Clarifying and expanding the social complexity hypothesis for communicative complexity

Variation in communicative complexity has been conceptually and empirically attributed to social complexity, with animals living in more complex social environments exhibiting more signals and/or more complex signals than animals living in simpler social environments. As compelling as studies highlighting a link between social and communicative variables are, this hypothesis remains challenged by operational problems, contrasting results, and several weaknesses of the associated tests. Specifically, how to best operationalize social and communicative complexity remains debated; alternative hypotheses, such as the role of a species’ ecology, morphology, or phylogenetic history, have been neglected; and the actual ways in which variation in signaling is directly affected by social factors remain largely unexplored. In this review, we address these three issues and propose an extension of the “social complexity hypothesis for communicative complexity” that resolves and acknowledges the above factors. We specifically argue for integrating the inherently multimodal nature of communication into a more comprehensive framework and for acknowledging the social context of derived signals and the potential of audience effects. By doing so, we believe it will be possible to generate more accurate predictions about which specific social parameters may be responsible for selection on new or more complex signals, as well as to uncover potential adaptive functions that are not necessarily apparent from studying communication in only one modality. Animals exhibit an astonishing diversity of communicative systems, with important variation in both the nature and the number of signals they produce. The roles of phylogenetic history, genetic drift, environmental factors, and sexual selection in shaping interspecific variation in communicative systems have long been acknowledged, whereas social complexity has only more recently emerged as a potential fundamental factor determining signal diversification. While a number of comparative studies support the key prediction of this hypothesis, i.e., that individuals living in more complex social environments exhibit more signals and/or more complex signals, we discuss several gaps in the current state of the art concerning this hypothesis and point out what we believe are neglected perspectives. By emphasizing the importance of the multimodal nature of communicative systems and the social context in which signals are exchanged, we hope to stimulate the development of new tests and specific questions based on this expanded framework.