Do exaggerated chelicerae function as weapons or genitalia in a long- jawed spider? Functional allometric analysis yields an answer

Abstract

From the elongated neck of the giraffe to the elaborate train of the peacock, extreme traits can result from natural or sexual selection (or both). The extreme chelicerae of the long- jawed spiders (Tetragnatha) present a puzzle: do these exaggerated chelicerae function as weapons or genitalia? Bristowe first proposed that Tetragnatha chelicerae function as a holdfast because these spiders embrace chelicerae during mating. This hypothesis has remained untested until now. Here, we use functional allometry to examine how extreme chelicerae develop and perform in the long- jawed spider Tetragnatha elongata. Similar to other Tetragnatha species, chelicerae were longer in adult males than in adult females. Overall, we confirm Bristowe’s hypothesis: elongation only occurred in the adult stage. However, we propose that chelicerae function as more than a holdfast in T. elongata. Male chelicerae exhibited positive allometry, which suggests scaling as weapons rather than genitalia. However, fieldwork revealed that the operational sex ratio is female- biased and both adult male- male competition and sexual cannibalism were rarely observed. Consequently, we propose that the positive allometry of male chelicerae may result from sexual selection to mechanically mesh with larger and more fecund females. Evidence for mechanical mesh includes multiple traits ranging from apophyses and grooves to guide teeth on the basal portion of the chelicerae. In contrast, we propose that chelicerae of females are analogous to the female peacock’s tail: shortened by natural selection limiting the exaggeration of sexually selected traits. Indeed, females had increased foraging efficiency compared to males and exhibited negative cheliceral allometry. We discuss the implications for the evolution of elongated chelicerae in Tetragnatha.In the long- jawed orbweaver Tetragnatha elongata, chelicerae grow from a typical orbweaver length in penultimate spiders (bottom) to the extreme form only seen in the adult (top) spider. Sexual selection has shaped the chelicerae in both sexes for an optimal - mechanical mesh- because these spiders interlock chelicerae during mating.