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Many scarab beetles have sexually dimorphic exaggerated horns that are an evolutionary novelty. Since the shape, number, size, and location of horns are highly diverged within Scarabaeidae, beetle horns are an attractive model for studying the evolution of sexually dimorphic and novel traits. In beetles including the Japanese rhinoceros beetle  Trypoxylus dichotomus , the sex differentiation gene  doublesex  ( dsx ) plays a crucial role in sexually dimorphic horn formation during larval-pupal development. However, knowledge of when and how  dsx  drives the gene regulatory network (GRN) for horn formation to form sexually dimorphic horns during development remains elusive. To address this issue, we identified a  Trypoxylus -ortholog of the sex determination gene,  transformer  ( tra ), that regulates sex-specific splicing of the  dsx  pre-mRNA, and whose loss of function results in sex transformation. By knocking down  tra  function at multiple developmental timepoints during larval-pupal development, we estimated the onset when the sex-specific GRN for horn formation is driven. In addition, we also revealed that  dsx  regulates different aspects of morphogenetic activities during the prepupal and pupal developmental stages to form appropriate morphologies of pupal head and thoracic horn primordia as well as those of adult horns. Based on these findings, we discuss the evolutionary developmental background of sexually dimorphic trait growth in horned beetles.

plos genetics

Precise staging of beetle horn formation in Trypoxylus dichotomus reveals the pleiotropic roles of doublesex depending on the spatiotemporal developmental contexts