Animal communication is inherently spatial. Both signal transmission and signal reception have spatial biases-involving direction, distance, and position-that interact to determine signaling efficacy. Signals, be they visual, acoustic, or chemical, are often highly directional. Likewise, receivers may only be able to detect signals if they arrive from certain directions. Alignment between these directional biases is therefore critical for effective communication, with even slight misalignments disrupting perception of signaled information. In addition, signals often degrade as they travel from signaler to receiver, and environmental conditions that impact transmission can vary over even small spatiotemporal scales. Thus, how animals position themselves during communication is likely to be under strong selection. Despite this, our knowledge regarding the spatial arrangements of signalers and receivers during communication remains surprisingly coarse for most systems. We know even less about how signaler and receiver behaviors contribute to effective signaling alignment over time, or how signals themselves may have evolved to influence and/or respond to these aspects of animal communication. Here, we first describe why researchers should adopt a more explicitly geometric view of animal signaling, including issues of location, direction, and distance. We then describe how environmental and social influences introduce further complexities to the geometry of signaling. We discuss how multimodality offers new challenges and opportunities for signalers and receivers. We conclude with recommendations and future directions made visible by attention to the geometry of signaling.
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