Basics of animal communication : interaction, signalling and sensemaking in the animal kingdom [1 ed.] 9781443881708, 1443881708

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Basics of Animal Communication

Basics of Animal Communication: Interaction, Signalling and Sensemaking in the Animal Kingdom By

Dario Martinelli

Basics of Animal Communication: Interaction, Signalling and Sensemaking in the Animal Kingdom By Dario Martinelli This book was peer-reviewed by two referees through a double-blind procedure This book first published 2017 Cambridge Scholars Publishing Lady Stephenson Library, Newcastle upon Tyne, NE6 2PA, UK British Library Cataloguing in Publication Data A catalogue record for this book is available from the British Library Copyright © 2017 by Dario Martinelli Cover Image Conversation Species © Dario Martinelli, 2017 All rights for this book reserved. No part of this book may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without the prior permission of the copyright owner. ISBN (10): 1-4438-8170-8 ISBN (13): 978-1-4438-8170-8

TABLE OF CONTENTS

Preface ....................................................................................................... vii Chapter One ................................................................................................. 1 Introduction to Animal Communication Studies: Definition and Problems Chapter Two .............................................................................................. 13 Preconditions for Communication Chapter Three ............................................................................................ 25 The Very Substance Communication is Made of: Signals Chapter Four .............................................................................................. 37 Channels and Functions of Communication Chapter Five .............................................................................................. 49 The Complexity of Communication: Multimodality and the Language of the Bees Chapter Six ................................................................................................ 61 The Secret Weapon of Communication: Lies, Tricks and Deception Chapter Seven............................................................................................ 75 Playful and Aesthetic Communication Chapter Eight ............................................................................................. 89 Interspecific Communication (Part I) Chapter Nine.............................................................................................. 99 Interspecific Communication (Part II) Chapter Ten ............................................................................................. 111 30 Key-Figures Who Shaped Animal Communication Studies

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Glossary ................................................................................................... 125 Bibliography ............................................................................................ 141

PREFACE

This monograph was written with two main purposes in mind, both mirrored by my choice to employ the term “basics” for its title. On the one hand, after almost twenty years of professional interest for animal communication via various paradigms (zoosemiotics primarily, but also cognitive ethology, sociobiology and others) – an interest that verged into dozens of publications, presentations and lecture courses – I realized that I had never tackled the task of a plain, systematic introduction to the topic in such a way that both my personal research and the most significant works of my illustrious colleagues in the field would be reflected. The closest I had got to this idea was what I tend to regard as my most accomplished monograph so far, A Critical Companion to Zoosemiotics (Springer, 2010). Then again, that companion was “critical”, meaning that the implied mission was not only a description of existing theories (including my own), but also an attempt to develop alternative, lateral paths, not sparing a share of criticism to what I considered obsolete, biased or inaccurate approaches. Of course, and regardless of how successful I may have been in my arguments, that program produced a less consistent structure of the book, one where certain topics were more (sometimes much more) relevant than others, because their demand for problematization was higher. Through this monograph, on the contrary, I intend to pursue a more systematic, equalitarian approach, where description is not just a vehicle for prescription, but it is actually the main point. The basics of animal communication are in this sense intended as the bulk of essential, systematized information that I consider necessary in the study of this fascinating area of inquiry. The second goal of this monograph is also the most important one, at least in the economy of this enterprise’s genesis. The book was indeed started in accordance with Dr. Kristian Donner, professor of zoology at Helsinki University, a great scholar, a friend, and really one of the kindest people one could ever meet. Having given a few courses in animal communication for his department, I discussed with him the possibility of assembling some pedagogical material that would allow us to employ one single, general resource for my (or someone else’s) lectures, instead of gathering single articles and book chapters here and there, and to uniform

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the level of complexity to our target: BA and MA students. The project received the crucial support of the Ella and Georg Ehrnrooth Foundation, which sponsored all of my research and writing work for this book, and also, for a few months, the assistance of my talented former student Cecilia Calais. In the course of the project’s implementation, it must be said, I experienced quite a U-turn about the quantity and the quality of features that this particular “textbook” was supposed to offer to its readership. I had started with a full collection of services to the student (and to the teacher willing to employ the book in his/her courses): every chapter (or lesson) was planned to include an introductory summary (“in this chapter you will learn that...”), a schematic list of topics tackled, a series of slides (in case the teacher wanted to turn the lesson into a presentation), a final summary (“in this chapter you have learned that...”), a ten-question test that students could use to verify their understanding of each lesson, a glossary of the difficult terms and two kinds of bibliography: one with the references mentioned in the text, and one of the “For more on this topic, read this” type. Quite a package, huh? This was until I realized three important things (and I have to thank many of my students and colleagues, with whom I had the chance to discuss the structure of this book, for opening my eyes): one, students are not idiots; two, teachers are not my servants; three, academic courses are becoming a mess. Students are not idiots: in the last few years, many universities have started the practice of packing courses with all sorts of extra material that is supposed to take a student by the hand and guide him/her through the mist of... through the mist of what? Through the mist of a written text! We “map” the information because we assume that the students would find its “narrative” too difficult. We spare on the natural time-flowing of data, and instead we fill the space with notes, post-its, snapshots and “bites”. In other words, indeed, we assume they are idiots. The problem is: I would not mind betting that, on the contrary, we are making them so, by using this strategy. The demonization of narratives, in modern academic courses, has certainly produced some advantages (sparing time being one, indeed), but it is also endangering the fundamental development of information-processing and critical skills in the students. It is a strategy that roughly equals the decision to take artificial vitamin integrators instead of bothering to eat a balanced amount of fruit and vegetables. We save time but we kill the “narrative” of food intake, the one that suggests what to eat with what, how much, when, and so forth.

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Also, have we ever thought that, once in a while, students really do not mind sitting on a sofa with a cup of tea, to actually read a book, instead of a collection of schemes? Have we ever thought that, perhaps, if a book is well written, it is actually a more pleasant (therefore more effective) way of studying? Please, do not get me wrong here: I am not criticizing “mapping” in general (on the contrary: it is an extremely effective way of processing information). I am just suggesting that a textbook, as such, should adopt the “narrative” strategy, because (along with a teacher’s lectures, of course) that is the very source of information to process. If the latter is already processed, then we are simply providing bad quality information (the same way – keeping up with food metaphors – readymade frozen dishes will never be as good as those carefully prepared with fresh ingredients). Any phenomenon of “learning”, in reality, needs to go through this two-step procedure: we intake the information in a “narrative” way (which means, we see the whole picture, the natural flow of time, the many variables...), and then we process it in a schematic way, distinguishing the more from the less relevant, generalizing the single elements into groups or patterns, and so forth. Therefore, the main challenge here, is making this kind of book clear and engaging reading, clarity being the very reason why maps came to replace narratives, and engagement being the very reason why students are less and less eager to invest their time in studying. This monograph wants to be game for this particular challenge, and would like to leave the important mapping stage to its readership, in the conviction that this will stimulate even more their understanding of the concepts. Secondly, teachers are not my servants: I do not really have a right to tell them what are and what are not the relevant parts in my text, nor how they should phrase them. A monograph, if employed as a textbook, must be a tool for the lecturer, not a bible. The lecturer should be able to do whatever s/he likes with the book, including the total omission of parts that I would instead consider crucial, and vice versa. The lecturer too has to work on the book, and not rely on ready-made information. The abovementioned mapping stage is, or should be, a teacher-student cooperation Finally, academic courses are becoming rather messy. There was a time (not such a long time ago, since I was living in those days myself) when a course consisted in a) a teacher coming in a classroom and doing most of the talking; b) a bunch of books (not few, in fact) to read and study with great attention; and c) an exam. End of story. None of the people of my generation seem to suffer from serious traumas as a consequence of this system. What we have nowadays, instead, is a glorious parade of handouts, Moodle accounts, mid-term exams, mock-

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exams, workshops, discussion groups, self-assessment processes, and so on and so forth. Anything. Except, well, a clear and engaging textbook. My intention here is not a conservative one, in fact it is experimental: I am not particularly against the new system (although, I must admit, it adds a lot of extra-time to my work, and I am deeply convinced that I could use that time in ways that would be much more useful for society). I just would like to perform an experiment of re-establishment of a direct dialogue between course material and students. I suspect that a) the general decrease of interest and preparation we are witnessing in modern students has nothing to do with a “traditional” textbook, when the latter is clear and engaging; and b) a storytelling approach to teaching is not more difficult or time-wasting than the schematic approach. Rather, it is simpler. In conclusion, it is not my intention to make a case out of this choice. The reader will be informed about my opinion on the matter only in these preface pages, and nowhere else. The book is called Basics of Animal Communication, and the title says all I intend to convey here. No subtitle of the likes of “towards a comeback of good old textbooks” was ever intended for this project. Plus, truth be told, it is not as if I will erase each and every sign of “organization” of the information, here: - Each chapter is carefully pondered to be in the vicinity of 4500 words of length (sometimes a little more, sometimes a little less). That is the length that usually corresponds to one hour and a half of attentive reading. One hour and a half is the standard duration of an academic lecture. - A glossary of difficult terms is available. In the text, they are marked with asterisks at both sides (e.g. *adaptation*, *behaviourism*), and at the end one can find short definitions for each of them. - Of course the bibliography tries to be as rich and informative as possible. But, really, the main strength of this book is, or should be, its clarity. Animal communication is a very complex topic: I put all my efforts into making some key issues as understandable as possible. I would like to express my gratitude to Kristian Donner for believing in this project from the start; to Cecilia Calais for assisting me in the first months; to Gavin Stewart for language-editing and Aušra Berkmanienơ for proof-reading; to all my students in the various Animal Communication and Zoosemiotics; and most of all to the Ella and Georg Ehrnrooth Foundation for their generous support. I dedicate this book to my family.

CHAPTER ONE INTRODUCTION TO ANIMAL COMMUNICATION STUDIES: DEFINITION AND PROBLEMS

To study animal communication has very little (or at least, not only) to do with admiring dogs whenever they are able to execute our requests. To study animal communication rather means to understand as much as we can of the way those dogs’ brains work within their own “canine” experience and how they are able to project their cognition while relating to the environment – including us, other animal species or other dogs. The point in animal communication is not only our disposition, as humans, to consider other animals as actively involved in communicative processes, but to see if such a ‘thing’ like a communicative process exists, and is operative, within their minds. There are many ways, and not necessarily coherent, to define communication. If we take a rather standard approach to the concept, we can say that communication is a phenomenon that involves the transmission of information from one living organism (sender) to another (receiver). We may of course say “animal” instead of “living organism”, but scientists are more and more convinced that communication occurs among plants as well, so it is safer to affirm that “communication” is a chief characteristic of being “alive”. Information is processed within the mind of the organism and encoded into a signal. When received by the other organism, this information undergoes a decoding phase, while still retaining a more or less recognizable relationship to the encoded information. During this process of encoding and decoding – which may consist of many steps – the information undergoes many transformations (including the fact that it may be “misunderstood”, while encoded): the receiver puts many other elements into the process, not only the information received (expectations, previous experience, relationship with the sender, circumstantial events, etc.). From the combination of all these factors, the receiver produces a response of some sort (which may also not consist in another act of communication, although usually it does).

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This approach to animal communication is called “cognitive”, and is the one we are adopting in this book. To possess *cognition* means to actually have a “mental activity” and the capacity to “process” information. Needless to say, scholars have not always defined animal communication this way. There was a long period, particularly before the late 1970’s, when the scientific community was not exactly persuaded that non-human animals had “minds” that could process the information in any complex way (in the next chapter, we are going to deepen the question of “mind”).

Two traditional approaches to animal communication Somewhat generalizing, we could say that there used to be two important approaches to animal communication, in the past (and, partly nowadays, although there are fewer and fewer scholars who follow these outdated paradigms). In one approach, communication is considered the result of *instinct*, which is a very tricky notion. Instinct is generally defined as a genetically acquired force that drives animals to react to a stimulus in certain fixed ways. In other words, when it comes to communication, the animal would not be really “processing” the information that s/he receives from the environment or from another animal: rather, there is supposedly a predetermined mechanism that makes the animal respond in a certain way. In a sense, instinct is like an application in a smartphone: when we install an app, we know that it has certain functions and that the smartphone is not really “reasoning” but simply enacting those functions. We tap on the screen and we see something appearing (a “response”), but we know very well that this is not because the smartphone has “understood” us in the way a friend of ours would understand our request. By assuming that communication is regulated by instinct, scholars were paralleling communication among animals to a mechanical, automatic process. Criticism of the concept by other scholars started from various angles. First of all, from a methodological point of view, it was noted that instinct was a kind of a “black box”, where we can put most (or all) of the observations that we are not able to explain in a more elaborated way. We see an animal doing something particularly complex, and we think “oh, it must be instinct!”, because – in principle – we are not ready to admit that animals can indeed do complex things. Another remark was that the notion of instinct makes no allowance for environmental influences upon behavioral patterns. If everything comes is pre-determined by instinct, what about the external stimuli? Does it not make any difference whether an animal is communicating with an animal that is bigger, smaller,

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friendly, aggressive, predator, prey…? Does it make any difference whether the animal him/herself is healthy, sick, young, old, relaxed, angered…? Finally, as research on animal behavior progressed, increasingly accurate descriptions of several behavioral patterns, originally thought of as “instinctive”, started appearing. It turned out that such patterns were not “smartphone apps”, but rather the result of different categories of motivation and cognition. In the second main pre-cognitive approach to animal communication we have an accent to the environment and external stimuli (exactly one of the critiques advanced to the promoters of instinct: what about the context?). This approach was heavily bound to a methodological school that developed in various fields of science (mainly in psychology): behaviorism. Behaviorism is founded on the idea that all actions performed by an organism do not have to be interpreted as the result of internal physiological or mental processes, but are rather the result of a direct contextual conditioning. When we talk about behaviorism, most of us think about “Pavlov’s dog” (from the famous behaviorist Ivan Pavlov) – a dog hears a bell sound and salivates in hunger, because, in a previous phase of the experiment, s/he was repeatedly given food right after the bell sound, so now the sound alone was enough to stimulate the salivation. The dog, in other words, was exposed to an external stimulus, and that (not some inner predetermined mechanism) was the reason why s/he reacted in a certain manner. Instinct would have made the dog salivate only after the food had appeared, but Pavlov’s insistence to announce food with a bell sound made the dog hungry at the sole announcement, even if no food would appear. Before being challenged (and partly overcome), in the second half of the 20th century by cognitive approaches, behaviorism was widely popular in the study of animal behavior. The objections to behaviorism were opposite-but-equal to those raised about instinct. In both cases, the process stimulus-reaction is automatic: either the reaction comes from the inside of the organism (instinct) or it is the stimulus that provokes and determines the reaction: in either case, no “mind” is conceived in the middle of the two steps. Also, if instinct risked becoming a “black box”, similar fears could be applied to behaviorism: we again see an animal doing something complex, and we go “oh! It must be the environment conditioning the animal!” Once again, albeit for different reasons, we deny the animal the possibility of “reasoning”. Finally, if instinct totally neglected the possibility that the animal could be influenced by the context, behaviorism neglected the

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possibility that there could be anything pre-determined at all in what the animal could do. For several decades, in the 20th century, scholars were divided into these two schools: communication by instinct, and communication by external stimuli.

The cognitive approach Then, as we said, the cognitive approach appeared in the second half of the 1970’s, provoking something of a revolution in the interpretation of animal behavior (communication included). It is frequently maintained that the turning point was the publication of a book called The Question of Animal Awareness, by Donald Griffin (1976), where issues of complex cognition in animals were finally raised in a very articulated way (there had been several scholars reflecting on this point in the past, including Darwin, but nothing had gone beyond the level of hypotheses and speculations). However, even if this approach soon became the most reliable of the three (and is an approach that is in constant progress: almost everyday we read in the news or in popular science magazines about discoveries on the complexity of animal behavior), criticisms were not spared in this case, either. First and foremost: how do we really know that animals have minds? And, once we (hopefully) establish that, how do we know how those minds operate? How do we go inside an animal’s brain to see what this animal is thinking? In fact, such questions are not easy even when studying humans (particularly certain communities or certain groups). How will this be possible with other animals? Everything we say is based on human criteria, which we are just arbitrarily applying to other animals, since nobody could ever say if animals really think in the ways we think they do. So, what do we do?

Inside and outside perspectives This problem was actually already emphasized in the field of linguistics (particularly after Pike 1954), as an opposition between the socalled “etic perspective” and the so-called “emic perspective”. Etic stems from “phonetics” – the study of linguistic sounds without regard to their significance in a language; while Emic comes from “phonemics” – the study of speech sounds that are meaningful in a language. In brief, by the emic perspective we mean that we are interested in what the insiders of a

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certain community think, believe, etc. By the etic perspective, on the other hand, we are rather interested in the outsiders’ perspective: what we think when we see a certain community. Clearly, the same principle is applicable to other animals: can we afford to be “emic” and understand how they think, or shall we be “etic” and project our own interpretation on what they do? Simplifying a bit, we can say that the two traditional approaches to animal communication were quite happy to be just “etic”, while the cognitive approach has a pretention to be “emic” too, at least to an extent. Evidently, however, the latter task is not an easy one. Even in the study of human behavior, cultures, etc., there is still an ongoing, quite animated discussion on the etic-emic issue. One of the main points is that a totally emic perspective is impossible to take, especially when there is no way to establish linguistic interaction with the culture observed, so that its members could describe their own views. How can we understand the insiders’ perspective, if we cannot even speak the same language with them? The problem becomes even more serious with non-human animals, where not only there is no language in common, but there is a whole gap based on physiological, behavioral and psychological differences. This problem was very well posed by a scholar called Thomas Nagel, who possibly wrote the most famous essay on this topic, “How does it feel to be a bat?” (Nagel 1974). Indeed: how does it feel? To Nagel, we will never know, because, simply, it is impossible to be a bat for someone who is not a bat. The scholars who were more eager to accept the cognitive approach to animal communication (and behavior in general), generally had three types of reaction: 1. From a common-sense perspective, that thesis sounds a bit simplistic, because it represents a comfortable way of facing a problem that is in fact more complex. Yes, it is true that there is no way to fully understand another subject, but such a problem exists in principle in every field of science, so we can only cope with it. For instance, pharmacology is possibly far more inexact than is animal communication studies. When a doctor prescribes a drug for a patient affected by a given disease, s/he knows perfectly well that that particular drug, made for that particular disease, does not necessarily work on that particular patient. In other words, an objective interpretation of a given disease does not allow one to grasp all of its possible individual implications. But, of course, this is not a good reason to dispense with pharmacology and not give any medicine at all: we still need to take some responsibility, the doctor cannot just dismiss the patient because s/he is not totally sure whether the drug will work or

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not. The doctor knows that there are “more chances” that the drug X will work on the patient A, because X worked much better than Y on the patients B, C and D, so there is a good reason to believe that it may work on A as well. 2. Secondly, everything, or almost everything, that can be grasped about phenomena occurring within any organism, is basically the result of observations made from the outside. When we see a teenager with long hair, black leather trousers, various metal accessories, and a T-shirt with an Iron Maiden album printed on, well, we cannot be 100% sure, and we do not want to have prejudices, but it really looks like this teenager is a heavy metal fan, does it not? Nobody could blame us if we made that assumption. If, on the contrary, we said that we have no clue what this guy is, then perhaps someone would look at us and think that we are not exactly bright-minded people. Now, in many cases, a given non-human animal pattern recalls a human one. Modes of production and correlated behaviours of that pattern are also impressively similar, and the same applies to emotional experiences. In addition, we know that this animal has a brain, we know that s/he interacts socially, and so on. So, if there are all these similarities, it is not that crazy to assume that maybe also what goes on inside the animal’s mind is very similar to what goes on inside our mind. Again: we are not 100% sure, but… 3. We actually do have clues as to how we might “emically” study non-human species. To start with, we can scientifically study the sensorial organs of animals – and we know that the way an organism interacts with the environment is largely due to the way s/he perceives it. By now, we have reached enough knowledge in research that is possible to wire a computer to an animal’s brain and create videos of what the animal is actually seeing. By recording the electrical activity of nerve cells in the thalamus (that region of the brain which receives signals from the eyes), the researchers can create images of what the animal sees. In other words, technology is little by little helping to be more “emic”. So, is the suggestion here that emic is good and etic is bad? Of course not. The best solution is a balanced combination of etic and emic. What we need to understand is that an etic approach is not necessarily “external” to the subject observed. When we observe another animal species, we are external at the very moment we take a “human” eye to look at it, but we should not forget that we are also animals, so our etic view is not only external. If we observe a community of chimpanzees, we are external as specimens of the Homo sapiens species, but we are insiders in many other categories. Taxonomically speaking, indeed, there are only three taxonomic

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rings that make us different from the Pan troglodytes: the species indeed (we are Homo sapiens and they are Pan troglodytes), the genus (we are Homo, they are Pan) and the subtribe (we are Hominina, they are Panina). This is it. In all other taxonomic groups, there is no “we” and “they” anymore: there is only “we”. We are the same tribe (Hominini), the same family (Hominidae), the same suborder (Haplorhini), the same order (Primates), the same class (Mammalia), the same clade (Synapsida), the same phylum (Chordata), and finally the same kingdom (Animalia). So, we are united by eight groups and separated by three only. If we were able to take, for instance, a “mammal” eye, when looking at chimpanzees, we would be taking their same eye. Is this possible? Of course it is: it just takes the right knowledge that will allow us to distinguish between what we do as “humans”, from what we do as – say – “primates”, “mammals”, “animals”, etc. We learned to do this very well within the human culture, so, let us say, an English person who comes from London, lives in the Whitechapel district, and comes from a certain family, knows very well what characteristics make him/her a member of that family (as opposed to other families), of that district (as opposed to other districts), and so forth. The way we are, obviously, does not come from a single source: we are shaped by dozens of factors, and we have to learn that these factors do not come only from our being “human”, but also from our being Hominini, Hominidae, Haplorhini, Primates, Mammalia, Synapsida, Chordata, and Animalia. Our style of parenthood, for instance (which is known as “Kselection” strategy: we make few offspring, but we take great care of them, as opposed to the “r-selection”: many offspring, but less care, like fish, amphibians, insects, etc.), is something that makes us much more “mammals” than “humans”: so, if we look at the way chimpanzees take care of their offspring (and communicate with them), we can relate to them much more than we think. Our position of “etic” observers, in other words, is not an unfair one, because it is not entirely external: we can recognize chimpanzees’ parenthood as belonging to the same style of parenthood that we have. This is a very important lesson that we need to learn, when we approach other animals. They are not just “others”: they are also something that we are. The more we accept that, the more we will fight the many prejudices we still have about these topics.

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Fig. 1 – A summary of the three approaches to animal communicaiton. In (1) instinct determines the animal’s response to the environment, in (2) the environment affects the animal’s response, in (3) there is an intemediary stage corresponding to the cognitive processing of the information

The main areas of inquiry concerned with animal communication Animal communication has been studied under a wide range of disciplines and fields of inquiry. Occasionally, it has been named in different ways, too, so it is not rare that scholars discuss the same thing by using different terminology. To make just one example, since Tembrock 1971, a popular term among scholars is “biocommunication”, used to describe various types of communication between different specimens of the same species, or even across different species (since the prefix “bio-”, from ancient Greek, stands for “life”, biocommunication is a term adopted not only to describe animal communication, but any other living forms like plants or fungi).

Ethology At any rate, a short overview of (some of) the many disciplines that deal with animal communication cannot but start from ethology, the most important field of the category. A branch of zoology, ethology is the

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scientific study of animal behavior, and that, of course, involves a lot of research on communication patterns and processes. The term stems from the Ancient Greek “ethos” (“custom”, “behavior”), and was first coined in 1762, by the Academie Française des Sciences. John Stuart Mill employed the word in his System of Logic (1843), but in a different sense, proposing the establishment of a field of inquiry, “ethology” indeed, would account for individual and national differences in character. It was the American Myrmecologist William Morton Wheeler, in 1902, to popularize the term in the modern sense. Leaving aside the word, however, traces of, so to speak, proto-ethology can be already found in studies on instinct and innate behavior, such as Pernau 1716, Reimarus 1773 and Spalding 1873. Before ethology, the main field devoted to the study of animal behavior was comparative psychology: however, besides being very different fields in themselves, comparative psychology prefers to perform empirical observation in artificial experimental contexts, while ethology concentrates on behavior in natural situations. Because of this, the two approaches may be considered either complementary or competitive, and so they acted all through the 20th century (collaborating in some cases, and disputing over their respective findings in others). Another difference is of geographical type, as comparative psychology developed most strongly in North America, while ethology was more followed in Europe. One of the main focuses in ethology is the study of the evolution of behavior and the interpretation of behavior in terms of natural selection, and that of course reveals the instrumentality of Charles Darwin to the foundation of modern ethology (particularly, the book The expression of the emotions in animals and men, 1872). Early ethologists, like Oskar Heinroth and Julian Huxley, focused primarily on the notion of instinct, as an explanation for animal behavior. An important step was the construction of the concept (and research related) of *ethogram*, which allowed significant collection of data about behavior. The turning point was the generation of Konrad Lorenz (whose name is still the first one we associate to the term “ethology”), Karl von Frisch and Nikolas Tinbergen, who 1) established unmistakeably ethology as an autonomous discipline; 2) set a number of methodological standards that are still used nowadays, and 3) engaged in sophisticated investigations of animal behavior that led to important, and sometimes revolutionary, discoveries (like von Frisch with the bee dance – which we shall talk about later in this book). Through the work of this generation, ethology became strong particularly in Europe before World War II, and also in the UK right after, when Tinbergen moved to Oxford University, and figures like William

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Thorpe, Robert Hinde and Patrick Bateson came onto the scene. In continental Europe, meanwhile, another generation of ethologists, pupils of the old masters, was growing fast, providing new names of historical importance, such as Irenäus Eibl-Eibesfeldt and Martin Lindauer. During the 1970’s, ethology underwent that cognitive “restyling” we mentioned, witnessing the birth of cognitive ethology, but also of another branch we shall soon talk about, sociobiology. Cognitive ethology was pioneered by Charles Darwin in his last works of zoological nature (1871 and 1872), and established by Donald Griffin’s The Question of Animal Awareness (1976). It is a rapidly growing field with a strong inclination for interdisciplinary work. Currently, the list of eminent cognitive ethologists includes Colin Allen, Marc Bekoff, Frans De Waal, Dorothy Cheney, Robert Seyfarth, Dale Jamieson, Remy Chauvin, and the great majority of scholars in interspecific communication experiments (which we shall discuss later on in this book). Sociobiology, on the other hand, specifically studies the social behavior in animals, therefore with communication on the frontline. Its main goal is to prove that social behavior is also the result of natural selection, even in cases where it seems clear that context and stimuli play a prominent role (like aggressive behavior, for instance). Wilson (1975) is generally regarded as the landmark in the development of the field, although it must be pointed out that protosociobiological approaches to animal behavior had existed since the end of the 19th century already (e.g., in Kropotkin 1902), while the term itself “sociobiology” had allegedly been circulating since the 1940’s in biological and psychological environments. At present, it is safe to say that ethology consists of at least the following branches: 1) Field ethology: the classical investigation of animal behavior in natural conditions; 2) Applied ethology: the practical exploitation of ethological knowledge in activities like breeding, veterinary medicine, companion animals, etc.; 3) Cognitive ethology: the study of the mental processes of acquisition, representation and use of information in animals; 4) Human ethology: the application of ethological theories to the study of human behavior; 5) Sociobiology: the systematic study of the biological bases of social behavior.

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An excellent introduction to classical ethology, before the cognitive breakthrough, is Eibl-Eibesfeldt (1970).

Other fields Ethology is of course not the only field of inquiry that is interested in animal communication. There are also more specific fields that focus on particular aspects of animal communication. One of these is bioacoustics, an interdisciplinary field that mainly combines biology and physical acoustics, devoted to the investigation of the neurophysiological and anatomical mechanisms of sound production, spreading and reception in all animals. Its foundation is generally attributed to Slovenian biologist Ivan Regen, who started studying sound communication in insects around the 1920’s. In bioacoustics, the study of acoustic signals is also aimed at practical and applied use, such as agriculture and biological pest control. Methodologies and equipment are varied. Perhaps more than any other field mentioned in this chapter, bioacoustics employs a lot of hi-tech tools, including software for sound recording and analysis, special microphones, infrasound and ultrasound detectors, laser vibrometers, recording machines and others. Methodologically, the research work can be performed both in natural and artificial contexts. An excellent overview of the field is Hopp et al. 1998. It is not only natural sciences that investigate animal communication. Important contributions also come from humanities, and zoosemiotics certainly deserves a special mention. The study of animal communication is precisely the main concern of this discipline, which uses methodologies such as semiotics, linguistics, information and communication theory. The term and the first definition were established in 1963 by Thomas Sebeok and Rulon Wells, and were proposed for a discipline that would cross the paths of semiotics and ethology, focusing on the study of signalling behavior in and across animal species. Like ethology, zoosemiotics too went through a transformation in the late 1970’s, adopting the cognitive paradigm for a more complete understanding of animal communication. An overview of different approaches and theories in zoosemiotics appears in Martinelli 2010. A particular sub-field of zoosemiotics is zoorhetorics, introduced by semiotician Stephen Pain in the 1990’s (see Pain 2002 and 2009), which interprets animal communication as “natural argumentation” that employs some kind of tropes and figures like in classical rhetorics. Still within the humanities, an important, and more recent, field is certainly anthrozoology. Partly an umbrella term, and partly a discipline

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with a specific paradigm, anthrozoology is an interdisciplinary field of inquiry originating within social sciences, and that nowadays comprises studies in anthropology, art, education, ethology, history, literature, philosophy, psychology, sociology and veterinary medicine. It is defined quite simply as the scientific study of the relationship between human and nonhuman animals – a relationship that, of course, is very often based on communication (as we can for instance see in the case of human interaction with dogs or cats). Among the topics most often investigated by anthrozoology, there are human-pet relationships and their link with physical and psychological health; animal assisted therapy; abuse and cruelty to animals; and companion animals as social facilitators. Institutionally, anthrozoology took its first steps in 1987, with the first issue of the journal Anthrozoos, and in 1991, with the creation of the International Society for Anthrozoology (ISAZ) at Cambridge University. Among the foundational literature on anthrozoology: Serpell 1996 and DeMello 2012.

CHAPTER TWO PRECONDITIONS FOR COMMUNICATION

Sometimes, when we think about “communication”, we tend to be a bit superficial. We imagine this phenomenon as a mere exchange of information, as if this exchange was a simple, mechanical “movement” of a signal (or word, sentence, or otherwise) from a point A to a point B, that is, from one interlocutor to another. For centuries, the main assumption (best embodied by the notion of “mechanism”, promoted by philosopher Rene Descartes) was that animal brains were functioning in a very simple, machine-like, manner. For most of the 20th century, the dominant theoretical trend (originated in psychology) was *behaviourism*, founded on the idea that all actions performed by an organism do not have to be interpreted as the result of internal physiological or mental processes, but are rather the result of a direct contextual conditioning. Key behaviourist scholars were Ivan Pavlov, Edward Lee Thorndike, John B. Watson and Burrhus F. Skinner. Before being challenged (and partly overcome), in the second half of the 20th century, by different approaches, behaviorism was widely popular in the study of animal behavior, affecting the early developments of animal communication studies. Of course, the reality turned out to be much more complex than this stimulus-reaction scheme: two (or more) subjects engaged in a communication process have a lot of “problems” to solve and “requirements” to meet in order to make that process possible. In fact, realizing this complexity has been one of the major progresses within animal studies as a whole. To summarize this development in one sentence, we could say that “animals” emancipated from their status as Cartesian machines, to become actual “subjects” (“persons”, if we like: in fact, the status of personhood in non-human animals is now becoming an important legal issue, as well). Two *phenomenologists* (San Martin and Pintos, 2001), following the indications emerging from Edmund Husserl’s work, have very clearly explained what it means to be a “subject”, in a list of eleven points:

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1) Every animal is a living bodily entity, that ‘feels’ and experiences life in an intentional way; 2) Every animal rules its own body. The animal is the entity that moves its own body, that ‘decides’ to move; 3) Every animal experiences a mental life articulated in time units;Every animal experiences its own body in a direct way. Whether aware or not, the animal ‘feels’ its own body (heart beating, illness, etc.); 5) Every animal – to the letter – is alive. In phenomenological terms, this means that the animal is situated in its body first, then in the surrounding environment. The animal lives in the environment because it lives in itself; by consequenceEvery animal is in material relation with the world. Its body is made of matter, like the surrounding environment. It is on a material basis that we are open to something which is not only ‘ourselves’;Every animal perceptually feels itself as the “starting point” of the world. Every animal is egocentric in principle, and perceives the world starting from itself. Things ‘are’, starting from what ‘I’ am and am not. The relation with the world is based on affirmations and negations of our identity, thus starting anyway from ourselves;Every animal perceives the world from one point of view. To mention a trivial example, changing physical position, sensorial perception changes; 9) Every animal experiences a common animal world, and a social horizon of its own community/group/species. The animal knows to be part of a given category and understands who is a member of the same category, and who is not. The way a dog relates with a cat is clearly different from the way s/he relates with a tree, but has several points in common with the way s/he relates with a human being; 10) Every animal has to confront the surrounding environment and its inhabitants: it communicates, understands, misunderstands, etc.; 11) Every animal interacts with each other primarily on an affective and emotional basis. In this chapter, we shall get acquainted with some of the most important conditions that allow communication to happen. Some of these conditions need to occur simultaneously, some may or may not happen (depending on the situation), and some, finally, may occur in different moments of the process. For these reasons, we shall deal with them in no other order than an alphabetical one.

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To learn more about these topics, at least the following books are recommended: Box 1973, Griffin 1976, Beck 1980, Bonner 1980, Fagen 1981, Akimushkin 1988, Bekoff-Jamieson 1990, Gardner et al. 1994, De Waal 1996, Dennett 1996, Vauclair 1996 (if you can read French), AllenBekoff 1997, Rogers 1997, Cimatti 1998 (if you can read Italian) and Baber 2003.

Abstraction Abstraction, or generalization, is the ability to represent an event/entity in terms of general qualities/characteristics, independently from concrete realities, specific instances or actual objects. To abstract means to be able to consider a given object as a case within a category, to mentally represent that category, and finally, on the basis of that representation, to recognize other cases of the same category. For example, abstracting David Copperfield as part of the category “Dickens’ novels” allows us also to recognize A Tale of Two Cities within the same whole. The process applies also to the capacity of spotting the category even when the second case is merely an alternative example of the first one, or is manifested in another form. In the previous example, not only an actual hard copy of David Copperfield can be recognized as Dickens’ novel, but also an electronic file of it, or even a picture of the cover appearing in a magazine. In psychological studies, abstraction is defined as a “higher brain function”, and therefore has been long considered a *species-specific* human feature. Already in the first decades of the 20th century, however, and even more effectively starting from the 1950’s, an increasing number of ethological studies, in both natural and experimental contexts, found out that abstract representations are extremely common among non-human animals. Subjects of such studies included pigeons, toads, elephants, parrots, ravens, squirrels, rats, monkeys, cats, and of course great apes.

Code When we communicate, we use various different signs (words, gestures, displays…) that, in principle, do not make any sense, but they do just because we “decided” that they do. In English, for instance, we use the word “book” to describe that particular object with letters written on, pages, cover, etc. But, how come that thing is called “book”? Does this particular word look like, smell like, sound like, and so forth, that particular object? Definitely not: nothing in a book justifies the word

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“book”. It is just a decision that we took collectively, as English-speaking people. An Italian-speaking community took another decision (“libro”), and that other decision, too, has nothing in common with the object itself. We could now gather a few people, and agree that, from now on, that object will not be called “book”, but – say – “train”. That would be just as legitimate as “book”, as long as there is this agreement among us. Now, a system of “agreements” and “decisions” of this kind is called “code”. A code is a set of rules of transformation, shared by at least two subjects (usually a community), through which signs are more or less deliberately associated to one or more meanings. A code requires social interaction in order to be created, and – after being established – enriches such interaction. Within the same community, let alone species, codes may be subject to progressive variations and adjustments, due to different factors: increasing efficiency of a new sign-meaning association, as compared to an old one of the same type; introduction of new subjects from a different community, who carry a different sign-repertoire, etc. We know, for instance, that languages evolve, so these agreements evolve too. If, in the time of Shakespeare, we would say “Thou art”, now we say “You are”, and the meaning is the same: it is the code that changed. Needless to say, non-human animals must use codes as well. They also have to “agree” on certain conventions, and these conventions change depending on the species and also on the communities within the same species.

Cognition “Cognition” is the way we call the totality of mental activities and information processing occurring within an animal, in a fashion partly or totally untied from instinct (a subject we shall also discuss later). The study of animal cognition focuses on at least the following aspects: 1) Attentive skills – The capacity of focusing on and discriminating between different stimuli; 2) Classification and categorization – The capacity of finding similarities across single items, and grouping them into so-called “semantic fields”: for instance, we see a birch, an oak and a pine, and we categorize them in the semantic field of “trees”; 3) Temporal cognition – The capacity of learning, retaining and transferring past information for (immediate and/or remote) future use; 4) Spatial cognition – The capacity of orienteering and navigatin;

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5) Production and use of tools; 6) Problem-solving – The capacity of pursuing a goal by avoiding especially complex and unusual obstacles; 7) Language (which we are about to discuss); 8) Consciousness (which we are about to discuss); 9) Counting and computing – The capacity of discriminating between quantities and perform basic mathematical tasks.

Consciousness “Consciousness” is an umbrella-term that may refer to a vast scale of mental states, from subjective experience to simple awareness. When it comes to non-human animals, there are not many data that scholars can use to “scientifically” assess the presence of consciousness: for this reason, scientific inquiry is regularly accompanied by philosophical conjectures. The range of hypothesis goes from a pessimistic, very famous essay entitled “What is it like to be a bat?”, whose author, Thomas Nagel (1974), practically answered “No idea!” to the title’s question, to the optimistic Cheney-Seyfarth 1990, who instead offered a few hypotheses to their question (ironically referring to Nagel himself) “What is it like to be a monkey?”. A central theme in modern animal studies, consciousness was very interestingly approached in Mortenson 1987, where different anatomicphysiological-behavioral clues for consciousness are gathered as indications for the presence of the phenomenon. Summarizing that whole book we can say that: 1) Indications of consciousness come from anatomy (particularly the structure of the brain), physiology (particularly the function of the muscles) and ethology (particularly the consequences of a given behavioral pattern); 2) In relation to brain, we have three forms of expression: the structural expressions (Cortex, Cortical and sub-cortical areas, Hemisphere, etc.); the electric expressions (Cerebral waves, Induced potentials, Peaks); and chemical expressions (transmitters like adrenaline or dopamine, hormones like hypophysis or thyroid, endorphins, etc.); 3) When it comes to the function of muscles, we have voluntary and involuntary ones; 4) When we consider behavior, we see patterns like sensibility, exploration, learning, communication, and others.

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Now, all these elements, together or separately, indicate the existence of consciousness. Cerebral waves, muscles, sensibility… each and all of these get activated only when a specific status of “awareness” exists: the animal is “alerted”, “reactive”, “sensible”, and so forth. It goes without saying, in order to communicate, one must be at least positive that there is something to communicate about and someone to communicate to.

Culture “Culture” can be defined as the totality of information acquired and developed by a community and transmitted non-genetically from one generation to another. Phenomena of this kind have been observed in some insects, fish, amphibians and reptiles, and in most birds and mammals. With few exceptions, the presence and the quantity of cultural processes are directly proportionate to the degrees of sociality (we shall discuss this concept later) displayed by each species. At the core of any cultural process, there is the capacity of social learning (this concept, too, to be explained later), which implies the transmission of some information from one or more “demonstrators” to one or more “observers/listeners”. Several factors, at different (and interchangeable) stages, intervene in the process. These are (in no specific order): 1) *imprinting* (see later); 2) attention; 3) imitation; 4) memory; 5) *social facilitation*; 6) creativity; and of course 7) communication. One famous case in which all of these factors are operating at the same time is that of the macaques Macaca fuscata, closely observed during the 1940’s and 1950’s by a group of researchers led by Masao Kawai in Koshima, Japan (Kawai 1965). Researchers had the idea of introducing a “novelty” in the life of a group of macaques, by leaving some sweet potatoes along the beach. One macaque began to wash the potatoes in the water, instead of brushing the sand off with her hand as other macaques did. In short time, this habit became popular among the other members of the community and was passed on from generation to generation. In addition, the macaques proved to make a cognitive use of the innovation, as further modifications to this pattern were eventually provided (for instance, they started washing the potatoes in salty, rather than fresh, water, in order to enhance their flavor).

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Imprinting “Imprinting” can be defined as the capacity to learn specific types of information at certain critical periods in development. Although anticipated by the English biologist Douglas Spalding in 1873, the concept became central in science when the famous ethologist Konrad Lorenz had an insight of this phenomenon while observing young geese and chickens spontaneously following their mothers from almost the first day after they were hatched. He then reproduced the procedure with eggs that were incubated artificially, discovering that the stimulus can come from any source and still serve as imprinting, as long as it is presented during a critical period (the few days after hatching). To an extent, imprinting is the first “official” form of communication between parents and their offspring.

Intentionality In a general sense, intentionality represents the conscious planning or performance of a given action, aimed at a goal. In philosophy, the expression has been used in a broader sense, designating any mental state provided with a content (e.g. thinking, willing, longing), but in ethology, the concept has been practically overlooked until the coming of cognitive ethology, when – on the contrary – it became central. In this book, the role of intentionality in communication is very important: as we said at the beginning of this chapter, we should not think about “communication” as a trivial “movement” of a signal from one interlocutor to another. It is a much more complex phenomenon, for which it is also essential to “have intentions”. The notion that animals act only by instinct is an obsolete one: for the last fifty years, at least, scholars have explored more interesting explanations.

Learning The term defines the process of acquiring and retaining information through *ontogenesis*, i.e., the consequence of interaction between an animal and its environment, rather than through *phylogenesis*. While in the past it was often claimed that almost all non-human behavior is due to instinctive predispositions, the current scholarly position generally holds that:

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1. All animal species (including protozoa) are provided with so-called “non-associative” learning abilities (i.e. learning based on constant exposure to a unique event or stimulus); 2. Most of them (except for protozoa) are provided with “associative” learning abilities (i.e. learning based on constant exposure to numerous causally related events or stimuli); 3. In many species, particularly the so-called higher ones, there is “complex” learning (i.e. learning based on cognition, rather than simple association). Further categorizations of the concept concern a) the specific learning processes associated with singing abilities in bird and mammal species (song learning); b) the form of learning that occurs without an explicit stimulus, with the animal acquiring information on his/her own environment in an exploratory way (latent learning); c) the learning processes specifically tied to an organism’s experience (perceptive learning); d) the information received and stored by an organism in its prenatal life (prenatal learning); e) the acquisition of a (subjectively or objectively) “correct” reaction to a stimulus, after repeated attempts (trialand-error learning); and f) the type of learning specifically emerging from social interaction and parent-to-offspring transmission (social or cultural learning).

Mental Representation A mental representation is an inner sign of an external object (which functioned as its stimulus), and that can be reactivated also in absence of the original stimulus. The most typical example of mental representation is the mental map, that is, the capacity of orienting oneself in space and heading towards a certain destination, by using signs available in the environment, therefore being able to solve a consistent number of spatial and temporal problems. Back to communication in a strict sense, the process is more easily enacted (and enhanced in its potentials) when the interlocutors are able to create a mental representation of each other. When we can mentally represent the subject we are communicating with, it becomes easier to understand him/her, to expect certain reactions, to find the best strategy for a successful communication, etc.

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Mind Leaving aside the endless philosophical, psychological and neurological implications of such a concept, and focusing only on its relevance within our topic, a “mind” can be defined as a network of biological and physiological activities and relations (including brain functioning, sensorial perception, nervous system, etc.) that allow organisms to acquire, store and elaborate the information they are, were, or might be exposed to, and produce an adequate response (which includes not producing any response whatsoever), after a consideration of the variables involved at any stage of this process. A network of this kind is essential in a great number of communication processes (most of which we shall discuss in this chapter or in general in the book), including deception, play, aesthetics, mental representations, etc. As such, it must be considered an integral part of any member of the Animal Kingdom. To put it simply: in order to be understood, a signal must be interpreted. This requires taking several variables into account, including the fact that a signal does not necessarily look like (or smell like, or sound like, etc.) the message that is being displayed at that particular point. When we think about it, the action of – say – wagging the tail for a dog does not look at all like “happiness”, “eagerness to interact”, “excitement” or else. We need that particular network to connect that very signal to a number of variables (our past experience of dogs wagging the tail, the fact that it is a dog and not a cat – for whom, as we know, wagging the tail is rather a sign of discomfort, and so forth): once we do that, as quickly as possible, we can conclude that, yes, that dog is happy to see us.

Ritualization Ritualization is defined as a process through which some behavioral patterns are modified, in terms of intensity, recurrence, articulation and dynamics, in a way that increases and/or varies its effectiveness to other specimens. Forms of ritualization are the stereotyping of certain patterns, incompleteness and repetition in putting them into action, and – mainly – the fact that some signals end up referring to other signals rather than to specific meanings (we shall discuss this in Chapter Four). For instance, a dog growling to another dog may not want to fight at all, but this signal is a precise and “ritualized” message, and the other dog reacts consequently: the signal does not mean “I will bite you now”, but becomes a “ritual warning” that displays an intention of biting.

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Ritualization has been detected in practically all species, especially in the so-called superior ones (birds and mammals). A famous ethologist (whom we shall mention at length in Chapter Twelve), named Irenäus Eibl-Eibesfeldt (1970: 100-1) discusses ritualization as a process that produces the following changes: 1) Function – After ritualization, a pattern previously serving a certain function in the animal *ethogram* comes to serve another one; 2) Motivation – The ritualized pattern can emancipate from its original motivation, and be now animated by a different one; 3) Redundancy and exaggeration – Ritualized signs are generally more redundant and exaggerated than non-ritualized ones; 4) Threshold values – The ritualized pattern is generally more easily released than the non-ritualized one; 5) Postures – In the ritualization process, several bodily movements tend to be framed into still or quasi-still postures (as in the case of the dog grinding his/her teeth, who “freezes” his/her aggression into a simple warning); 6) Orientation – Elements in space orientation may be changed during the ritualization process; 7) Intensity – The intensity (and its variations) of a sign may become more stereotyped (and therefore more constant) in a ritualized display; 8) Complexity – A ritualized sign reduces the element of complexity of the non-ritualized sign it refers to; 9) Body structure – Selection may favour variations in the body structure for the ritualized processes (typically, through the addition of ornamental elements in an animal’s body). More on ritualization will appear also in Chapter Three.

Sociality “In the last analysis, all animals are social beings, each species with a characteristic set of communication problems to solve.” (Sebeok 1963: 465) This sole quotation should be enough to give the notion of sociality the role of quasi-synonym to communication. Sociality, not to be confused with socialization (which is a concrete action), is the condition or quality of being social (tendency to intersubjective interaction and association, to form groups, to establish social roles, to mentally represent other entities, etc.). Sociality is at the basis of nearly all processes and activities related

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to animal communication, up to the point of becoming indistinguishable from the latter (sure, one can still speak with him/herself, to an extent, but the basic reason for communication remains the existence of social relations). The two phenomena, in particular, are in a mutual causal relation, that allows the reciprocal increase of specialization and complexity. The act of communication, for instance, may increase the level of an animal’s sociality, by taking a cognitive role and therefore producing and exchanging more complex forms of behavior (conveyed through an increasingly sophisticated information level). Such an exchange is possible when the subjects involved are able to create mental representations (already discussed) of the environment and recognize the entities within it (including other subjects). This process is in turn possible thanks to the sharing of a similar set of mental representations (that is, a code – another concept we have illustrated already).

Syntax of Perception All animals perceive the environment as *gestalts*, i.e. through a topbottom or, better, figure-background framework. For instance: when we attend a class, we are in a place (the classroom) that is full of objects: chairs, desks, hangers, pictures on the wall, windows... yet, most of time, we are paying attention to very few elements: our teacher, the screen where the slides are being shown (or the blackboard, or similar devices), our notebook (or laptop, or both), a couple of classmates with whom we exchange some comment, and – sadly – more and more often, our smartphone. Roughly speaking, these five are the main elements of the classroom, the rest is not so relevant, is it? In Gestalt theory, we call the relevant elements figure and the less relevant background. As a consequence, the communication of this experience is also gestaltic. Among all data available in the environment, animals will select only those they are most interested in (e.g. a bee searching for nectar will perceive flowers as a figure and the rest of the meadow as background). This is the very first level of syntax in natural languages, which can be detected both in the simplest interactions of some insects and in the most complex conversations of human intellectuals. In Chapter Five we shall speak about the so-called “multimodality”: that – we shall see – provides a bunch of perfect examples of this gestaltic organization of the perception, within the area of communication.

CHAPTER THREE THE VERY SUBSTANCE COMMUNICATION IS MADE OF: SIGNALS

In general terms, a *signal* is any pattern (behavior, appearance, organic process…) that has the purpose to transmit information from one animal to another. In physical terms, a signal is also a form of energy: it can be chemical, mechanic, electric and so on. This is why we think of communication also in terms of *channels*. A signal is really the currency of communication: if money is used to trade goods, signals are the “coins” used to trade information. To study communication means to study how signals are constructed, organized, assembled, distinguished, interpreted, codified, and so on.

Analogical versus digital Signals can be digital or analogical. By digital (or discreet) we mean a signal that does not vary in quantity (length, strength, intensity…), or, if it does, that is not relevant to the reception of the message. In other words, a *digital signal* is either present or absent (“yes” or “no”, “on” or “off”, “1” or “0”). The appearance of a certain body color in some animal species is a good example of digital signal. The males of the Three-spined stickleback Gasterosteus aculeatus display a characteristic red color on their chin and belly during the mating period. This color is either there or is not, so it is really an “on/off” type of signal. In the case of analogical (or graduate) signals, variability is the issue. In *analogical signals* the contents of the message depend on the degrees of its emission. Here, therefore, we have a case of “more/less”, “darker/brighter”, “higher/lower”, etc. The threatening signals displayed by many species possess this kind of relationship: the more aggressive the animal feels, the more intense the signal becomes. Obviously, given these premises, analogical signals tend to be richer than the digital ones, that is, their communicative potential is higher. An analogical signal carries many meanings, not just one (that may or may not

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be there, as in the case of digital signals). Insisting with the example of the threatening signal, we could say that analogical signals may tell the receiver that the sender is puzzled, irritated, angry, very angry, furious, ready to fight, ready to kill… Having said that, it must be also added that the distinction analogical/digital pertains to the stages of perception and reaction in the receiver, and not to the emission of the signal: only machines are capable to deliver signals and data that are truly digital. Otherwise, living beings can only emit analogical signals, which sometimes may be perceived and categorized in a digital manner (on/off, yes/no, up/down, etc.).

Context and dynamics Sometimes, signals evoke a fixed, specific response (in which case they are called *releasers*). But most of the times the meaning of a signal depends on the context. Time, place, subjects involved, and so on, make it possible that the same signal means A in one situation and B in another.

Interception There are also circumstances where signals are encoded or decoded in slightly unusual manners. For instance, in a conventional sense, communication is a process that produces an advantage for the sender, first and foremost, and then it may or may not produce it for the receiver. However, when the emitted signal produces an advantage only for the receiver, we have a case of *signal interception*. This phenomenon occurs in several contexts, including intraspecific and interspecific competition, predator-prey relation, parasitism, etc. In signal interception, the specimen who receives (“intercepts”) the message is actually not the one whom the sender wanted to communicate with, but a third subject, who indeed benefits from “meddling” in someone else’s business. A typical case is that of territorial fish species: the male sends courtship signals to the female (the “designated” receiver), but another male (the “interceptor”) gets the message too, and enters in mating competition with the sender. Other frequent cases include intraspecific signals of a given species that are intercepted by a predator of that species, who therefore get useful information on the presence of a potential prey. And so on. Among other things, the existence of this phenomenon is one important reason why very “extrovert” (visible, loud, etc.) exhibitions in intraspecific communication are performed in limited time and limited

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contexts. The goal is to diminish the chances of being intercepted (and therefore damaged in some form).

Self-promotion Signals can also be used for conveying specific information about the sender. Signals that indicate the sender’s position, identity, characteristics and else are called *self-promotion signals*. For example, several forms of singing, as appearing in birds, insects, amphibians, are forms of selfpromotion signals, in that they inform about the singer’s species, sex and individual qualities. A particular case of self-promotion signal is the *social status signal*, that informs co-specimens of the sender’s position in the group hierarchy. Several species, for instance, present visual displays to identify the dominant subject/s in the respective communities: a white-silver color on the back for the Mountain Gorilla Gorilla beringei; a dark plumage under the beak for the Zonotrichia querula, and so forth. Self-promotion signals, in a way, are also the signals emitted by young or very young subjects. *Infant signals* have the primary goal of letting the adults know that the sender is indeed an infant, with all that this information implies (the subject is weak, not self-sufficient, etc.). Generally, an infant is not recognized by a single characteristic, but by a combination, an “overall appearance” that Konrad Lorenz named *infant scheme*, and which includes such features as a certain morphology of the head (proportionally bigger than an adult’s), certain behavioral patterns, etc. For evolutionary reasons, the recognition of a subject as “infant” (whether or not co-specific) tends to inspire a more compassionate behavior in the observer, as if “it would not be fair” to take advantage of the situation. Such, for instance, might have been the case with human beings, when they became hunters: a certain (not systematic, but significant) tendency to spare infants from their preys was surely instrumental in preventing the latter from premature extinction, ensuring a wise exploitation of the resources.

Ritualization Almost all signals evolved from basic patterns that were not born with a “communicative” intention. These patterns were simply phenomena that were happening naturally, as a consequence to given stimuli (muscle movements, physiological processes…). At some point, in the evolution of a species, these patterns underwent a process of *ritualization*. That is,

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they were modified and stereotyped, becoming something that “signify” a certain meaning, not simply that is “produced” by it. In Chapter Two, where we also mentioned the question of ritualization, we were using that signal that canids display when they are reacting to an intruder: the pattern of growling, the notorious grrrrr. Now, there must have been an initial point, in canids’ evolution, in which growling was simply a natural consequence (a reflex) of distress and anger. At this point, the pattern was not a form of communication, it did not “stand for” or “represent” anger: simply, it was “part” of being angry, just like yawning is part of feeling sleepy. A bit later (hundreds, or maybe thousands of years later), some dog ancestor must have, so to speak, realized that growling alone was often enough to intimidate the intruder. Why then engage in a fight, if that pattern was perfect to “give the impression” of wanting a fight? Growling, at this point, became “a ritual”, a threat detached from the fight, in that the fight could or could not occur.

Complexity The process described above was, in other words, a more complex form of signal, that (in disciplines such as linguistics, semiotics and others) is generally referred to as a *sign*. A sign, in a classic formulation provided by philosopher St. Augustine, is an aliquid pro aliquo, something that stands for something else. To introduce a distinction between “signals” and “signs” may or may not be useful for the purposes of this book. Let us just say that signals can range from very simple and invariable to very complex and flexible, and a process of ritualization is certainly a form of complication of signals (up to making them become symbols, as we shall see later). In any case, it is clear that communication is a mediated phenomenon, it needs a currency, as we said above, something that both sender and receiver trade for representing what they have in mind. Sounds, gestures, visual displays, words… these are all forms of currency that are used to signify a given event/circumstance/state. The word “flower” is not the actual flower, but a way to represent it. So is its smell, which is a property that indicates the presence of a flower. So is something “looking like” a flower (an experiment made with honeybees showed that if they are exposed to different geometric figures, they are most attracted by those with many corners, shaped like stars, that indeed remind of flowers). The implication is that there are different forms of relation that a signal establishes with its meaning. Exploring at least the most important ones is a good way to realize how rich animal communication is (and perhaps also

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to understand that human beings are not the only species that communicates in very varied manners). In this monograph, we shall explore five different types of signal-meaning relation: Indexes (or Indices), Icons, Symbols, Syntax and Names. Definitions and concepts of these relations were mostly provided in the fields of Philosophy of Language and Semiotics, by the likes of Charles Sanders Pierce and Thomas Sebeok.

Indexes An *index* is a signal that appears as a “natural consequence” of its meaning. The presence of smoke is an index for fire. Smoke is not fire, yet it “means” fire. A toothache is not a cavity, but an index of it. And so on. Now, as one can easily imagine, indexes are the signals that the vast majority of animal communication is concerned with. A great number of markers, symptoms and indicators produced through any of the available channels are in one way or another indexical, from the tail-wagging of a dog (index of its excitement) to the wheel display of a peacock (index of its health conditions). Let us not make the mistake of inferring that this high recurrence of indexes is an indication that mental processes do not play a major role in animal communication, as, after all, an index is most often a natural reflex, an instinctive reaction, a physiological display. This conclusion would be as inaccurate as saying that a human display of joy as the smile is alien to any cognitive implication, ignoring that elaborate path that goes from joy to smiling and that includes the reasons for joy, the specific causes of a smile, the frequent presence of receivers of the “smile” message, the different degrees of display (a smile can be emphasized, hidden, or else, depending on the situation and on the subject), and others, that obviously require the type of mediation that a mind can provide. The same applies to a similar non-human manifestation of joy as the tail-wagging of a dog, who has the same cognitive ‘problems’ to solve, and normally provides an analogous range of solutions. What instead is true is that, although requiring mental processes in most cases, indexes do not necessarily require complex and multi-layered mental processes, and that is primarily because an index tends to require temporal presence and spatial continuity, and its encoding is strongly bound to the hic et nunc of communication. The decoding may still happen later or further (e.g. some chemical signals may be carried by the sender for days and kilometres), but, what we might call the “packaging” of the

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signal is an immediate process that does not require too much cognitive articulation. Finally, it would not be entirely true to say that the number of indexes in animal communication is higher than in human communication, not least if we interpret this statement literally (i.e. non-human animals use more indexes than humans), perhaps thinking that human language was able to replace indexes with more efficient symbolic descriptions. What has happened is that human language added a series of communicative and cognitive elements on top of the existing ones, not in place of them. Indeed, natural communication is still intact in human beings, and goes on roughly to the same extent as it used to before language appeared. What has crucially changed is the decoding stage: precisely because human language is admittedly a very efficient strategy for informing (or deceiving) each other about several topics and conditions, human beings learned to pay attention mostly to that, losing – or better: stiffening up – their abilities in non-verbal communication.

Icons *Icons* are signals that bear some “physical resemblance” with their meaning. That is, they may “look like” it (like a portrait looks like the person it represents), “smell like” it (like rose water smells like a real rose), etc. In one word, the signal imitates the meaning in some form. As a phenomenon, iconicity is slightly less recurrent than indexicality in animal communication, but still very easy to find. The many fascinating forms of *mimicry* are normally the first category of examples mentioned. In a more general sense, a great deal of signals related to deception is of iconic type. The various forms of imitation, however, do not serve exclusively deceptive purposes: the enormous area of birds’ vocal imitations, for instance, qualify in most of the cases for iconicity (and quite often for symbolicness, too, as we shall soon see). Other examples of iconic signals are recurrent in play (for instance, when a canid intends to mock-fight, s/he will not rarely offer an iconic representation of someone sleepy, or dead – therefore defenceless – by lying on his/her back), in ritualized social behaviour, in aesthetic signals, and in several other situations.

Symbols *Symbols* are signals that relate to a meaning only by convention. They are not similar, they are not caused, they do not relate to their

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meaning if not in an arbitrary sense. That means that, in order to understand each other, both sender and receiver have some kind of “agreement” that associates a given signal to a given meaning. Such an agreement can be established in different ways, including communication itself, habit, and learning. The concept of symbolicness has for a long time been considered species-specific of human beings, mostly because it was maintained that only human language possesses the characteristics for, indeed, “establishing an agreement”. On the contrary, evidence of symbolicness appears already in insect communication. For example, in the dipterans of the family Empididae, one of those species that practice cannibalism, the male, before the copulation (the act following which he is normally eaten by the female), offers the female an empty balloon. Biologists have unraveled the evolutionary origins of this gesture, noticing that it originated from an actual “food” offer that was eventually ritualized. The basic reason why food was offered (“eat this, not me!” might have said the male) was lost, and the idea of a “symbolic gift” took over: something was offered anyway in order for the male not to be eaten (the “meaning”), but this empty balloon (the “signal”) is not related to that message if not in an arbitrary way. Another example, often used by ethologists to describe symbolic communication, is that of the dance of the honey bee Apis mellifica (which we shall deal with in Chapter Five), a phenomenon that is in fact often called language. Among the evidence they bring in support of this view, a rather interesting example is the geographical variability in the significance of the various signals. Different communities of the same species Apis mellifica may for instance use the very same articulation of the distance-signal (that is, the time employed to cover the middle axis of the 8 figure) to represent completely different distances. Let us insist a bit more on symbolicness, because it is a feature of communication that many regard with skepticism when it comes to nonhuman animals. Two more examples. One is, of course, the singing activity of many bird species, which presents features of cultural learning and development, geographic dialectal variation, and – most of all – conventional and abstract signal-meaning codification, that makes it hard to consider these songs as merely composed of indexes and/or icons. The other example, perhaps more interesting as it is less obvious, is the system of alarm cries developed by species like the ground-squirrels of the genus Spermophilus, the prairie dogs Cynomys, the monkeys Cercopithecus and others, to inform their co-specific of the arrival of a predator. These

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species emit differentiated cries depending on the type of predator, and it is highly specialized in the case of the vocalizations of the Vervet Monkeys Cercopithecus aethiops and of the Greater Spot-nosed Monkeys (Cercopithecus nictitans). When one vervet monkey sees a predator, s/he produces a vocalization that alerts the rest of the group. This type of alarm is of three different kinds, and depends on the specific kind of predator in the vicinity. A loud barking call is given for leopards (therefore, it means “leopard”, or whatever the monkeys call that animal), a short, double syllable cough stands for eagles, and a “chutter” sound stands for snakes. The calls are not similar to the sounds that those predators utter or produce, they are therefore not of the iconic type, and the response of other monkeys to a given call is appropriate for escaping the corresponding predator (i.e. when the leopard call is heard, the monkeys run to the trees; when the eagle call is heard, they look up into the air and seek shelter; and when the snake call is heard they stand up on two legs and look in the grass). In many communities, in addition, specific cries were also noted for two more possible predators, which evidently the vervet monkeys perceive as rather similar to each other: baboons and human beings (no offence meant, hopefully). Once again, it is the strong dialectal variation (plus the fact that the cries are learned, and it takes a while before the young specimens provide an adequate response to them), to persuade scholars of the symbolic character of this catalogue of signals. At a distance of few kilometers, in another community, a leopard is signaled with a completely different cry. One may argue that the dialect variation does not exclude a priori the possibility that these cries remain of indexical or iconic type. However, the notion can be accepted only if fairly traded with the hypothesis that most of human language, too, is therefore only indexical or iconic, despite the existence of different languages and dialects. If we take the example of onomatopoeias, it is clear that the creation of such words stems from iconic logics. Yet, words describing the same acoustic phenomenon can be completely different, depending on the language, as in the case of animal vocalizations (a dog goes “Woof” in the UK and the USA and Australia, three geographically separated places, yet s/he goes “Bau” in a country, Italy, which is much closer to the UK than any of the other Englishspeaking countries). Or, alternatively, we could accept the hypothesis that no signal is actually 100% symbolic, as a link (even a remote one) with a physical entity is sooner or later detectable: even street signs themselves, which are normally taken as the most extreme example of symbolicness, show different degrees of this relation (an arrow remains after all iconic, to some

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extents; the use of the red color for interdictions and dangers is easily related to the recurrence of this color for blood and poisonous substances in nature, etc.).

Fig. 2 – Indexes, icons and symbols

Syntax An even more complex feature that we might find a bit surprising, as applied to non-human animals, is the existence of forms of *syntax*. In general, scholars distinguish between Phonological Syntax (a combination of signals that, taken alone, do not necessarily have a specific meaning, or at least not a qualitatively different one from the combination), and Lexical Syntax (a combination of signals that are also meaningful as single units, and that also mean ‘something else’). Predictably, no particular effort is required for a language scholar to find examples of phonological syntax in non-human animals: the examples of wedge-capped capuchins (Cebus olivaceus) and male gibbons were widely studied already in the 1980’s. The difficult task is to find evidences for lexical syntax. In the insects Hymenoptera, already, we have an example of this kind. Pheromones carry a given message if emitted singularly, and another one

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if presented together (that is, a meaning that is not simply the sum of the single pheromones. Exactly like blue plus yellow makes green, an entirely different color). However, one may object, the communicative potential of *pheromones* is so little flexible that there is simply no alternative: if one wants to convey more than one message with pheromones, the only chance is to add more of them, making different quantities signify different things. Imagine to be forced to communicate only by writing a “X” on a piece of paper: what you will do is creating a little code, where one X means – say – “Hello”, two X mean “Yes”, three X mean “No”, etc. More interesting (because there are alternatives, in this case) is the case of the Greater Spot-nosed Monkeys. Like vervet monkeys, these animals too have alarm sounds for signaling predators. In this case, there are two of them: something like ‘pyow’, which designates a leopard, and something like ‘hack’, which stands for “eagle”. The use of these sounds proceeds in the same vein as with vervet monkeys. Recent research, however, (Arnold–Zuberbühler 2006) has revealed an interesting phenomenon with regard to the Greater Spot-nosed. Apparently, they form sequences with these sounds (sort of “pyow pyow hack hack”), sequences whose meaning is not simply “A leopard and an eagle” (or two leopards and two eagles), but mean something entirely different, that is, “Let us move to another place”. This is clearly a case of lexical syntax, where existing meaningful units are combined in order to produce a new meaning, that is not simply the sum of its constituents. Keeping up with the hypothesis that languages are never fully symbolic, one might guess that this new meaning developed from an indexical formulation of the type “eagles everywhere, leopards everywhere”, which syllogistically turned into a kind of “as dangers are everywhere, this is definitely not a safe place”, and refined into the more practical “let’s go somewhere else”. This type of interpretation can reasonably be applied to all languages including the human one, and can easily explain why, for instance, we went from the red of blood and poison to the use of red colors for warnings and interdictions in street signs.

Names The last category of signals here analyzed is that of the *names*. A name can be defined as a signal that identifies individually a subject or an object (humans and other animals, of course, but also events, things, and other inanimate or abstract entities). Normally, the giving of a name has to do with an arbitrary, symbolic procedure (if a person is named Karl, it is not because he possesses some physical property called “Karlness”), but

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there are also indexical properties (the fact that a name “refers to” a subject, and it is very often a consequence of ethnic origins – making most of the times “Karl” a name given to a German, “Carlo” a name given to an Italian, etc.) and iconic properties (in those cases where the name “portraits” some characteristic of the subject – this being often a case for pets, whose names are often onomatopoeias of the sounds they make). Given such properties, can we also advance the hypothesis that nonhuman animals name themselves/each other (rather than being simply “given” names by humans)? Once again, the answer seems to be “yes”, at least when it comes to birds and cetaceans, the two categories that have received the most attention in this respect. Very interesting is the case of Bottlenose dolphins (Tursiops truncatus), to which specific attention has been devoted in a number of investigations (Janik et al. 2006). Dolphins apparently possess individually-distinctive signature whistles that are used to maintain group cohesion. While identification signals are present in many other species, dolphins’ names have the characteristic of being influenced by vocal learning. By using a sample of 14 dolphins at Sarasota Bay, Florida, a group of researchers recorded and then played the signature whistles to the animals, who recognized their names, and did so (in 9 cases out of 14) also when they heard an artificially-synthesized version of the whistle. The conclusions of the research team are that dolphins extract identity information from signature whistles, even after a complete artificialization of the signal, where all voice features are removed from the signal and only the time-frequency relation is preserved (which corresponds more or less to a complete stylization of a colored painting into a simple pencil contour drawing), showing that dolphins seem to be a very rare case of animals, besides humans, that transmit identity information regardless of the caller’s voice or location. In other words, dolphins “recognize” each other through their names. Therefore, “to recognize” is something different from to simply “discriminate”. Recognition means to perceive something as identical with something previously experienced. Discrimination may, but does not require to, use such experience, and it is based solely on detecting differences between two or more stimuli. In other words, discriminating implies that one can tell that A and B are different from each other. Recognizing implies understanding who A and B are. Finally, the choice of the personal name, which is up to the carrier him/herself and occurs during the first year of life, is the result of a combined imitation of several other names the young dolphin is exposed to – more or less as if a woman chooses the name Lina because her uncle is named Linus, her friend is called Tina, and her teacher is called Linda.

CHAPTER FOUR CHANNELS AND FUNCTIONS OF COMMUNICATION

As we humans know very well, communication does not only occur with words. All our five senses fully intervene in making our communication rich and varied: we detect information and produce signals by seeing, hearing, touching and so on. In other words, communication occurs through sensory modes, that we call *channels*: for instance, when we make or hear a sound, we are using an “acoustic” channel for communicating. It would, however, be a mistake to think that channels merely “correspond” to the five human senses. As a matter of fact, there are at least eight channels we have knowledge of: olfactory, gustatory, thermic, electric, tactile, acoustic, visual, magnetic (we shall see, however, that it is more practical to gather them in five groups). An analysis of the channels includes their classifications, their constitutive parts, the occurrences they operate in, the species making use of them, and their characteristics.

The chemical channel The most elementary and common channels are the olfactory and the gustatory, which can be grouped under the single label of *chemical channel*, as they both use chemical substances as vehicles of specific or general messages between two or more organisms. There are species who have a clear distinction between taste and smell, and others who do not, and basically have the same receptor organ for both functions (as a matter of fact, there is always a close collaboration between these two senses: in humans, the taste of food is always provided by a combination of the olfactory and the gustatory channels). The chemical channel is also the most ancient mode of communication ever, in any form of life. The earliest forms of bacteria communicated when one cell released a chemical that was picked up by a receptor on the

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membrane of another cell. That was the very first exchange of information that appeared on this planet, long before anything else. Among the most important messages that is possible to convey through chemicals, we must count territorial marking and defense, mating, and some forms of description, like for instance showing the location of a food source. As we mentioned in Chapter Three, the chemicals that send messages are called pheromones, and are present in fish, insects and most mammals (more in Wyatt 2003). Birds do not seem to communicate through pheromones. Whether humans use them or not, it is still a debated question: it has been shown that we possess the proper gene (so-called V1RL1, present also in rodents, who on the contrary largely rely on pheromones for their communication), but it is not clear whether we make use of it, or in fact totally replace it with other cues such as visual ones. However, and obviously, that does not mean that we do not produce or detect information in a chemical way. Smelling a person can reveal a great deal of messages, as we know very well by positive or negative experience. There are three main advantages of the chemical channel: it is relatively permanent, it can act at relatively long distances, and it can be decoded at a given time distance. In contrast, this channel has a slow variability and a high susceptibility to interferences. The territory signal emitted by dogs through their urine is a perfect example of these pros and cons: the message is quite permanent, it can be smelt at a certain distance, and can be decoded after several days by other dogs. At the same time, there is not much information spread (except a “this is my land!” kind of message), and the next dog will easily invalidate the message by urinating over it.

The tactile channel Another important channel that we can use for grouping a few of the existing ones is the *tactile channel*, that includes also the thermic and the electric one. The messages transmitted through these channels all occur by direct body contact or by vibrations produced in the close vicinity of the receiver (see more on Chapter 5 of Halliday 1998). Of the three, the human animal possesses the tactile, and – possibly – the thermic channel (in the sense that a certain degree of information is conveyed also through body temperature). Vibrations also produce sounds, so there are instances in which tactile communication overlaps with (or is reinforced by) the acoustic one.

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The main advantage of this channel is variability in time and intensity, which is indeed very high (animals can touch each other for all kinds of purposes and in all kinds of ways). The main disadvantage is the total absence of distance transmission: unless the sender and receiver are at touching distance, no message can be transmitted. The case of thermic communication, widely documented in ecosystem, is still not empirically studied among animals (although it appears obvious that body temperature is a carrier of many different messages), while for the electric one, there are several examples among fish families, like the Mormyridae, the Gymnotidae, and, famously, the Torpedinidae. Through the electric signals, besides predatory purposes, these fish can exchange a very high variety of messages, including detection of objects, social status, conflicts, courtship and others. Species like the Atlantic torpedo, Torpedo nobiliana, which is the biggest Torpedo still in circulation (it can reach up to 90 kilograms of weight), is capable of delivering a 220-volt electric shock. A very good and significant example of tactile communication is the so-called *allogrooming*, or social grooming. It is that pattern, generally performed by two specimens, of carefully removing dirt, parasites, or specks of other matter from each other’s fur, skin, etc. It has an important social function, in that it reinforces the relationships between group members, and helps maintain the hierarchy in a group. In certain cases, allogrooming occurs even if there is no specific need to clean each other: it is another case of ritualized behavior that is specifically performed as a “friendship” signal.

The acoustic channel The sensory mode connected with the production, emission and reception of sounds is called the *acoustic channel*. Acoustic signs can be produced through the vocal apparatus, or through parts of the body or the environment (by hitting them, making them vibrate, and so forth). Every species can perceive a specific range of frequencies, and that (besides determining the potentials of acoustic communication in each of them), creates also several *soundscapes* in nature: not everybody can hear everybody else. As humans, for instance, we can perceive a range of 20 to 20,000 Hz: anything below and above is not audible to us (and it is called respectively “infrasounds” and “ultrasounds”). To make a few other examples dogs go from 15 to 50,000 Hz, porpoises from 75 to 150,000, bats from 1,000 to 120,000, mice from 1,000 to 91,000, and cats from 60 to 65,000.

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Forms of communication that adopt the acoustic channels include all kinds of utterances, calls and songs (mating, territorial, social…), and somehow they ended up being interpreted as quintessential forms of communication. Of course, this is not the case, but it is undeniable that a great deal of information goes through this sensory mode, which – as we shall see now – has great potential. There are indeed many advantages of the acoustic channel. Among them are the broadcast transmission (i.e. the possibility of communicating in darkness, around corners, etc.), directional reception, rapid fading (which makes a fast continuation and response possible, thus increasing the communicative potential), immediate feedback, and a very high degree of specialization (as the example of language, developed first of all as a vocal-acoustic sign system, clearly demonstrates). The main limitation of the acoustic channel lies in the rapid fading, which – besides its aforementioned advantages – also implies that a message does not last in time, and must therefore be replaced by other means of communication (typically, the chemical channel) in those cases where an enduring message is needed.

The visual channel The sensory mode connected with the production, emission and reception of visual signs is called *visual channel*. From a cognitive point of view, it perhaps represents the main perceptive vehicle for the human species, even though it is hardly the best-specialized (we do not see as well as many other species, yet visual signals are the most common form of communication in human societies). Visual signals can be divided into extrinsic and intrinsic. An *extrinsic signal* is produced in the organism’s environment (tracks, nests, traces, etc.), while a signal that is part of the organism’s body or behaviour (orientation towards the receiver, body shape, colour, movement patterns, etc.) is called *intrinsic signal*. Several forms of animal communication are to be considered intrinsic visual signals (for the same reason why a lot of human nonverbal communication occurs through gestures, facial expressions, postures, and so on). The body and the behaviour of an animal can convey a great deal of information, including specific forms of social or individual status (e.g. the predisposition to mate could be revealed by a change in color in the animal’s body; or a high position in the social hierarchy may imply that certain gestures/actions are allowed only to a dominant specimen). The main advantages of the visual channel are the specialized orientation in space, and the great (virtually endless) quantity of

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information that may be conveyed. The biggest limitation is the need for light (which, contrary to common belief, is a handicap also for nocturnal species, whose perceptive qualities are anyway limited in comparison with daylight conditions).

The magnetic channel To conclude, relatively little is known about the *magnetic channel*, that is, that kind of sensory mode related with the perception of magnetic fields. First and foremost, as one can detect from the definition, this channel applies only to the field of reception of information, not the production. In this case, we speak of *magnetoception*, the capacity to use magnetic fields to gather information about direction, altitude or location of given objects or places. Magnetoception is important, if not central, in the navigational skills of many animal species, particularly migratory ones. It has been observed in many birds (first and foremost), and in flies of the family Tephritidae, honeybees Apis mellifera, turtles of the species Caretta caretta and Dermochelys coriacea, lobsters of the family Palinuridae, crocodiles of the family Crocodylidae, sharks of the superorder Selachimorpha, cartilaginous fish of the order Chimaeriformes, stingrays of the family Dasyatidae, common cows Bos primigenius, and deers of the family Cervidae. In humans, too, the presence of magnetic deposits of ferric iron in the ethmoid bone, allows, to a certain extent (not fully explored, yet), magnetoceptive abilities. Given its peculiar nature, however, the question remains open whether magnetoception, a sensorial ability that seems to function only in the phase of reception of information, should or should not be considered a channel in all respects. In conclusion, just to have a little summary of the pros and cons of the four main channels (we shall keep the magnetic one out, on this occasion), here is a little table to check how well or how bad each of them performs in their main tasks: RANGE SOURCE LOCATION GOING AROUND OBSTACLES SPEED PERSISTANCE

VISUAL Good Very good Poor

ACOUSTIC Very good Good Very good

CHEMICAL Very good Variable Very good

TACTILE Poor Very good Poor

Very good Poor

Very good Poor

Poor Very good

Very good Poor

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The functions of communication What is communication for? How do we use it? What are the contexts in which it is employed? To answer these questions means to understand and classify the “functions” of communication. There are of course many ways to do it. For instance, William Searcy and Stephen Nowicki (2005) have organized animal communication in three main groups: 1) when interests overlap (e.g. food calls), 2) when interests diverge (e.g. mating signals), and 3) when interests oppose (e.g. aggressive signals). A more thorough classification, which we shall use in this book, comes from the study of human communication. In the late 1950’s, the Russian linguist Roman Jakobson (1960: 353-7) formulated a theory on the functions of communication that became a sheer “standard” in the different fields of communication studies. Almost half a century later, an Italian semiotician, Felice Cimatti (1998: 59-105, unfortunately in Italian only), noticed that the very same model could also be very useful for interpreting animal communication. Jakobson classified six main functions within a communication system, each concerned with a particular element of the process of message production: expressive (referred to the sender of the message), conative (referred to the receiver), phatic (referred to the establishment of contact between sender and receiver), referential (referring to the context of the message), metalinguistic (referring to the code used in a given communication act), and aesthetic or poetic (referring to the form of the message). None of these functions is to be considered in isolation or as mutually exclusive. A message is usually the intersection between two or more functions, even if one of them is often dominant and more evident.

The expressive function By *expressive function* is meant the use of communication in which the most relevant part is the display of the emotional state and the identity of the sender. All kinds of messages more or less display some feelings or emotions: it is sufficient to look at the way we humans talk, or to take a look at our pets, to realize how crucial this component is in communication. All animals, when delivering a message, are not only conveying a given content, but they are also saying something about themselves. A word like “Hello!” does not just greet somebody: it can assume endless nuances related to the way we feel at that very moment. It can be the casual “Hello!” to a colleague, a heartfelt one to somebody we have not seen for a long time, a sarcastic one to somebody who is very late

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to an appointment, and so on. All these nuances represent the “expressive function” of communication. An interesting example is that of the Australian magpies Cracticus tibicen, performing their whole repertoire of songs throughout the entire year. For this species, singing is a ritualized way to give instructions about food, migration, territories, etc. Yet at the same time, and according to intensity, speed, loudness and other parameters, these songs are also an indicator of anxiety, joy, alarm, enthusiasm, attention or other emotional states (Kaplan 2009).

The conative function By *conative function* is meant the use of communication in which the sender uses the message in order to make the receiver have a certain reaction and act consequently. A sentence like “Close the window, please!” is conative because it implies that we want our interlocutor to react in some form. Conative functions stand out clearly in all mating and all territory signals, and that, of course, means a great deal, because it implies that a very large portion of the communication that occurs in nature is of conative type. The sender wants the receiver to do something (mate, leave a territory, etc.), no matter how ritualized such signals may be. Conative are usually also those signs used for deceptive purposes, and in general all those aimed to some form of manipulation of the receiver. It may be curious, but also very significant, to notice that the traditional definitions of animal communication, as provided by classical ethology, tended to emphasize only the conative aspect of sign exchange: “Communication between animals involves the giving off by one individual of some chemical or physical, that, on being received by another, influences its behaviour” (Frings and Frings 1964: 3); “Animal communication evokes a change of behaviour in another individual” (Cullen 1972: 103); or “Communication occurs when an animal, the actor, does something which appears to be the result of selection to influence the sense organs of another animal, the reactor, so that the actor’s behaviour changes to the advantage of the actor” (Dawkins and Krebs 1978: 283). That, possibly, shows how important the conative function is, within the whole economy of animal communication.

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The phatic function The *phatic function* refers to the use of communication aimed at establishing and/or keeping contact between sender(s) and receiver(s). An expression like “Good evening!”, besides being a more or less sincere wish that our interlocutor indeed has a nice evening, is most of all a way to establish a contact with him/her. Through this signal, we started “interacting” with this person, and now we have his/her attention. The phatic function is probably best illustrated by the way Konrad Lorenz was famously “translating” the ducks’ calls: “I am here, where are you?”. That call was a constant search for reassurance that, indeed, the contact between them and him was still operative. To communicate in a phatic way does not only mean to “establish” contact, but also to keep or even reinforce it. An example of this more complex kind concerns wolves, Canis lupus, and other Canidae. After hunting and rallying together as a pack, wolves act very excited and joyfully howl together. The meanings of this practice are of celebrative type: celebrating the success of the hunt (i.e. the presence of food), the wolves’ safety, and the joy of reuniting after some time of separation and difficulties. In addition, howls serve the purpose of practically rallying the wolves within a pack, before and after hunting, and of preventing packmates from wandering off on their own. This range of meanings somehow covers the entire spectrum of a message’s phatic function (and of course, of the conative, aesthetic and expressive ones, too, as we shall see later, but it was in any case anticipated that it is only very rarely that a function appears alone). In howling, one finds the most simple (the establishment of a contact) and the most complex (the strengthening of comradeship) extents of phatic communication. In particular, the act of celebrating the reunion of the pack is a most revealing example of how close cultural, ritual, aesthetic and phatic elements, gathered in a single behavioural pattern, can be. Similar conclusions can also be made for such patterns like the previously mentioned allogrooming, and in general for all those displays of kinship that characterize all those species with a complex social organization.

The referential function The fourth function of communication systems is the *referential function*. In this case, signals refer to the context, to the surrounding environment, i.e. including the sender. Referential signals are thus descriptive, and may refer to places, times, people, sensations and so on.

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“My name is Leonardo”, “Franz Kafka was born in 1883”, and “It’s summertime, and the living is easy” are all referential messages. Referential communication includes some of the classic cases in animal communication studies, such as the dance of the honey bees (which we shall see in detail later on, in this book) and the alarm calls of vervet monkeys, described in the previous chapter. It can also be added that several displays of an animal’s health, identity, social status, etc., share an expressive and a referential dimension. Finally, several birds use acoustic communication in a referential fashion for showing migration routes. A few others, like the Herring Gull Larus argentatus and the Greater Honeyguide Indicator indicator may use sounds also to indicate sources of food. The latter case is also a very interesting one, because it shows that a referential form of communication can provide a great advantage for survival also at interspecific level. The honeyguide, indeed, is not only eager to guide its fellow specimens to a food source (a beehive, from which this bird eats all sorts of things, including wax), but can also do that with other species, such as the Honey badger Mellivora capensis (the names of both species suggest a common food interest), creating an effective form of *mutualism* (on this topic, see also Hoeksema-Bruna 2000). The badger gets to know the location of a beehive and can have its honey, the honeyguide spares the (often painful) trouble of emptying the hive and gets all the badger’s leftovers (some honey and all the wax, which the badger does not eat).

The metalinguistic function The fifth entry in Jakobson’s list is the *metalinguistic function*. In this case, signals are used to refer to other signals, as occurs in playing, deception, or several forms of ritualization. In playing, particularly, the adoption of metalinguistic signals is crucial, in that several forms of play are nothing but unserious imitations of “real life” situations, most of which are rather dangerous, such as hunting or fighting (we shall see this in Chapter Seven). The signals of aggression, therefore (running towards the receiver, displaying anger, etc.), must always be accompanied by other signals, which refer to the former, indicating that all seemingly serious actions taken from that point on, must be interpreted only in a playful way (for instance, a dog might physically attack another dog and wag the tail at the same time, thus showing a friendly disposition). For this reason, the *code* that the animals are using on that particular occasion is the actual focus of metalinguistic communication (we spoke about codes in Chapter Two). The difference between a serious attack and

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a playful one is not in the two signals themselves (which may easily be identical), but in the fact that the former is performed within the “code” of aggressive signals, and the latter within the “code” of playful signals. How does the receiver understand that this time the code is playful? Exactly through the other, metalinguistic signal, the tail-wagging signal, that reveals that all that is being done in that moment is playful and not serious. Another very interesting example comes from the vast catalogue of sound imitations (particularly through singing). The imitation of another species’ or group’s singing style is an iconic (and generally referential) sign of that style, but at the same time another important intention is again playful: the animal finds it pleasant and challenging to imitate another specimen. In other cases, imitation serves very precise functions, one of which is related with territoriality: by imitating a potential enemy belonging to another species (including predators), the imitator pretends to belong to the species imitated, and practically “says” that the given territory is already occupied by a member of the potential enemy’s species. Finally, imitation in singing may also recall the – very metalinguistic – use of columns in medieval cathedrals. Not only do they hold the church up, but variously styled columns coming from different places, placed throughout the cathedral, would serve ostentatiously, as a sign of richness. As the mating season approaches, the male of the Superb Lyrebird Menura superba, whose usual mating task is to impress the female, adorns his singing with a huge set of short imitations of different species and objects. The rule is “the richer the better”, thus signs made for totally different reasons (like a humanly-produced sound) are here used metalinguistically as a display of the creativity and musical skills of the Lyrebird.

The aesthetic function Finally, now singing has been mentioned, it is also clear that several signals related with mating (and not only with mating), serve an *aesthetic function* in communication, that is, they concentrate on the message itself, elaborating on its representation and form. Any kind of message can be produced in more than one manner, even if the contents are exactly the same. What can make a difference is the form of this message, the “way” we say it. To pay particular attention to the form means once again to somehow ritualize the message, make it more (or also less) appealing, more (or less) ambiguous, more (or less) direct, and so on. In fact, as it is probably clear by now, using communication in an aesthetic way also means bringing the message into an “artistic” dimension. We can say “I used to be happy”, and that is one

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thing, but we can also say “Yesterday, all my troubles seemed so far away”, and that is something else, even though we have basically expressed the same concept. Yet, in the second case, we have produced something more artistic, and this was possible thanks to a message that was more elaborated in its form. When we think of mating, we think exactly of a situation where form can make the difference. The courtship process requires a lot of “art” and “beauty”: a shining plumage, a complex song, an articulated dance… The Hungarian semiotician Thomas Sebeok (the very scholar who invented and elaborated the paradigm of zoosemiotics) suggested that in animal communication we can find the rudiments of art itself, and proposed the classification of aesthetic signals into four categories (Sebeok 1981): architectural (related with the construction and the decoration of artifacts), pictorial (related with the material production of visual signals), kinaesthetic (related with bodily movements) and musical (related with the production of sounds). To make just one example out of many, the Sandhill Crane Grus canadensis conveys a few messages (including mating, other social relations, territorial claims and play) through an articulated and charming dance, often accompanied by loud and repeated sounds. Both sounds and movements contrast with the normal behavior of cranes, which is very cautious and quiet: dances, on the contrary, represent an explosion of energy articulated in dozens of different carefully-elaborated steps.

Fig. 3 – Features and functions of communication, according to Roman Jakobson

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A cognitive function? As a bonus, it may be worthwhile to mention a hypothetically seventh function of communication, not suggested by Jakobson in his model, but possibly useful to trace a more accurate picture of the role of mental processes in communication. We may call it *cognitive function*. It relates to how an animal’s inner and outer behaviour changes in consequence of the possession itself of a communication system. To be able to communicate, indeed, is not only a useful tool for interacting with other specimens: it also implies a progress in our mental capacities: in order to convey a given message, indeed, we need to think about it, isolate it from the rest of the possible messages, relate it to some element of reality, perceive and classify that element, etc. It is a complex process that calls into question several aspects of our brain activity, and – as such – it is fair to think that communication serves a specific function in this respect. In other words: communication affects us in our entirety, as much as we affect communication. Let us take again the example of the monkeys’ alarm calls that we discussed in Chapter Two. When danger appears, the monkey’s mind is probably activated in two ways: on the one hand, the mental representation of the perceived object (what kind of danger, how close, how fast…); on the other hand, the activation of a signal, i.e. that specific alarm call that stands for that specific danger. Here one already notices a “change” in the animal: if the monkey did not possess a communication system, its reaction would have been of another type (e.g. escaping from the danger). In addition, a second characteristic of the cognitive function is that the call does not only act as a referential signal describing the danger, and as a conative signal for the monkey’s community, but it is also directed to the sender itself, in that it increases the self-control skills of the animal (who is now expressing its emotional status and guiding its own behaviour). In other words, not only the mental complexity of an animal functions as source for communicating. The process works the other way round too: the richer and more complex the communication, the more sophisticated the mental life. The active participation to communicative processes opens up for further – and increasingly articulated – cognitive abilities.

CHAPTER FIVE THE COMPLEXITY OF COMMUNICATION: MULTIMODALITY AND THE LANGUAGE OF THE BEES

When we take communication in a superficial way, we usually tend to think that every time we communicate, we emit one message per time, maximum two (for instance, when we are playing or joking we know that we often need a serious message along with an unserious one: we call a friend in a not very complimenting manner, but we do it while laughing, so that our friend knows that we are joking). Yet, the cases when only one message (or even two) are being produced are extremely rare, possibly inexistent. As a matter of fact, every second we produce several messages at once. Our whole organism, from the words we utter to the very position of our body, is a constant factory of signs that reveal a consistent amount of information about ourselves: what we want to say there and then, what we really want to say (as we know, the latter may differ from the former), what we think, what we feel, the state of health we are in, some habits/attitudes typical of our culture, of our family, of ourselves as individuals, and so forth. It is a very long list! In fact, even when we take a single message, we rarely find that it is really only one. Let us make an example: a car driven by some tourists stops by a pedestrian to ask for directions. The pedestrian says: “Go this way for about 5 km, and then you will see another bunch of buildings like these here”. Here it is: one single message. But wait a minute: which way is “this way”? How are the buildings “like”? Is this sentence all the pedestrian said? Obviously not. Obviously we have to assume that this person was not only sending messages with words, but with hands too: first s/he must have pointed towards a specific direction, showing the tourists which way to go, and second s/he must have indicated some visible buildings nearby, in order to provide a point of reference. So: one message, many messages. As we have seen in this example, many channels too: the pedestrian employs both acoustic and visual information, in this case.

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Multimodality The name that describes this particular complexity in communication is *multimodality*, and this shall be the topic of this chapter. Multimodality is a very interesting concept, which became rather fashionable in science during the last couple of decades. It refers to those instances of communication when different patterns cooperate (or compete) to display one or more texts. It operates in almost every communicative context, and it is particularly relevant in instances like play, linguistic or pseudo-linguistic interaction, aesthetics, deception and others that we shall deal with in the next chapters. Communication researchers, as we have seen, tend to categorize signals by the primary sensory channel involved, but the reality is rarely so straightforward, as multiple channels are regularly engaged simultaneously. This is especially true in highly social, group-living animals. As anticipated by the great Charles Darwin (1872), multiple, concurrent stimuli are very important in order for a signal to be fully efficient. However, the influence of such stimuli on the signal and its meaning has only recently been closely analyzed (see, for instance, Horn 1983). Behavioral neuroscientists have found that many species (and particuarly primates, birds, and insects) process information from multiple sensory channels, in the fields of both attention and perception. In particular, posture and movement are the stimuli most frequently associated with the production of the signal. When we think about it, this is also what we human beings do, more often than not: if someone is talking to us, we not only “listen” to words, but we look at gazes, gestures, we pay attention to tone, mood, volume, timbre... The golden rule of multimodality is that the appearance of different signals at the same time may either aim to convey the same message (that is, they are redundant), or different ones (non-redundant). For instance, if we say “look there!” and at the same time we point a finger towards a certain direction, obviously our mouth and our hand are “saying the same thing”. However, if we say “look there!” and at the same time our hand is, let us say, caressing our interlocutor’s head (for whatever reason we might want to do these two things together), we are clearly witnessing two different messages.

Types of multimodal communication Partan and Marler (1999) provided a synthetic but exhaustive account of the most important points in multimodal communication. When the

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diverse signs are redundant (i.e. they all concur to convey the same text), there can be: 1) Equivalence – The multimodal signal provokes the same exact reaction as the signs emitted separately; 2) Enhancement – The multimodal signal produces a reaction that is increased in intensity. When the multimodal signal is based on the combination of nonredundant signs (i.e. they convey different texts), there can be a wider range of possibilities: 1) Independence – The different signs are independent and produce distinct reactions that are not in relation with each other, although they are combined; 2) Dominance – One of the signs prevails on the other(s); 3) Modulation – One non-redundant sign affects the other(s), by modulating its/their effect; 4) Emergence – The multimodal signal provokes an entirely new reaction that has nothing to do with the separate signs. Their classification may be summarized in the following scheme:

Redundancy

Nonredundancy

SEPARATE COMPONENTS

MULTIMODAL COMPOSITE SIGNAL

Signal

Response

Signal

Response Category

a

x

a+b

x

Equivalence (intensity unchanged)

b

x

a+b

X

Enhancement (intensity increased)

a

x

a+b

x and y

Independence

b

y

a+b

x

Dominance

a+b

X or x

Modulation

a+b

z

Emergence

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The left side of the scheme depicts redundant (above) and non-redundant signals (below) as separate components (a and b) with consequent responses (x and y – the same letter indicates the same qualitative response; different letters indicate different responses). The right side depicts the responses to combined multimodal signals (a+b). The meaning of a single signal may be either redundant or non-redundant, in that different signals may produce the same message, or two different ones. The advantage of redundancy is the reinforcement of the message and the reduction of the risks of interference (not by chance these signs are also called “backup signals”). Non-redundant signs have the advantage of providing more information per time unit. Empirically, the two typologies of sign can be distinguished by the reaction of the receiver. When emitted separately, redundant signs should provoke the same or a very similar response from the receiver, whereas non-redundant signs should provoke different reactions. The picture may change quite radically when the signs are combined simultaneously into a multimodal signal (see the right side of the scheme). With redundant signs, as we have seen, we can have equivalence (the multimodal signal provokes the same exact reaction as the signs emitted separately – in the scheme we have a small x letter) and enhancement (the multimodal signal produces a reaction that is increased in intensity – in the scheme we have a capital X). Let us exemplify: in the case of equivalence, let us imagine that “a” is pointing the finger towards a given direction, and “b” is the sentence “Look there!”. If a alone, or a+b produce in us the same understanding of the message, then we can talk about equivalence. If the finger alone was a bit confusing and the sentence helped us to focus on that particular message, then we have enhancement. Among non-human animals, a case of equivalence is that of the courting male moths Cycnia tenera, that “elicit equivalent responses from females regardless of whether their pheromones and ultrasonic sounds are presented separately or together” (Partan and Marler 1999: 1273). As for enhancement, we can mention the Aphaenogaster ants, which “recruit help for carrying prey by emitting pheromones, but with large prey they also stridulate, producing a substrate-borne vibrational signal. The stridulation has a small effect alone, but both components together recruit more workers. Similar multiplicative effects occur during the neural processing of simultaneous visual and auditory stimuli in the superior colliculus of cats” (Partan-Marler 1999: 1273). A multimodal signal based on the combination of non-redundant signs, as we have seen, offers us no fewer than four options: independence (the

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two signs are independent and produce distinct reactions, unrelated with each other, although combined), dominance (one of the two signs prevails on the other), modulation (one non-redundant sign affects the other), and emergence (the multimodal signal provokes an entirely new reaction). Here are four examples provided by Partan and Marler themselves: 1) Independence: “Pheromones from female Cupiennius salei spiders alert males to the presence of a potential mate. Concomitant vibrational signals from the female then direct males to her location. These two components function independently whether they are perceived simultaneously or not” (Partan-Marler 1999: 1273); 2) Dominance: “Dogs signal play behavior visually with a bow, and sometimes also growl, normally a threat. Separately, these signals are contradictory, but their combination elicits play, the visual component taking precedence” (Partan-Marler 1999: 1273); 3) Modulation: “Male Alpheus heterochaeli shrimp respond aggressively to visual cues alone, such as an open claw, but do not respond to chemical cues alone. When the two are combined and the pheromone is from a female, male aggressive responses are suppressed” (Partan-Marler 1999: 1273); 4) Emergence: “Aromatic pyrazines and red and yellow coloration are commonly associated with noxious insects. Presented alone, neither cue produces aversion in chicks; aversion appears only when the odor and color occur simultaneously. Here, multimodal stimuli evoke a response not elicited by the unimodal components” (Partan-Marler 1999: 1273).

Effects of multimodality on the receiver When a multimodal message is produced (as we have seen, that occurs much more often than we think), the receiver is of course affected in a different manner, as compared to a basic process of reception of the information. A multimodal message, in most cases, can establish a new context for communication, by adding, replacing, reinforcing elements. This in turn creates altogether a new meaning for the receiver. Guenther Kress (2010), one of the major scholars in multimodality, made for instance extensive research on how students understand information differently when the teacher uses different media together for his/her lectures: we know very well, by now, that a traditional lesson when the teacher is only speaking to the student has become rather obsolete.

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Already in the past, multimodality was assured by a combination of speech and visual messages (on a blackboard). Then transparency projectors came onto the scene, and then slide presentations, and so forth, up to nowadays, where a standard lecture can easily be the combination of different exposures of the same concept (video, audio, image, etc.). The point is that the receiver can now get a basic information item (in a lecture, that would be the words that the teacher pronounces) and various forms of recontextualization of it. The meaning switches from one medium to the other, and that creates a different quality of understanding, certainly a more complete and a better one. Multimodality is also good for giving a specific argumentative “direction” to the receiver’s interpretation, especially when simple information, presented in a single mode, proves not to be particularly rich (in definition, values associated, effect, etc.), or – as multimodal scholars would say it – the text is “static”. Multimodality intervenes on this stasis, provide the text with many more “tools”, and make it more productive (that is: not only is the information delivered clearly and successfully, but it can also stimulate continuation, reactions, extension in a more effective way). Again, if we take the example of teaching, students know very well that when a lecture is delivered with various modes together (possibly offering interactive opportunities), they are facilitated to create their own meaning, in other words to absorb the information in a more personalized manner. In conclusion, there is no doubt that communication, when organized in a multimodal way, is a) more successful (we understand better), and b) richer (we understand more). Imagine now having to very precisely communicate the following amount of information: 1) a certain place where you can find something useful; 2) what kind of useful thing is there; 3) how much of it is there; 4) what precise direction you have to take; 5) how far that is; 6) will you be against or in favor of wind; 7) how tiring is to reach this place. That is a lot of information with very critical nuances (if the place is – say – 10 kilometers away, and you point in a direction that is only one degree further south, that single degree will become a greater distance meter after meter, and after 10 kilometers you will end up in a totally different place). These seven problems were solved by the honeybee Apis mellifica with one of the most amazing forms of multimodal communication that nature has to offer.

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The language of bees One of the most impressive examples of animal communication in general, and of multimodality in particular, is certainly the language (or dance) of the honey-bee Apis mellifica (or, as you will see sometimes written, Apis mellifera). By means of this dance, one or more bees inform the hive about the presence of food, nesting sites or water sources in the more or less immediate vicinities. It is an extremely fascinating and very complex form of communication that adopts at the same time tactile, olfactory and acoustic channels; that produces data such as source type, distance, direction and type of route; that uses the sun as a compass, and that contains elements of iconicity, indexicality and symbolicness in the way we illustrated in Chapter Three. If “dance” is a term used in reference to the circular movements that bees perform, the employment of the expression “language” has a lot to do with the richness of this communication system. Its discovery is mostly due to the work performed, during the 1960’s, by the Austrians Karl von Frisch and his then student Martin Lindauer, who carefully observed and described the behavior of the Apis mellifica. Von Frisch and Lindauer went down in history as two of the most important ethologists of all times. For this particular work, von Frisch ended up winning the Nobel Prize in Physiology or Medicine in 1973, sharing it with two other giants of ethology, Konrad Lorenz and Nikolas Tinbergen: his work included studies on the bees’ sensorial mechanisms, their various communication systems, their sensitivity to polarized light, and the pheromones emitted by the queen bee and her daughters.

How does bee language work? Back to the bee language, as such, we can safely summarize the process in the following five stages: 1. One bee (called scout) goes in perlustration of the neighbourhood of the hive, generally in search of three useful things for its community: water, food or a nesting site. For the sake of our explanation, let us say that this scout finds a rich food source. 2. Impregnated with the pollen of the flowers found there, the scout flies back to the hive, and – surrounded by other bees, by now anxious to learn the “news” – it starts acting frenetically, repeatedly performing a movement that is either in a round shape or in a figure of eight (as you can see in the picture), very close to the other bees

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of the hive. The scout also produces sounds by vibrating its wings, therefore the other bees can receive information via the tactile channel, the olfactive channel and the acoustic channel (not the visual, though, as the hive environment is too dark for allowing this type of communication). Let us see these three channels in detail.

Fig. 4 – The two types of bee-dance, for short and long distances respectively

3. By smelling the scout, the other bees get indexical information on the quality of the object found (in our example, the bees smell pollen, therefore it is a food source, and not – say – a nesting site that the scout has found). As a matter of fact, in the beginning, this part of the process was not considered particularly relevant. It was after Wenner 1969, and the discussion followed, that it became clear that the olfactive channel plays a more important role than previously suspected. 4. By touching the scout, thus by perceiving what kind of dance it is performing, the bees get symbolic information on the location of the place. If the dance is round, it means that the place is very close and does not exceed a distance of about 100 metres. A dance in figure of eight represents a distance that ranges between circa 100 metres and 13 kilometres. When the distance is so great, it is clearly of considerable help to have precise indications on the route to follow, as well. That is provided by the orientation of the 8 figure in relation to the position of the sun. The middle part of the 8 (the one where the two lines cross each other, as you can see in the

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left image of the picture) is the moment when the scout indicates the direction. That part is performed very carefully and at a slower pace, and creates an ideal angle with the sun’s position: from that very angle, the other bees understand what direction to take. In the same part of the figure, the scout also provides information on the distance of the site. 5. By hearing the scout, the bees get additional (or reinforced and more precise – as we know it is one possibility in multimodal communication) information on distance and direction of the site. For a long time, the presence of acoustic communication was not noticed by the researchers, but after Esch 1967 it became clear that the scout can create a tridimensional acoustic field through the wings’ vibrations, and that the other bees can perceive the sound through their antennas (specifically on the so-called Johnston’s organ).

The many functions of bee language When showing the distance of the site, von Frisch noticed, the scout provides an accurate and directly proportional correspondence between the time spent on the middle part of the “8” and the distance of the site. In other words, the slower the bee goes, the further the distance is. In the community studied by von Frisch, the scout would spend a period between half and one second if the site was within 500 meters, less than 2 seconds if the distance was within 1500 meters, up to 3 seconds when the site was within 3 km of reach, and so on. The axis of the very movement in the “8” middle part is also progressively varied by the bee. As the directional aspect of the dance is crucial for saving time and energy to the other bees, the scout rotates approximately 15 degrees per hour, in perfect accordance with the position of the Earth towards the Sun. This is possible due to the bees’ endowment of the so-called “solar chronometric orientation”. Even in weather conditions that limit the sun’s visibility, the bees equally succeed in conveying and learning the right information, as they are sensitive to polarized light. Additional information on the type of route and the travel conditions, are detected by the rhythm of the dance. From this element, the bees understand how much energy is spent reaching the place. Matched with the data received concerning distance, this piece of information reveals whether the scout was flying against or with the wind, or if the route was particularly demanding.

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Finally, the approximation in specifying the exact distance of a site requiring a round-dance (in point 4 of the list in the last paragraph) is not due to uncertain data, but to the existence of a local *dialect*. Depending on their geographical origins, bees have a different conception of “close” and “far”. For certain communities, it is important to perform a rounddance strictly if the site distance does not exceed 50 meters. Other communities seem to have a better sense of orientation, and can even afford a round-dance (that is, a dance without specific indications on the direction) if the site is 150 meters far.

Symbolic aspects: why it is a “language” What makes the use of the word “language” rather appropriate for this kind of communication system is its symbolic property: the fact, that is, that some aspects of this communication are established by convention within a single community of bees, and not because they resemble or are a consequence of the meaning they intend to convey. We can use the very last example we have made as a demonstration: some bees do not use the figure of 8 dance unless the distance is at least 150 meters, to some others 50 meters are enough. It is not as though the circle tries to “imitate” the distance (for instance, it is not wider if the distance is longer: the circle is always the same; it is the convention within this or that community that changes), but each community conventionally assigns a certain distance to the same round movement. For instance, in human communication, one can say “matto” in both Finnish and Italian and mean totally different things (“carpet” in Finnish and “crazy” in Italian), just because different groups of people (Finns and Italians) agreed on a conventional way to call certain objects, always using a sign (a word, in this case) that has no resemblance with the objects it represents (the word “matto” does not look like a carpet or like a crazy guy at all!). You may need to brush up on Chapter Three to remember what makes a symbolic sign different from the others, but perhaps this explanation of the bee-language provided by Thomas Sebeok may help: It is common knowledge that if its food source is farther away than 100 m., the bee’s tail-wagging dance conveys, among other bits of information, the direction of the goal, the sun being used as a reference point. Now if the bee dances on a horizontal surface, von Frisch tells us that “the direction of a waggling run points directly to the goal”, that is to say the display is indexic. If, however, the dance takes place on a vertical comb surface – as is the case, normally, in the dark hive – then “the dancer transposes the solar angle into the gravitation angle”, according to von Frisch. In other

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words, if a vertical honeycomb is involved, when an angle with respect to gravity is substituted as the orientation cue, the indexical aspect of the display attenuates to the extent that, temporarily, its symbolic aspect comes to rank predominant. (Sebeok 1990: 43)

To clarify this aspect, we may think of the positioning of an arrow in a street, for signifying the direction “forward”. Now, as we all know, a sign for forward (which is most often bi-dimensional) may appear in at least three configurations: 1) pointing forward, in a horizontal position, as for instance when the arrow is painted on the asphalt of the road; 2) still pointing forward, but in a vertical position, as it may appear on a street sign, on the right side of the street; and finally, more significantly 3) pointing up, still on a street sign, when it appears positioned in front of the driver. Now, not only do configurations 1 and 2 not provoke any significant changes in the behavior of the driver, but neither does configuration 3, that is, the driver does not start flying with his/her car. This is possible when the appearance of the arrow is perceived neither as indexical nor as iconic, but simply as the result of a convention, untied to any physical connection (resemblance or consequence) with the object represented. It is all three configurations which have a symbolic value, not just the third one. It would be too laborious, even for a human mind, to separate the notion of indexicality, iconicity and symbolicness as applied to the same sign. Much more economic would be to establish one single convention (still a convention, but at least “one”), and accord one’s behavior on that basis. Among the evidence scholars bring in support of this interpretation of bee-dance as symbolic language, a rather interesting example is the geographical variability in the significance of the various signs. Different communities of the same species of Apis mellifica may for instance use the very same articulation of the sign that indicates the distance in the figure of 8 dance to represent completely different distances (as you may remember from the previous paragraph, this sign consists in the time that the bee spends in the middle axis of the figure of 8: the greater the time, the longer the distance). The same amount of time will signify 5 metres for an Egyptian Apis mellifica, 25 for an Italian, and 75 for the predictably more efficient German bee (see Mainardi 1992: 455). Of course, Germans are always more laborious! In conclusion, if you are interested in studying the language of bees in detail, Lindauer 1961 and Frisch 1967 remain two “classics”, but the subject is still very popular among biologists and communication scholars,

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and new publications appear regularly: worthy of mention are at least the works of Thomas Seeley (1995) and Fred Dyer (2002).

CHAPTER SIX THE SECRET WEAPON OF COMMUNICATION: LIES, TRICKS AND DECEPTION

Umberto Eco, the famous Italian writer and semiotician, once said that semiotics “studies everything that can be used to lie”. This is a great definition, and not only for semiotics, which is in any case a methodology that we use very often in this book. When we talk about semiotics, of course, we talk primarily about the way communication is used through signs and codes. Now, the beauty of signs and codes is that you can essentially do whatever you want with them. A sign is associated to a meaning, and that meaning can be assigned via a set of conventions (a code): if I say “bicycle” you would normally think about a means of transportation with two wheels, pedals and so on, because that is what the conventional code of the English language has assigned to this particular word. But if I point at a car and I say “that’s my bicycle”, I am manipulating both the sign (by deliberately associating it to another meaning) and the code (by breaking that convention). I am, in other words, lying. Never mind semiotics in particular: the presence of a lie shows us how rich and complex communication can be. If we witness a lie in a communication system, we are witnessing at least four important phenomena: 1. There is a code, which is socially shared, that associates a sign to a meaning. Since lies break this association, the latter had to exist previously. Calling “bicycle” a car constitutes a lie only because there is already a code (English language) which associates a bicycle to the means of transportation with two wheels and pedals, instead of the one with four wheels and an engine. If we were using a language where neither “bicycle” nor “car” make any sense as words, there would be no lie, but simply an absence of sense. 2. The sender of the sign must be able to break this association. If I choose to lie by deliberately calling “bicycle” a car, then I must

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know what a bicycle is and what a car is. Otherwise, there would be no lie, but simply ignorance on my part. 3. In order to lie, the sender must be able to mentally represent not only the receiver’s behavior, but also his/her mind. That is why we can only lie to animals, not to plants or objects. We lie, for fun or for other reasons, many times to our fellow human beings, and we can lie to other animals too (I can fake throwing a ball to a dog, just to tease him/her, I can shake the crackers box to call a cat’s attention, without wanting to feed him/her, etc.): these lies are possible because the other subject possesses a mind, so they can be lied to. Let us try now to call the attention of a stone, show it our car and say: “this is my bicycle!” How smart does that make us feel? 4. The sender must be able to take advantage of the broken link, making the receiver believe something that is not happening in the way s/he thinks. If I tell a friend that my car is in fact a bicycle, very probably it is because I want to have a laugh, so my “advantage” is amusement. If a seller tells us that a given product makes us slimmer (which is always a lie, no exception), his/her advantage is the sale, i.e. the financial reward. And so forth. This means that lying is a cognitive act, and cannot be seen as the result of instinct or of a simple behaviorist stimulus-reaction process. In Chapter Two we have seen a few prerequisites for communication that are related to a complex cognitive activity: when we talk about deception, we need at least seven of such prerequisites (see also Cimatti 1998: 179-190): 1) Sociality; 2) Arbitrariness of categorizations; 3) Mind; 4) Principle of complexity; 5) Ritualization; 6) Self- and Hetero-perception; 7) Syntax of perception. Now that we saw how rich a lie can be, let us also discuss about those forms of deception that do not display such a complexity. The bicycle-car lie is clearly a lie that requires all the cognitive processes we have described above. However, there are plenty of “lies” that occur outside our control or which are part of the way we are. If, to make a silly example, I happen to look like a famous football coach (which I do: people tell me I look a bit like Rafael Benitez), there is a chance that I will be mistaken for

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that person. Am I lying in that complex sense we were talking about? Of course not: I was born like that, and I do not go around claiming I am that person. Yet, the information people receive by looking at me is still of a deceiving type: there is deception going on, although I do not try to deceive anybody. Unless, of course, I explicitly intend to use my natural resemblance to a famous coach in order to get some advantage (getting a free ticket to a game, or something). In this chapter we shall start from “simpler” forms of deception and get little by little to more complex ones.

Biological Mimicry When we think about animal deception, the first context that comes to our mind might be survival. For instance, we know that animals adopt several tricks to escape being preyed upon, by pretending they are not prey: they may lie “above the threshold” when their action metaphorically corresponds to a sentence like “Look at me – I’m not your prey!”, but if their action metaphorically corresponds to “Let’s hope the predator doesn’t see me”, then they are probably lying “below the threshold”, in a less complex fashion. These two cases, along with many others, are known as biological *mimicry*, an umbrella term that covers several forms of similarity (appearance, behavior, messages…) of one species to another. Mimicry involves all channels (not only the visual, as is sometimes implied), and occurs largely (but not only) in insects. It occurs when an organism (called the *mimic*) evolves to share common characteristics with another organism (the *model*), while a third party (the *dupe*) is the receiver of the mimicry sign (e.g. a predator common to the mimic and the model). Sometimes, model and dupe are the same organism. Mimicry is, in the majority of cases, advantageous to the mimic and harmful to the receiver (“harm” includes also the idea that a hungry predator will not find its prey). Usually, models and mimics are closely related organisms, but sometimes the phenomenon occurs among biologically “distant” species. A particular case of mimicry is *camouflage*, occurring when the model is a non-living part of the environment (as in the famous case of chameleons). Occasionally, mimics may have multiple models during the stages of their life, or they may be polymorphic, i.e. different individuals imitate different models. Models, also, may have more than one mimic. The main typologies of mimicry are “defensive”, “aggressive”, “automimicry”, plus one, “reproductive”, which however occurs only in plants.

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Defensive mimicry Defensive (or protective) mimicry is the most important and widespread typology of mimicry. It occurs when mimics avoid/deceive predators or other enemies by resembling other species or (in the case of camouflage) inanimate parts of the environment. At least four forms of defensive mimicry have been observed in animals (additional ones have been observed in plants, but of course we are not concerned with those): 1. The first one is called Batesian mimicry (after English naturalist Henry Walter Bates): in this case, the mimic shares signals similar to the model, without being as unpalatable as the latter. It is observed, among others, in Lepidoptera, Hymenoptera, some snakes, and Octopuses of the genus Thaumoctopus. For instance, the species Clytus arietis and Aegeria apiformis are two perfect imitations of a wasp. A predator knows that eating a wasp may be a very painful experience, so if other insects look like wasps there are more chances that the predator will spare them. When Batesian mimicry is intraspecific (that is, model and mimic belong to the same species, while dupe still belongs to another) it is known as Browerian mimicry (after biologist Lincoln P. Brower). 2. The second form of defensive mimicry is called Müllerian (after German naturalist Fritz Müller): in this case, two or more species have very similar signals, and both are “unpalatable”. It has been observed mainly in Lepidoptera and Hymenoptera. This is a curious case of mimicry that Bates (who was operative at the same time as Müller, in the second half of 19th century) could not explain. One may ask what the point is of imitating each other if both animals are unpalatable. We can say that this is a kind of “marketing strategy” that reinforces the “business” of both species. We may compare it to the way certain shops selling similar items end up looking very similar, even if they belong to totally different companies. One may think that this is not a good strategy, because customers may end up thinking that they will find exactly the same things, the same brands and the same models. As a matter of fact, the actual result is that these shops become very recognizable as “types”, so in fact customers get more attracted by both, since they see both as shops of interest. Let’s say: two different shops for youngsters’ trendy clothes may end up having the same appearance (urban-industrial interiors with fake graffiti) and the same background music (loud techno). These features will immediately

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put a “mark” on these shops as places where youngsters can find interesting things, knowing that young customers can very well tell the difference between one brand and another. This situation of reciprocal help is called *mutualism*, and that is exactly what happens in Müllerian mimicry. If two species are confused with one another by a common predator, both mimics and dupes increase survival chances (mutualism, indeed). The dupe is advantaged because s/he knows that both mimics are unpalatable. The co-mimics are advantaged because their unpalatableness is “reinforced” by the other species.

Fig. 5 – An example of Batesian mimicry

3. The third form of defensive mimicry is named after a German herpetologist called Robert Mertens (although other scholars had been working on it before him), and it is called Mertensian. Mertensian mimicry is a little less common than others, and it describes those (apparently paradoxical) cases where deadly prey mimics a less dangerous species. Why such a suicidal tendency? Because, if there is some other species that is harmful but not

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deadly as well as *aposematic* (that is, colored or constructed in a way that indicates special capabilities for defense), the predator may learn to recognize its particular warning colors and avoid such animals. A deadly species will then profit by mimicking the less dangerous aposematic organism, because, to put it trivially, it does not want to be chased either. Mertensian mimicry has been observed only among coral snakes: the harmless Milk Snake is a mimic of the deadly Coral Snakes (this being a common case of Batesian mimicry), but both are mimics of the moderately toxic False Coral Snakes (genus Erythrolamprus): they all have a red background color with black and yellow stripes, with minor differences. There is even an old American nursery rhyme that teaches how to distinguish the deadly snake from the harmless one: “Red touch yellow, kill a fellow; red touch black, friend of Jack” (in other words: in the coral snake the sequence of colors of the skin is red-yellow-black-yellowred, so “red touches yellow”; in the milk snake the sequence is redblack-yellow-black-red, so “red touches black”). You may want to remember it!

Fig. 6 – An example of Müllerian mimicry

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Fig. 7 – An example of Mertensian mimicry

4. The last form of defensive mimicry is known as Wasmannian (after the Austrian entomologist Erich Wasmann). It refers to those cases where the mimic resembles a model of which it is a co-habitant, in a nest or a colony. Not surprisingly, most of the models here are social insects such as ants, termites, bees and wasps.

Other typologies of mimicry Besides the defensive one, there are three more important typologies of mimicry: aggressive, reproductive and automimicry. Aggressive mimicry is of course the opposite of the defensive one. In this case we have predators (or parasites, which can be considered predators too), which share characteristics with a species harmless to their prey or host, so that the latter cannot detect them. The mimic may resemble three types of organisms: one that is neutral to the prey/host, one that is beneficial to it, or even the prey/host itself. The model of such mimicry may or may not be affected by this “exploitation” of its characteristics. When it is affected, there may be negative or positive

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consequences alike – it depends on the case. It is also sometimes referred to as Peckhamian mimicry after American entomologists George and Elizabeth Peckham. Unlike defensive mimicry, there are no particular nuances that may lead to distinguishing different forms, so aggressive/Peckhamian mimicry is really one big category of mimicry. There are many different species that practice aggressive mimicry: spiders of the species Nephila clavipes and Argiope argentata, fireflies of the genus Photuris, crickets like the Chlorobalius leucoviridis, fish like the Aspidontus taeniatus, and others. Reproductive mimicry occurs when, thanks to its imitation, the mimic is helped by the dupe in the act of reproduction. The phenomenon mostly concerns plants, but there is one case where animals are involved, and another one where animals and plants are interacting, so we shall mention both of such cases: 1. Inter-sexual mimicry. It happens when the members of one sex in a species mimic specimens of the opposite sex. An interesting example comes from the marine isopod Paracerceis sculpta. *Alpha males* are the largest individuals of the species and, as it often happens in these cases, they are in charge of a sort of harem of females. In order to mate with some of these females, *beta males* mimic them and manage to access this harem without being detected by the alpha males. Also, the *gamma males*, the smallest ones, manage to enter the harem by mimicking juvenile specimens. Similar phenomena can also be spotted in other species, most notably the side-blotched lizards of the genus Uta. 2. Pseudocopulation. This is the “mixed” animal-plants case we mentioned. It occurs when a flower mimics a female of a certain insect species, inducing the male of that species to copulate with the flower. Orchids, for instance, tend to mimic females of the Hymenoptera order (bees and wasps). It is important to point out that the mimicry process here is not only visual (in fact, the visual part is of little relevance): the real imitation occurs at chemical and tactile level: the flower mostly “feels like” and “smells like” the insect female. Finally, there is auto-mimicry: one part of an organism’s body resembles another part. This is a rather well-known form of mimicry among wildlife documentary enthusiasts, because it is shown very often as one of the visual marvels offered by Nature. Most of us will certainly remember fish species like the Foureye butterflyfish Chaetodon

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capistratus displaying a big spot on the tail that resembles a big eye, and therefore makes them look like bigger fish. Or we may remember seeing the Pygmy owl Glaucidium californicum displaying eyespots behind its head, or those on the torso of the larva of Deilephila elpenor. There is a wide range of literature on the subject of mimicry for those who want to study this topic a bit further: Brower 1988, Owen 1980 and Wickler 1968 are certainly three “classics” that deserve to be singled out.

The complexity of deception Mimicry is without doubt a fascinating topic in all of its forms. Yet it is far from being the only manifestation of deception that animal communication has to offer, and further still from being the most complex one. In the second part of this chapter, we shall explore a few of the various ways animals lie to each other while communicating. To better frame these different typologies, it is worth underlining that the word and the concept of “deception” in fact cover a wide range of behavioral patterns. For instance, we might mention the classification provided by Jean-François Bouvet in a popular book called La stratégie du caméléon (The strategy of the chameleon, 2000). According to Bouvet, a lie may occur through: 1) 2) 3) 4)

Performing (as in staging, exhibiting some action); Hiding (as in putting something out of sight); Simulating (as in imitating an action); Cheating (as in causing to accept as true or valid what is false or invalid); 5) Disguising (as in changing the usual appearance of something); 6) Dissimulating (as in hiding under a false appearance); 7) Pure lying (as in making a false statement). Most of these categories have been already made clear through the case of mimicry: we have seen animals “in disguise”, “dissimulating” their appearance, etc., but there are more varied and complex examples to mention. First of all, one can use the same lie for different purposes, not just for one. When, as kids, we used to pretend we were sick, we could use that lie for various goals: avoiding our school homework, avoiding going to school altogether, skipping a not-too-tasty dinner, and so on. We may all

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remember that lying about our health could bring a lot of immediate advantages. Let us therefore start from a case, in animal communication, that shows how a simple deceptive sign can be used in a very flexible manner: the food-call uttered by a rooster in order to signal the presence of food to hens. It is a situation that anybody who has grown up in the countryside might have experienced: the rooster walks in exploration, and as soon as he finds some food, calls the hens with a distinctive sound. Usually, the longer or the faster the vocalization, the better and more plentiful the food. Very often, however, the rooster emits the call even when there is no food available, and that usually happens when the male needs an excuse to call a female with whom he intends to mate. This is the basic lie of the rooster, but in fact this is not the end of the story, as there are no less than five variants to this theme: a) The rooster utters the call when there is no food, but instead there is a new hen in the fowl-yard: the call is long and intense (as if there was plenty of food), but of course the reason behind it is that the male wants to mate with the new female; b) There is a new hen and some food is available, but this time the rooster may not utter the call: in this case, he simply does not want to share the food, neither is he interested in the new female; c) There is a new hen and there is food, but there is also another rooster: no call is uttered (the rooster, so to speak, is “chickenhearted” and does not want to face the new male); d) There is a new hen, there is another rooster, but there is no food: a food-call may be uttered and then a fight with the rival occurs (the rooster wants to establish hierarchies); e) There is a new hen (or an old-but-still-attractive one), and there may or may not be food: the rooster may choose not to emit any call, but rather move himself to the female. In fact, this is not really a lie, but admittedly a nice coup de théatre. Put simply, the rooster seems to do exactly what he pleases with the food-call, being very aware of how to use it for deceptive purposes. Incidentally, the same applies to the hens, who very often guess the mating purposes of the food-call and simply ignore it – unless, of course, they like the idea.

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A few case studies Lies, in non-human animal communication, occur more or less in the same contexts as human life; once again, there is no surprise in that since humans are animals too. What follows are a few well-studied examples that will surprise you for how much they seem like us, in the most embarrassing or noble situation. To start with, animals may lie for the express purpose of survival. There are situations when a well-placed bluff can really save our life. This may be the case in predator-prey interactions. Gazelles are known for playing such wild cards, on occasions. If a gazelle is sick, or young, or weak in general, and a lion or a cheetah attacks her herd, this very gazelle will know that she is the most likely to be caught, because her running speed will be clearly slower than her mates. What may happen in this situation, is that the weak gazelle may decide to stand still and wait for the predator to approach. Once the latter is close, the gazelle, instead of running for her life, may display that playful jumping behavior known as *stotting*, just in front of the predator. This is nothing but brave showingoff aimed at impressing the predator and persuading her to give up chasing prey that must be very healthy and fit, if she even has the nerve to happily jump when her life is in danger. Keeping something safe and/or hidden (food, for instance) is also another typical motivation for lying. Again, memories of our childhood may spring to mind, at this point. There is a very amusing case reported about two chimpanzees named Belle and Rock, subjects of a training by the famous scholar in *primatology* Dr. Emil Wolfgang Menzel, Jr. (documented in Menzel 1974), which not only illustrates this kind of lie, but also displays that lies can evolve and be improved to better fit the purposes. Rock was an alpha male, and as such he would often harass Belle in different ways: for instance, if she was given sweets by Dr. Menzel, Rock would often steal and eat them himself. For this reason, he had taken the habit of checking on Belle everyday to see if she had any treat. After a while, Belle began to hide her sweets in a secret place: everyday, after hiding them, she would go to Rock, let him search her (mark this moment: this is where Belle is actually lying to Rock), and then, as he left, she would run towards the hideout and enjoy her treats. After some time, Rock understood the trick, so he, too, prepared his own trap. He searched Belle, as usual, then pretended to move, but then instead he followed Belle, catching her redhanded with the sweets. So, now, it was Belle’s turn to change strategy: she started changing her hideout every day, but neither that trick worked

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for long: by now, Rock had learnt that the signal for the “hidden sweets” was not a specific place, but was the moment in which Belle would move away from him after being searched, in no matter what direction. Finally (because this story has a happy ending) Belle found the perfect strategy: not only did she change the hideout every day, but now, after the inspection, she would also go to the wrong place, waiting for Rock to come very close, pretending that he had caught her red-handed once again, but then, as Rock would start searching around in this place, she would run towards the real hideout, and consume the sweets while Rock searched in vain. You have to admit: that is quite a story! But wait to hear this next one. Material objects are not the only things one can hide and lie about. Actions can be hidden as well, and there is no need to mention that sexual “cheating” is probably the most recurrent action to be hidden/disguised/manipulated in human history (the destiny of entire kingdoms was altered by these matters, as perhaps the case of Henry VIII and England’s reformation best exemplifies). Then again: only in human history? Let us see an example that may remind us of morbid stories we read in literature or history books. In baboons’ communities, for instance, dominant males are in charge of what can be defined real harems, that is, groups of females who are only supposed to mate with the boss. These females are expected to “report” every now and then, in order to prove that they are not mixing with lower ranked males: the show up is a signal sort of standing for “I’m here, master, I’m doing nothing I’m not supposed to do”. Females, of course, do not necessarily like this condition, but, as occurs with humans, flirtations with other males must be kept secret, especially when they are “dangerous liaisons” like in this case (alpha males being alpha males for a reason). What a couple of young lovers may decide to do, in this case, is to couple in a discreet place, far from the alpha male’s sight, to actually take a break, let the female report to the alpha male, and then restart the process. Not the most romantic strategy, of course, but an effective one. Another simple knack to avoid the consequences of one’s (forbidden) actions is to accuse someone else. Again: how many times do we see that in human actions? One good example (since we shall talk about this study also later in this book) comes from one of those apes trained with Sign Language, the gorillas Koko and Mike (female and male respectively). Their trainer Francine Patterson reports that Koko often indulges in stealing food from the refrigerator, but when questioned on the matter, she resumes her competences in sign language and happily makes the sign for “Mike did it!”, accusing her mate in a cowardly way.

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Let us now mention something a bit more noble, in the vast catalogue of lies that nature offers. Indeed, altruism may be another good reason for lying. There is a well-known pattern exhibited by many animals, known as *distraction display*. We see it particularly in birds and fish. By distraction display, it is meant any form of behavior that is employed to divert the attention of an enemy (rival, predator...) from a threatened object (typically, a nest or a young specimen). The most famous example is probably the “broken wing act” performed by birds like the killdeer Charadrius vociferous. As her nest is attacked by a predator, the killdeer mother may move in a clumsy way as if she had a broken wing, trying (often with success) to focus the plunderer’s attention to herself rather than to the young chicks (who, being weak and slow, would be a better prey if the mother was in good health). Something interesting that emerges from these altruistic displays is that the advantages of deceiving communication may also not be immediate. The perspective for a, personal or not, later advantage provides enough motivation for deceptive actions. One, in other words, can also use lies as an “investment” for a future purpose. There is another account from a sign language-trained ape, this time the chimpanzee Washoe (arguably, the best-known case). After months of intensive training, Washoe had acquired the ability to cheat with sign language. If any of her trainers would punish her for any violation of the rules, she might use her communicative abilities for getting her revenge. Here is a full account of one of the most amusing episodes: When I did report Washoe’s obnoxious behaviour to Trixie [Gardner, leader of the Washoe Project, together with her husband Allen] I could always count on Washoe trying to get even with me for making trouble. Once I told Trixie that Washoe had tried to bite me, and Trixie gave Washoe a rare scolding. That very night at dinner, Trixie was cooking at the stove. Washoe was in her high chair, at the head of the table, acting like a little angel. I didn’t believe this act for a minute, so I was sitting as far from her as I could get. COME ROGER, Washoe signed to me. PLEASE COME. I shook my head no. There was no way I was getting near her. PLEASE PLEASE COME ROGER, she tried again. I signed an emphatic NO. At that moment Trixie turned around and saw Washoe making these very sweet and perfect signs. ROGER, Trixie implored me, WASHOE WANTS YOU! I was trapped. I began edging around the table, ever so slowly. Trixie went back to her cooking. I kept sliding over one inch at a time. Finally, Washoe couldn’t contain herself any longer. She lunged out of her high chair and

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To conclude, one may also lie out of pure ambition. As we have seen in Chapter Two, the vervet monkeys Cercopithecus aethiops have a wide range of sounds to warn the rest of the group about the presence of plunderers (sounds are different according to the kind of predator – eagles, snakes, felines; each different predator corresponds to a different strategy of escape). Sometimes when the only specimen in danger is the alpha male, the vervet monkey who has seen the plunderer may not emit the right sound, or make no sound at all, hoping to set the conditions for a perfect murder and climb up in the troop’s hierarchy.

CHAPTER SEVEN PLAYFUL AND AESTHETIC COMMUNICATION

As we are heading towards the end of this monograph, this chapter will be used to group two different types of animal communication, that are only partly related to each other, and that – in principle – consist of separate topics: communication for playful and for aesthetic (or even “artistic”) purposes.

The handicap principle Something you may have noticed in more than one of the examples we listed in the last chapter, is that the mechanism of lying often occurs through the giving of some kind of “advantage” to the receiver. We have seen gazelles risking their life to show off in front of their predators when they could have run away, we have seen birds pretending to have a broken wing... What is all this “playing with fire” about? Is it an important component in animal communication? There is a very worthwhile concept to be illustrated that can answer a question or two, in this respect. It is called the *handicap principle*, and it was elaborated in the early 1970’s by the Israelian biologist Amotz Zahavi (Zahavi-Zahavi 1997). Deception as such is only one of its uses, and in fact it serves as a perfect bridge from the last chapter and this one, which – as you can read from the title – is about play behavior. Zahavi’s main question was: why do animals produce so often costly, apparently useless and extravagant displays (the typical example being the peacock’s rich and colorful plumage) or behavioral patterns (like the above-mentioned gazelles’ stotting)? Zahavi maintained that such patterns are unusual forms of communication to other individuals that provide important information about the sender: the “handicap” says (or pretends to say) how much the sender can afford in terms of physical qualities, health status, social position and relation with the receiver. The peacock exhibiting its luxurious wheel is kind of saying “Look! I am so strong and healthy that I can afford to carry this big plumage”; the gazelle tricks the lion with the same message (although, as we have seen, in this case it is a

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bluff): “Look! I am so fit and fast that I can afford to stay here jumping before you, instead of running for my life”. There are four different types of handicap (see Grafen 1990): 1) Survival handicap – In this case, the possession of the handicap puts into serious danger the sender’s life, so the risk of this lie is very high. The receiver is the one who somehow has to establish the quality of the handicap: if the sender survives, then s/he has in fact proven the efficiency of her/his handicap. The bluffs we have examined in this chapter fall under this category; 2) Revealing handicap – Here, the sender makes an effort in order to allow the receiver to judge her/his overall quality. This is of course the case with the peacock’s tail, and – in fact – of many forms of courting (we shall see that in Chapter Eight); 3) Condition-dependent handicap – In this case, only certain senders (e.g. the high-quality ones) are capable of producing a given handicap. For instance, shiny feathers in birds are a handicap of this type, because their presence implies good health (nutritious diet, low parasite load, etc.). 4) Strategic Choice handicap – The sender chooses what type (and size) of handicaps to display, taking the receiver’s expected reactions to the signal into account. Since the cost of the signal and/or the benefit of the receiver’s reactions may vary from one sender to another, different senders normally choose differentlysized handicaps. The act of “teasing” someone is a good example: the sender wants to see how far s/he can go with the receiver. As we shall see, “handicaps” are an important component (though not the only one, of course) of both types of communication we shall focus on here.

Playful communication The last form of handicap listed above is a good opportunity to inaugurate this chapter, as “teasing” is nothing more than a very common form of playing (in fact, we shall see that handicaps can be used in other playful manifestations as well). Play behavior is another widely-discussed issue within animal-related studies, and another one where communication is central. Generally speaking, it can be defined as one or more behavioral patterns that imitate, in a non-serious fashion, patterns that, in normal situations, would be serious.

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Animals, including humans, usually play when primary needs do not have to be met. In particular, as biologist Robert Fagen has shown in primates’ behavior (1981 and 1993), the fact itself of playing reveals that animals are capable to use their energy not just for utilitarian purposes, but also for frivolous ones. Although it is not a fixed rule, young specimens tend to be the most interested in playing (but of course there are many exceptions, as anybody who has an adult dog may testify). This is one reason why scholars think that play serves two basic functions in an animal’s life: 1) amusement and 2) the learning of rules of social behavior. These functions are in strict causal connection (that is, to have fun makes it easier to learn). As we said, normally (we shall see exceptions later), playing is an unserious imitation of serious situations: that means that it uses and represents already existing behavioral patterns: hunting (whichever role, predator or prey) and fighting are the most recurrent “serious” patterns that get represented. Now: clearly, such representation has to be previously established between the two or more participants. One important difference between play and deception, apart from very few exceptions, is exactly that all the subjects involved in the game “share the same lie”, i.e. they agree on the type and the characteristics of the representation. From an evolutionary point of view, to cheat on this aspect (i.e. to lie in the proper sense, manipulating the code without any agreement with the receiver) is definitely not convenient, for the liar would be immediately excluded both in the game and in real life. Imagine two puppies playing “catch me if you can”, as a playful imitation of hunting, and then one really catches and kills the other: it would not be easy to explain that to the parents and to the rest of the pack! In fact, it goes even further than this (and here we are talking about the handicap principle again): in typical conditions, the players voluntarily handicap themselves (an action also called *play inhibition*) and establish a situation of balance, so that every participant has equal opportunities to win the game (see the very interesting Aldis 1975). Such silent agreement is usually interpreted as the result of a typical trial-and-error process. Young specimens learn very soon that the game will not last long if the stronger/older ones are too energetic towards them. The latter will soon escape or (successfully) invoke help from adults. Thus, in order for the game to succeed, it is convenient to create a situation of equality: the stronger participant will bite more softly, show with apparent indifference weaker parts of its body, and so forth (you may have seen two pets mockfighting: the bigger one often tends to stay on the ground, letting the

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smaller one play the part of the aggressor). In this sense, it is no exaggeration to speak of playing as an important factor for creating a sense of morality in animals, mostly through the development of intersubjective empathy. Back to the main point, it should be clear by now that in play behavior, too, communication is crucial, and must be correctly articulated. In order to activate a given game, animals need at least three types of communicative patterns, more or less corresponding to the following sentences: “I want to play”, “What I am about to do is playing”, “What I just did was playing”. Such communicative patterns can be either innate or learned. The sign repertoire is exclusive of play (i.e. not used for any other activity), and that is another reason why play is interesting within our discourse: we find here forms of communication that we do not find elsewhere. If you have a dog, you must have seen the so-called “play-bow” hundreds of times: the dog, acting very excited, stretches the anterior part of her/his body down and leaves the posterior one up. This is an exclusively playful sign, that dogs (and other canidae) will not perform in other situations, and it is their own “I want to play” announcement. There are many examples of this type in other species as well: pigs announce fight-simulations with a circular dance; great apes display the so-called “game face”, which is similar to a human smile, etc. (see Fig. 8).

Fig. 8 – A dog’s “play bow” and an orangutan’s “game face”

However, despite these examples, game signals are far from being only visual: they can also be chemical, acoustic and tactile. The mongoose Helogale parvula, for instance, points out her/his playful intention through

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a vocalization; the field-mouse Microtus agrestis through a pheromone, and so on. Also: although typical, the use of playful signs only before playing, is not exclusive. Species like the squirrel-monkey Saimiri sciureus display their playful intentions all over the game-session, the end of those signs actually meaning the end of the game itself.

Symbolic aspects Wanting to engage in the technical jargon we mentioned in Chapter Three, playful communication serves very often a “metalinguistic function” (remember? It is when signals are used to refer to other signals), since they have the double goal to point out the unserious intention of the sender, and refer to the serious pattern they are imitating. Moreover, and still referring to what we illustrated in Chapter Three, this communication is more often than not of a “symbolic” type, since it has no physical resemblance to the object represented, and it was more likely shaped in consequence of a code agreement during the evolution of each species. If we again take the case of the play-bow, we see no specific resemblance between that particular position and the act of playing itself: it is rather that kind of process (again, mentioned in Chapter Three) where some kind of natural connection between a sign and its contents, gets ritualized little by little and indeed becomes a symbol. When we think about it, the bow reminds a bit of a position dogs may find themselves in when they mockfight, or anyway when they do something “unusual”, as compared to ordinary life. The same applies with the pigs’ dance: it is not like they fight by running in circles, but the movement generated by a mock-fight can be stylized and ritualized by a representation of that kind. And so forth (I hope, by now, to have demonstrated how central is symbolicness in animal communication, despite our arrogant prejudice that only human beings are able to communicate by symbols).

Aesthetic communication “Aesthetics”, from the ancient Greek aisthetikos, is a concept that relates generally to sensibility and perception, but the modern use of it is strongly associated to the production and the enrichment of art. So, are we going to talk about “art in animals”? Of course, this is another of those topics where our anthropocentric view of nature generates more skepticism than understanding. In fact, for a long time, art and aesthetics were used as the main threshold to distinguish humankind from the rest of

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the animals. “Art is what makes us humans”: you must have heard it a few times. Like with play behavior, the main problem lies in our difficulty to acknowledge that something so “unnecessary” like aesthetics can be part of animal behavior. If, at least, play has its fair share of possible interpretations suggesting an important role in the individual’s growth and development (most playful activities are imitations of serious ones), with art (and aesthetics in general) we do not have this possibility, so we need to search somewhere else. Even more problematic is the paucity of knowledge about such topics (very few biologists are experts on art, and very few art researchers are experts on animals), and quite often, the history of animal studies has provided embarrassing, yet revealing, examples of this lack of mutual understanding. A famous case is that of some paintings drawn by two chimpanzees, named Congo and Betsy, which were exhibited anonymously at a London gallery. No one knew the artists were not human, and – as it happened – they were received by enthusiastic comments by the critics, except that these same people got very offended, and quickly revised their comments, when the true identity of the painters was eventually revealed. So, it seems clear that if we want to talk seriously about art and aesthetics in non-human animals, we need to free ourselves from anthropocentric barriers, and focus on what we have (theory and practice alike) in an unbiased manner. Let us start by listing what we know about aesthetics: 1) We know (particularly after Hartshorne 1973) that, at a very biological level, an aesthetic behavior is not merely or directly utilitarian, that is: there is no specific connection with some primary need; 2) We know that, when we review the classic philosophical works on “aesthetics”, the word is always connected with concepts, such as “pleasure”, “perception”, “taste”, “beauty”, etc. That is: an aesthetic action is something to be “liked”, “enjoyed”; 3) We know, after what we have seen in Chapter Four, that Roman Jakobson (1963) finds “aesthetic” that kind of message that is more concerned with form than with contents. Yes, we can “say” something, but it is “how” we say it that may determine its aesthetic dimension; 4) We know, from Aristotle, that aesthetic messages have to be at the same time parà tèn dóxan (i.e. unexpected, contrary to common

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beliefs) and katà tò eikòs (i.e. likely, believable). When we see a work of art, in other words, we sort of have to be able to say, almost at the same time, “Wow!” and “Right, I get it!”: surprise and recognition; 5) We know, from Umberto Eco (1968), that aesthetic messages have to be “ambiguous”, not too obvious. For instance, in poetry we may not express things directly, but we may use metaphors. We can safely say that the more we find these characteristics in a certain work, gesture or activity (be it human or not), the more we can qualify it as aesthetic, or specifically artistic.

The case of the satin bowerbird When we turn the attention to animal communication, few examples explain aesthetics better than the case of the satin bowerbird Ptilonorhynchus violaceus. During the mating period of the year, the male specimens (different from females because of their more uniform and bluish plumage – as we shall see, this is an important detail) have to inform the female that they are available for mating. To pass this admittedly important message, they build a sort of nest, called a bower, that functions as a space where the two of them can, so to speak, spend a romantic time together. Several males in the same area build these bowers, which will all compete for the attention of the female, who will eventually choose the one she likes best. What is really amazing in bowers is the precision and meticulousness with which the bird decorates the inside and the outside of his construction. All types of objects are utilized, from those already available in Nature (twigs, shells, flowers, etc.) up to small human artefacts (fragments of bottles, pens, and pieces of clothing). The objects are usually brightly colored, preferably dark bluish, which seems to be a favorite color for this species. More blue can be added to some areas (particularly in correspondence to the entrance of the bower) by actually painting them with a special paint that the bird obtains by chewing berries, and distributes with a chip of wood held in his beak. The choice of where to place each object seems not to be random. Very often, the bird can be seen putting a given element in a given place, then taking a couple of steps backwards, carefully surveying the whole thing, then perhaps not content with the general effect, deciding to change the position of that element. Withered flowers, in particular, seem not to be tolerated, for as soon as one dries up, the bird soon replaces it with a new

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one. Now, all this description may sound a bit anthropomorphic, but it is well-backed by the following, definitely more authoritative one, provided by Karl von Frisch: Every time the bird returns from one of his collecting forays, he studies the overall color effect. He seems to wonder how he could improve on it and at once sets out to do so. He picks up a flower in his beak, places it into the mosaic, and retreats to an optimum viewing distance. He behaves exactly like a painter critically reviewing his own canvas. He paints with flowers; that is the only way I can put it. A yellow orchid does not seem to him to be in the right place. He moves it slightly to the left and puts it between some blue flowers. With his head on one side he then contemplates the general effect once more, and seems satisfied. (Frisch 1974: 243-44)

It should be underlined that the bower does not need all these decorations in order to stand. The principle of its construction is similar to an ordinary nest, whose stability is basically due to interlaced twigs. The presence of such decorations is thus absolutely unnecessary (or “not utilitarian”, as we have seen before) in practical terms. After finishing their job, the males stand awaiting the female, and the latter, once chosen the “best” bower, heads towards the “winner”. As a grand finale, before starting a courting dance (another aspect that qualifies as “aesthetic”), the male may also put the final touch by welcoming the female with a flower in his beak. Now, let us all connect this amazing display with the theoretical reflections we listed before: 1. A bower is not a “merely utilitarian” construction. Even if it is conceived for mating (a very “utilitarian” action, obviously), it features a lot of “extras” that usually courtship rituals do not have. The majority of birds, when courting a female, perform dances and/or songs: the bowerbird does them too, but beforehand it creates this outstanding construction full of decorations. 2. The bowerbird aims at constructing something “beautiful” and “enjoyable”. Not only does he decorate the bower with extreme care, but he makes adjustment to it if the final result is not satisfying. Also, let us not forget that the female makes a choice among many bowers: the female must “like” one bower more than another. As the anthropologist Theodosius Dobzhansky once said, “It is undeniable that a well-decorated bower gives the bird a pleasure that can only be defined as aesthetic” (1962: 215).

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3. The bowerbird gives special attention to form, not just contents. Decorations, as we said, have nothing to do with the stability of the construction or with other “meaningful” functions: they are, indeed, “decorations”. 4. As we already said, the female selects a bower among the many offered by the males around her. It is important, say ethologists, for her to be “impressed” and “pleasantly surprised”, by a construction that stands out from all the others. At the same time, she has to recognize what she sees it for what it is. Here, then, is a typical, Aristotelian example: unexpected but believable, bowers are parà ten dóxan and katà to eikòs at the same time. 5. Finally, the bowerbird adds elements of “ambiguity” to his construction. For instance, he uses the metaphor of the blue colour. Males, as we said, differ from females by their bluish plumage. By putting so many blue objects on the bower, the male is sort of representing himself to the female: it is almost a gesture of “virility”, if we like.

Fig. 9 – A satin bowerbird constructing a bower

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The many forms of aesthetic communication The case of the bowers is just one (very outstanding) example of aesthetic communication. We cannot review them in detail one by one, of course, so we shall rather classify them in typologies, hopefully making you curious enough to do some further research on your own. The aforementioned zoosemiotician Thomas Sebeok proposed a very interesting categorization of animal aesthetics in his book The Play of Musement, in a chapter entitled “Prefigurements of Art” (Sebeok 1981, 210-59). Sebeok classifies aesthetic activities of non-human animals into four main categories: architectural signs, pictorial signs, musical signs, and kinaesthetic signs. In other words, animals (including us) tend to be “artistic” in four basic ways: by constructing an object of variable size (as we have seen with the bowerbird), by tracing an image, by producing sounds, and finally by moving their body (e.g. dancing, most typically). Let us see these categories a bit more in detail, with some examples. Architectural signs – Under this category, we include any big or small object that is “constructed” in some way. Animals are known for being excellent constructors of nests, shelters, traps, dams and – as we have seen with bowerbirds – meeting places, but they can also create smaller objects that can be used for various purposes. In 1995, a Finnish architect named Juhani Pallasmaa made an extensive study on animal architecture, and made a very interesting list of “reasons” why animals engage into architectural work, and besides predictable functions like “protection”, “food gathering” and others, he also included “communication” and “decoration”: “Some animal behavior related to nest construction looks like superfluous decoration, lacking any other obvious function in the structure” (Pallasmaa 1995: 55). Given our previous illustration of satin bowerbirds, it is worth mentioning works by other avian architects. The decoration of nests with diverse objects is a pattern displayed by numerous species, such as the Ploceus benghalensis (who plasters its nest with mud, and then decorates it with brightly colored flowers) and the Ptiloris paradiseus (who decorates its nest with snake skins). Among insects, bees show signs of superfluous decorations: after building their cells in the traditional hexagonal shape, bees attach a small hexagonal decoration on the external surface of the queen’s cell (which has a different shape from the other cells). This small hexagon serves no apparent purpose. The category of “architecture”, as we said, also involves smaller artifacts, not just big constructions. For instance, there are tools fabricated

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in order to be used as gifts. A somewhat disgusting but interesting case is that of the males of the fly Hilara sartor. Many of them perform a courting dance around one lone female. Each carries a sort of big balloon produced by an anal secretion and offers it to the female as a gift. The balloon is a totally useless artifact: it cannot be eaten, nor does it serve any other purpose. In fact, it is just as ephemeral as another type of balloon: the one we humans give to our kids. Some species of spiders, on the other hand, offer “useful” objects, but with a useless accessory: packaging. They kill a prey and they give it to the female carefully wrapped in their cobweb. Pictorial signs – Compared to other forms of aesthetic communication, pictorial art constitutes a little studied category in nature, but extensively studied in captivity. The biologist Desmond Morris (1963) made extensive research on pictorial art, and identified nine categories of perception that seem to provoke aesthetic pleasure in animals: saturated colors; primary colors; brilliant colors; rhythmical repetition of equal components; bilateral and radial symmetry; steady curves, such as circles, spirals, wave-lines, and so on; conspicuous lines or shapes; a certain balance/symmetry between the left and right halves of a picture; and a polarized preference for either the same colors or for conspicuously different colors, when two objects of different color have to be combined (as opposed to coupling two colours that are almost the same). These categories set a solid basis for the choices that animals make when they themselves are given the opportunity (in nature or in captivity) to draw. Many experiments were made with great apes, by providing them with paper, pencils and brushes. In observing them, scholars took the most notice of two things: a) apes clearly display a wish to draw, and not only to “play” with paper and pencils; b) apes are not conditioned or forced to draw – drawing seems to be their own choice. As we already mentioned, apes like Congo and Betsy ended up being exhibited in art galleries, and in fact there is nowadays a whole business based on apes’ paintings, which get sold at rather high prices, via the web and even via some art dealers. Besides apes, a well-known story is that of the Indian elephant Siri. Her artistic skills were totally ignored before 1980, when a new keeper, David Gucwa, was employed to look after her. Gucwa noticed that Siri would trace signs on the sand, using sharp stones as pencils, and then indicating the final product with her proboscis. The keeper then provided Siri with a pencil and some big sheets of paper that he would hold steady.

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The result was a long series of abstract drawings (again, no reward was given). Once spread through the media, the news surprised everyone except other zookeepers, who claimed to have often seen “their” elephants drawing the same way. Predictably enough, a sort of trend began, and various elephants were provided with paper and pencil (and even brushes and canvas), with some of them undergoing specific training that made them capable of more complex drawings, and thus soon made them big tourist attractions in their respective zoos. Another celebrity was Ruby, an elephant from Phoenix, who was particularly devoted to painting colored canvases. She became famous for changing the choice of her colors depending on circumstances offered by real life. Once, for instance, she noticed an orange-colored van, belonging to the maintenance service of the zoo, parking right in front of her “atelier”. Immediately, Ruby took the color orange from her palette, and started using a lot of it. Another time, after an injury in which she bled a bit and was medicated by veterinarians dressed in blue, she resumed painting and produced a canvas dominated by red and blue. Kinaesthetic signs – Aesthetic communication can of course be performed also through the use of one’s body. The most common form is obviously dancing, which is a type of behavior that is widespread across many different species, particularly (but not only) when it comes to courtship and mating. Examples of dance range from very basic and individual to extremely complex and collectively-coordinated bodily movements. Humpback whales, for instance, when they perform their very articulated songs (which we shall talk about later), tend to assume the same position: head down, flippers outstretched, and body inclined at a 45degree angle. Same goes with wolves, who display the typical “head up” position when they howl. And so on. Such postures, have both a utilitarian function (they are positions that can better exploit the vocal resources), but they also have a ritualized-aesthetic component. This is how they “look like” proper singers, just like famous singers who have their own way of standing while singing. But, as we said, these are just basic forms of bodily movement. There are others that can more properly be defined as “dance”. The classic example is the many species (birds most of all) who perform courtship rituals during the mating season. Generally these are individual performances, but there are also several examples of collective and coordinated movements, from simple circle figures (a pattern that we find

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in many species) to very complex choreographies, like the ones displayed by cranes and flamingos. But it would be a mistake to assume that animals only dance for mating purposes. A most interesting case, and somehow a most unexpected one, comes from the geese of the species Anser anser, and it is called “ceremony of triumph”. The simplest form occurs when a couple of mates meet an unknown male of the same species, who may or may not be invading their territory, but surely may represent a threat for the female. The male of the couple then furiously attacks the intruder, arching his neck over him. Usually no fights occur, for the intruder does not have aggressive purposes. Yet even in the case of challenge, for the same reasons, he does not keep up the fight for long, and soon leaves. The aggressor soon turns toward his mate, then both start an apparently euphoric dance, by quickly moving their heads and necks up and down, and uttering their typical nasal sounds gang, gang, gang. Fish and insects also display an incredibly wide range of dancing performances, most of which are closely related to mating. Among the most interesting and complex examples, there are nymphalid butterflies, species of spiders like the very small Attulus, of ca. 3.5 millimetres in length, green grasshoppers, cichlid fishes such as Tilapia and Haplochromis, and others. Finally, another interesting aspect is that some dances are also integrated with bodily decorations (objects, colors, simple artifacts), just like theatre or circus human actors may do. Forms of “make-up” have been observed in apes and birds primarily. Musical signs – To conclude, we have of course singing, a practice that is possibly the best-studied of all forms of aesthetic communication. Many bird species, insects like cicadas, frogs and whales like the Megaptera novaeangliae, are certainly very popular (and evokative) examples of singing animals. Who has not enjoyed, at least once in their life, the performance of a nightingale or of a blackbird, or has been in the middle of a lake, exposed to the beautiful concert of several frog species performing together from all around? Practically speaking, every species that can communicate through sounds have some form of singing: some can be very simple, some other very elaborate; some can be pleasant to listen to, from a human perspective, some can be less pleasant (but that does not mean anything, as we should have learned from Chapter One and the etic-emic topic). Most of such singing activities have, again, mating purposes (which is easily comparable to human beings, since the majority of the songs we write and

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sing are “love songs”), but – as with the other forms of animal art – there can be other functions as well, in particular territorial defence, reinforcement of relationships (either couples or entire groups), play and entertainment, relaxation, and finally display of emotions and feelings. In each of these cases, the communicative potential is very high: animals use sounds for all the six functions of communication that we have seen in Chapter Four. Songs have often a cultural component, meaning that they are transmitted generation after generation. In birds, for instance, one can witness a very early stage of rudimentary singing in small chicks, and then – with the guidance of the adults – the songs become more refined and elaborate. There are also the geographical variations within the same species which form a cultural identity: specimens of a given area sing different songs from those who live in other areas. When two groups mix – for instance, one specimen has moved from one group to another – we see that the new community “imports” some of the characteristics of the group from which the new member is coming. A common misconception is that every specimen always sings the same song: on the contrary, songs can vary by several degrees, up to the quite astounding case of the humpback whales, who basically change their songs all the time, continuously, like jazz musicians. There is also a specific field of inquiry that was born for studying “animal music”: it is called zoomusicology, and it was pioneered by the French composer François-Bernard Mâche in 1983 (see Mâche 1992, for the English translation of his book).

CHAPTER EIGHT INTERSPECIFIC COMMUNICATION (PART I)

In Chapter Three we discussed the complexity of animal signals, and perhaps, throught the course of this entire book, we have understood that animal communicaiton offers a very rich variety of situations and phenomena, all of which deserve attention. Something we mentioned only indirectly (particularly in Chapter Six) is that signals are not only used to communicate within the same species, but can also be interspecific. Any of you who have a dog or a cat will know very well that the communication with a different species is a bit like those Sci-Fi movies like Close Encounters of the Third Kind, where humans attempt to communicate with aliens. Especially at the very beginning of our relationship with a pet, we really have to re-invent our communication strategies in order to understand and be understood. What does our dog mean when s/he barks, or howls, or whines? In addition: how shall we address him/her: what words, what tone, what volume? Interspecific communication is, more than any other type of animal communication, a labour of patience and trials and errors. But finally, yes, we do find a common ground with our pet. We understand when s/he is hungry, when s/he wants out, when s/he is happy or sad or angry... and the pet also understands us, to several extents. All animals, at some point, can find themselves in a situation where interspecific signals are needed. However, some interactions are circumstantial and improvised, some others are more systematic and display the existence of a long-standing interspecific relationship: in the latter case, the signals are codified and clear. The most important forms of interspecific communication are the following: 1. Predator-Prey – As much as we may imagine that predators and prey do not really communicate, but one is simply trying to catch the other, as a matter of fact several signals are exchanged. We saw important examples in Chapter Six, with various cases of mimicry (maybe the best known form of interspecific communication) and

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other forms of deception based on the “handicap principle” that we introduced in Chapter Seven (birds pretending to have a broken wing to mislead a predator, gazelles jumping in front of a lion to show that they are healthy and fit, and so forth). Another important example is *aposematism*, which consists in warning signals, that inform the predator that the prey may be dangerous or unpalatable. Aposematism is generally characterized by visual signals (like for instance a particular body coloration of the prey), but more recently forms of “acoustic aposematism” have been investigated in caterpillars and ground squirrels. 2. Mutualism – Sometimes two different species can help each other for a common benefit. For instance, two different species of monkeys may develop common alarm signals, as they have the same predators. This of course increases their chance to avoid being predated. Another example of mutualism is the uncommon but not rare phenomenon of an older specimen of a given species adopting a young specimen of another: when this happens, the two animals develop a common communication system that significantly facilitates the relationship. 3. Parasitism – Nnot to be confused with mutualism, parasitism in communication occurs when one species “exploits” the signals of another, rather than sharing them. If, as we said, some species develop common alarm calls, there are others that simply learn the other species’s calls, and make use of them for their own advantage (for instance, they know that, when a certain call is emitted, a predator is coming). Since the advantage is not reciprocal, we cannot call this “mutualism”.

Interspecific communication between human beings and other animals The topic of interspecific communication is very rich and varied, and it is not possible to discuss in detail all the related phenomena. So, for this and the next chapter we shall focus on a very particular case of interspecific communication, the one occurring between human beings and other species. Even more specifically, we will take the case-study of language, that is, those attempts that various researchers made to train non-human animals to communicate with human language. First things first. Let us see how many types of interaction exist between humans and other animals that would require, at any point of the process, some communication:

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1. The human being can be a predator of other animals, and vice versa. Human beings exploit, through killing it, the lives of other species: they do it for food, clothes, research, etc. At the same time, the human being also happens to be a prey of certain non-human species: such is the case of sporozoites (those protozoa responsible for the malaria disease, transmitted to humans through the mosquito Anofele), but also of those occasional instances when specimens of different species attack humans in order to eat them or – more often – defend themselves. 2. The human being can also be a partner of other animals – Partnership relations between humans and other animals date back to ancient times. There are four particular typologies of partnership (see also Sebeok 1998: 68-9): a) host-guest relation, as in the case of pets; b) mutual dependence, as in the case of bee-breeding or dogs for blind people; c) sexual relation, as in the case of certain communities or certain pornography; and d) relations aimed at social facilitation, as in the case of pets used to facilitate the approach with other people or for therapeutic purposes. 3. The human being can exploit other animals in sports and hobbies – This category includes both very cruel examples like bull-fighting (corridas) and circuses, as well as very innocent ones like feeding pigeons or bird-watching. 4. The human being can be a parasite of other animals, and vice versa – This category includes cases like reindeer-breeding, where humans exploit every single resource offered by the species in question; or – in the opposite relationship – cases like the hairfollicle mite acari demodex, fleas, lice, etc. 5. The non-human animal can establish relations of conspecificity with the human being – This happens mostly with pets, who tend to identify their human host as part of their own species (or, more rarely, themselves as part of the human species). We do not have clear proof of whether humans can see other animals as conspecific, as well: however, such could be the case with children, for instance, or with humans bred in the wild by some non-human community, in the same way that inspired Edgar Rice Borroughs’ famous character of Tarzan. 6. The non-human animal can establish relations of insensibility with the human being, and vice versa – Some non-human animals may consider human beings as an inanimate object (like those birds who perch on a human body as if it were a tree). Human beings, too, can be physically insensitive towards other animals. This is mostly due

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to ignorance, so that some species (especially marine ones) are often mistaken for plants or others. 7. The human being can domesticate other animals – Domestication is defined as the “reduction or possibly total elimination of the reaction of escape in a non-human animal from a human one” (Sebeok 1998: 70). That also means that it is fair to state the opposite as well, as very often the process of domestication is mutual between the two species. 8. The human being can train other animals, and vice versa – The non-human animal is more or less forced to learn a given behavioral pattern on the basis of quite drastic options (e.g. grant or punishment). According to Heidiger (1968: 120), there are two types of training: a) apprentissage, i.e. laboratory training; and b) dressage, or circus-like training. The difference consists basically of the typology of relation established between trainer and trainee: almost inexistent in the first case, quite intensified in the second case. Sometimes human beings, too, can be trained by other species. In domesticated animals, for instance, it is quite clear that the relation is conducted by both parties in terms of reciprocal and constant training. People who take their dogs outside for the daily walk precisely when the dogs approach them holding the leash in their mouth, are rather evidently humans trained by dogs dealing with dogs trained by humans. 9. The human being can also be a manipulator of non-human animals – Cases like Dolly the sheep suggest that human beings now have quite a strong control of the genetic heritage of non-human species (in fact, of the human species as well). However, without only referring to very recent times, the domestication process of many species occurred and still occurs through a constant manipulation/selection. Finally, even single parts of characteristics of the non-human animal (especially physical and somatic ones) are manipulated for essentially aesthetic purposes (ear-pointing, tailcutting, etc.). 10. Finally, the human being can be defender/protector/promoter of other species, and vice versa – Such is the case of animal rights organizations, wildlife protection, and so on, but it is also the case of different individual attitudes. Non-human animals can also be at least defenders and protectors of the human being: typical is the case of a dog barking at or attacking an individual who is potentially dangerous for his/her human host.

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All these types of interaction require communication: in some cases the communication is extemporaneous and improvised – sometimes even involuntary (if we imagine facing a shark while swimming, there are certainly many signals exchanged, that indicate “danger”, “food”, “escape” and so forth, but we may well imagine that many of these signals are not intentionally produced, from neither of the two parties). In some other cases (like the aforementioned relationship with a pet) there is a much more codified interaction, with signals that are repeatedly exchanged, and that are perfected with time. With this in mind, we can now focus on our (very particular) case-study: the so-called “interspecific language training programs”.

The interspecific language training programs Do animals speak “language”, or do they simply “communicate”? Or, is human language something else (perhaps, something more) than other forms of communication? Here are a couple of questions that have been buzzing scholars for many years, in fact centuries. There is scarcely a more critical topic in the study of animal communication than the notion of “language”. To start with, scholars (across different disciplines, or even within the same one) do not agree on the definition itself. To some, “language” is a name we give to a communication system, so in that sense every animal species has its own language. To others, “language” is only the specific communication system adopted by the human beings: every communication system has (or should have) its own name, but “language” is the name we give to the human one. To yet others, the human language is something entirely different from any other communication system, and as such it cannot even be compared with anything else: in fact, some claim, language also serves more complicated functions than mere communication, like for instance the function of shaping and expanding human cognition (just to get an idea, through language one can “invent” and “describe” abstract or inexistent entities, like a god, a unicorn or the vampires of the Twilight saga. Through this invention, those entities become “real”, that is, people can share information about them, like writing a sacred text, shooting a movie, making a toy or a sculpture of one of these entities, etc.). As a trend, the fields of humanities are very eager to consider language as a species-specific human feature, radically separated from other animals (or even as the characterization of human identity: the very thing that distinguishes human beings from other animals. Or, even, the proof that we are not just animals), while natural sciences are often in favor of an

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extension of the concept, suggesting that, if anything, a difference between the human and other communication systems is only a matter of degrees. Generally speaking, and save several exceptions, natural sciences do not think that other functions (like the “cognitive” one we mentioned in Chapter Four) precede in importance the “communicative” one, as instead suggested in some fields of humanities, such as linguistics and semiotics. Secondly, most natural sciences do not agree that certain characteristics of language (for instance, the organization of signals in syntax and the use of symbols) are exclusive property of human language, as instead most fields of humanities propose. Counterproofs offered include phenomena like birdsong, bee dance (both often referred to as “language”), alarm calls, and else. Most of all, in order to check whether or not a gap exists between human communication and that of other species, some scholars thought of simply “training” other animals to use human language, in one form or another: the making and the result of these attempts, which we shall call Interspecific communication experiments, are the subject of this chapter. The expression “Interspecific Language Training Program” (ILTP, from now on) refers to a range of experimental programs conducted by biologists, psychologists or linguists, aimed at teaching human language to other animal species, particularly great apes. Generally, the scholars working on ILTPs advanced the basic claim that there is no unbridgeable gap between human language and other communication systems. Such a claim relied on the Darwinian idea of evolutionary continuity between Homo sapiens and other species, where language plays only the role of a more refined system for communication, not a unique one. Together with the Darwinian one, another stand, based on behaviorism, defined language as a conative tool, whose function is basically that of transmitting thoughts from a sender to a receiver. In this sense, language was seen as a “normal”, albeit complex, behaviour, that could be learnt as any other. The first experiments were affected by the (eventually proven) wrong assumption that the vocal and acoustic channel was the only reliable source for a communication system to be considered true language. In other words – the pioneers of ILTPs thought – the real nature of language is speech: other uses of language (writing, gestures, reading...) would not prove the point. The idea probably originated from writings by Samuel Pepys (The diary of Samuel Pepys, 1661), Johann Conrad Amman (Surdus loquens, 1692 and Dissertatio de loquela, 1700) and Julien Offray de La Mattrie (L’homme machine, 1748), who all shared the opinion that apes and monkeys could easily be taught to speak.

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Among the first experiments of this type, we have the chimpanzees Peter, raised by clinical psychologist Lightner Witmer around 1909, and most of all Joni, raised by ante-litteram cognitive primatologist Nadezhda Ladygina-Kohts (better known as Nadia Kohts) between 1913 and 1916. Born in Russia in 1890, Kohts can be considered a forerunner of modern *primatology*. Between 1913 and 1916, she raised a young chimpanzee, Joni (see Kohts 1935), in her home in Moscow, conducting different types of cognitive test (she checked Joni’s ability with tools, his capacity of recognizing himself in the mirror, his inclination to art, and so forth). Dr. Kohts worked specifically on emotional responses in Joni, providing indepth studies of such feelings as jealousy, guilt, empathy and loyalty, and describing facial expressions in detail. She would then compare Joni’s behavior to her own infant son, Roody. In general, her achievements were quite remarkable, but indeed language training ended up being her major failure: convinced that Joni had to speak language, she did not manage to obtain any acceptable sound from her chimpanzee’s mouth. Later relevant attempts to teach spoken language to chimpanzees were performed on Vicki, raised by Keith and Catherine Hayes between 1952 and 1958, and Gua, raised by Winthrop and Luella Kellog during the late 1960’s. All the chimpanzees – after years of intensive training – learned a maximum number of 3-4 words, of the likes of mama, papa, cup and up. On the other hand, however, they proved to understand dozens of words: they might have not been able to pronounce certain sounds, but they seemed to be able to figure out what they meant, and act by consequence. Similar attempts were also made with other primate species, like the orangutans trained by William Furness in 1916, and the gorilla Toto (trained by Maria Hoyt in 1942). Perhaps we can already guess what was the real problem, here (and it was later discussed and demonstrated in Liebermann 1968 and 1972). The problem laid in the neurological and anatomical incapacity of non-human bodies to articulate sounds in the way humans do. The difficulties of these apes were expressive, not cognitive. So, although these experiments failed, a breakthrough was actually made: the problem now was to find other ways to teach language, not relying exclusively on the spoken form, or, alternatively, working with animal species that do not have this expressive limitation. In the 1960’s, things started to get more serious, and more accurate experimental programs were designed. The first relevant one, in chronological order, was initiated by David Premack in 1966, as a chimpanzee named Sarah was trained to communicate with coloured plastic symbols, instead of “talking”. Importantly, the symbols were in no

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way related with the object represented (e.g. an apple, which is normally a red, yellow or green spherical item, was represented by a blue triangle): “symbols” was therefore the correct denomination of these objects, as they were not icons or indexes (see Chapter Three). The project, however, achieved mixed results: Sarah showed ability in understanding and representing concrete objects, plus a couple of abstract concepts like “name of” (a symbol used to ask the name of a certain object), symbols representing colours, big-small, equal-different, square-round oppositions, etc. She could also answer questions like “what is the color of…”, and in that sense everything looked quite encouraging. However, there was a problem: since day one, Sarah gave the feeling that she was not really “understanding” the symbols, but she had learned that by using a certain item she would get a certain result, without making the necessary mental connection that would show that the item is meaningfully related to that result. To understand the difference, let us imagine that some aliens land on our planet and want to destroy it. While they are about to do it, a young girl says “Look Mum! A real UFO!”, which makes the aliens stop for a minute. Witnessed the scene, everybody starts screaming “Look Mum! A real UFO! Look Mum! A real UFO!”: the aliens pack and go away. Why would they do that? The earthlings have absolutely no clue, but they understood that with those very words the aliens do not destroy the planet. They have no clue, for instance, that “Look Mum! A real UFO!”, in alien’s language, means “We are very close friends with Luke Skywalker: either you leave or we’ll call him!”: they just understood that there was a causal connection between those words and the response they all hoped for. In our story, the chimpanzee Sarah would be the earthling who has no clue of the signs’ meaning but learns that it is beneficial to perform them, and Premack would be the alien who believes that Sarah knows what she is “talking” about. What had happened was a phenomenon called Clever Hans Effect, whose explanation requires a little step back in time.

The Clever Hans Effect Der Klüge Hans (German for “Clever Hans”) was a German horse owned by a gentleman named Wilhelm von Osten, a math teacher and horse trainer, between the 19th and the 20th century. Hans made the headlines of local newspapers for allegedly being able to perform arithmetic and other complex intellectual tasks. During increasingly popular shows arranged by von Osten himself, throughout the whole of Germany, Hans amazed the spectators, answering

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diverse questions by hitting his hoof on the ground (he was known for adding, subtracting, multiplying, dividing, working with fractions, telling time, keeping track of the calendar, distinguishing musical tones, and reading and understanding German). As a problem was presented to him (either orally or in written form), Hans would start tapping his foot and stop when the exact figure was reached. For instance: one could ask him to calculate the square root of 81, and Hans would hit the hoof exactly nine times, and then stop, to everybody’s amazement. Obviously, not everybody was ready to believe that Hans was such a genius, so investigations started. A first analysis on Hans’ skills was performed in 1904 by psychologist Carl Sumpf and the so-called “Hans committee”, that is, 13 collaborators from different fields, including veterinarians, circus managers and zoo directors, only to conclude that no trick was involved in the horse’s performances. In 1907, another psychologist, Oskar Pfungst, introduced a few variables that turned out to be crucial (for instance, he prevented Hans from seeing the questioner, or made sure that the questioners did not themselves know the answer to the problem, etc.). This way, Pfungst demonstrated that Hans was not actually cognitively able to perform the tasks, but was instead very skillful in watching the reactions of his observers to his “counting”, until some involuntary cues in their body language would alert him that the task had been correctly performed. Which is understandable, of course: imagine asking a horse to calculate the square root of 81 and seeing that this horse is actually giving the right answer. Obviously, we would all stand openmouthed or say something like “ooohh!” or “wow!” Those reactions, Pfungst found out, were exactly what Hans was paying attention to: they were the actual signals that would confirm that he had given the right answer, so at that point he would stop hitting his hoof. After all, he was still a very clever horse: he might have not been the Einstein-type, but it seems he was very much the Freud-type! At any rate, the case of Clever Hans became paradigmatic for those experimental contexts where a) the subject of the experiment ends up manipulating, rather than being manipulated by, its trainers, and b) the trainers, driven by the enthusiasm of an apparently remarkable performance of their trainee, fall victim of excessive anthropomorphism in their interpretation. The phenomenon became known as “Clever Hans Effect”, and became the main criticism moved to ILTPs by some researchers. The fear of falling prey to this methodological bias encouraged some researchers to adopt the so-called emotion-free experimental contexts, loosely based on Pfungst’s principles of investigation in the Hans’s case, that is: removing all possible clues to the subject of the experiment. As we

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shall see, all the ILTPs that took place from the 1960’s onwards (after Premack’s Sarah project) could be roughly divided into two categories: those that arranged the “emotion-free” conditions, fearing the Clever Hans Effect, and those that did not, convinced that the presence of “emotions” (interactions, affection, cosiness...) is a necessary requirement for communication to operate. In the next chapter, we shall see, specifically, what the most historically-significant ILTPs were following the Sarah project, and what their main results were.

Fig. 10 – Wilhelm von Osten and the horse Hans

CHAPTER NINE INTERSPECIFIC COMMUNICATION (PART II)

Training with ASL Let us now return to the 1960’s. As it became clear that not only verbal communication had to be considered ‘language’, the interest in ILTPs increased, also because of the growing fame of behaviorism. In 1966, almost at the same time as Premack started his Sarah program, Allen and Beatrice Gardner initiated another program, this time based on *American Sign Language* (i.e. gestures, not spoken words) with the chimpanzee Washoe, with remarkable results. The success of this project paved the way for other programs employing ASL as interface. The most significant of these occurred in 1972, when Francine Patterson trained a female gorilla named Koko, in 1973, when Herbert Terrace trained a male chimpanzee called Nim Chimpsky, and in 1978, when the anthropologist Lyn Miles trained an orangutan named Chantek.

Washoe Washoe was a female Pan troglodytes born in 1965. The program was designed and initially carried out by the Gardners, then taken over by their former student Roger Fouts. Over the years, the project accumulated more chimpanzees, until a whole colony was formed (also consisting of Washoe’s direct descendants), that allowed the program to continue, in spite of Washoe’s death in 2007, at age 42. The Washoe project was a real breakthrough in the history of ILTPs. It was the first one to be generally acknowledged as a success, it set several methodological indications eventually followed in other programs, and it inaugurated a series of experiments based on linguistic sign systems alternative to speech. Washoe was trained with an ad hoc version of the American Sign Language (ASL, from now on). The Gardners wanted to prove that chimpanzees are able to learn a human language, and that real interspecific communication between humans and other animals is possible. The training was organized so that Washoe could imitate her

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trainers’ gestures, and – at the same time – through direct manipulation of her arms. The training environment was set in a cosy, relaxing, nonlaboratory-like atmosphere, an aspect which the Gardners and later Fouts considered very important in order for Washoe to be trained properly (Gardner-Gardner 1969: 666). The first signs taught were “more” and “come here”, both used for several purposes (“come here”, for instance, was not only a call for a trainer, but also a request for objects and actions: “come here food”, “come here tickle”, etc.). After the initial training stage, Washoe showed increasing curiosity in learning signs. Some were very close to the natural intraspecific gestures by which chimpanzees interact with each other. The sign for “come here”, for instance was similar to the *grooming* sign. Instead of considering that a problem, the Gardners encouraged such mixtures, in order to facilitate Washoe’s work, claiming that the same process occurs in human language, when abstract words or signals often carry characteristics of their meaning (as in onomatopoeic words: the word “splash”, when pronounced, makes a sound that reminds of the phenomenon it describes). Plus, their resemblance to “reality” does not make these signals less important than more abstract ones. The project’s outcome was very positive: Washoe spontaneously used the signs, even to name things, rather than just making requests, then started to associate the ASL signs to more complex meanings – for instance, the words flower or car were used not only for real flowers and real cars, but also for pictures portraying those items (Gardner-Gardner 1969: 667). Signs were no longer strictly related to the context in which they had first been produced: the sign “open” was used not only for doors, but also for refrigerators, windows, taps: Washoe, in other words, had become able to grasp the logic of these signs and their various characteristics (Gardner-Gardner 1969: 670). Furthermore, Washoe could also discriminate among the specific applications of a sign: in the beginning the sign “flower” represented all smelling objects, but later, when she learned the sign “perfume”, she understood when to use “flower” properly. Later on, Washoe showed interest in adjectives, and compiled metaphoric object-attribute relations (e.g. the term ‘dirty’ became an insult, not only the opposite of “clean”). In formulating sentences of the object-attribute or object-action type, Washoe had her personal rule on how to establish syntactic orders: with no exception, the sign that she perceived as most relevant or most dynamic was used at the beginning of her ASL sentences. So, it was “Roger good” and not “Good Roger”; “Bottle red” and not “Red bottle”; “Look there” and not “There look”, etc.

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This process appeared in most ILTPs with other animals, thus seems to suggest a basic perceptive syntax. Washoe was also given ASL-trained company (including equallytalented chimpanzees like Moja, Tatu and Ally, who used ASL quite regularly in intraspecific communication, as well), and was also allowed to form a family, with natural and adopted offspring. Her adopted infant, Loulis, by Fouts’ deliberate decision, was not taught ASL in the first 5 years of his life, but nonetheless acquired more than 70 signs by watching the other chimps (reflecting the manner in which human children acquire language). Later in time, Washoe taught ASL to her natural infant Sequoyah by her own initiative. Along with Koko, Kanzi and Chantek, whom we shall soon discuss, Washoe acquired over the years a wide popularity, becoming the “speaking ape” by definition. At the time of her death, thousands of condolence messages were sent to the website of the organization (www.friendsofwashoe.org), and several books and documentaries were made about her story (particularly Fouts 1999).

Koko Born in 1971, Koko (whose complete name was in fact Hanabi-Ko) is a female lowland gorilla (Gorilla gorilla) trained in ad hoc version of American Sign Language (a system nicknamed “Gorilla Sign Language”, GSL) by psychologist Dr. Francine Patterson. Like the Washoe program, this one too proved to display successful results. Trained since 1972, first in Woodside, California, then in Maui, Hawaii, Koko is reported to understand and use more than 1000 GSL signs, and to understand more than 2000 spoken words in English. Within the same training program, there were and are also other gorillas, namely Michael (Mike), a long-term “buddy” for Koko (the latter repeatedly signing GSL grief expressions when the former died in the year 2000), who reached a proficiency of about 600 signs (and also showed a strong inclination towards painting), and the younger Ndume, who was chosen by Koko herself in a kind of audition (Koko was exposed to some videos, and asked to choose her favourite companion from the gorillas displayed). Like Washoe herself, Koko too ended up being a sort of popstar, particularly after the internet came onto the scene. In 1998, indeed, Koko became the first non-human animal to “chat” through the Internet, in a specific event hosted by the AOL portal. One of the most important characteristics of Koko’s training program was a direct challenge to the Clever Hans Effect: Dr. Patterson was

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convinced that she and the other trainers had to establish a solid and emotionally-close relationship with the gorillas. Her theory was that children gain motivation in learning a language most of all because of the interaction with parents. To deprive them of such a fundamental element would mean to deprive them of language itself. Also, the apparent problem that the apes may not understand the signs they produce, but simply be aware of the causal connection with certain responses (much like the aliens-earthlings example we made before), was not something that would concern Dr. Patterson. In fact, her line of reasoning was rather plausible, as any parent could testify: when children start communicating actively, they go through a stage when they do not really understand the meaning of the things they say/do, but only their “power” to get them what they need. A relatively long period of time is spent refining this connection: it gets more precise in performance, it is refined in its application case by case, etc. Until, finally, with the development of the brain, something clicks, and the child understands that a particular sign actually “means” the very thing that was requested, and she/he realizes the enormous potential of communication. Not by chance, at this point, the intense (and occasionally exhausting, for parents) period of “what is this?”, “what does it mean?”, and “why?” begins. Back to Francine Patterson, the results achieved with her working philosophy proved very satisfying, just like Washoe’s case (who – as we have seen – received similar affective treatement). Koko became able to comprehend spoken words and use the correspondent GSL signs, she could create new signs to describe objects she did not know the name of (among the most revealing cases: “white-tiger” for “zebra”, “fingerbracelet” for “ring”, “drink-fruit” for “melon”, “animal-person” for “gorilla” and “water-bird” for “swan”). Such a skill is crucial in distinguishing a cognitive use of the signs from a mechanical one: to call a zebra “white tiger” means that there is a mental process determining this decision. It means that Koko thought that this animal, like a tiger, has stripes, but, unlike a tiger, is white. The referential use of the signs was also very important for Koko. She did not make only requests, but she would talk to Patterson about what she could see around her. Basically, she would engage in casual conversation, once more dismissing the widespread belief that language-trained apes could not go beyond a basic, very direct, use of the signs. Well-documented, as with Washoe, is also Koko’s extensive use of lies: a classic, in this category was her recurrent habit of stealing candies and then blaming her friend Mike when confronted by Patterson. Or, another reported case was the so-called “gorilla humour”: Koko would for

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instance ask to drink some juice, and Patterson (to encourage more elaborate conversations) would ask her where she wanted to drink it from. Koko would then start signalling the most improbable body parts (ears, nose, eyes...) and laugh after each suggestion. Only in the end, satisfied with the good time she had had, Koko would signal her mouth. Koko’s life and training is documented in both scientific and semiscientific literature. Among the former: Patterson 1977, 1978, 1981, and Patterson-Linden 1981. Interesting resources are also available at The Gorilla Foundation website (www.koko.org).

Nim Chimpsky One year after Koko, another ASL-based program was initiated by Dr. Herbert Terrace on a male chimpanzee named Nim Chimpsky (an obvious pun with the very popular linguist Noam Chomsky). After few years of apparent success, in which Nim apparently acquired knowledge of ca. 125 signs, Terrace had a very famous second thought: he realized that the whole training process was heavily affected by the Clever Hans Effect, and – with rare professional honesty – he admitted it. Nim, Terrace argued, had really understood not more than 25 signs, while the rest was mere result of that manipulative cause-effect trick that the Hans case had first revealed to the world. Terrace’s failure did not prevent him from becoming a very wellknown figure within the scientific community. He wrote a book called Nim Chimpsky, the Chimpanzee Who Would Be Human that brought him fame and admiration among those colleagues who also believed that nonhuman animals would never be capable of properly understanding human language. The zoosemiotician Thomas Sebeok, who was among such colleagues, even went as far as to say “In my opinion, the alleged language experiments with apes can be divided into three groups: one, outright fraud; two, self-deception; three, those conducted by Terrace”, implying that Terrace had been the only honest researcher in the bunch. The very sad part of the story was the tragic end suffered by Nim: by now useless to Terrace’s purposes, he was sent to the Institute for Primate Studies in Oklahoma, where he had to adapt to a totally different life. Terrace went to visit him only once, and on that occasion Nim showed considerable progress, spontaneously signing various words of welcome and friendship to Terrace. Unimpressed (or, more likely, uninterested at this point, since he had anyway achieved his success by becoming the hero of the sceptics), Terrace left without visiting Nim anymore, a factor that, among others, brought the ape into a growing state of depression. The

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Institute later sold Nim to a pharmaceutical animal testing laboratory, the Laboratory for Experimental Medicine and Surgery in Primates, managed by New York University. Nim became subject of heavy experimentation. Perhaps even more heartbreakingly, during this most difficult time, medical workers reported that Nim would make signs of “hug” and “play” to them. Finally, he was purchased by the Black Beauty Ranch, operated by The Fund for Animals, in Texas, where he lived in slightly better conditions, but was isolated inside a pen. His behavior became more aggressive, until he was joined by other chimpanzees, several coming from the same pharmaceutical lab in which he had been imprisoned a few years earlier. He continued to use ASL signs whenever a researcher from the Institute would go and visit him. He finally died in 2000 at the age of only 26, from a heart attack. It is important to mention this tragic story, not necessarily to point out that perhaps, after all, Nim learned more than Terrace thought, but rather to stress on the ruthlessness and cruelty of the majority of these experiments. Regardless of how successful they may be, they remain the result of our human arrogance to use and abuse other animals in any way we fancy. While, via this book, the hope is to get more and more students interested in animal communication studies, the plea is to always pursue this subject in a non-invasive, ethically sound way.

Chantek The fourth important ASL-based experiment was initiated in 1978, by anthropologist Dr. Lyn Miles, on a male orangutan of the species Pongo abelii named Chantek. Named after the Indonesian “Cantik” (beautiful, lovely), Chantek was born in 1977 and was raised at the Yerkes Regional Primate Research Center in Atlanta (Georgia), where the training program took place. As every new ILTP was trying to improve upon its predecessors, the Chantek program, too, presented a few innovations. Besides the fact that, this time, the trainee was an orangutan (something rarer, but – as we have seen with William Furness’s research in the early 20th century – not unique), the distinctive feature, here, was that the trainee was put in a context in which all trainers would communicate with ASL, even among themselves, and nobody was making specific attempts to train him directly. Simply, when Chantek was told something, he was told in ASL, and when he wanted to communicate, he had to use ASL. By consequence, and pretty soon, the orangutan learned to use the sign system

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spontaneously and rather efficiently, amounting to a vocabulary of several hundreds of signs, later enriched by a quasi-second language: the understanding of spoken English. His skills were so convincing that they had a political impact in the application of the ethic and juridical concept of personhood in Great Apes (as, for instance, promoted by Paola Cavalieri’s and Peter Singer’s Great Ape Project). Chantek is also known for his creativity, particularly expressed in toolmaking, painting, several forms of handicrafting (especially necklaces), and even in music-making – alone or in collaboration with Lyn Miles herself, who is also a professional percussionist (the “duo” has also released some recordings under the stagename Animal Nation). Furthermore, Chantek allowed significant advancements in the studies of self-consciousness in non-human animals, by displaying a great variety of revealing reactions to the famous *mirror test*. Besides several documentaries and “popular science” articles and books, Chantek’s efforts in ASL training are also academically documented in, among others, Miles 1990 and 1994.

Training with lexigrams One problem related with the employment of ASL in the training programs – it was perceived – was that it is not very easy to avoid the Clever Hans Effect, because one anyway needs to closely interact with the trainee. On the contrary, at this particular stage of ILTP history, scholars (particularly after the Terrace fiasco) were quite frightened at the perspective of falling into that trap, therefore they started elaborating ILTPs that would exclude, as much as possible, any interaction between trainer and trainee. Duane Rumbaugh, starting from 1975, attempted to teach language by employing symbols called “lexigrams”: not the type of plastic objects used by David Premack with the chimpanzee Sarah, but rather bi-dimensional abstract figures that could be printed on a paper or displayed on a screen or on a keyboard. The trainee was a female chimpanzee named Lana. Rumbaugh felt that computer automation could prevent anyone from cueing the animal, so a specific machine was programmed for performing certain tasks, like dispensing food or displaying an image in response to Lana’s requests made with the lexigrams. Lana was given a special keyboard which had these lexigrams in place of regular keys. The trainer was connected to Lana with another, equally equipped, computer. The interaction would thus occur only through the respective monitors.

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As in all programs maniacally concerned with avoidance of Clever Hans Effect, the results of this ILTP, too, were controversial: Lana learned to formulate requests and to engage in relatively complex conversations, but she would also do so only in response to a specific need (usually, hunger or thirst). Other than that, Lana looked quite unmotivated to interact. Which of course raises an important question: why would anybody find motivation in learning another communication system if there is nobody in the room to use it with? If there is a phenomenon on this planet that is quite useless in conditions of isolation, that is communication. It would be pretty bizarre (in fact, very sad) if parents expected their children to learn to communicate without being in the same room as them, but simply interacting via monitors. A bit of direct contact is not only useful: it is essential.

Kanzi Duane Rumbaugh’s program was however far from useless, as it proved to be a solid basis for a later one, conducted by Rumbaugh’s wife, Dr. Sue Savage-Rumbaugh, which resulted in one of the most interesting and successful experiments of the whole series. The protagonist was this time a male bonobo of the species Pan paniscus called Kanzi, born in 1980, who was trained with a technique that was still based on the use of lexigrams, although in a different manner. The program was (and still is) conducted at the Language Research Center at Georgia State University. Dr. Savage-Rumbaugh departed from a quite critical attitude towards other ILTPs: she felt that, rather than establishing whether apes understand the signs they produce, previous experiments were more concerned about finding out about grammatical capabilities. In all previous programs, she maintained, the primates were not using signs symbolically, but rather indexically: they learned to associate certain behavioral patterns with certain consequences (once again, our funny example of the aliens and the earthlings can be mentioned: Savage-Rumbaugh felt that all previous programs had led to that scenario). In her view, the main point of human language is that it uses symbols, according to a three-step process: 1) the physical external substance of a word, e.g. “door”, as either written down or spoken with a given linguistic pattern; 2) the relationship between that and a real door (when the word “door” is pronounced one conjures up a mental representation of a door); and 3) the capability of symbols to make one think about things that are not present, or even not existent.

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The experiment, performed on a small community of bonobos, among which Kanzi is the most famous, achieved impressive results. Kanzi, who was born when the project had already begun with other apes, took everybody by surprise from the very beginning of his own training, showing that he already knew some lexigrams and could already understand a remarkable number of spoken words (evidently, this competence was achieved by observing his fellow apes, especially his adoptive mother Matata, who was part of the initial stage of the project). Kanzi’s competences grew rapidly. Once it became clear that he did not need a “basic course” to grasp the meaning of spoken words and written symbols, the trainers abandoned the usual reward-based training, and went straightaway to the “advanced” level, by adding new symbols to his keyboard. Kanzi was not trained to associate the lexigrams with specific objects, but rather the trainers would naturally employ the symbols in conversations with him, generally helping him to understand the general sense (as is often done in certain techniques for learning a foreign language). This way, Kanzi’s vocabulary steadily increased to over 200 produced and 500 understood words. Kanzi, particularly, showed an unquestionable comprehension of spoken words, something that had always been a weak point of the other ILTPs. In the various tests he was given (including recorded words played to him through headphones), he was nearly 100% accurate on all words that were part of his vocabulary at any given age. He was also able to respond to speakers with different accents as well as artificially-produced words. Far from succeeding only with single words, Kanzi showed comprehension of sentence structures, word order and grammar rules. Although one may be tempted to think that Kanzi himself was a particularly gifted bonobo (something that is probably true anyway), the effectiveness of the whole program was proved by the fact that similar results were achieved with two other bonobos of Kanzi’s community, Panbanisha and Panzee. For more, see Savage-Rumbaugh et al. 1998.

Not only apes To conclude our review, the 1970’s also witnessed the appearance of ILTPs that involved different species from great apes. The ones that we shall mention here were conducted by Lou Herman on two dolphins, Phoenix and Akeakamai, and by Irene Pepperberg on the parrot Alex. The Phoenix and Akeakamai program started in 1976, and was particular not only because of the involvement of a species that was not previously

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considered within this type of research, but also because the two dolphins were in fact trained at the same time with two different methods. Phoenix received training on an acoustic language generated by an underwater speaker, while Akeakamai was exposed to an adapted version of the thenvery trendy ASL. In both cases, the words produced consisted mostly of concrete entities, actions, modifiers (like “right”, “left”, “bottom”, “surface”) plus important signs, such as “yes”, “no” and “erase” (the latter used as a signal to interrupt an activity). Herman and his collaborators were among those who were thoroughly preoccupied with the Clever Hans Effect. In order to avoid it, the trainers wore masks that prevented them from showing facial expressions that could influence the two dolphins. Moreover, one trainer would give the task, and another one, without knowing what it was, would record the dolphins' reactions. An important element was the syntax of the messages, which were organized in coherent temporal sequences, like “surface frisbee bring-to basket” (i.e. on the surface there is a frisbee: bring it into the basket: as we can see, a different order of the words would have produced a different task). When a task could not be performed, the dolphins would touch a panel correspondent to “no”; when it was, then they would perform it, and eventually go and touch a panel correspondent to “yes”. This project, too, achieved interesting results. Phoenix and Akeakamai also proved able to understand messages that referred to a distant time. They would memorize the instructions, and – as the conditions became favourable – perform the task.

Alex Although the Phoenix-Akeakamai project received consistent international resonance, it is probably safe to say that the most famous experiment involving a species that was not a great ape was the Alex project. In 1977, Dr. Irene Pepperberg initiated a program that pursued a very old idea: teaching animals “to speak”, in the phonetic sense – the very idea that was ruled out years earlier. When it became clear that, despite psychic faculties that are close to the human ones, great apes do not have adequate anatomic and physiological characteristics to learn spoken human language, Pepperberg thought of teaching them to speak to a species that does not have limitations in that department, such as parrots, and more specifically a male African Grey Parrot of the species Psittacus erithacus named Alex. The experiment was also meant to disprove the stereotype that parrots can only repeat things… like parrots (that is, mechanically, without a real cognitive activity underlying the whole process).

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Born in 1976, Alex was an abbreviation for Avian Learning EXperiment. The training was conducted in different American universities until 2007, when Alex died of unknown causes at age 41 (against a life expectancy for his species of about fifty years). This program, too, was conducted rather successfully. Alex was able to understand and correctly pronounce about 150 English words, including some that imply the knowledge of other words (or entire concepts) they refer to, or that they modify in some way: “bigger”, “different”, “same”, “zero”, “over”, “under”, “smaller”, and others, all of which Alex managed to learn properly. He also could master quantities of objects up to six items, and also, not long before his death, started to learn reading, in order to relate written and spoken words (at the time of his death, he had been able to learn reading sequences of two letters). He was also, much like Koko and Washoe, capable of inventing new words, when his vocabulary was short of proper terms. The most frequently reported example is the word “Banerry”, a combination of the known words “Banana” and “Cherry” for naming an apple (probably referring to the yellow pulp and the red peel). All in all, Alex was estimated to possess the intelligence of a 5 year old human being. Alex’s daily training consisted of different types of interviews, in which the trainers would ask him to name, request and describe objects and actions. When the answer was correct, instead of directly receiving a treat, Alex had to explicitly ask for it (e.g. “Want banana”, “Want nut”). The main criticism moved to the Alex program, and in fact to all ILTPs of the last half a century, is our by now familiar Clever Hans Effect. That is, it was argued that – in one way or another – the researchers were so anxious to get a positive result from the tests, that they would either “help” Alex give the correct answer (particularly with non-verbal clues, like an encouraging smile, some particular gesture of approval, etc.), or overinterpret the results, thinking that Alex made “on purpose”, something that in fact he had done at random. Aware of the risks, Dr. Pepperberg and her team took their precautions to avoid the effect. The experiments were indeed conducted through a method called “double-blind way”: one trainer would ask the questions to Alex, and another one, unaware of the questions, would interpret and transcribe the parrot’s answers. With this method, Alex had a score of correct answers of 80% of the questions he was asked (including rather specific ones), and his mistakes were basically down to two factors: 1) misunderstanding: sometimes Alex would react to words that are similar in sound to the ones actually pronounced, (cable/table, fork/cork, etc.); 2) interpretation: for instance, Alex would classify a piece of paper and a piece of leather in the same way. Possibly his criteria for categorization

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had more to do with the bi-dimensional and squared shape of the two objects, rather than the material of the two objects. In that sense, the request had to be considered inadequate as it contained more than one variable, and was therefore subject to different interpretations. Shortly before Alex’s death, the new “sensation”, in the field of speaking parrots, became another specimen of Psittacus erithacus named N’Kisi. By January 2004, N'kisi was reported to possess a vocabulary of nearly 1000 words, frequently combined in complete sentences, with appropriate verb forms and tense, and with a variety of applications that go far beyond the information collected during the training. After all, we may have underestimated parrots! For more, see Pepperberg 1999.

Fig. 11 – Three of the most recurrent methods employed in interspecific language training programs. From top, clockwise: five letters in American Sign Language (employed with Washoe, Koko and others), 25 lexigrams (employed with Lana, Kanzi and others), and four plastic symbols (employed with Sarah)

CHAPTER TEN 30 KEY-FIGURES WHO SHAPED ANIMAL COMMUNICATION STUDIES

There is no doubt that, in the course of your studies, you will come across many important scholars in animal communication: the topic is nowadays so widely investigated that new research and publications are emerging at a constant rate. In this sense, it would be a hopeless enterprise to try and list all these people, and summarise what they do in a few words: no matter how hard we try, we would end up omitting someone, and anyway such a list would soon be outdated by new research and new publications. However, there is perhaps something more feasible and useful we can do here: we could instead try to mention some important characters of the past, who were crucial in the development of what eventually became the field of “animal communication studies”. Some such characters would be easy choices, and certainly you need to get acquainted with them already from now (Charles Darwin or Konrad Lorenz being the most obvious examples), but there are also others that you may never have the chance to meet in the future, especially if you specialize in some area of natural sciences (as is predictable, if you have an interest in animal communication). These two chapters may easily be your only opportunity to learn what the likes of Aristotle or Pierre Bayle did to help develop the knowledge on how animals communicate, behave and operate cognitively. Also, it will introduce you to rather obscure figures that you may have never heard of. Hopefully, this and the next chapter will be an interesting journey into some key-figures of the history of human knowledge. You will get to know a few of the “classic” scholars in animal communication, and a few of those who, despite being specialized in other areas of investigation, managed to produce research and arguments that contributed enormously to the growth of this area of inquiry. The list of entries proceeds in chronological order, not in alphabetical one. Such a choice was made in order to give you a better understanding of the actual progress of the field.

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ARISTOTLE (384-322 BC) – Greek philosopher. One of the most important and influential figures in Western thought. His range of interests was so wide that it also reached the area of animal studies, where he performed both empirical observations (he was a forerunner in several zoological and ethological studies) and philosophical reflections. His contribution is however rather ambivalent. Aristotle was at the same time an attentive observer of Nature, but his interpretations of these observations were often a bit biased, mostly because – as a philosopher – he had already constructed a certain idea of Nature and life, and that ended up affecting his whole view on the various phenomena observed. Paradigmatic is his definition of Scala Naturae, a hierarchical scale from the most to the least “perfect” being, which – judging from a modern, scientific point of view – is full of inaccuracies and altogether rather naïve. Plus, what had an enormous impact over the whole Western thought, Aristotle considered non-human animals a category totally subordinated to humans (particularly to free men – women and slaves were also considered “inferior”). In Aristotle’s hierarchy, there is no real interaction between superior and inferior categories. The former act upon the latter, they impose an order and affect their nature and behaviour, without being affected in any way. There are five main works of Aristotle: History of animals, Parts of animals, Generation of animals, Movements of animals and Progression of animals. Exactly as mentioned, all these texts alternate brilliant intuitions with rather large mistakes: Aristotle observes muscles and nervous system but he seems incapable of understanding their functions; he sees veins and arteries but he does not distinguish them; he gathers information about the reproductive act, but he fails to grasp very basic aspects (to mention one, semen is to Aristotle merely aimed at sexual excitement); he sees the brain as an important organ of the body, but not as the central one, maintaining instead that its sole function is to chill blood, and so forth. Several other considerations are instead remarkable. The observations on the anatomy of octopuses, cuttlefish, crustaceans, and many other marine invertebrates are detailed and accurate. Aristotle distinguishes cetaceans from fish, describes the embryological development of a chick, the chambered stomachs of ruminants and the social organization of bees. Most of his observations would only be confirmed many centuries later. Aristotle groups together animals with similar features into genera (although the term is used in a much broader sense than nowadays) and then distinguishes the species within the genera. Animals are then divided into two types: those with blood, and those without blood (or at least

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without red blood): the distinction is closely correspondent to that between vertebrates and invertebrates. Blooded animals are organized into five genera: viviparous quadrupeds (mammals), birds, oviparous quadrupeds (reptiles and amphibians), fish, and whales. Bloodless animals are cephalopods (such as the octopus), crustaceans, insects, shelled animals (such as most molluscs and echinoderms), and “zoophytes”, or “plantanimals”, supposedly resembling plants in their form, such as most cnidarians. Altogether, there is not so much investigation about the phenomenon of animal communication in itself, but the amount of information that Aristotle produced in anatomy, physiology, taxonomy and other areas is so big that the whole area of animal studies received a potent boost that was beneficial for centuries. Plus, Western philosophy as a whole, has in Aristotle the most significant of roots, and as such his work simply cannot be disregarded. GAIUS PLINIUS SECUNDUS (24-79) – Roman philosopher and naturalist, also known as Pliny the Elder. His presence in this chapter is self-evident by his massive encyclopedic work Naturalis historia, which arguably comprises the entire knowledge on Nature and animals available at the time. It consists of 37 books, of which the 8th, the 9th, the 10th and the 11th are entitled “Zoology”, and contain several topics of interest for us. Among these: the detailed (and fairly complete) description of a few species (particularly the honey-bees), a few hypotheses on zoological phenomena and a rather poetic (and still scientifically interesting) description of the nightingale’s singing activity. PORPHYRY OF TYRE (234–circa 305) – Neoplatonic philosopher born in Tyre (nowadays Lebanon). He is one of the most important forerunners of animal rights philosophy, supported with rational arguments. On abstinence from animal food (Perì Apokhês Empsykhon, in English also known as On abstinence from killing animals) is a passionate ethical defense of non-human animals. Among the arguments used by Porphyry to support his beliefs, we find at least three that are very interesting for this book: 1 – Animals do think and communicate. In contrast to the philosophical currents of the stoics and the peripatetics, Porphyry maintains that we can find the logos, the discourse, among nonhuman animals, and that this discourse can also reach “perfection” (that is, a full state of completeness and efficiency).

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2 – Animals also have the “inner discourse”: the general organization of their organism is similar to the human one, e.g. they suffer the same pathologies. Animals are not less sensitive than humans. To maintain that a different physical constitution corresponds to the absence of reason and sensibility is like saying that gods are not sensitive either, because their physical constitution is also different from the human one. The difference between humans and other animals is (in a proto-Darwinian fashion) a matter of more/less, rather than presence/absence. 3 – Animals are intelligent and rational. Porphyry mentions the great amount of information collected by ancient philosophers on the topic. To be able to take care of one’s own interests is a first important sign of intelligence (“each animal knows where it is weak and where it is strong, and it protects the former and makes use of the latter, as the leopard uses its teeth, the horse its hooves and the bull its horns, the cock its spur and the scorpion its sting”). Ratio, to Porphyry, does not originate from learning, nor from memory, and that applies to all beings, including gods. The sole fact that we cannot see the world through their own senses and figure out their own way of reasoning is not a good excuse to state that a ratio is missing. Moreover, non-human animals understand our language to many extents, and perceive the diverse signals. For more, see Taylor 1823. KIRCHER, ATHANASIUS (1601-1680) – German scholar, “master of a hundred arts”. Often referred to as “the last Renaissance man”, Kircher cultivated the most diverse interests in the most diverse fields of knowledge. He provided a very intriguing contribution to the study of animal acoustic communication. In 1650, Kircher published the twelve books of Musurgia universalis (a work nowadays known as the forerunner of the so-called “theory of passions”). The first volume (Anatomicus de natura soni et vocis, chapter “De vocibus naturalibus in animalibus eorumque anatomia”) is, considering the times, one of the most complete dissertations on the subject of animal vocalizations, particularly “singing”. Kircher essentially works on three levels: 1) direct investigation of the anatomical and physiological aspects of sound production; 2) personal observation, which points out those features that are not detectable on empirical basis; and 3) anecdotes, collecting facts related to the subjects discussed. He elaborates a classification of singing species, using the (very limited) zoological knowledge of that time (Kircher 1650: 25), and

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noticing that the definition of “voice” is something that fits best to humankind and some birds, while nature has gifted other animals with rough vocalizations (“Boues mugiunt, balant Oues, Equi hinniunt, barriunt Elephantes” and so on – ibid.: 26), that are nevertheless still suitable for displaying emotions and feelings. Kircher transcribes different birdsongs on score, and does the same also with the American animal Pigritia (the sloth Choeloepus didactylus). Birds remain, however, the main focus, and are taken into consideration in sections 3-5 of the chapter. The species analyzed, and transcribed on a score, are the parrot (with their “amazing imitations of human speech”), the nightingale (with their “beautiful chant”, and their capacity to sing “planus, gravis, acutus, creber, extensus” – ibid.: 29), the magpie, the wren, the rooster and the hoopoe. The chapter is concluded with a small dissertation on insects (cicadas, locusts and crickets) and amphibians (frogs), of which Kircher provides only anatomical descriptions. BAYLE, PIERRE (1647-1707) – French philosopher, particularly instrumental in the development of agnostic and secular ideas in Western thought, by arguing that faith cannot be justified by reason. In his Dictionnaire historique et critique there is a very interesting entry “Beasts (souls of)”, which discusses matters regarding reasoning and other inner states in non-human animals (Bayle 1697/1825: 199-219), in an openly anti-cartesian fashion. The entry starts with a direct attack on mechanism (“a strange notion”), and then proceeds by defining the elements in perception and cognition in animals. Among the topics tackled (all gathered in the umbrella term “soul”), there is awareness, memory, intentionality, sensations, reasoning and morality. The general message conveyed is that of the biological continuity between humans and other animals, where differences are never of qualitative type, but just matter of degrees (as Darwin will later point out very clearly). DARWIN, CHARLES ROBERT (1809-1882) – English naturalist. Quite simply, the most important animal scholar of all times. All fields of inquiry pertaining to the study of animals owe an incalculable debt to his theories and observations, and there is certainly no point in listing all of them, besides those that have been instrumental for the development of animal communication studies (and still considering that Darwin’s many other contributions to animal studies were instrumental to this field anyway, although possibly in a less direct way). If the theory of Natural Selection is what brought Darwin most of his fame, it is in his later works that one finds the basis for the development of

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fields related with animal communication, such as cognitive ethology, zoosemiotics, sociobiology, and other fields (including research ethics: Darwin’s methodology, notoriously, relied upon personal observations and descriptions of an anecdotal and non-invasive nature). In The Descent of Man (1871), Darwin introduces reflections and observations about animal cognition, reasoning, the use and making of tools, consciousness, learning, aesthetics, sociality and morality. In the subsequent The Expression of Emotions of in the Man and Animals (1872) Darwin follows up the arguments of the previous book, and focuses on topics that are even more pertinent to modern research on animal communication: signalling behaviour, representation and interpretation. Darwin also meticulously describes the main features of what is nowadays known as multimodality. The importance of Darwin’s writings in the context of this course resides, among other things, in their modernity. Accurate descriptions of signalling behaviour, important methodological distinctions (he distinguishes between intraspecific and interspecific communication; he separates innate, learned and ritualized sign repertoires, etc.), and specific theoretical formulations. It is safe to affirm that Darwin is to animal studies as Bach is to music or Shakespeare to theatre. UEXKÜLL, JAKOB JOHANN VON (1864-1944) – German-Estonian biologist. Most of Uexküll’s fame revolves around the revolutionary notion of Umwelt. This word means “Environment” in German, but in Uexküll’s formulation it came to designate a much more complex notion, usually referred to as “subjective universe” (Sharov 2001: 211). The starting assumption is that the environment inhabited by an organism is not merely the actual environmental niche, but is a larger not purely physical ‘environment’, of which the niche is just a part, that is perceivable and meaningful in its entirety only from the perspective of that particular organism. Uexküll (1982: 29-30) exemplifies this concept by describing the completely different meanings that a flower may have to a young girl (an element of decoration), an ant (a path for reaching food), a cicada-larva (construction material), and a cow (fodder). An Umwelt is the result of a Merkwelt, i.e. the specific perceptive field of a given organism, and a Wirkwelt, i.e. the field of actual interaction (which means communication too, of course) between the organism and the environment. Perceptual and operational elements come to establish the specific Umwelt of the given organism, which is exclusive for each species, in particular, but also – broadening the concept – for each community, individual, class, family and so forth. The Merkwelt and the

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Wirkwelt are constantly in action, as the organism affects and is affected by the environment. In that sense, the process is described as an “Umwelt circle”. REGEN, IVAN (1868-1947) – Slovenian biologist and entomologist. He is generally regarded as the founder of the discipline of Bioacoustics. After World War I, he conducted natural and artificial observations of katydid and cricket stridulation (using also such devices as microphones and loudspeakers), finding that insects respond to acoustic stimuli from other individuals. He also provided the first description of the functioning of insects’ hearing organs, and later studied other topics of entomological interest (not directly related with communication), such as the mechanisms of breathing and hibernation. FRISCH, KARL VON (1886-1982) – Austrian ethologist and Nobel Prize winner in Physiology or Medicine in 1973. Along with Konrad Lorenz and Nikolas Tinbergen, with whom he shared the prize, von Frisch must be considered the most important figure in classical ethology. His main contribution to animal studies remains the extensive research conducted on the language of honey bees, together with his then student Martin Lindauer (as documented, among others, in Frisch 1967). Their work included studies on the bees’ sensorial mechanisms, their various communication systems, their sensitivity to polarized light, and the pheromones emitted by the queen bee and her daughters (these topics are also discussed in Chapter Five). KOHTS, NADIA (1890-1963, full name: Nadezhda Ladygina-Kohts) – Russian primatologist, she had a significant impact on the study of chimpanzee communication. Between 1913 and 1916, she raised a young chimpanzee, Joni (see Kohts 1935), in her home in Moscow, conducting different types of cognitive tests (including tool, mirror, art and discrimination tasks). She invented the matching-to-sample paradigm, and worked specifically on emotional responses in Joni, providing in-depth studies of such feelings as jealousy, guilt, empathy and loyalty, and describing facial expressions in detail. She would then compare Joni’s behavior to her own infant son, Roody. Among her experiments, one of particular interest is her failed attempt to teach spoken human language to the chimpanzee. LORENZ, KONRAD (1903-1989) – Austrian ethologist and Nobel Prize winner “for discoveries in individual and social behavior patterns”.

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Possibly the most influential figure in classical ethology (see Chapter One), and certainly the most popular one. Following the steps of his teacher and mentor, German biologist Oskar Heinroth, Lorenz contributed crucially to the foundation of ethology, in its modern form. Among the (many) concepts he helped define and/or develop, one must mention at least instinct (a.o. in Lorenz 1949, 1963, 1965), imprinting (a.o. 1949 and 1979), human-dog co-evolution (1950), aggression (a.o. 1965), plus an extensive discussion on humanity and its role within and towards Nature, especially in the last years of his life (a.o. 1973 and 1974). Among the species he observed closely, there are the jackdaw Corvus monedula, the greylag goose Anser anser and other *nidifugous* birds. BATESON, GREGORY (1904-1980) – Primarily an anthropologist, Bateson’s work covered a range of interests that included biology, psychology, semiotics, social sciences and cybernetics. He offered several important insights for animal communication studies, in terms of both methodology and analysis. Worth mentioning are at least his reflections on such topics as instinct, play, aesthetics, mind, communication and intelligence. His reflections, and writing style too, are often provocative and unconventional, and possibly best embodied by his famous “metalogues”, imaginary dialogues between a father and a daughter over controversial topics of the most diverse nature. In the article “Problems in Cetacean and other mammalian communication” (Bateson 1972: 364-78), Bateson addresses the difficulty, as humans, in approaching communication processes in non-human species, with the hypothesis that the latter might use communication in a consistently different manner. The communication of relationship statuses (affection, disaffection, dependency and so on) is to Bateson the predominant communicative function in non-human (that is, to him, nonlinguisitic) mammals. Among humans, the communication of relationship is less explicit when it comes to language, and mostly found in that protolinguistic communication (such as gestures and proxemics) that is common to all mammals. TINBERGEN, NIKOLAS (1907-1988) – Dutch ethologist, ornithologist, and Nobel Prize winner in 1973 together with Karl von Frisch and Konrad Lorenz. One of the most important figures in modern ethology, Tinbergen’s interest covered a wide range of topics of interest for our course, including studies on the notion of instinct (a.o. 1951) and social behavior (a.o. 1953 and 1974). His contribution extends to the very foundations of methodology in ethological sciences, often influencing

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other fields of inquiry like sociobiology and social sciences, as in the wellknown case of the so-called “Four Questions” (Tinbergen 1963), a methodological model articulated into two categories (including two questions each): Proximate mechanisms and Ultimate mechanisms. Proximate mechanisms include 1) Causation (or Mechanism), investigating the stimuli eliciting the response, and the modification of the latter after recent learning; 2) Development (or Ontogeny), investigating the modes of behavioral change in time units (age), in relation with earlier experiences. Ultimate mechanisms concern 1) Evolution (or Phylogeny), investigating the animal behavior in relation with other species performing similar patterns, and in general the phylogenetic aspects of such behavior; 2) Function (or Adaptation), investigating the impact of the given behavioral pattern/s on the animal’s general survival and reproduction chances. HEDIGER, HEINI (1908-1992) – Swiss zoologist, director of Zürich zoo and founder of the field of “zoo biology” (i.e. the ethological observation of captive animals). Hediger had a crucial impact on the study of human-animal interaction and communication in contexts of captivity. He closely studied the *proxemics*, social and territorial aspects of animal behaviour, with such focuses as: 1) Animal expression, defined as “variable nonpathological phenomena of the animal, which may help to an understanding of their situation” (Hediger 1968: 143): face-displays, general posture, *kinesics* are organized into acoustic, optic, olfactory and “internal” expressions (the latter including reactions of secreting tract, vomiting, urinating, level of adrenaline, etc.); 2) Animal’s managing of interpersonal distances (e.g. 1968) in social or generally interactional contexts; 3) Different forms of human-animal relationship (e.g. 1965); 4) The Clever Hans Effect (e.g. 1981). GRIFFIN, DONALD (1915-2003) – American zoologist. He is generally considered the founder of cognitive ethology, or rather his 1976 book The Question of Animal Awareness is considered the main transition

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point from classical to cognitive ethology. “Historical” role apart, Griffin performed extensive research on several topics related with animal communication, including navigation, acoustic orientation and sensory biophysics, delivering a major contribution to the discovery of *echolocation* in bats (see Griffin 1958). LINDAUER, MARTIN (1918-2008) – German ethologist, specializing in animal communication. A pupil of Karl von Frisch, with whom he collaborated for extensive research on the bee dance, Lindauer was not only a field researcher on communication, but also a remarkable theoretician on the subject. Works like Lindauer 1990 are crucial for the meticolous systematization of biocommunication research (see Chapter One), including a definition of it, a classification of the channels, and an investigation of its social dimension. TEMBROCK, GÜNTHER (1918-2011) – German ethologist. Professor at Humboldt University in Berlin, he there founded the first German research facility for ethology, in 1948 (in times when the field was still known as Animal Psychology), and the largest European Animal Sound Archive (Tierstimmenarchiv) in 1951 (the archive currently consists of some 120,000 recordings). His contribution to ethology is enormous, and his interests intimately keen on animal communication, most notably in the study of acoustic signals. In this area of inquiry, Tembrock provided an extensive body of research, including theoretical and methodological innovations. He postulated the term and concept of Biocommunication in 1971, while his promotion of the field of bioacoustics is also notable. MAYNARD SMITH, JOHN (1920-2004) – Evolutionary biologist with a strong interest in animal communication. Maynard Smith was particularly instrumental in establishing a contact between traditional biological sciences (particularly evolutionary ones) and fields like semiotics and linguistics, by for instance working on the concepts of “signal” and “meaning” (Maynard Smith-Harper 2003), the concept of “information” (Maynard Smith 1982, 1999 and 2000), and several others. SEBEOK, THOMAS ALBERT (1920-2001) – Hungarian-American semiotician. He was chiefly responsible for introducing the study of animal communication within the linguistic and semiotic areas. He was the founder of zoosemiotics (in 1963 – see Chapter One), and defined its theoretical foundations, creating an important following among semioticians and non-semioticians. He was particularly important for

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setting the zoosemiotic agenda, for nearly all of its main topics (either by discussing them extensively or by pioneering them and opening doors for other scholars), but he also contributed to specific discoveries, such as the presence of symbolic signals in animal communication. EIBL-EIBESFELDT, IRENÄUS (1928) – Austrian ethologist, particularly known for founding Human ethology. Currently Professor at, and Head of, the Max-Planck-Institute for Behavioural Physiology, in Andechs (Germany). Pupil of Konrad Lorenz, Eibl-Eibesfeldt is one of the keyfigures in the development of the ethological disciplines, focusing on such topics as morality, sociality and communication (a.o. 1970 and 1971), and providing specific contributions to the study of human behavior. MARLER, PETER (1928-2014) – English ethologist, with a strong interest in animal communication. He directed the Animal Communication Laboratory at the University of California, Davis. Initially trained as a botanist, Marler developed a strong interest towards animal acoustic communication, giving a major contribution to the research on the social functions of birdsongs (Marler-Slabbekoorn 2004) and primates’ vocalizations (Marler 1977). MORRIS, DESMOND (1928) – British sociobiologist and popular science writer. His work has been influential for animal communication studies in many ways, particularly in the field of animal aesthetics and human-animal relationship. A pupil of Nikolas Tinbergen, in the late 1950’s Morris started a thorough investigation on the painting skills of chimpanzees, resulting in Morris 1963. The research led also to the curious extravaganza of an exhibition in incognito of chimpanzees’ paintings (see Chapter Seven), mixed with human ones, that was very favourably reviewed by the unaware critics. Morris, himself a painter, intended to make a point that – when assessed unbiasedly – animal art can be compared to human art. Later, his work focused on the animal components of human behaviour, resulting in his best-known (and most controversial) works, including The naked ape (1967). Morris is still active as a writer and TV personality. WENNER, ADRIAN (1928) – American biologist. Wenner is very well known, besides his scientific merits, for raising a long-lasting controversy around Karl von Frisch’s theory on the bee dance, by maintaining that bees actually locate foods exclusively through the olfactory channel. Officially the “loser” in the challenge (in 1973 von

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Frisch was awarded a Nobel Prize, and Wenner withdrew from bee research), Wenner recently had some compensation in the findings of a study conducted in 2005 by the Rothamsted Research (consisting in beetracking with radars), which confirmed that odor is also a guide for the bees. Besides this episode, Wenner has been active also in zoosemiotics, by contributing to one of the classics in the field (Sebeok-Ramsay 1969) with a detailed overview of animal communication studies. In Wenner 1969, he proposed a classification of communication typologies that, among other things, include an intra-individual, inter-individual and animate-environmental level of signaling, and specific elements/parameters such as the variability of the signals, the redundancy within/between signals, the ontogenetic development and role of learning, and others. ZAHAVI, AMOTZ (1928) – Israeli evolutionary biologist and Professor Emeritus at the Zoology Department of Tel Aviv University. Zahavi formulated the so-called “handicap principle” (a theory according to which reliable signals must be costly for the organism), an alternative to Darwinian theories in the explanation of evolution, which had a significant impact in the description of communication processes. The theory was originally formulated in the 1970’s, but is presented in its most complete fashion in Zahavi-Zahavi 1997. FOSSEY, DIAN (1932-1985) – American zoologist and primatologist. Together with Jane Goodall and Birutơ Galdikas (the three are also known as Leakey’s angels, for being all pupils of famed anthropologist Louis Leakey), the most important scholar in great apes’ communication in the wild. Inspired by Prof. Leakey, Fossey became fascinated with the rare mountain gorillas of the Rwandan wilderness, and decided to go studying closely their lives and habits. Fossey achieved the gorillas’ acceptance in their community by developing a means of communication with them (a.o. Fossey 1972 and 1974). Her study interest developed into a passionate commitment for the gorillas’ welfare and preservation, up to becoming a persona non grata to Rwandan natives, who extensively hunt gorillas for their skin (sold to rich tourists in form of various trinkets). After years of campaigns and fights with local institutions, Fossey was found murdered in her tent, in officially mysterious circumstances. Her legacy remains big, and her autobiography (Fossey 1983) is still the best-selling book about gorillas.

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GOODALL, JANE (1934) – English primatologist and ethologist, and founder of the Jane Goodall Institute. Pupil of anthropologist Louis Leakey (along with the other two Leakey’s angels, Dian Fossey and Birutơ Galdikas), for nearly 50 years Goodall studied chimpanzees’ sociality, cognition, tool making and communication in Gombe Stream National Park, Tanzania (a.o. Goodall 1964, 1968, 1971, 1986). Similarly to the other two “angels”, Goodall too accompanied her scientific research with a strong ethical commitment for primates’ conservation and awareness. In this activity, she also managed to achieve specifically scientific results, as for instance the taxonomic inclusion of chimpanzees, bonobos and gorillas in the Hominids’ family, and the promotion of the so-called “critical anthropomorphism” (she deliberately chose to give names to her chimpanzees, challenging the contemporary conventions that would regard this practice as anti-scientific). This and other methodological aspects of her work generated some criticism and accusations of biased research. Goodall’s environmental and humanitarian work was recognized in several ways by several institutions: most notably, she is a UN Messenger of Peace. KAPLAN, GISELA (1944) – Australian cognitive ethologist, with a specific interest in animal communication. Kaplan is Professor in Animal Behaviour at the University of New England. Her education (she has one PhD in Arts and one in Animal Behaviour) led her to animate her approach to animal studies in a very interdisciplinary manner. A prolific writer of over 300 research articles and 20 books (and multiple awardwinner for her research publications), Kaplan is a specialist in vertebrate cognition, primate and avian behaviour, particularly birdsong (benefiting also from her first career as a musician). Since the 1990’s, she has specialized in Australian songbirds’ vocal communication, cognition, and the functions of song and mimicry (focusing mostly on the Australian magpie Gymnorhina tibicen). Also worthy of mention is her long collaboration with Prof. Lesley Rogers, a very relevant figure to the study of animal cognition and brain development (a.o. Rogers-Kaplan 1998 and 2004, particularly the first – an innovative research on animal communication). BEKOFF, MARC (1945) – American cognitive ethologist. Currently Professor Emeritus of Ecology and Evolutionary Biology at the University of Colorado, Boulder, Bekoff is one of the most prominent figures in cognitive ethology. His range of interests includes the study of mental and emotional aspects of animal behavior and different ethical subjects. Some

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of his publications (particularly Bekoff 1975) constitute an important theoretical bridge between ethology and communication studies. Interdisciplinarity, and particularly collaboration with human sciences, is to Bekoff an important goal to achieve in the process of building a theoretical paradigm for cognitive ethology. GALDIKAS, BIRUTƠ (1946) – German-Lithuanian primatologist and ethologist. The youngest of the three Leakey’s angels, together with Dian Fossey and Jane Goodall, Galdikas is currently Full Professor at Simon Fraser University in Burnaby, British Columbia. As per anthropologist Louis Leakey’s plan (he wanted to perform field research on the three closest “relatives” of the human being, the chimpanzees, the gorillas and the orangutans), Galdikas focused her work on the orangutans of the Tanjung Puting Reserve, in Indonesian Borneo, expanding scientific knowledge on their behavior, habitat and diet (Galdikas 1995), with a special attention to communication, of course. Like the other two “angels”, she then devoted most of her attention to the advocacy of social and ethical causes related to her research subject, achieving great recognition and founding the Orangutan Foundation International, of which she is currently president. DE WAAL, FRANS (1948) – Dutch ethologist and primatologist. Professor of Primate Behavior in the Psychology Department at Emory University in Atlanta, Georgia. A major expert in primate behavior, De Waal plays a central role, in modern ethology, in the study of morality and sociality in great apes, offering an approach that he openly ascribes to the recent tradition of cognitive ethology, of which he is one of the leading exponents. It is particularly through studies such as De Waal 1996 that his research achieved world recognition. A central thesis in that work is that most animals, and particularly great apes are not only, and fully, “moral patients”, but also provide evidence of “moral agency”, becoming thus, like humans, insiders of the ethical discourse.

GLOSSARY

Here is a short list of terms and concepts that you may occasionally find challenging throughout the book. In the text, they are marked with asterisks at both sides (e.g. *adaptation*, *behaviourism*). I kept the definitions very short, not more than a couple of sentences, in order to keep this glossary as a quick tool for consultation. For a more lengthy discussion (or, of course, for the terms missing), you will have to rely on the book as such, on the ones recommended in the various bibliographical suggestions, or – and why not? – on your personal research efforts. ABSTRACTION (or GENERALIZATION) The ability of representing an event/entity in terms of general qualities/characteristics, independently from concrete realities, specific instances or actual objects. ACOUSTIC CHANNEL Channel related with the production, emission and reception of sounds (e.g. mating calls). ADAPTATION The result of a process of phylogenetical and ontogenetical adjustment of an organism to its environment. AESTHETIC FUNCTION OF COMMUNICATION The message focuses on itself, taking particular care to its form and appearance. ALARM CALL (or ALARM RESPONSE) A sign or signal emitted by an animal in order to warn others of a danger that s/he has spotted or perceived. ALLOGROOMING Pattern of social significance, where two specimens tend each other by removing dirt, parasites, or else from the fur, skin, feathers, etc.

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ALPHA (MALE OR FEMALE) In social animals, it is the individual in the community (male or female) with the highest rank, that is, who has gained (by physical strength, or other factors) a preferential access to items and activities, such as food, reproduction and else. ANALOGICAL SIGNAL A signal that varies in quantity, and therefore in meaning too. ANALOGY Two structures or patterns are considered analogous if they are similar in function or constitution, although resulting from different evolutionary paths. ANIMAL A living organism of the kingdom Animalia bearing the following characteristics: a) being multicellular; b) being heterotroph; c) developing from an embryo (that derives from gametes produced in specialized organs). In most cases, animals are motile (in one or more stages of their life cycle) and provided with one or more sensory apparatuses. ANTHROPOCENTRISM The philosophical or commonsensical attitude of considering the human animal at the center of and above other species (or other elements of reality). ANTHROPOMORPHISM The ascription of human characteristics to things not human. In animal studies, anthropomorphism is in general considered a scientific bias, although, in the last few decades, particularly within philosophical discussions, there has been a revision of the perception of the pros and cons of this attitude, and some positive aspects have been emphasized. APOSEMATIC An organism that is constructed in a way and may emit signals that indicate special capabilities for defense from predators. APPEASEMENT Behavioral pattern/s, generally of submissive type, displayed by an animal to another in conflict situations, in order to reduce the aggression shown

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by the latter to the former. Appeasement is normally used for avoiding the need to escape from the aggressor. BEHAVIORISM A theoretical approach (mostly in psychology, but in other fields as well), the main focus of which is behavior as such, with no particular consideration for elements that are not observable (such as cognition, emotion, etc.). BETA (MALE OR FEMALE) In social animals, it is the second-in-command, after the Alpha (see entry). BIOSPHERE The globality of the Earth’s environments where living organisms live and operate, interacting with the lithosphere, the hydrosphere, and the atmosphere. The birth of the biosphere, by means of biopoesis, is dated at some 3.5 billion years ago. CAMOUFLAGE A strategy (which can be of different types) aimed at avoiding an animal’s detection by mimicking the surrounding environment in some form. CHANNELS The sensory modes used to transmit a message (acoustic, visual, chemical, etc.). CHEMICAL CHANNEL Channel that includes the olfactory and the gustatory modes, and that adopts chemical substances to communicate (e.g. courtship through pheromones). CODE Rule, or set of rules, for converting messages into signals, which may or may not resemble the messages themselves in their form. COGNITION The totality of mental activities and information processing occurring within an animal, in a fashion partly or totally untied from instinct. The study of animal cognition focuses on such aspects as communication; attentive skills; classification and categorization; temporal and spatial

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cognition; production and use of tools; problem-solving; consciousness and computing skills. COGNITIVE FUNCTION OF COMMUNICATION The way communication affects an animal’s mental processes. COMMUNICATION The process of transmission and reception of a message. In communication, at least two animals (a sender and a receiver) take part in the process, and therefore the “sense” is exchanged, understood or misunderstood. In a particular case, proprioception, sender and receiver may be the same subject. CONATIVE FUNCTION OF COMMUNICATION The message focuses on its receiver, soliciting (or even manipulating) some kind of reaction. CONSCIOUSNESS Umbrella-term that may refer to a vast scale of mental states, from subjective experience to simple awareness. CULTURE The totality of information acquired and developed by a community and transmitted non-genetically from one generation to another. DEAR ENEMY Ethological phenomenon that describes an animal’s particular reaction to the invasion of its territory on the part of a neighbor. The aggressive response, in this case, is shorter and less intense, as compared to the case when a total stranger is invading the territory. DECEPTION A communicative action where a sender 1) breaks the association between signal and meaning established in one or more codes, 2) mentally represents the receiver’s mind (for instance, mapping his/her expectations), and 3) takes advantage of the broken association. DIALECT A variety of a sign system that is distinguished from other varieties of the same system by features of sign-object association, structure, social use,

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and by its use by a community of users that are set off from others geographically or socially. DISCONTINUITY The idea that, from an evolutionary point of view, human beings constitute a radical deviation from the rest of the animal kingdom, and therefore the differences between the human and the other animals are of a qualitative type. DISTRACTION DISPLAY A strategy (which can be of different types) used to attract the attention of an enemy (predator, rival…) away from a threatened object or subject. DIGITAL SIGNAL A signal that does not vary in quantity. It is either present or not. DOMESTICATION The process of taming an animal, normally by generations of breeding, to live in close association with another animal (usually, a human one), in such a way that the former becomes dependent on the latter’s care and, in most cases, unable to live in its original conditions again. DUPE In mimicry (see entry), it is the very subject (e.g. a predator) that is deceived by the mimic (see entry). ECHOLOCATION Form of proprioceptive communication by means of which animals like bats, cetaceans and some birds can receive information about the surrounding environment. ECTOTHERM (often referred to as COLD-BLOODED) An organism that does not manage to maintain a constant heat-level by means of internal bodily functions, and has to rely on external factors as well. EMOTION-FREE EXPERIMENTAL CONTEXT In interspecific communication experiments, an emotion-free context is an experimental setting arranged in order to avoid possible unwanted inputs from the experimenters that might compromise the credibility of the test. In an emotion-free context, researchers might wear masks, observe the

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experiment from another room, leave the subject alone in the laboratory, double-check data, etc. ENDOTHERM An organism that manages to maintain a fairly constant heat-level by means of internal bodily functions. ETHOGRAM The whole catalogue of behavioral patterns exhibited by an animal. EXHIBITION A set of displays which, through a process of ritualization, become simplified, stereotyped and exaggerated. Generally, but not exclusively, exhibitions are species-specific and occur in courting- fighting- or feelingrelated contexts. EXPRESSIVE FUNCTION OF COMMUNICATION The message focuses on its sender, revealing its emotional status and/or identity. EXTRINSIC SIGNAL Signal produced in the animal’s environment (e.g. a track). FOOD REQUEST A form of intraspecific (and occasionally interspecific) interaction, aimed at obtaining food. It usually occurs between parents and offspring, but also in other forms of relationship, such as between mates or between pets and human beings. FOUR QUESTIONS An ethological model for interpreting animal behavior. It includes Causation (the study of the stimuli eliciting the response, and the consequent modification of the latter); Development (the study of behavioral change in time units, in relation to earlier experiences); Evolution (the study of the animal behavior in relation with other species performing similar patterns); and Function (the study of the impact of behavioral pattern/s on the animal’s survival and reproduction chances). GAMMA (MALE OR FEMALE) In social animals, it is the third-in-command, after the Alpha and the Beta (see entries).

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GESTALT (or GESTALT THEORY or GESTALTISM or GESTALT PSYCHOLOGY) A theoretical approach (mostly present in psychology, but in other fields as well) devoted to the understanding of the way mental processes form meaningful perceptions in the apparently chaotic environment. GRADUALISM The idea of an evolutionary adaptive continuum among animal species, with more and less specialized characteristics/patterns. GROOMING The ritual action of removing dirt, parasites, or specks of other matter from the fur, skin, feathers, etc. It can be performed by one animal on him/herself or on another. HANDICAP PRINCIPLE A theoretical approach suggesting that evolution favors extravagant/demanding/uncomfortable traits as an indication of strength, status or health. ECHOLOCATION The biological sonar used by animals like dolphins, bats, shrew mice and others, to detect objects by emitting sounds that reflect off the object and return to the animal’s ears or other sensory receptors. ECTOTHERM An organism that manages to maintain a constant heat-level by means of internal bodily functions HETEROTROPH A living organism that is not able to chemically produce its own food, and must therefore consume other organisms. Animals are heterotrophic, while other organisms like plants are autotrophic. HOMOLOGY Two structures or patterns are considered homologous if they share a common origin and basic structure, even if they may evolve into substantially different functions.

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ICON A signal that physically resembles its meaning (like a mimic with its model, a mock-fight with a real fight, etc.). IMITATION A paricular case of mimicry consisting of taking from a model (i.e. another individual or species) one or more patterns not present in the mimic (i.e. the imitating individual/spiecies). IMPRINTING The capacity to learn specific types of information at certain critical periods in development. INDEX A signal that is a natural consequence of its meaning (like a symptom for an illness, a bright plumage for good health, etc.). INFANT SCHEME A theoretical formulation by Konrad Lorenz that suggests that there are common and consistent features across the infants of different species. INFANT SIGNAL A type of self-promotion signal that informs of the sender’s young age. INSTINCT A genetically acquired force that drives animals to react to a stimulus in certain fixed ways. INSTRUMENTAL PLAY A form of playing in which some tool is needed. INTENTIONALITY The conscious planning or performance of a given action, aimed to a goal. INTERSPECIFIC COMMUNICATION EXPERIMENTS Expression that refers to a range of experimental programs (conducted usually by biologists, psychologists or linguists), aimed at testing the possibility of teaching human language to other animal species, particularly great apes.

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INTERSPECIFIC COMMUNICATION Any form of communication occurring between two or more different animal species. INTRASPECIFIC COMMUNICATION Any form of communication occurring within the same animal species. INTRINSIC SIGNAL Signal produced on the animal’s body (e.g. a facial expression). K-SELECTION The reproductive strategy by which an organism produces few offspring, but (via parental care, mostly) increses their probability of surviving to adulthood. KINESICS The study of body movements, gestures, facial expressions and otherwise, as a means of communication. KINETIC PLAY A form of playing that is solely based on bodily movement, with no tool needed. LANGUAGE To some scholars, “language” refers to a particularly complex and articulated communication system (such as that of human beings, honeybees or cetaceans). To some others, the expression refers only to the human species-specific communication system. LEARNING The process of acquiring and retaining information through ontogenesis, i.e. the consequence of interaction between an animal and its environment, rather than through phylogenesis. MAGNETIC CHANNEL Channel related with the perception of magnetic fields (e.g. for locating an object). MAGNETOCEPTION The capacity to use magnetic fields to gather information about the environment.

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MENTAL MAP (or COGNITIVE MAP) The capacity of orienting oneself in space and heading towards a certain destination, by using signs available in the environment. To possess a cognitive map means to be able to solve several spatial/temporal problems: finding the shortest way for a familiar destination, finding an alternative way when the usual one is not available, going back to familiar places from unknown ones, etc. MENTAL REPRESENTATION A mental representation is an inner sign of an external object (which functioned as its stimulus), and that can be reactivated also in absence of the original stimulus. METALINGUISTIC FUNCTION OF COMMUNICATION The message focuses on the code shared by senders and receivers, so that a signal refers to another signal. MIMIC In mimicry (see entry), it is the very subject (e.g. prey) that deceives the dupe (see entry) by imitating a model (see entry). MIMICRY Phenomenon that includes several forms of similarity (appearance, behavior…) of one species to another (or, more rarely, within the same species). MIND A network of biological and physiological activities and relations (including brain functioning, sensorial perception, nervous system, etc.) that allow an organism to acquire, store and elaborate the information it is, was, or might be exposed to, and produce an adequate response, after a consideration of the variables involved at any stage of this process. MIRROR TEST A behavioural technique consisting of determining whether a non-human animal possesses the ability of self-recognition in front of a mirror. MODEL (in reference to MIMICRY) In mimicry (see entry), it is the very subject that is imitated by the mimic (see entry).

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MODELLING A process through which a phenomenon/entity is executed and/or multiplied on the basis of a virtual or real general model. MULTIMODALITY Instances of communication when different patterns cooperate (or compete) to display one or more signal or text. MUTUALISM A form of cooperation between two organisms of different species, in which each individual derives a benefit. NAME A signal used to identify and recognize a specific subject or object. NIDIFUGOUS An organism that leaves its nest shortly after hatching or birth. ONTOGENESIS (or ONTOGENY or MORPHOGENESIS) The origination and development of an individual organism. PHATIC FUNCTION OF COMMUNICATION The message focuses on the contact between senders and receivers, in order to establish or even reinforce it. PHENOMENOLOGY A theoretical approach (particularly in philosophy, but in other fields as well) focused on the study of experience and consciousness. PHEROMONE Any chemical substance released by an animal that works as a signal (generally a self-promotion one). PHYLOGENESIS (or PHYLOGENETICS) The origination, development of, and relations among, groups of organisms. PLAY (or PLAYING) A set of behavioural patterns that non-seriously imitate otherwise serious patterns and situations. The two basic functions of this behavior seem to be amusement and the learning of rules of social behaviour.

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PLAY-BOW A characteristic “head down, bottom up” position assumed by canidae (wolves, dogs, etc.), used to announce their intention to play. PLAY INHIBITION In playful situations, the action by which the stronger/older players provide themselves with a handicap (see HANDICAP PRINCIPLE), in order to establish a condition of equality with the weaker/younger ones. PRIMATOLOGY The field of biology that studies the physiology and the behavior of primates (apes and monkeys). PROPRIOCEPTION A message/signal is proprioceptive when the sender and receiver are the same subject, that is, when the animal sends messages for its own use and benefit. A typical case is echolocation. PROXEMICS The study of the space management and interpersonal distance in humans and other animals. R-SELECTION The reproductive strategy by which an organism produces many offspring, each of which has a relatively low probability of surviving to adulthood. REFERENTIAL FUNCTION OF COMMUNICATION The message focuses on the context, describing one or more parts of it. RELEASER An external stimulus to which an animal reacts in a fixed, specific manner. REPRESENTATION The process through which a message is presented or depicted in some way. The meaning, therefore, is produced first and foremost by the sender of the message. RITUALIZATION An evolutionary process that progressively modifies and turns noncommunicative patterns into actual signals.

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SELF-AWARENESS The explicit acknowledgment of one’s own existence as individual, that is, separated from other subjects, with an individual mental activity (including, possibly, the acknowledgment that other subjects possess selfawareness too). SELF-CONSCIOUSNESS The reflective mental activity thanks to which a subject becomes aware of him/herself and of his/her identity, and from which a process of investigation of the self may be activated. SELF-PROMOTION SIGNAL A type of signal that informs about the sender (identity, sex, etc.) SIGN Something that stands for something else. In this book, a sign is also a more complex and articulated form of signal. SIGNIFICATION The process by which signs and meanings are produced. Signification occurs independently from a specifically identifiable source: animals make sense out of each other and out of their environment whether or not this sense is explicitly codified and conveyed. The act of producing meaning, therefore, is first and foremost performed by the receiver of the message. SIGNAL Any pattern that transmits information. SIGNAL INTERCEPTION A form of “intrusion” where the communication between sender and receiver is decoded by a third subject (predator, parasite, competitor, or else) who takes advantage from it. SOCIAL FACILITATION The process by which a certain behavioral pattern, performed by a given animal, increases the chances that another animal will perform the same pattern (or increase its intensity, if it is already being performed).

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SOCIALITY The condition or quality of being social (tendency to intersubjective interaction and association, to form groups, to establish social roles, to mentally represent other entities, etc.). SOCIAL STATUS SIGNAL A type of self-promotion signal that gives information about the sender’s role in its social group. SOUNDSCAPE The totality and the combinations of sounds within a given area. It is the acoustic equivalent of “landscape”. SPECIES The last taxonomic rank before the “individual” (with the exception of those cases where a “subspecies” is identified). SPECIESISM Form of discrimination based on species difference. In the human-animal relationship, speciesism is the equivalent of racism in the relationship between two different ethnic groups. SPECIES-SPECIFIC Any feature/pattern/event that occurs typically and exclusively in one species, up to the point of typifying it, in a certain respect. STOTTING (or PRONKING or PRONGING) A behavioral pattern, typical of quadrupeds like gazelles, consisting into repeated jumps with all four feet lifting simultaneously. SYMBIOSIS Umbrella-term that describes various phenomena, like mutualism, commensalism, amensalism, or parasitism, which share the common denominator of two dissimilar organisms sharing a number of complementary and reciprocally useful interests. SYMBOL A signal that is related to its meaning only in an arbitrary and conventional way (like a monkey’s alarm call, the dance of the honey bee, etc.).

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SYNTAX A string of signals, ordered in a certain manner, in order to convey a certain meaning. It may be composed of signals that individually have no specific meaning (phonological syntax), or signals that individually mean something else than their sum (lexical syntax). TACTILE CHANNEL Channel that includes the thermic, the electric and the tactile modes, and that require physical proximity to allow communication (e.g. allogrooming). TOKEN An object that is taken as an example within a certain category (called “Type”): “red” is a token for “colors”, “Helsinki” is a token for “Finnish cities”, etc. TOOL-USE (or TOOL-MAKING) The capacity of using material elements from the environment, in order to reach an otherwise unattainable goal. TRAINING The procedure of teaching one or more animals specific responses to specific stimuli, generally by using the methodologies of behaviorism, based on reinforcement and/or punishment. TROPHALLAXIS The practice of exchanging liquid food by means of regurgitation within the members of a community of social insects, or between these and the so-called guests. It serves as an important communicative function for enhancing a group’s cohesion. TYPE A general category of objects (called “tokens”) related to each other through one or more characteristics. The type “colors” includes the tokens “red”, “blue”, “yellow”, etc.; the type “Finnish cities” includes the tokens “Helsinki”, “Tampere”, “Turku”, etc. UMWELT In Jakob von Uexküll’s formulation, it means a “subjective universe”, that is, an environment that is not merely physical, but that includes the

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information that is perceivable and meaningful in its entirety only from the perspective of the particular organism that inhabits it. VISUAL CHANNEL Channel related with the production, emission and reception of visual signals (e.g. body displays) ZOOMORPHISM The attitude of considering human beings as animals like all others, giving more emphasis to the characteristics they share with other species, than to the species-specific ones.

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