A review of paravian phylogeny with new data

Abstract

Paraves are theropod dinosaurs comprising of living and fossil birds and their closest fossil relatives, the dromaeosaurid and troodontid dinosaurs. Traditionally, birds have been recovered as the sister group to Deinonychosauria, the clade made up of the two subclades Dromaeosauridae and Troodontidae. However, spectacular Late Jurassic paravian fossils discovered from northeastern China - including Anchiornis and Xiaotingia - preserve anatomy that seemingly challenges the status quo. To resolve this debate we performed an up-to-date phylogenetic analysis for paravians using the latest Theropod Working Group (TWiG) coelurosaur data matrix which we supplemented with new data from recently described Mesozoic paravians from Asia and North America (e.g., Zhenyuanlong and Acheroraptor). This includes data from the unnamed dromaeosaurid IVPP V22530 and Luanchuanraptor, which are included in a phylogenetic analysis for the first time. We also incorporate new data from iconic paravians such as Archaeopteryx and Velociraptor based on firsthand study. The analysis adopted the maximum parsimony criterion and was performed in the phylogenetic software TNT. Our preliminary results support the monophyly of each of the traditionally recognized paravian clades. The Late Jurassic paravians from northeastern China (e.g., Anchiornis and Xiaotingia) are recovered as avialans rather than deinonychosaurians, at a position more basal than Archaeopteryx and other derived avialans. The traditional sister group status of Troodontidae and Dromaeosauridae is reaffirmed and is supported by a laterally exposed splenial and a characteristic raptorial pedal digit II. Recently reported Early Cretaceous dromaeosaurids from northern and northeastern China, including Zhenyuanlong, Changyuraptor and IVPP V22530, are closely related to other microraptorines as expected. Luanchuanraptor, a dromaeosaurid from the Late Cretaceous of central China is recovered as a more advanced eudromaeosaurian. By tracing character evolution on the current tree topology we report on the latest insights into the adaptive radiation amongst early paravians, including the origin of flight and changes in body size and diet.