Lujiatun Psittacosaurids: understanding individual and taphonomic variation using 3D geometric morphometrics.

Psittacosaurus is one of the most abundant and speciose genera in the Dinosauria, with fifteen named species. The genus is geographically and temporally widespread with large sample sizes of several of the nominal species allowing detailed analysis of intra- and interspecific variation. We present a reanalysis of three separate, coeval species within the Psittacosauridae; P. lujiatunensis, P. major, and Hongshanosaurus houi from the Lujiatun beds of the Yixian Formation, northeastern China, using three-dimensional geometric morphometrics on a sample set of thirty skulls in combination with a reevaluation of the proposed character states for each species. Using these complementary methods, we show that individual and taphonomic variation are the joint causes of a large range of variation among the skulls when they are plotted in a morphospace. Our results demonstrate that there is only one species of Psittacosaurus within the Lujiatun beds and that the three nominal species represent different taphomorphotypes of P. lujiatunensis. The wide range of geometric morphometric variation in a single species of Psittacosaurus implies that the range of variation found in other dinosaurian groups may also be related to taphonomic distortion rather than interspecific variation. As the morphospace is driven primarily by variation resulting from taphonomic distortion, this study demonstrates that the geometric morphometric approach can only be used with great caution to delineate interspecific variation in Psittacosaurus and likely other dinosaur groups without a complementary evaluation of character states. This study presents the first application of 3D geometric morphometrics to the dinosaurian morphospace and the first attempt to quantify taphonomic variation in dinosaur skulls.