Satan’s Skeleton Revealed

Satan eurystomus Hubbs & Bailey 1947, the widemouth blindcat, is endemic to the deep Edwards Aquifer below San Antonio, TX. Monotypic Satan is one of four subterranean ictalurids, Trogloglanis pattersoni, Prietella pheatophila and P. lundbergi, that all exhibit common features of stygomorphs: loss of eyes and pigmentation, hypertrophy of some chemo and mechanosensory systems, small size, and variously reduced musculoskeletal system. Each species is distinctive in its own ways, and hypotheses about their phylogenetic positions range from separate ancestries of each scattered among the lineages of epigean ictalurids to exclusive monophyly of a strictly subterranean clade. Specimens of Satan are rare, thus we used highresolution CT scans to develop the first detailed, richly illustrated descriptive and comparative study of its skeleton. Satan exhibits typical and singular reductive features plus complex structures, e.g. 3 novel symphyses closing the posterior cranial fontanel; an unusually deep temporal fossa; and an ornately shaped dorsal fin locking spinelet. Satan shares 15 synapomorphies with other ictalurid troglobites: the stygomorphisms plus bone and joint reductions. Satan shares 11 synapomorphies with Pylodictis, including increased numbers of cephalic sensory pores and paired fin rays, and several features associated with predatory suction feeding: wide gape, depressed head, expanded branchiostegal and opercular membranes and anterior extension of epaxial muscle. Incomplete character information, including lack of molecular data for Satan and Trogloglanis, poor quality of available skeletal preparations for Trogloglanis and Prietella, and uncertain identifications of some specimens of Prietella impede construction of a complete dataset for phylogenetic analysis.