding. Turning again to the examples in mimicry, there are clearly cases where the existing physiological or anatomical properties of an organism turn out to be decisive factors in channelling the evolutionary process in a way that enables the particular mimicry resemblance to emerge. In his overview of mimicry in plants, Delbert Wiens has described succulents that imitate ground, stones or dead grass and branches (Wiens 1978: 384–385). The imitation can be based on gathering loose sand particles between hairs, as it is in cactus Ariocarpus kotschoubeyanus in Mexico, by elongated central spines that make Pediocactus papyracanthus resemble dried grass, or on complex changes in plant coloration and form (Lithops and other species of Mesembryanthemacea, see figure 9.1), where even the “shape, size, color, texture and fracture lines” (Wiens 1978: 385) of specific rock types can be imitated. Although mimicry in plants is a very complex phenomena, it can be assumed that imitations of dead organic matter depart from the characteristic of many plants growing in an arid environment to have partially dried branches or greyish leaf coloration because of different drought-related effects and adaptations. Under suitable conditions, these features can have an effect of making the plant less attractive to herbivorous animals, thus closing the semiotic loop of the mimicry system. 139 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. [Enter figure 9.1 here] Figure 9.1. Mimesis of the living stone Lithops fulviceps on the stony background (Collections of the Tallinn Botanical Gardens, photo courtesy of the author). In fact, in most cases of mimicry, the resemblance does not start from an empty place, but in the framework of already existing structures of organisms’ bodies. From a semiotic viewpoint, it would be pertinent to acknowledge that there is no such neutral starting point for mimicry, as biological organisms are always equipped with some body shape and appearance that has affordances and cues that have potential to become involved in semiotic processes. The initial physical similarity of the forms and patterns that gives ground for mimicry resemblance may have developed due to chance, being induced by evolution in similar physical or ecological environments (e.g. convergence), having a similar genetic basis in the case of closely related groups, etc. As an example of convergence, a close resemblance of bodily forms between snake-eels Leiuranus, Myrichthys and sea snakes Laticauda (Pernetta 1977; Randall 2005) appears to be based on the fact that both groups have slim and streamlined bodies that are evolved and adapted for quick movements in huge water masses. Many species of snake-eels (e.g. the saddled snake-eel, Leiuranus semicinctus) have, in addition to their snakelike bodies, a specific warning coloration of black and pale (white, light blue) stripes that is a characteristic feature of sea snakes of the genus Laticauda. The bodily resemblance formed an operating mimicry system, where avoidance behaviour between the predatory species (sea-birds) and sea snakes emerged and predators started to erroneously recognize snake-eels for sea snakes (cf. Caldwell and Rubinoff 1983). The close similarity of colours between snake-eels and sea snakes can be considered to be a secondary evolutionary adaptation. In its physiological dimension, scaffolding comes close to the phenomenon of exaptation (Gould and Vrba 1982: 6), where certain characters or properties have “evolved for other usages (or for no function at all), and [are] later "coopted" for their current role”. At the same time there is a considerable difference—in the case of exaptation, the given biological property obtains radically new functions, whereas scaffolding describes step-wise development in which earlier stages channel the emergence of the following stages. In exaptation, such continuity is not presumed and very different forms and usages can become related (see a biosemiotic discussion in 140 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Weible 2013). Furthermore, in Gould’s and Vrba’s understanding, the intermediate gap or lack of functionality of the biological form is an essential characteristic of exaptation. In the context of mimicry, we have specified exaptation for the semiotic realm by introducing the concept of semiotic co-option, where existing conspicuous meaning-relations in nature create conditions for new forms to emerge (Maran, Kleisner 2010: 196): Semiotic co-option, as we understand it, focuses plainly on remodeling through meaning relations, that is, on cases where a new function and an evolutionary path is given to an existing structure through the attribution of new meaning. There are several attributes that are specific for that type of reuse: 1) objects of semiotic cooption are structures of animal appearance (in a wide sense, as all perceivable forms, sounds, smells, movements etc.); 2) semiotic co-option is based on previously existing meaning complexes and operates in the framework thereof [...]; 3) semiotic cooption establishes meaning relations and connections throughout the biological realm (from intraspecific relations to interspecific ecological relations); 4) in semiotic cooption structures are objects of more specific targeted shaping and tend to have limited variability based on existing meanings. It can be said that if scaffolding is a step-wise change in a biological system, and in exaptation existing structures obtain new functions, then semiotic co-option (Kleisner 2010, 2011) takes advantage of the existing meaning complexes of the biosemiosphere and uses these to foster the development of the given biological structure. Mimicry could also be considered as an important step in cognitive scaffolding, namely as an aid for certain psychological properties and capacities to develop. In making this claim, we would focus on the position of the receiver in the mimicry system, who is constantly encountering confusingly similar mimics and models, which, at the same time, have different meanings or applicability. If the receiver is able to recognize some mimics correctly as mimics and make use of them—for instance, if a bird recognizes some hoverflies as “not wasps” and eats them—then this usually means eliminating these individuals and their genetic information from the population. The mimics that more closely resemble models will 141 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. be preserved. Inasmuch as the receiver’s feedback of correct or erroneous recognition initiates change in the following generations of mimics (and models), the receiver as a species is not able to get rid of the confusing similarity on an evolutionary timescale. The receiver is capable of shifting the characteristics of mimics (and models), but in most cases, the receiver is not able to break the mimicry system of which it is part. Ideally, as much as the receiver’s perceptual and cognitive abilities develop, so develops the deceptive similarity between the mimic and the model, becoming more detailed or complex. From this perspective, the mimicry system posits repetitive cognitive challenges to the receiver, stimulating the development of its cognitive skills to differentiate between the confusingly similar organisms. 27 It needs to be pointed out that mimicry resemblances do not only challenge the perceptual skills of the receiver, but also its ability to memorize certain colours and forms, and their indexical linkage to certain meanings. This is due to the fact that in very few mimicry systems does the receiver actually have a possibility to compare mimics and models in the same timeframe (as it is in the case of camouflage coloration and egg mimicry of brood parasites). In most cases, mimics and models are not simultaneously present and the receiver needs to rely on its memory when encountering a suspicious object. Cognitive capacities of the receiver are tested in different ways in many mimicry systems. This aspect is well exemplified in the concept of satyric mimicry (Howse and Allen 1994; Howse 2013), which points to the mimicry system where approximate similarity in appearance is combined with different behaviour and escape strategies. In satyric mimicry, “imperfect mimics that blend some components of the model’s signal with irrelevant stimuli confuse signal receivers, resulting in better protection than imperfect mimics that do not” (Kikuchi and Pfennig 2013: 300). Howse (2013) discusses the concept of satyric mimicry in the context of mimetic resemblances of hoverflies Syrphidae, saturniid moths, and nymphalid and danaid butterflies. Another interesting example is aide memoire or recall mimicry, a concept that was proposed by Miriam Rothschild (1984). She points out that no one to one resemblance needs to exist between the mimic and model species: “defense is adequate if the aggressor is forced to recall the attributes of such species, or the disagreeable sequela to a previous assault upon them.” (Rothschild 1984: 311). 27 Dalziell and Weldbergen (2016) have recently pointed out that learning can fundamentally influence the evolutionary dynamics of the mimicry system. They note that besides receivers, learning by mimics and even by models (to find ways of reducing mimic’s interference) also has an effect on the mimicry. 142 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Mimicry systems can challenge the receivers to enhance their cognitive skills, thus being a scaffold toward developing better accuracy of perception, greater cognitive competence and overall development of the neural system. Thus it seems reasonable to propose the hypothesis that deceptive resemblances in nature can function as a scaffold towards more complex communication systems and towards greater semiotic freedom (Hoffmeyer 1998, 2010; cf. Kampis 1998). The distinction between physical and cognitive scaffolding in mimicry also relates to the difference between the steadiness of past events and openness to future options for semiosis. To make a paraphrase reference to Charles S. Peirce, the past becomes filled with fixed and habitual structures that can be used in the present as objects for interpretation in learning, thus creating new sign relations and developing novel ways of life for the future. Learning both presumes and overcomes the uncertainty of the environment. 9.3 Evolution of mimicry in the bio-semiosphere I have described different dimensions of scaffolding processes as ontogenetic, evolutionary, physiological and cognitive scaffolding with reference to mimicry systems. It would be more correct, however, to consider these as aspects or dimensions and not as types, due to the fact that ontogenetic, evolutionary, physiological and cognitive scaffolds can combine in different ways in the actual mimicry systems. The understanding of this connecting faculty of scaffolds is also evident in Hoffmeyer’s writings: “semiotic scaffolding systems painlessly bridge the mind-body gap, being in their function as controllers, essentially somatic and social in one and the same process” (Hoffmeyer 2014b: 95–96). Let us return for a moment to the example of the eye-spots in butterfly wings. We saw how ontogenetic scaffolds have an important role in the development of eyespots on butterfly wings. However, eyespots also depend on evolutionary scaffolds. From an evolutionary perspective, the eyespots co-opt to an earlier use of the eyes that have vivid coloration and sign value in the intraspecific communication of many animals and birds (for discussion, see Kleisner 2011; Kleisner and Maran 2014). There has been a recent discussion about whether it is resemblance to the eye or conspicuousness in general that makes eyespots effective as a deterring device (Stevens et al. 2009; Stevens and Ruxton 2014). Here we should note that eyespots may well be involved in both functions 143 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. simultaneously, but it is very difficult to consider eyespots without any reference to the interpretative act (the signification) on the part of many vertebrates, which gives the meaning of “eye” to the visual perception organs. In other words, with their reference to an active animal, eyes function as biological archetypes, and any later similar structure tends to become grounded on the meaning that vividly coloured irises have. In regard to physiological scaffolding, it needs to be mentioned that in many cases, the development of eye-spots is based on the general structure of the butterfly wing-plan and the way in which homological serial development in different segments contributes to the creation of the over-all image. The segmented wing-plan creates conditions for the genes to be expressed in a similar way in multiple neighbouring compartments, which, being influenced by local regulatory forces, create the eyespot or eyespots as a complete visual structure (Hombría 2011; Oliver et al. 2014). Also, the development of eyespots appear to make use of some pre-existing network of developmental genes/program, as for instance ventral appendage development, wound healing or wing margin development were co-opted to function in eyespot development (Monteiro 2014). Stepwise change as described by the concept of scaffolding is an important mechanism for the regulation and development of mimicry systems. Different from the communication system of a single species, mimicry often includes several species through ecological and semiotic relations. Due to the interrelations between these aspects and the recursiveness of the mimicry system, as well as the fact that there are several participants involved, it seems plausible to consider mimicry as a system that scaffolds itself. This means that at least in principle, scaffolding in mimicry may cross the boundaries of species in such a way that the genetic, developmental or ecological property of one species becomes a scaffolding device for another species and so on. The example of brood parasitism of indigobirds on weaver finches, given above, fulfills this criterion. While such examples might not be very common, they demonstrate how effective mimicry is in influencing the open-ended nature of evolution. Let us consider another example of brood parasitism — that of the common cuckoo and small passerine birds (see Figure 8.3, left). Whereas the primary mimicry relation takes place between the eggs and nestlings of the cuckoo (mimic) and those of the host (model, receiver), there is a secondary resemblance present between the body shapes and plumage of adult cuckoos (mimic) and a local hawk species (model) 144 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. (Welbergen and Davies 2011). Secondary resemblance has been described, for instance, between the common cuckoo and the sparrow hawk Accipiter nisus. It is believed to reduce the mobbing behaviour of the host species, as they appear to be less aggressive to hawks than they are toward cuckoos (Davies and Welbergen 2008). Due to the local diversity in both the cuckoos’ host species and the hawks, cuckoos are the subjects of different selection pressures through semiotic means that increase the local variety of the cuckoos (Thorogood; Davies 2013). It also shapes the conditions of their own intraspecific recognition and communication. Mimicry has been shown, at least in some cases, to catalyse the emergence of new species (Mallet et al. 1998). Mimicry can increase the local diversity of communication in mimetic species and create similarities between different species in the region (as in the case of so-called mimicry rings). This kind of local variation and constrained change (partly due to semiotic scaffolds) inevitably brings along changes in species. Chris D. Jiggins (2008: 542) describes the importance of local variation for the speciation of Heliconius butterflies: “It appears that color pattern and color based assortative mating are the first traits to diverge between adjacent populations. [...] Selection for phenotypic convergence due to mimicry is undoubtedly an important force in the evolution of Heliconius patterns.” In addition to Heliconius butterflies, this connection between mimicry resemblance and speciation has been described for instance in poison frogs (Twomey et al. 2014), parasitic indigobirds (Sorenson et al. 2003) and fly orchids (Ayasse et al. 2010). From a semiotic viewpoint, it also needs to be emphasised that mimicry systems are not closed systems, but exist in the rich and open environment of signs. The semiotic understanding of the systems of mimicry, usually described with the triadic representation of mimic, model and receiver, is heuristically useful but still an idealization. In the actual environment, mimicry systems are open; that is, they can occasionally include individuals of many species beyond the fixed model of the mimicry triad. These individuals are, to a certain extent, underdetermined in their interpretations, as they can perceive and interpret the resemblances between mimics and models in many ways. As a most primitive distinction, the receiver can recognise: a) the mimic to be a model (erroneous recognition); b) the mimic to be different from a model (correct recognition); c) mimic to be something resembling a model (recognition based on analogous interpretation); d) mimic to correspond simultaneously to a mimic and a model (recognising two sign sets simultaneously). 145 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Receivers may also make interpretive connections between signs in the mimicry systems and other semiotic structures in the biosemiosphere as principally new analogy-based sign relations. In cases where mimicry is underdetermined and a space of different interpretations becomes existent, the subjective properties of individuals as interpreters start influencing the future of the mimicry system. The notions of semiotic selection and semiotic scaffolding fit well with and support the understanding of mimicry as a two-layered structure. In addition to the ecological composition of the species in predation, symbiosis, and parasitism, mimicry also includes the layer of semiotic processes. Whereas the first layer is composed of specific species and individuals, the second layer consists of characteristic cues of the species or their signals, properties of the environment or more abstract signs. In semiotic relations, different organisms are bound together through perception, recognition, communication and action, and it is in this layer that the organism’s interpretation starts shaping the future stages of the mimicry system. The interplay between the ecological and semiotic layers is important for the dynamics of mimicry as an open system. The combinations of species fixate mimicry in an ecosystem but also open it up to new ecological relations in the surrounding ecological network, whereas the sign processes enable mimicry to relate with existing meaning complexes, generalized images and conventions of communication. 146 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 10 THIRD EXCURSION: AN EPISTEMOLOGY OF THE UNCERTAIN Modern science is generally designed to eliminate uncertainty and ambivalence. There are many theoretical tools and practices in science to distill solid knowledge from the occasional and unpredictable dynamics of the world. Thinking with the aid of theoretical postulates and paradigmatic cores, the requirement for repeatability of the experiments and observations, and conventional use of statistical methods (that restrain minorities and exclude unique entities) are some means that are used to build reliable and solid knowledge in the sciences. Such inclination towards hard data is probably an inevitable aspect of the biological sciences, but it certainly has its cost. Namely, downplaying or ignoring processes that are occasional, unique or uncertain, which becomes problematic in cases where the ambivalence and fuzziness itself are characteristic properties of an object that is under study. For semiotics, or at least for post-structuralist semiotics (in a literary sense, semiotics that has surpassed the limits of structural thinking), this is an essential topic. There are many semiotic processes that rely heavily on the non-determinism or fuzziness of culture or nature. All truly creative behaviours and artistic usages of signs are, in principle, nondeterministic. Miscommunication, misunderstanding and partially shared knowledge play an important role in the dynamics of culture, as they bring along new communication codes and practices. Juri Lotman, an establisher of the Tartu-Moscow Semiotic School, repeatedly emphasised the role of limited knowledge, diversity of languages in culture and the partial understanding deriving from this basis: You could say that, ideally, an identical addresser and addressee would understand each other very well, but they would not be able to talk about anything. […] In normal human communication and, most of all, in the normal functioning of language, a pre-supposition is made as to the initial non-identity of speaker and hearer. (Lotman 2009: 4–5). The same appears to be true for living biological systems—at a fundamental level, they are non-deterministic and their complexity is far beyond our capacity of measurement and computation. For instance, Stuart Kauffman (2012) claims that the 147 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. complexity of living systems (e.g. the number of all possible variants of a biological protein structure) is so enormous that it cannot be based purely on mechanical combinatorics and selection; rather, it is chance, history and local self-organisation that become mechanisms to shape the development and evolution of living structures. Correspondingly, the quantitative approaches work well in research describing relatively simple living systems, but if the complexity rises, then at a certain point, a complete description of the system and its behaviour become impossible. Retaining the quantitative methodology will, in such a case, be achieved at the cost of simplifying the system, e.g. by limiting the system through setting boundaries or by limiting the variability of the system by statistical means. An alternative possibility to study complex living systems would be by using qualitative description and modelling. In qualitative modelling, the object is described with the help of a number of sign systems (languages, registers) to represent the complexity of the model. The complexity and uniqueness of the system can be celebrated through different and partly contrasting descriptions that correspond to the behaviour of the object. These different viewpoints enter into complementary relationship with one-another by providing a multi-faceted and thorough understanding of the object. Such a qualitative model presumes the involvement of the perceiving subject and it connects the object and subject’s viewpoint into a one of a kind experience. Semiotic modelling of life is necessarily unique by embracing both the living system under description as well as the observer with its specific umwelt and cultural context. Another problem with the appreciation of solid knowledge in the sciences is a tendency to argue for discrete types and distinctions on occasions where the object tends to be fuzzy by nature, or its borders ambivalent. This topic was partly discussed in an earlier chapter in relation to evolutionary views of mimicry as a criticism against the tendency to treat mimicry as a distinct phenomenon. To take a more balanced position, I will discuss another example from semiotics, as my concerns are at first methodological and only thereafter connected with the specific subject-matter. There is an old dispute in semiotics over the distinction between matter and living entities and the related criticism of the pansemiotic approach. Pansemiotic thinkers, which is by no mean a unified group, generally consider triadic sign processes to take also part in physical, chemical and cosmic realms of non-living matter (Deely 2001; Taborsky 1999). Most biosemioticians oppose pansemiotism by seeing life and semiosis interconnected, thus following Sebeok’s biosemiotics maxim. According to the 148 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. biosemiotics maxim, there is a semiosic activity in very simple unicellular organisms, but there cannot be semiotic processes without a living interpreter. This is a solid formal position, but at the same time, we appear to run into problems when trying to apply the same distinction in the semiotic studies of ecological processes. Several ecologists have showed from different viewpoints that the demarcation line between living organisms and physical matter is extremely vague. John Turner (2000, 2015) talks about extended organisms in the case of termites and other animals that relocate and shape matter and energy in their surrounding as a part of their metabolic processes. John Odling-Smee (1988) points out in his concepts of niche construction and environmental heritage that organisms are able to make use of the surrounding environment for passing specific information across the generations. Scott Gilbert (2016) demonstrates in the framework of ecological developmental biology how environmental properties (e.g. chemical conditions of the water) can trigger different developmental tracks in daphids, amphibians and other groups. In regard to environmental problems, I have proposed the concept of “semiotization of matter” to indicate that matter is often not neutral, but may include semiotic traces imprinted on it by humans or other living beings (Maran 2014b). An example of this could be the conservation issue of waterbirds (and also crocodiles) caused by leaden shots left on the seashores and waterways by hunters (Martinez-Haro et al. 2011). Characteristics of leaden shots may be triggered in the interpretation by waterfowls who peck the shots instead of pebbles for using these as grit or gastroliths. The result of swallowing shots instead of stones is often serious lead poisoning. This semiotic encounter, which often brings along deadly results, is rather similar to biological mimicry in that it rests upon an inability of the organism to recognize and correctly categorize the matter semiotized by other species—in this occurrence, by humans. My case here is not to defend the pansemiotic view, but to point out that biosemiotics also holds potential to problematize the organism-matter divide. This is a paradoxical outcome of Sebeok’s maxim: if we define a living organism through its semiotic activity and take relationality of the semiosis seriously in Peirce’s sense, then the borders of life become dissolved. Although the distinction between the living organisms capable of semiosis and passive matter appears to be formally important for biosemiotics, the same distinction does not appear to work well in many practical applications in semiotics. Biosemiotics and especially ecological semiotics would benefit from more a subtle epistemological position that would allow tracing semiotic 149 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. processes that surpass the limits of different domains (as between living and nonliving, human and animal, culture and nature), or in which the fuzziness is an intrinsic property. We saw in an earlier chapter that probable mistake or ambivalence is an important property of mimicry. This is true for the receiver, who could mistake a mimic for the model, but also appears to be a systemic property of mimicry systems. From an evolutionary perspective, mimicry balances on the borderline between perception and non-perception, or between interpretation and misinterpretation. I have also demonstrated how mimicry studies could lead to the description of a new sign type—an ambivalent sign—where two similar signs are entangled together because of the receiver’s activity. Therefore, we can state that ambivalence is an essential characteristic of mimicry and would thus require full attention. Based on these arguments, I would like to point out the necessity in biosemiotics to develop approaches for working with uncertain and ambivalent objects. This requires not only acknowledging the existence of such objects, but also shifting the epistemological position and methodological approach, as well as developing conceptual tools that would be suitable for describing such objects. In biosemiotics and in zoosemiotics, the need to take uncertainty seriously arises from the otherness of the object due to the inclusion of the animal’s Umwelt, which is always partly different from our subjective worlds. Such semiotic duality or hybridity is most evident in human-animal interactions, as part of animal meaning making activities remains beyond our perceptual reach (cf. Magnus 2016). In cultural semiotics, the necessity to take into account the subjectivity of the objects is also recognised. This derives from the fact that cultural objects are language-based and their symbolic character gives ground for their own subjectness and potential for dialogue. In such a case, an open epistemology and dialogue-based research methods would allow the semiotic potential of the cultural object to unwrap itself. Peeter Torop, a leading cultural semiotician of the Tartu Semiotic School, has underlined the necessity of being responsive towards the research objects and the importance of adhoc approaches. “Culture as an object of analysis often dictates its analysability, and therefore ad-hoc theories as object-based theories have a special place in the disciplines that study culture” (Torop 2009: 38) Ad-hoc in this sense is not considered to be an aid to support an otherwise unsteady theory, but in a positive sense, as a methodological stance that uses temporary conceptual tools to develop a framework, 150 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. within which the semiotic activity and dynamics of the object itself would unfold. Adhoc approaches should not be self-sufficient, but should leave room for explorative and speculative thinking. For biosemiotics as well, using ad-hoc approaches appears to be a very reasonable methodological choice in cases where the research object is complex or has subjective properties. In regard to living systems, uncertainty is as important as knowledge, although the former is largely ignored and down-played in modern philosophy and science. 151 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 11 FROM ABSTRACT MIMICRY TO ECOLOGICAL CODES Developing a broader ecological perspective is both a big challenge and a necessity for biosemiotics. Without an ecological account, biosemiotics as a paradigm would remain incomplete. Furthermore, the semiotic approach could in turn offer a fresh perspective to the natural sciences for understanding ecological processes. In 1981, system ecologists Bernard C. Patten and Eugene P. Odum described an informational layer in the ecosystem with a local regulatory capacity, without which the ecosystem would fall into a mass of chaotic processes. In 2007, theoretical ecologist Søren N. Nielsen proposed that this sphere of semiotic functions in the ecosystem could be called semiotype, referring to the parallel with genotype, phenotype and envirotype. Kalevi Kull, in his several writings (Kull 1998, 2008, 2010), has expressed the view that the ecosystem is semiotic in its nature, and that semiotic processes have much to do with the integrity of ecosystems. To analyse what the effects could be of mimicry as a semiotic phenomenon on broader ecological communities, we would need to take a viewpoint that foregrounds semiotic processes in the ecosystem. One starting point for such a view could be Jakob von Uexküll’s “Theory of Meaning” (1982), where he describes relations between animals as correspondences between their body plans and Umwelten through counterpoints of meaning. The different Umwelten are mediated by functional cycles, where animals obtain the positions as meaning utilizers and meaning carriers for each other through their perceptual and effectual activity (cf. Uexküll 1926: 270–280). Relations that are described in classical ecological theory as predation, parasitism and herbivory, are deducted in Uexküll’s work to be qualitative relations of meaning, where animals attribute specific meanings to one-another. At the ecological level, such cycles of qualitative meanings can be considered as minimal regulatory hubs that make the connection and feedback between different parts of the ecosystem available. In addition to direct physiologically-based correspondences of meaning between two species, Uexküll observes interspecies communication as mediated by meaning carriers which are distinct from the animals’ body, such as the squeaking sound standing for a bat in the moths’ Umwelt (Uexküll 1982: 56), as well as a completely distinct organism who acts as a meaning carrier. With respect to mimicry, Uexküll mentions two examples: the angler-fish Lophius piscatorius who uses a long and movable appendage to lure prey fish, and butterflies that carry colourful eye- 152 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. resembling spots which scare off insectivorous birds. Uexküll sees these examples as extensions of meaning rules that organize forms in nature. The form of the prey is, in these cases, not directly connected to the form of the predator, but correspondence is achieved due to some other image or shape-schemata present in the animal’s Umwelt (Uexküll 1982: 58–59). Thus Uexküll’s “Theory of Meaning” opens up a significant aspect to understand ecological relations, which could also be considered as a biosemiotic ground for interpreting mimicry. Namely, ecological relations in general can be based on or regulated through some image-schema that connects different animal Umwelten. Correspondingly, the common description of mimicry as a resemblance between two species covers only limited cases among many possible mimetic similarities. In addition, there may be images that do not have any direct or strong relation to any specific physical form, but can still mediate relations between animals. In such a case, it is the animal itself that attributes meanings to a variety of different objects or situations that match the character of the meaning. Such complexes are not limited to the visual sense, but can also relate to many signals, such as auditory communication (hissing, alarm calls) and other modes of communication. In principle, my main argument is that on some occasions, meaning complexes that have gained relative independence from the underlying physical forms can themselves become entities that organise the relations in ecosystems, and can also act as factors in the evolutionary process. Paying attentions to such meaning complexes is a relevant topic for ecological semiotics as it allows for a better comprehension on the semiotic regulation of ecological communities. In this chapter, I analyse possibilities for this kind of semiotic phenomena to emerge by drawing on two notions: abstract mimicry and ecological codes. In the final part of the subchapter, I make a parallel between ecological codes and Jungian archetypes to emphasise their analogical image-based nature as well as the connection between ecological codes and biological universals in human culture. 153 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 11.1 Abstract mimicry: When the meaning comes first28 The most general level of abstraction in mimetic messages is probably present in warning displays. In his book “Defence in Animals. A Survey of Anti-Predator Defences” (1974) Malcolm Edmunds describes these behavioural patterns under the term ‘deimatic behavior.’ For example, cryptic stick-insects such as Pterinoxylus spinulosus and Metriotes diocles expose colourful areas on their wings when fleeing because of disturbance (Edmunds 1974: 153–155; cf. Edmunds 1976). The sudden emergence of glaring colours will cause the predator to stop and seek more information about the situation, showing signs of unpredictability (model), thus giving the insect enough time to escape. This kind of deimatic behaviour could be considered mimicry, as stick-insects do not hold any chemical toxins or other defences that would justify their vivid signalling. Another well-known defence strategy that uses abstract resemblance is the behavioural adaptation of many reptiles and amphibians to make themselves appear larger in the presence of danger. For example, upon noticing a snake, the common toad Bufo bufo lifts itself up from the ground and emits strange growling sounds while simultaneously becoming bloated from the inhaled air. The same type of warning display is common in many lizards (e.g. Brodie 1977; Bustard 1967). In the case of body-lifting in toads or lizards, the object of imitation (model) may be related to the categorisation of organisms in the receiver’s Umwelt. By raising itself up from the ground, a toad shifts itself in the receiver’s eye away from the sign complex corresponding to prey animals and closer to such animals that are too large to catch or that may even be dangerous. In the case of flash marks in stick insects, the object of imitation is not so obvious at all. To rationalise this, we first need to accept that there are sign complexes in nature that relate primarily to meanings and only secondarily to physical features. Quite often, such abstract and fuzzy resemblances are not considered to be true examples of mimicry. Georges Pasteur excludes the previous cases from his speciesbased classification on the grounds that “the model is not an actual species” (Pasteur 1982: 191) and describes them under the terms “semi-abstract and abstract homotypy.” As I have indicated before, the question of resemblance to a model not 28 This subchapter is partially based on Maran, T. (2011). Becoming a sign: The mimic's activity in biological mimicry. Biosemiotics (Springer), 4(2), 243–257; and Maran, T. (2007). Semiotic interpretations of biological mimicry. Semiotica (DeGruyter Mouton), 167(1/4), 223–248. Used with permissions. 154 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. belonging to any concrete species may actually go deeper than just classificatory issues and pertain to a common biological understanding of relations between species, which focuses on physical forms and properties and largely ignores perceptual features and meanings for the animals themselves. We can revert to Pasteur’s description of abstract mimicry by stating that in real nature, the model is seldom an actual species, as “species” as such is a human taxonomical construct and not present in the Umwelten of receivers. Cases where the model can be determined down to the specific species or object should rather be considered as special cases and not as a general condition in mimicry. The major difference between abstract and semiabstract mimicry lies in the generality of meaning on which the imitation is based. In abstract mimicry, certain universally perceptible features, for example a change in the situation, unexpected movement, or indications of possible danger, are expressed, whereas in semiabstract mimicry, the model can be related to a group of organisms or its features, but not to a concrete species. Being based on the expression of meaning, resemblance in semiabstract mimicry between species participating as mimic and model remains approximate, and if many species are involved, then the resemblance may vary to a remarkable degree. A good example of a semiabstract sign complex in nature is ‘snakeness.’ Rather than being connected with a representation of a particular snake species, the meanings such as dangerousness, poisonousness and lethality relate to a complex set of snake-specific features, such as vermiform body shape, specific reptilian type of movement on the ground and hissing sounds. The meaning complex of ‘snakeness’ can be considered a biological universal or archetype, as the practice of avoiding snakelike features is common to many different animal groups. Therefore it also becomes useful for many harmless species to adopt a snakelike appearance and perform snakelike movements and sounds. Snakelike behaviour is performed by various organisms, for example the chicks of woodpeckers, storks and wrynecks; large caterpillars with eye-spots of many genera as well as lizards and eels. Hissing sounds are used for deterring potential enemies by various snake groups (Aubret and Mangin 2014), warblers (Sibley 1955), bees (Moushumi et al. 2002) and bumblebees (Kirchner and Röschard 1999). Krams et al. 2014 show with an experimental study that hissing calls of the female great tit (Parus major) prevent predator attacks and can increase the survival of the incubating females or their offspring. The abstract meaning complex of ‘snakeness’ also has a role in human culture; for example, in 155 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Estonian folklore, the legless lizard slow-worm Anguis fragilis has been considered to be a very poisonous and even magical creature in some places. Another well-known example of a semiabstract meaning complex is related to eyes. Visible eyes are a characteristic of most vertebrates. According to the common biological understanding, the perception of round eye-like objects can give two kinds of information to the receiver. First, it can be a straight indication of the presence of a large and active animal. And second, it can give information about the position of this animal, indicating the location of the head and thus allowing it to make conclusions about the direction of movement as well as the placement of the animal’s vital body parts. In mimicry, accordingly, two kinds of eye imitations have been described. First, many butterflies, moths, caterpillars, frogs, fishes and other small animals have large round eye-like areas on their body surface—a feature which is often combined with the behavioural adaptation to demonstrate these eye-marks in cases of disturbance. And second, many fast-moving animals such as butterflies or fish have smaller eyespots on their wings or fins, which are considered to confuse the predator and direct the attack away to the nonvital body areas (Blest 1957; Stevens 2005). Eyespots appear to be especially diverse and abundant in tropical rainforests, in which case the specific mimic-model relations is very difficult to construe (Janzen et al. 2010). Eyespots belong to the category of semiabstract mimicry, as it is not possible to point to the exact species of the model. Rather, eyes are imitated as abstract objects on the basis of their conspicuousness, common meaning and function in nature. There seems to be, however, little justification for drawing a sharp border between mimicry and non-specific resemblances (or between abstract and semiabstract mimicry). There is a functional connection between the threat display of eyespots in moths and the anti-predatory behavioural displays of the toads, and an attempt should be made to describe these within the same theoretical framework. From a semiotic viewpoint, I would rather argue that the object of resemblance is related to the structures present in the receiver’s Umwelt. Thus, in the example of eyespots, it is not owl eyes that are the object of imitation but the insectivorous bird’s mental image (or search image) of its natural enemy’s characteristic marks. By definition, the mental image is a generalisation of eyes of various predators that a bird has encountered or could potentially encounter in its habitat. One way to evaluate a novel theoretical approach is to consider whether it helps to rationalize some problematic phenomenon better than traditional explanations 156 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. do. It may be that the concept of abstract mimicry and the corresponding biosemiotic view can help us analyse cases in which the similarity between the mimic and model species is approximate or diffuse. An example of such ‘imperfect mimicry’ (Sherratt 2002), characteristic of Holarctic, is the combination of yellow-black warning coloration of many Hymenoptera (wasps, bees, bumblebees) and their imitations on many levels of exactness by hover-flies Syrphidae, as well as some moths, beetles, dragon-flies and other insects. Most authors regard imperfect mimicry as some deviation from the ‘normal situation’ of drive toward absolute similarity. Such approaches seek to explain imperfectness with specific environmental conditions or ecological relations, or try to find some other factor that would compensate for deviation. Having reviewed a large amount of science literature, Francis Gilbert has distinguished ten directions of argumentation for solving the problem of imperfectness in hover-fly mimicry: 1. The occurrence of mimicry: denial that poor mimics can be mimics at all; 2. The different perception of the predator: poor mimics appear perfect to bird predators; 3. Poor mimics match the Müllerian complex rather than Batesian mimicry (because of flight agility or weak unpalatability); 4. The wasp models of poor mimics are exceptionally noxious, so mimics do not need to be perfect; 5. Mimics of different size have different predators (poor mimics who are smaller are hunted by invertebrates with poor vision); 6. Poor mimics compensate their visual discriminability with some behavioural adaptation toward predators or other mimics; 7. The effect on the predator: poor mimics confuse rather than deceive; 8. The speed of evolution: poor mimics are still evolving their mimetic resemblance; 9. Disturbance by man: poor mimics have recently become abundant, causing mimetic degradation; and 10. Selection for perfection is opposed by other forces (kin selection; the existence of multiple models; cost for producing perfect mimicry patterns) (Gilbert 2004; see also Edmunds 2000). Most of these explanations are based on the assumption that the object of imitation is the model species and that the mimicry should normally progress towards maximizing the similarity. From a biosemiotic perspective, we can make a principally different suggestion: hover-flies do not imitate any concrete species, but rather a certain combination of colours that have the meaning of unpalatability or danger for a large group of animal receivers. In other words, the attention of the receiver is focused on the relations between the insect and the conspicuous colour pattern with its possible meaning, not on comparing different insects and typifying these. Richard I. 157 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Vane-Wright exemplifies the nature of such identification by using the example of two unpalatable species. “An insectivore which has learned to avoid viletasting lycid beetles by their black and yellow colour may also avoid similarly coloured pompilid wasps, even on the first encounter. If so, it makes this decision through identifying, on the basis of its similar black and yellow signal pattern” (Vane-Wright 1980: 3). In such a case, it is not the exact resemblance between the wasp and the beetle that becomes decisive, but whether they expose their common colour pattern recognizably enough and whether the receiver is familiar with the meaning of the pattern. According to the semiotic explanation, different model species in Müllerian complex also do not need to be similar to each other, but rather have to be identifiable as species that carry the ‘colors of uneatability.’ Let us consider in parallel the human ability to recognise different letters in the alphabet. If we know the meaning and phonetic equivalence of the letter “a”, the font or emphasis used to display it: a, a, a, a have secondary importance and all single instances still have the same meaning for us. If the recognition of different insects for the birds is not form-based but meaning-based we can assume the same logic also exists in biological mimicry. A similar explanation of imperfect mimicry is given by Miriam Rothschild, who speaks of aide mémoire mimicry—resemblances to noxious or dangerous prey that have evoked earlier unpleasant experiences in the receiver (Rothschild 1984). The problem with relying on strict formal features is that in actual environmental situations, they may be unreliable. The perceptual accessibility, behaviour and environmental context varies from encounter to encounter, and animals still need to recognise, categorise and adequately react in these non-stereotypic situations. The phenomenon of abstract mimicry conveys and thus highlights some properties of semiotic structures in nature that could be significant for a semiotic interpretation of ecological systems. First, sign structures in nature seem to operate on different levels of generality of meaning. This is enhanced by the existence of interpreters with different body structures, sense organs, activities and Umwelten in nature compared to the uniformity of humans. There are specific meanings that are present only for some species in their Umwelten, and there are also more general meanings that are shared by various organisms. Related to this issue is the question of association between physical form and meaning that can vary from case to case in nature. There appear to exist semiotic structures in which form and meaning are 158 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. tightly bound together and are inseparable, but there also exist meaning complexes that have become quite independent, capable of inhabiting nature’s many forms of different descent and properties. The more general and independent semiotic structures could also be expected to have an active role as agents in evolutionary processes. 11.2 Connecting Umwelten, sharing codes 29 In this subchapter I discuss ecological codes as a possibility for comprehending meaning structures in nature that surpass the limits of one species and thereby organise species relations in ecosystems. Kalevi Kull has suggested ecological code as one among four concepts relevant for developing semiotic ecology (others being consortium, Umwelt and biophony, Kull 2010). Kull states (with reference to Russian ecologist Alexander P. Levich) that: “ecological code [...] can be defined as the sets of (sign) relations (regular irreducible correspondences) characteristic to an entire ecosystem, including the interspecific relations in particular” (Kull 2010: 354). It is, however, not entirely clear what the relationship between “the sets of (sign) relations” and local semiotic processes is, or to put it in other words, how exactly semiosis and communication can take part in large-scale ecosystem regulation. My initial suggestion is that the properties and functioning of codes on the ecosystem level are rather different from the ways in which codes regulate human communication or any other intraspecific communication. Codes and coding are a frequently discussed topic in biosemiotics (e.g. Barbieri 2008; Sebeok 1972; Hailman 2008). This tendency can be seen as a residue of linguistic heritage in biosemiotics (Maran 2010c; Cobley 2014). The understanding of the concept of code has been derived from the studies of human communication; more specifically, a special type of human communication—linguistic and technically mediated communication, which is thus an idealization even in the context of the human species. In animal communication and especially in interspecific relations, codes cannot act in a similar manner to human linguistic codes. This is so first because of the lack (or minimal presence) of lexical syntax in animal sign systems, 29 This subchapter is partially based on Maran, T. (2012). Are ecological codes archetypal structures? In T. Maran, K. Lindström, R. Magnus, M. Toennessen (eds.), Semiotics in the Wild. Essays in Honour of Kalevi Kull on the Occasion of His 60th Birthday. (pp. 147–156). Tartu: Tartu University Press. Used with permissions. 159 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. which bars the use of code and coding in the specific sense of combining elements of sign systems to produce new meanings. According to another possible interpretation, a code is a system of correspondences between messages and their significance or behavioural outcomes (Sebeok 1991a). But here emerges a problem: messages in animal communication are not as systemic as in human language. For instance, for birds it is not one and the same code that operates in the phatic calls of migrating flocks and courtship songs, as these utterances cannot be used in the same spatiotemporal context. Basically, if there is no choice between alternatives for the animal, then there is nothing to code. Especially problematic are instances of interspecific communication, such as warning coloration, mimicry, communication in symbioses, etc. A prerequisite for the concept of code appears to be that it is shared by the participants of communication. In interspecific communication, this requirement is generally not met. For instance, in the warning coloration of the ladybird Coccinella sp. there is no shared code between the insect and the insectivorous bird, as the ladybird does not have perceptual access to the link between its red-and-black warning pattern and unpalatability. Therefore, when using the concept of code in regard to interspecific communication that is based on ecological relations, certain concessions need to be made about its scope and applicability. To theorise on the properties of ecological codes, a few premises should be accepted. First, different species live in their specific Umwelten and do not have direct access to one-another’s sign systems. They also generally lack access to the meta-level of communication that would allow perceiving and communicating about the rules of the sign system apart from the specific usage of messages of this sign system. One way to proceed from here would be to assume that animal communication has the capacity to self-arrange in a way that messages emitted by single individuals become organised at a community level. This appears to be the case in birds’ morning chorus, when individuals look for the pauses to start their song which results in a “coordinated interspecific chorus” (Malavasi and Farina 2013). In many other cases, it is plausible to assume that codes on the ecological level are not formal or of strict regulations, but rather ambiguous and fuzzy linkages based on analogies and correspondences. In my understanding, the basic properties of the ecological code can be proposed as following: 1. Ecological codes are distributed and open. Ecological codes involve different species, for some of which the involvement being obligatory and for some, 160 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. occasional. The involved species have different perceptual organs, Umwelten and relation to the environment. Therefore, no single individual or species has full perception of an ecological code. Instead, an ecological code forms as the sum of memories and experiences of corresponding perceptions. Every single species and organism involved in an ecological code has a partial variation of the convention. Having once emerged, an ecological code is open to new species becoming involved. 2. An ecological code is built upon and incorporates the consistencies, constraints and habits existing in a particular ecological community. An ecological code rests on indexical relations, as it is in these that representamen—object relationships surpass and remain independent of any specific interpreter. An ecological code also relies on the habitual semiosis, behaviour and action of animals. 30 With regard to living agents and environment, ecological codes are communal and disperse. Cognitive capacities of organisms (i.e. semiotic thresholds) act as gateways to an ecological code but do not include or determine the content of the code. 31 3. An ecological code uses different memory types (following Jablonka and Lamb 2005), that is, an ecological code has both cognitive and non-cognitive (or conscious and unconscious) aspects. A regulation can be fixed in different memory types simultaneously: for instance, it can be fixed partially in physical regularity, partially in the genetic memory of a species, and partially in the cultural memory of another species. For the regulation to become effective, the different types of memory need to come into contact and operate together. To sum up the three proposed characteristics, ecological codes do not resemble human linguistic codes or algorithms, but are rather like archetypal imagery32 or patterns—dispositions in animals to establish certain types of meaning relations in ecological communities and to link sign processes with actions in a particular ways. Ecological codes are distributed and without a fixed centre or structure. There appears to be, however, different bases from which ecological codes 30 The first possibility is expressed in John Maynard Smith’s concept of indices (Maynard Smith, Harper 1995: 306), the second in Hoffmeyer’s concept of semethic interactions (Hoffmeyer 2008a: 189). 31 This is so because of the third property: ecological codes use different memory types including evolutionary regulations. In some cases, semiotic thresholds can also be bypassed or counterfeited. The story of Clever Hans is, among other things, an example of how limited cognitive capacities do not restrict an animal from becoming involved in complex semiotic phenomena. 32 I use the concept of “imagery” here to stress the analogical fuzzy nature of ecological codes. 161 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. depart as well as different functions that they have (see table 11.1). In general, we can distinguish: 1) distribution codes, where animal activities and communication through the process of self-assembly organises the spatial and temporal organisation of the animals; 2) identity codes, where the ecological code is centred on the species or group that has significance or that it is charismatic to a broad number of species in the given ecological community; 3) symbolic codes, where the code is centred on the specific patterns of colour (or other modality) that have a shared meaning for the number of species; and 4) archetypic codes, where the ecological code is centred on the meaning relation that is valid for a broad number of different species due to general physiological, ecological or behavioural constitution of the organisms. Table 11.1. Common types of ecological codes Type of ecological Basis code Function Examples Distribution codes Self-assemblage of Organising the animal distribution and activities movement in space or time Acoustic codes, birds’ morning choir, animal tracks in landscape, territorial arrangements Identity codes Semiotically dominant species or group Representing or misrepresenting identity Myrmecomorphia, hissing signals, snake body shape. Symbolic codes Shared patterns Signifying specific meaning Aposematic patterns (e.g. yellow-black stripes) Archetypic codes Metonymic generalisation Conveying properties, actions or intentions Thanathosis, kindchenschema, eyespots, deimatic displays In her excellent paper, philosopher Annabelle Dufourcq has recently discussed my notion of ecological codes by pointing out that expressing “meaning in animal life as simply fuzzy or vague, or even archaic” is done “from an exclusive rationalist perspective that can only define what is beyond it in a negative privative and pejorative way as not completely clear, not rigid enough” (Dufourcq 2016: 61–62). She further points out that in this context, the link between animal studies and ‘the 162 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. imaginary’ can appear as a handicap as it could restrict the fuzzy meanings to become objects of scientific investigation. The way forward, as Dufourcq suggests, would be to focus on the ubiquitous mode of being in animals, where animals’: “do not possess the mode of being of an innert, self-contained, well-circumscribed thing: [but] they consist in an open meaning” that is a characteristic of subjects, thus being a property that would allow animals to be not just objects of thinking, but also as subjects that can participate in creative imagination; that is, to provide new images and fantasies to human culture (Dufourcq 2016: 61–62). Dufourcq’s comments allows me to specify some aspects of ecological codes as I understand them. On the one hand, I also consider the relationship between image-based (iconic) communication and creativity to be an important aspect to consider in animal communication. Iconic communication is necessarily qualitative and analogous, and not regulated by communicational codes. As also demonstrated by mimicry studies, images can easily cross borders of different sign systems, cultures and even species. On the other hand, my attention to the fuzziness and ambiguity in animal communication is primarily on the object level. This is not about arguing for or against the fuzzy status of animals in human discourse or blurring the humananimal distinction, but I do indeed want to make a strong ontological claim that there are many instances of animal communication in ecosystems where distributed and ambivalent sign processes play a crucial role. This understanding comes from a few relatively simple observations: the iconic mode of communication is much simpler and evolutionarily more wide-spread then syntax-based communication; in iconic communication, variants of a more general image-schema are often used, whereas the correspondence between variants and invariant (token-type relation) is approximate and not absolute. On this basis, my attention to conceptual and epistemological questions has practical grounds. It is rather evident that abstract mimicry and other similar fuzzy phenomena have been excluded categories (described through exclusion form mimicry typology, e.g. Pasteur 1982) in biology. Here, mimicry studies would indeed benefit from a conceptual shift that would allow taking such ambiguous phenomena more into the spotlight. 163 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 11.3 Ecological codes and archetypal structures 33 Distributed over many species, archetypic codes appear to have some similarity to conventions. Yet they are not cultural conventions as in the case of linguistic signs and cultural symbols. Rather, we can say that ecological codes are natural conventions based on the fact that many species have similar entities or meaning relations in their otherwise different Umwelten. Ecological codes are rooted in the structural and functional similarities between different animal physiologies (e.g. kindchenschema as a general characteristic of juvenile mammals) as well in the fundamental similarities in the physical world (e.g. gravitation determining the upward-downward axis and light coming from above, initiating counter-shading and many other principles of animal coloration). There appears to be some parallel between the ecological codes as described here (and especially archetypic ecological codes) and the notion of archetypes in the humanities as it derives from the works of Carl Gustav Jung. Jung writes in “The Archetypes and the Collective Unconscious” that: “the archetype is essentially an unconscious content that is altered by becoming conscious and by being perceived and it takes its colour from the individual consciousness in which it happens to appear” and that “so far as the collective unconscious contents are concerned we are dealing with archaic or [...] primordial types, that is, with universal images that have existed since the remotest times” (Jung 1981: 4–5). Further, Jungian scholar Andrew Samuels presents a list of main features of the archetype theory: (a) archetypal structures and patterns are the crystallisation of experiences over time, (b) They constellate experiences in accordance with innate schemata and act as an imprimatur of subsequent experience, (c) Images deriving from archetypal structures involve us [animals] in a search for correspondence in the environment. (Samuels 1986: 22) 34 33 This subchapter is partially based on Maran, T. (2012). Are ecological codes archetypal structures? In T. Maran, K. Lindström, R. Magnus, M. Toennessen (eds.), Semiotics in the Wild. Essays in Honour of Kalevi Kull on the Occasion of His 60th Birthday. (pp. 147–156). Tartu: Tartu University Press. Used with permissions. 34 My replacement in square brackets in the quotation points to the essential difference between Jung’s archetypes and ecological codes. Jung’s theory is originally aimed to describe the psychological content of the human species, whereas in the study of ecological codes, archetypes should be widened to include Umwelten of other animals as well as interspecific semiotic and ecological relations. 164 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. These charactersistics appear largely suitable for describing ecological codes. Similar to Jungian archetypes, ecological codes are collective or connecting different living beings, and similar to archetypes, ecological codes are fuzzy and general, being able to take various forms in specific animal Umwelten. Thus I would like to explore in the present chapter, whether ecological codes are archetypal structures, and what this connection can tell us about semiotic processes in ecosystems. I am aware that referring to Jungian psychology in the context of contemporary biosemiotics and mimicry theory is walking on thin ice, so we need to proceed cautiously here. Apparently, there are unsurpassable differences between these domains, for instance the concept of “collective unconscious” not being readily applicable in biosemiotics. Jung’s understandings of specific archetypes and their use to interpret human dreams is much too speculative for being applicable in biosemiotics. Let me state clearly that applying the archetype concept in analysing ecological codes needs to be done without including any causes (supernatural or other) to the argumentation that remain beyond the reach of our discernment and analysis. At the same time it appears that the notion of archetype can be reinterpreted for biosemiotics in a way that it could elaborate our understanding of interspecies communication. What is fascinating about the archetype concept is its radical difference from any language-based or logical content of the mind. When trying to analyse semiotic processes of other species, most available concepts (language, code, communication) are inevitably encumbered by anthropomorphic or sociomorphic modelling. For reinterpreting Jung’s citation given above in biosemiotics terms, we could replace “collective” with “interspecific” and “individual” with “speciesspecific”; we could take “unconscious” in a Sebeokian sense, as a reference to many non-linguistic layers in the semiotic self; we could interpret “altering” and “taking colour” in an Uexküllian way as references to the conditions of the Umwelten of specific animal species. In such an interpretation, the archetype notion denotes the relationship between an invariant or common theme and its expressions in specific animal Umwelten. My rephrasing of Jung’s definition of archetype given above would be the following: The archetype is a content of the pre-linguistic modelling layer of the organism that is altered by becoming actualised and expressed. The archetype takes its Umwelt-specific form from the type of mind of the species in which it happens to appear. 165 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Making a connection between Jungian psychology and biosemiotics may appear more justified when one considers that there already exists a tradition in theoretical biology that has found inspiration in Jung’s works. Swiss zoological philosopher Adolf Portmann refers to Jung, interpreting, for instance, rituals and instinctual life of higher animals as an archetypal imprint (Jacobi 1959: 41). Czech historian of biology and polymath Stanislav Komárek specifies that “the psychological content of animals is composed of archetypes—think only of the [...] Lorenz baby schema or animals’ innate images of predators” (Komárek 2007: 22). Uexküll’s use of the archetype concept (Urbild) is quite similar as it refers to the subtle image-based linkages between different species (e.g. spider and fly). Uexküll’s archetype concept does not appear to have, however, a direct connection to Jung’s archetype (Archetyp), but derives probably from a much earlier use of primal image by W. Goethe (Amrine 2015: 51). 35 Uexküll’s archetype is closely related to his concept of an animal’s general meaning plan (Bedeutungsplan), but it appears to be limited to specific image-based linkages between two related species. The concept of seme, proposed by Karel Kleisner and Anton Markoš (2005) to decipher a certain type of mimicry in which the characteristic signs of a charismatic species (for instance, ants) are imitated by several symbionts, can also be interpreted as a special kind of archetype. Kleisner and Markoš write: “Seme should be understood as a sign originally developed by one species or group of organisms and consequently extended to the other often unrelated groups that were able to receive (or imitate) and built it up on their bodies or environment” (Kleisner and Markoš 2005: 218). In a seme, it is predominantly one species that defines images or signs that are later used in interspecific communication (therefore I would list seme under the identity codes in the table 11.1). But there are many other examples in which the origin and placement of a sign complex is not so clear, and which can be considered to be true ecolocical codes floating in the intermediate semiotic space between participating species. Most likely, one of the strongest archetypes in interspecific communication is related to fear. After all, all animals in ecological networks are in danger of getting 35 Louise Westling (2016), a specialist of Maurice Merleau-Ponty’s philosophy, has recently suggested how the concepts of ecological code and archetype could be actualized for reinterpreting human-animal communication. Historically, there is an interesting thread of thought stressing the role of general images in sense making that runs from C.G. Jung to Adolf Portmann to Maurice Merleau-Ponty. 166 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. consumed by some other animals, and therefore anticipating and perceiving signs of potential predators are vitally important. Signs of fear cannot be rigidly defined (except in close co-evolutionary relations), as there exist various possibly dangerous species, communication contexts are always different and hence signs of fear remain ambiguous. We can distinguish general characteristics of the fear archetype, such as unfamiliarity, unexpectedness and a sudden change or movement; and specific characteristics that are preferably used in certain relations and animal groups. Specific characteristics can include: the image of eyes, fangs and other means of attack, large body size, low and loud sounds, and fast-moving shadows. For many ground-living mammals, for instance, an essential part of the fear archetype is a “shadow from above” 36. This sign relation can also be put to practical use, as I have found out thanks to the wisdom shared by an old nature observer. To avoid confrontation with an angry dog or another mammal of a similar size, waving a jacket or some other garment above one’s head is an effective strategy. Seeing a fast-moving shadow above, the animal develops an irresistible urge to hide, since a fast-moving shadow is a sign standing for an aerial predator in most mammalian Umwelten. Fear is a biological universal, and most higher vertebrates show signs of fear on some occasions. Thomas A. Sebeok has used the concept of anxiety to denote the same idea—anxiety is an induction device that is activated when some cue of danger is perceived in the environment; the purpose of anxiety is to increase the probability of the continuation of the organism’s self (Sebeok 1991c: 39). Although the majority of organisms have specific natural enemies with their characteristic features and signs, judgement about the nature of threat can often be done after the encounter post factum. Therefore some sudden move, flashing colour, shadow from above, quick unexpected change in the environment are often the only signs of a life-threatening encounter that require quick reaction and withdrawal. Impreciseness is exactly the characteristic of this meaning complex—rather than being based on some specific signs, threat displays are fearful exactly because they are vague, unfamiliar and unexpected. They do not represent any specific model organism or environmental element but the potentiality of the receiver to be attacked, to become eaten and its fear of that. 36 See also Jakob von Uexküll’s example of the “shadow from above” in the sea urchin’s Umwelt (1992: 345–346). 167 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. Signs and properties that accompany the archetype of fear are employed by nature in phenomena known as deimatic displays, as discussed earlier. The human version ofthe archetype of fear can be seen in a gamut of signs that are used in literature, art and movies to depict monstrous creatures. Scales, mandibles, fangs, slimy skin, hissing and other properties of reptilian or insect origin are not meant to trick our cultural knowledge but refer back to much more ancient and general conventions. As shown in an analysis of Gómez-Moreno (2014), such archetypal images can also be connected and analysed as image schemas using methods that derive from cognitive linguistics. It is important to note here, however, that archetypal meaning complexes are also means for connecting humans and other animals. We share meaning of the snake’s body shape with many other animals as something potentially dangerous and are attracted by the big eyes and round face that are common signs of juvenile animals in most vertebrates. We recognise an animal by the eyes and we also interpret physical affinity and distance in a similar way to many other species. There appears to be a layer of zoosemiotic modelling underlying our language-based consciousness that, at least potentially, can act as a basis for relation to other species. In conclusion, I turn back to the question posed earlier: are ecological codes archetypal structures? I have treated the topic in a somewhat explorative and playful manner, but the possibility seems to be worth considering. Archetypal analogy-based imagery is structurally simpler, and thus its existence in ecosystemic relations is more probable when compared to the type of codes derived from strictly regulated intraspecific communication. To argue for a code-based communication, one should be able to show that the given species have cognitive capacity to operate with these codes, and that there are means by which codes can be communicated and become accessible between different species. In many Neo-Darwinian interpretations, as for instance in game theory, animals appear to be depicted as little book-keepers or strategists, calculating the cost-and-benefit of the given action or deducing the best available strategy for the given encounter. Such interpretations appear to carry anthropomorphic modelling as they presume animals rationalise at the level that would require access to linguistic sign systems. Very seldom the question is asked what codes or rules are actually accessible for the animal and by which cognitive and communicative means these rules are shared between the different participants. We 168 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. should be, however, very aware of the danger of transposing our own Umwelt structures to the animals that we intend to study. An alternative and in my understanding more realistic approach to interspecies communication would be to recall the claim of Gregory Bateson (1969: 21–30), according to whom the closest resemblance to animal communication in humans can be found in dreams. What Bateson had in mind here, was that like the human dream, animal communication also lacks linguistic structure, and that it is not possible to express negation, abstraction or make complex logical operations in a dream (as also in animal cognition). Rather, the perception in dreams is organised by a sporadic flow of images. Interspecies communication and regulation also quite probably take place through the exchange of a fuzzy flow of images. Building blocks of ecological code could very well be single but intense images that are shared by many species in different variations and are, at the same time, motivated or constrained by surrounding ecological networks. 169 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. 12 CONCLUSIONS Mimicry studies in biology have a long history, and they are conceptually related to the understandings of mimesis and imitation in human culture that reach back much farther. Acknowledging this background, I have treated mimicry in this book as simultaneously a cultural-scientific construct and as a real phenomenon—a result of semiotic activities of the participants of the mimicry system. Thus my analysis has two reference points: the mimicry concept as it is recognized and conceptualised in modern biology, and mimicry as a confusing communicative encounter in the Umwelten of the participants. A central theoretical model of mimicry in biology is the tripartiate mimicry model that includes a definite mimic, model and receiver species. This is a coarse simplification of the diversity of real mimicry cases in nature, but it is also a useful heuristic or modelling devise that helps explicate the structural properties of mimicry and carry out a comparative study of different mimicry cases in nature. At the same time, the triadic mimicry model conveys a bias of contemporary evolutionary biology that focuses on species and their genetic heritability and largely ignores any non-tangible relation of nature; that is, the organism’s knowledge, cultural traditions of populations and semiotic relations between different species. A biosemiotic view of mimicry, on the contrary, puts the communicative and behavioural activity of the organisms into the foreground and considers mimicry to be an outcome of their combined action. This means, that from a biosemiotics view, mimicry is a semiotic relationship that can easily emerge between various species and on many different occasions. Mimicry can appear in any communication system, given that the positions and interests of the participants differ and that there is a potential for the messages to become interchanged and misinterpreted. This understanding is in good correspondence with the diversity of different mimicry cases in nature, as well as the diversity of case-specific research traditions in biology: e.g. study of brood parasitism, Heliconius’ mimicry rings, coral snake mimicry, and so on. If the semiotic activity has a central role in mimicry, then mimicry as a systemic phenomenon is able to modify its structure on an evolutionary timescale through the balance of the semiotic activities of different participants. Putting emphasis on the role of the organisms’ semiotic activity brings along the necessity for using research methods that would be sensitive to the subjects participating in the mimicry. My analysis also demonstrates that the understanding of mimicry differs a lot depending 170 Manuscript of “Maran, Timo (2017). Mimicry and Meaning: Structure and Semiotics of Biological Mimicry. (Biosemiotics 16). Dordrecht: Springer.” Final version with pagination, images and index is at http://www.springer.com/gp/book/9783319503158. on the specific viewpoint taken: whether the analysis focuses on the communicative relation between model and receiver, deceptive relation between mimic and receiver or resemblance-based relation between mimic and model. This is the main reason why I have advocated an ad-hoc approach and tool-box like conceptual methods for studying mimicry, as these allow taking into account both the specifics of the participants’ Umwelten as well as cultural meanings and connotations related to the specific mimicry case. From a semiotic view, the basis of mimicry is both narrower and broader than the similarity between different species. Similarity seldom exists between full physiologies of animals, but is more often focused on the specific features or messages of the participating species. This means that the existence of the mimicry resemblance presumes the underlying meaning structures—some features or messages that need to be taken as representations of the animal as a whole. For instance, for eyemarks to function as a deterring devise, there needs to be an underlying metonymic generalization between an eye and a predatory animal for the participants of the communication. Such relations cannot arise mechanically, but need to be constructed through the meaning making activities of the animals. The bases of meanings in mimicry can exist on many different levels and have different degrees of generality. Through different argumentations, three basic mimicry categories were distinguished in this respect, the first two being camouflage that is related to the emergence of iconic relations in nature, and Batesian type mimicry that I have connected with the novel concept of ambivalent signs. As a third type of mimicry, I have discussed abstract mimicry, based on more general meaning relations and in connection to ecological codes, opening up possibilities for complex interspecies regulation in ecosystems. Mimicry is necessarily related to the semiotic qualities of the living world—to the fuzzy web of meanings that connect different species and arrange their behaviour, ecological relations and distribution. 171 Manuscript of “Maran, Timo (2017). 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