Evolutionary trends in Triceratops from the Hell Creek Formation, Montana.

The placement of over 50 skulls of the well-known horned dinosaur Triceratops within a stratigraphic framework for the Upper Cretaceous Hell Creek Formation (HCF) of Montana reveals the evolutionary transformation of this genus. Specimens referable to the two recognized morphospecies of Triceratops, T. horridus and T. prorsus, are stratigraphically separated within the HCF with the T. prorsus morphology recovered in the upper third of the formation and T. horridus found lower in the formation. Hypotheses that these morphospecies represent sexual or ontogenetic variation within a single species are thus untenable. Stratigraphic placement of specimens appears to reveal ancestor-descendant relationships. Transitional morphologies are found in the middle unit of the formation, a finding that is consistent with the evolution of Triceratops being characterized by anagenesis, the transformation of a lineage over time. Variation among specimens from this critical stratigraphic zone may indicate a branching event in the Triceratops lineage. Purely cladogenetic interpretations of the HCF dataset imply greater diversity within the formation. These findings underscore the critical role of stratigraphic data in deciphering evolutionary patterns in the Dinosauria.