A comparative study of the male genitalia of the Cacodemoniini (Pseudoscorpiones: Withiidae)

Propositions of homology are fundamental in systematics, since they provide the basis for supporting clades. Consequently, such phylogenetic propositions rely on correct character and character state definitions. Although male genital morphology is a key source of information for understanding the phylogeny and classification of the Withiidae (Pseudoscorpiones), they have only been subjected to examination in six of the 170 species of the family. The suprageneric classification of the Withiidae is unstable, as subfamilies and tribes are not well supported by morphological characters, and only the unranked group of genera Cacodemoniini is currently accepted. The aim of the present work is to characterize the male genital armature of the Cacodemoniini and propose homology statements for these structures based upon their morphological correspondence. Through direct examination and literature review of 12 of the 13 genera of the Cacodemoniini, we provide the first structural correspondence statements and descriptions of variation for the dorsal apodemes, the ejaculatory canal, the lateral apodemes, and the lateral rods; we also conclude that unlike other pseudoscorpions, the Cacodemoniini have paired, independent lateral rods and a long ejaculatory canal formed not by the dorsal apodemes exclusively, but by a fusion of the dorsal and the lateral apodemes. The proposed interpretations lay the groundwork for phylogenetic testing of homologies and may allow a better understanding of the formation of the spermatophore, given that it is molded by the genital armature.