title: On the size and flight diversity of giant pterosaurs, the use of birds as pterosaur analogues and comments on pterosaur flightlessness.
abstract: The size and flight mechanics of giant pterosaurs have received considerable research interest for the last century but are confused by conflicting interpretations of pterosaur biology and flight capabilities. Avian biomechanical parameters have often been applied to pterosaurs in such research but, due to considerable differences in avian and pterosaur anatomy, have lead to systematic errors interpreting pterosaur flight mechanics. Such assumptions have lead to assertions that giant pterosaurs were extremely lightweight to facilitate flight or, if more realistic masses are assumed, were flightless. Reappraisal of the proportions, scaling and morphology of giant pterosaur fossils suggests that bird and pterosaur wing structure, gross anatomy and launch kinematics are too different to be considered mechanically interchangeable. Conclusions assuming such interchangeability--including those indicating that giant pterosaurs were flightless--are found to be based on inaccurate and poorly supported assumptions of structural scaling and launch kinematics. Pterosaur bone strength and flap-gliding performance demonstrate that giant pterosaur anatomy was capable of generating sufficient lift and thrust for powered flight as well as resisting flight loading stresses. The retention of flight characteristics across giant pterosaur skeletons and their considerable robustness compared to similarly-massed terrestrial animals suggest that giant pterosaurs were not flightless. Moreover, the term 'giant pterosaur' includes at least two radically different forms with very distinct palaeoecological signatures and, accordingly, all but the most basic sweeping conclusions about giant pterosaur flight should be treated with caution. Reappraisal of giant pterosaur material also reveals that the size of the largest pterosaurs, previously suggested to have wingspans up to 13 m and masses up to 544 kg, have been overestimated. Scaling of fragmentary giant pterosaur remains have been misled by distorted fossils or used inappropriate scaling techniques, indicating that 10-11 m wingspans and masses of 200-250 kg are the most reliable upper estimates of known pterosaur size.
authors.affiliation: School of Earth and Environmental Sciences, University of Portsmouth, Portsmouth, United Kingdom. mark.witton@port.ac.uk
authors.forename: Mark P
authors.initials: MP
authors.lastname: Witton
authors.forename: Michael B
authors.initials: MB
authors.lastname: Habib
mesh_terms: D000818:Animals; D005075:Biological Evolution; D001696:Biomechanical Phenomena; D001717:Birds / Q000033:anatomy & histology / Q000502:physiology*; D049628:Body Size / Q000502:physiology*; D001842:Bone and Bones / Q000033:anatomy & histology / Q000502:physiology; D025061:Dinosaurs / Q000033:anatomy & histology / Q000145:classification / Q000502:physiology*; D005426:Flight, Animal / Q000502:physiology*; D005552:Forelimb / Q000033:anatomy & histology / Q000502:physiology; D005580:Fossils; D012863:Skeleton; D013045:Species Specificity; D014921:Wings, Animal / Q000033:anatomy & histology / Q000502:physiology