Feather-like development of Triassic diapsid skin appendages

Of the recent sauropsid skin appendage types, only feathers develop from a cylindrical epidermal invagination, the follicle, and show hierarchical branching. Fossilized integuments of Mesozoic diapsids have been interpreted as follicular and potential feather homologues, an idea particularly controversially discussed for the elongate dorsal skin projections of the small diapsid Longisquama insignis from the Triassic of Kyrgyzstan. Based on new finds and their comparison with the type material, we show that Longisquama's appendages consist of a single-branched internal frame enclosed by a flexible outer membrane. Not supporting a categorization either as feathers or as scales, our analysis demonstrates that the Longisquama appendages formed in a two-stage, feather-like developmental process, representing an unusual early example for the evolutionary plasticity of sauropsid integument.