The perinate skull of Byronosaurus (Troodontidae) with observations on the cranial ontogeny of paravian theropods

摘要

The skulls of two perinate paravians from Ukhaa Tolgod, Djadoktha Formation, Mongolia, are described here. The skulls are nearly unique in their combination of ontogenetic age and preservational quality and provide us with the first look at the morphology of such important anatomical regions as the rostrum, palate, and braincase at or near the onset of postnatal development in a nonavian paravian coelurosaur. Based on a number of derived characters, the skulls are allocated to a derived position within Troodontidae that is outside the clade consisting of Saurornithoides mongoliensis, Saurornithoides junior, Troodon formosus, and probably Sinornithoides youngi. A single synapomorphy, presence of a lateral maxillary groove, supports the Ukhaa perinates as Byronosaurus. The comparative morphology of the Ukhaa perinates with adult troodontids indicates a number of significant postnatal transformations (e.g., elongation and flattening of the rostrum, increase in the number of maxillary and dentary teeth, restructuring of the occipital plate and paroccipital process). These comparisons demonstrate that many characters historically considered important for phylogenetic and taxonomic assessments of adult maniraptorans are present at a relativel+y early stage of ontogeny. Differences in the developmental timing of various cranial characters have important implications for interpreting the fossil record as well as for understanding the role heterochrony has played in the evolution of derived coleurosaurs, including birds. The ontogenetic information provided by the Ukhaa perinates also allow us to comment on the enigmatic paravian Archaeornithoides deinosauriscus, which has been considered both the sister taxon to Avialae and a juvenile specimen of the troodontids Saurornithoides mongoliensis and Byronosaurus jaffei. We found no unique characters that support a priviledged relationship of Archaeornithoides deinosauriscus with avialans and only weak character support for this taxon as a basal troodontid-there is no known character evidence supporting it as a juvenile of either Saurornithoides or Byronosaurus.