A Paleozoic stem hagfish Myxinikela siroka - revised anatomy and implications for evolution of the living jawless vertebrate lineages

Hagfishes and lampreys comprise cyclostomes, the earliest branching and sole surviving Glade of the once diverse assemblage of jawless crown-group vertebrates. Lacking mineralized skeletons, both of the crown cyclostome lineages have notoriously poor fossil record. Particularly in the hagfish total group, dagger Myxinikela siroka Bardack, 1991 from the Late Carboniferous estuarine system of Illinois (USA) represents the only definitive stem taxon. Previously known from a single specimen, Myxinikela has been reconstructed as a short-bodied form with pigmented eyes but otherwise difficult to distinguish from the living counterpart. With a new, second specimen of Myxinikela reported here, I reevaluate the soft tissue anatomy and formulate diagnosis for the taxon. Myxinikela has a number of general features of cyclostomes, including cartilaginous branchial baskets, separation between the esophageal and the branchial passages, and a well-differentiated midline finfold. In effect, these features give more lamprey-like appearance to this stem hagfish than previously assumed. Myxinikela still has many traits that set modern hagfishes apart from other vertebrates (e.g., nasohypophyseal aperture, large velar cavity, and cardinal heart) and some intermediate conditions of modern hagfishes (e.g., incipient posterior displacement of branchial region). Thus, Myxinikela provides an important calibration point with which to date origins of these characters.