Theropod trackways as indirect evidence of pre-avian aerial behavior.

Body fossils set limits on feasible reconstructions of functional capacity and behavior in theropod dinosaurs, but do not document in-life behaviors. In contrast, trace fossils such as footprints preserve in-life behaviors that can potentially test and enhance existing reconstructions. Here, we demonstrate how theropod trackways can be used as indirect evidence of pre-avian aerial behavior, expanding the approaches available to study vertebrate flight origins.

Preserved soft anatomy confirms shoulder-powered upstroke of early theropod flyers, reveals enhanced early pygostylian upstroke, and explains early sternum loss.

Anatomy of the first flying feathered dinosaurs, modern birds and crocodylians, proposes an ancestral flight system divided between shoulder and chest muscles, before the upstroke muscles migrated beneath the body. This ancestral flight system featured the dorsally positioned deltoids and supracoracoideus controlling the upstroke and the chest-bound pectoralis controlling the downstroke. Preserved soft anatomy is needed to contextualize the origin of the modern flight system, but this has remained elusive.

Response to Serrano and Chiappe.

In the recent study in Current Biology by Pei and colleagues, we used two proxies - wing loading and specific lift - to reconstruct powered flight potential across the vaned feathered fossil pennaraptorans. The results recovered multiple origins of powered flight. We respectfully disagree with the criticism raised by Serrano and Chiappe that wing loading and specific lift, used in sequence, fail to discriminate between powered flight and gliding.

Potential for Powered Flight Neared by Most Close Avialan Relatives, but Few Crossed Its Thresholds.

Uncertainties in the phylogeny of birds (Avialae) and their closest relatives have impeded deeper understanding of early theropod flight. To help address this, we produced an updated evolutionary hypothesis through an automated analysis of the Theropod Working Group (TWiG) coelurosaurian phylogenetic data matrix. Our larger, more resolved, and better-evaluated TWiG-based hypothesis supports the grouping of dromaeosaurids + troodontids (Deinonychosauria) as the sister taxon to birds (Paraves) and the recovery of Anchiornithinae as the earliest diverging birds.