Multimodal signals: ultraviolet reflectance and chemical cues in stomatopod agonistic encounters

Complex signals are commonly used during intraspecific contests over resources to assess an opponent's fighting ability and/or aggressive state. Stomatopod crustaceans may use complex signals when competing aggressively for refuges. Before physical attacks, stomatopods assess their opponents using chemical cues and perform threat displays showing a coloured patch, the meral spot. In some species, this spot reflects UV. However, despite their complex visual system with up to 20 photoreceptor classes, we do not know if stomatopods use chromatic or achromatic signals in contests. In a field study, we found that Neogonodactylus oerstedii meral spot luminance varies with sex, habitat and, more weakly, body length. Next, we conducted an experimental manipulation which demonstrated that both chemical cues and chromatic signals are used during contests. In the absence of chemical cues, stomatopods approached an occupied refuge more quickly and performed offensive behaviours at a lower rate. When UV reflectance was absent, stomatopods performed offensive behaviours more frequently and contest duration trended towards shorter fights. These results provide new evidence that UV reflectance and/or visible spectrum luminance is used to amplify threat displays. Our results are the first to demonstrate that chemical and chromatic cues comprise a multimodal signal in stomatopod contests.