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10 Where Is the Information in Animal Communication? Julia Fischer Abstract Communication is a central topic in animal behavior studies and yet the dispute over what constitutes communication is far from settled. Over the last few years, a number of papers have revisited the core issues in this field and have advanced divergent views regarding the explanatory power of the concept of “information.” After a review of this debate, an integrative framework is proposed that conceives communication in its elementary form as an interaction between two individuals (sender and receiver) and involves the use of signals by the sender as well as the processing of and responses to those signals by the receiver. Signals are structures or behaviors that have evolved because their effects on other individuals benefit the sender on average, irrespective of whether or not the behavior of the receiver has evolved to be affected. Receivers have been selected to make inferences about the environment, including the behavior of conspeci fics. Signals may in this sense be informative to the receiver, because they can be used to assess the state, identity, or subsequent behavior of others. Thus, signals contain “potential information,” which turns into “perceiver information” once processed by a receiver. The value and amount of this information can only be defined from the receiver ’s perspective. This framework thus defends the concept of information but rejects the notion that senders have generally been selected to “provide” that information, and that information is “encoded” within a signal. The notion that animals process information also creates a bridge from studies of communication to those assessing the cognitive underpinnings of communicative behavior. Introduction Explaining the evolution of communication is a major challenge, and despite many years of research, a number of conceptual issues remain unresolved. This has led to both confusion and sometimes unproductive friction. Some of the disputes appear to stem from diverging initial points in the analyses: some focus on signal evolution, others on responses to signals. Apparently, these different foci have profound implications for the conceptualization of communication . While each approach has its merits as well as shortcomings, the real 152 J. Fischer challenge is to incorporate insights from both to develop a full understanding of the complexity of communication. In this chapter, I examine accounts that focus on explaining signal evolution and contrast them with accounts that have been adopted to explain the processing of signals. I review the recent critical discussion of the term “information” in animal communication and argue that this concept should be retained. I believe that the concept of information is indispensable for understanding not only the cognitive mechanisms which underpin the responses to signals, but also the selective pressures operating on receivers. What Is Communication? Definitions of communication commonly involve the use of signals and incorporate at least a signaler and a receiver. Notably, in one of the most influential contributions, Maynard Smith and Harper did not even bother to provide a definition of communication in “Animal Signals,” but instead restricted the discussion to the evolution of signals (Maynard Smith and Harper 2003:388). Indeed, the term “communication” is not even indexed in the book. Because a number of researchers who seek to explain the evolution of communication refer to that text, they in turn put more emphasis on the sender’s side than on the receiver’s (Stegmann 2005; Scott-Phillips 2009). Analyses which follow Maynard Smith and Harper’s emphasis on the sender have been labeled “adaptationist ,” because they have a strong focus on identifying the selective pressures that shape signal design and affect the costs associated with signaling. A broad definition was put forward by Todt (1986), who characterized communication as “interactions with signals.” This definition stresses the notion of communication as an integral part of social behavior. Moreover, it facilitates the application of insights from pragmatics, a field in linguistics that has to date only played a minor role in animal communication studies (see Wheeler et al., this volume). One complicating issue in any analysis of communication is that most communicative interactions do not only involve two individuals but rather several subjects, hence the concept of “communication networks” (McGregor and Peake 2000). This is particularly important when different receivers have divergent interests and exert different selective pressures on signalers (Skyrms 2010). For the sake of simplicity, however, this aspect will not be further elaborated here. What Is a Signal? An “adaptationist” account seeks to explain the evolution of signaling behavior...

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