Joermungandr bolti, an exceptionally preserved microsaur from Mazon Creek, Illinois, reveals patterns of integumentary evolution in Recumbirostra

The Carboniferous Pennsylvanian-aged (309-307 Ma) Mazon Creek Lagerstätte produces some of the earliest fossils of major Paleozoic tetrapod lineages. Recently, several new tetrapod specimens collected from Mazon Creek have come to light, including the earliest fossorially-adapted recumbirostrans. Here we describe a new long-bodied recumbirostran, Joermungandr bolti gen et sp. nov., known from a single part and counterpart concretion bearing a virtually complete skeleton. Uniquely, Joermungandr preserves a full suite of dorsal, flank, and ventral dermal scales, together with a series of thinned and reduced gastralia. Investigation of these scales using Scanning Electron Microscopy (SEM) reveals ultrastructural ridge and pit morphologies, revealing complexities comparable to the scale ultrastructure of extant snakes and fossorial reptiles, which have scales modified for body based propulsion and shedding substrate. Our new taxon also represents an important early record of an elongate recumbirostran bauplan, wherein several features linked to fossoriality, including a characteristic recumbent snout, are present. We utilized parsimony phylogenetic methods to conduct phylogenetic analysis using the most recent recumbirostran-focused matrix. The analysis recovers the new taxon within the Cocytinoidea clade as sister taxon to all brachystelechids. Finally, we review integumentary patterns in Recumbirostra, noting reductions and losses of gastralia and osteoderms associated with body elongation and, thus, likely also associated with increased fossoriality.

Methods
Data Matrix.