By-product information can stabilize the reliability of communication.

Although communication underpins many biological processes, its function and basic definition remain contentious. In particular, researchers have debated whether information should be an integral part of a definition of communication and how it remains reliable. So far the handicap principle, assuming signal costs to stabilize reliable communication, has been the predominant paradigm in the study of animal communication. The role of by-product information produced by mechanisms other than the communicative interaction has been neglected in the debate on signal reliability. We argue that by-product information is common and that it provides the starting point for ritualization as the process of the evolution of communication. Second, by-product information remains unchanged during ritualization and enforces reliable communication by restricting the options for manipulation and cheating. Third, this perspective changes the focus of research on communication from studying signal costs to studying the costs of cheating. It can thus explain the reliability of signalling in many communication systems that do not rely on handicaps. We emphasize that communication can often be informative but that the evolution of communication does not cause the evolution of information because by-product information often predates and stimulates the evolution of communication. Communication is thus a consequence but not a cause of reliability. Communication is the interplay of inadvertent, informative traits and evolved traits that increase the stimulation and perception of perceivers. Viewing communication as a complex of inadvertent and derived traits facilitates understanding of the selective pressures shaping communication and those shaping information and its reliability. This viewpoint further contributes to resolving the current controversy on the role of information in communication.