The phylogenetic affinities and morphological peculiarities of the bird-like dinosaur Borogovia gracilicrus from the Upper Cretaceous of Mongolia

Abstract

Borogovia gracilicrus is a small-bodied theropod dinosaur from the Maastrichtian (Upper Cretaceous) Nemegt Formation of southern Mongolia. The taxon is based on a single fragmentary specimen preserving only the distal part of the hindlimbs. The morphology of Borogovia shows a peculiar combination of features, some of which are traditionally considered troodontid synapomorphies and others which are unusual for Troodontidae but are shared with other maniraptoran clades. In particular, the second toe of B. gracilicrus differs from other troodontids in lacking some of the features which contribute to the specialized ‘sickle-clawed’ second toe, here termed the ‘falciphoran condition’, shared with dromaeosaurids and some other paravians, such as the strongly compressed and falciform ungual. Phylogeny reconstructions intended to explore the affinities of Borogovia consistently support its referral within a subclade of troodontids including all Late Cretaceous taxa. The placement of Borogovia is not significantly affected by its unusual combinations of hindlimb features or by the homoplasy of the elements forming the falciphoran condition. Borogovia is supported as a valid taxon and is distinct from the other Nemegt troodontids, Tochisaurus and Zanabazar. The lack of a falciform ungual, and the distinctive morphology of the second toe in B. gracilicrus are interpreted as a derived specialization among Troodontidae and not as retention of the plesiomorphic condition of non-paravian theropods.