Vocal morphology of the Physalaemus pustulosus species group (Leptodactylidae): morphological response to sexual selection for complex calls

Abstract

Most male frogs in the genus Physalaemus produce a whine-like advertisement call. Male P. pustulosus, however, add chucks to the call. This enhances the attractiveness of the call to females, and has evolved under the influence of sexual selection despite the increased predation risk from the frog-eating bat (Trachops cirrhosus). This complex call is unusual, if not unique, among anurans because the two call components overlap in time. Here we investigate the morphological changes responsible for the production of complex calls. The Physalaemus purtulosus species group consists of four species. Physalaemus pustulosus and P. petersi are sister species, and recently it has been shown that P. petersi produces chucks. Physalaemus coloradorum and P. purtulatus are sister species and neither is known to produce chucks. Two laryngeal characters vary within the species group. Physalaemus pustulosus has a large fibrous mass (FMI), whose vibration is responsible for production of the chuck. This mass is much smaller in the other three species. In P. pustulosus and P. petersi the FMI is anchored dorsally, deep within the bronchial process, the attachment is more extensive in P. pustulosus. Neither P. pustulatus nor P. coloradorum have such a dorsal attachment associated with their FMI. This character is responsible for allowing the FMI to vibrate independently of the vocal cords, that is, for the production of the complex call. Thus the morphological changes responsible for the evolution of this unusual behavioural innovation, the complex call, are gradual, and almost trival, in nature. This study also shows that the primitive condition of the larynx of the P. pustulosus and P. petersi ancestor, was predisposed to the production of complex calls. We also document ontogenetic and sexually dimorphic patterns in larynx structure.