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To explore the phylogenetic position of the new Chinese luminescent click beetle, we newly generated 14 mitochondrial genes (13 protein-coding genes and 16S) and two nuclear rRNA genes (18S and 28S) for this species and merge them with the available four-gene Elateridae dataset (18S, 28S, 16S, cox1) by Kundrata et al. (2016, 2018) and Douglas et al. (2018) plus all publicly available Elateridae mitochondrial genomes downloaded from the GenBank (altogether 179 terminals representing 13 subfamilies). Four species of Phengodidae were used as outgroups following Kundrata et al. (2016, 2018) (Suppl. material S1, File 1).
In Lampyridae, bioluminescence was first gained by larvae as an aposematic warning display, and subsequently gained by adults and co-opted as a sexual signal; the overall trend of courtship is the use of pheromones in ancestral species, then pheromones used in conjunction with photic signals, then the sole use of photic signal (Branham and Wenzel 2001, 2003; Ohba 2004; Stanger-Hall et al. 2018). Males of S. schimmeli have a large luminous organ on abdomen plus very long antennae (longer than in females, hypothesized to be for pheromone detection) and large eyes (larger than in females, and hypothesized to be for bioluminescence detection), which suggest the use of multiple communication channels for mate attraction (Stanger-Hall et al. 2018, and references therein). It is hypothesized that in elaterids, bioluminescence of the abdominal lantern is an optical signal for the intraspecific sexual communication, while the signals from the prothoracic lanterns (if present) serve to warn predators and may also provide illumination in flight (Lall et al. 2000, 2010). Additionally, antennae are usually relatively longer in species which rely greatly on pheromones as sexual signals (Stanger-Hall et al. 2018). Therefore, the external morphology of S. schimmeli suggests the combined sexual communication of both light signals and pheromones. The relatively larger sexual signal sensors of S. schimmeli males in comparison to conspecific females are in agreement with situation in many other insect groups, in which males are the more actively searching sex, with more sensitive sensors (Steinbrecht 1987, Stanger-Hall et al. 2018).
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