The clitoris is a part of the genitalia of female amniotes that typically functions to stimulate sensory arousal. It usually consists of a small organ that is dimorphic and homologous to the penis. The developing amniote embryo forms a genital tubule, then sex hormones initiate a developmental cascade to form either a penis or clitoris. In squamates (lizards and snakes) the genital tubule develops into a paired hemiphallus structure called the “hemiclitores” in the female and the “hemipenes” in the male. The complex evolution of squamate hemipenes has been extensively researched since early discoveries in the 1800’s, and this has uncovered huge diversity in hemipenis size, shape, and ornamentation (e.g., protrusions of spines, hooks, chalices, cups). In contrast, the squamate hemiclitoris has been conspicuously under investigated, and the studies that describe this anatomy are fraught with inconsistences. This paper aims to clarify the current state of knowledge of the squamate hemiclitoris, providing a foundation for further research on its morphology and functional role. We show that while several studies have described the gross anatomy of hemiclitores in lizards, comparative information is entirely lacking for snakes. Several papers cite earlier authors as having reported discoveries of the snake hemiclitores in vipers and colubrid snakes. However, our examination of this reveals only erroneous reports of hemiclitores in snakes and shows that these stem from misinterpretations of the true anatomy or species involved. An especially problematic source of confusion is the presence of intersex individuals in some snake populations; these form reproductively functional ovaries and a single hemipenis, with the latter sometimes mistaken for a hemiclitoris (the intersex hemipenis is usually smaller and less spinous than the male hemipenis). Further research is recommended to identify the defining anatomical features of the squamate hemiclitores. Such studies will form a vital basis of future comparative analyses of variation in female genitalia in squamates and other amniotes.
Female genitalia are conspicuously overlooked in comparison to their male counterparts, limiting our understanding of sexual reproduction across vertebrate lineages. This study is the first complete description of the clitoris (hemiclitores) in female snakes. We describe morphological variation in size and shape ( n = 9 species, 4 families) that is potentially comparable to the male intromittent organs in squamate reptiles (hemipenes). Dissection, diffusible iodine contrast-enhanced micro-CT and histology revealed that, unlike lizard hemiclitores, the snake hemiclitores are non-eversible structures. The two individual hemiclitores are separated medially by connective tissue, forming a triangular structure that extends posteriorly. Histology of the hemiclitores in Australian death adders ( Acanthophis antarcticus ) showed erectile tissue and strands/bundles of nerves, but no spines (as is found in male hemipenes). These histological features suggest the snake hemiclitores have functional significance in mating and definitively show that the hemiclitores are not underdeveloped hemipenes or scent glands, which have been erroneously indicated in other studies. Our discovery supports that hemiclitores have been retained across squamates and provides preliminary evidence of differences in this structure among snake species, which can be used to further understand systematics, reproductive evolution and ecology across squamate reptiles.
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