Intraspecific communication in frogs plays an important role in the recognition of conspecifics in general and of potential rivals or mates in particular and therefore with relevant consequences for pre-zygotic reproductive isolation. We investigate intraspecific communication in Hylodes japi, an endemic Brazilian torrent frog with territorial males and an elaborate courtship behavior. We describe its repertoire of acoustic signals as well as one of the most complex repertoires of visual displays known in anurans, including five new visual displays. Previously unknown in frogs, we also describe a bimodal inter-sexual communication system where the female stimulates the male to emit a courtship call. As another novelty for frogs, we show that in addition to choosing which limb to signal with, males choose which of their two vocal sacs will be used for visual signaling. We explain how and why this is accomplished. Control of inflation also provides additional evidence that vocal sac movement and color must be important for visual communication, even while producing sound. Through the current knowledge on visual signaling in Neotropical torrent frogs (i.e. hylodids), we discuss and highlight the behavioral diversity in the family Hylodidae. Our findings indicate that communication in species of Hylodes is undoubtedly more sophisticated than we expected and that visual communication in anurans is more widespread than previously thought. This is especially true in tropical regions, most likely due to the higher number of species and phylogenetic groups and/or to ecological factors, such as higher microhabitat diversity.
Since it is clear that studies involving species of Hylodes are critical to better understanding frog communication, we investigate how the Brazilian torrent frog Hylodes japi communicates intraspecifically. Hylodes japi was recently described as endemic to the montane Atlantic forest of Serra do Japi, in southeastern Brazil [26]. This small frog is mainly diurnal, however it may also exhibit crepuscular and nocturnal reproductive activity [26, 27]. The species is rheophilic, has territorial males and an elaborate courtship behavior, as do other members of the family Hylodidae. The reproductive biology of H. japi is associated with fast-flowing montane streams where males construct underwater chambers for egg deposition; the tadpoles are exotrophic [26]. Our goals were to: (1) characterize visual displays and acoustic signals performed by males and females of H. japi, and to identify their respective roles and associated behavioral contexts; (2) investigate communication during their elaborate courtship ritual, including analyzing potential multimodal compositions and their roles; and (3) characterize the ways by which a sender male controls signal emission according to the position of the receiver male. We discuss the behavioral diversity in the family Hylodidae in light of current knowledge on visual signaling in Neotropical torrent frogs.
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Among hylodids, the currently known repertoire of visual displays is most complex in H. japi (Table 3). In fact, Hylodes japi has one of the most diverse repertoires of visual displays known within the order Anura. The five new visual displays that we described and categorized here correspond to 20.8% of the visual displays recognized for members of the family Hylodidae and around 13.5% for anurans. We trust that our results on visual communication are not an exception among hylodids (and anurans in general), but a consequence of the time invested to understand the behaviors. Among hylodids, the most studied species have more diverse repertoires, such as C. schmidti, H. phyllodes, and H. japi ([21, 34, 37]; present study; see Table 3). For Hylodes species, some behaviors, such as arm lifting and arm waving, are only distinguishable via video analysis. Moreover, other visual displays are performed only during specific situations, making them difficult to observe (because they are rarely executed). Head snaking, for example, was recorded only twice among all studies on hylodids; during courtship, once in H. phyllodes [34] and in the present study. The accepted male is the only individual that performs head snaking and only during courtship. From the set of information presented here, it is plausible to expect that the complexity observed in the visual communication of H. japi is similarly widespread within the family Hylodidae, or at least among Hylodes species. Complexity of the visual communication system may be a pattern for the Brazilian torrent frogs (Hylodes species), and most likely as a phylogenetic trait of the genus. Neotropical torrent frogs (i.e. hylodids) still deserve attention, since new studies on their communication have potential to help clarify behavioral patterns and multimodal compositions, and even uncover other new behaviors. Behavioral patterns tend to be similar within families and within genera [2].
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