found that the pterosaur with jaws most similar to those of Thalassodromeus was the smaller Rhamphorhynchus, although they believed that it would have had limited skimming ability. In 2004, palaeontologist Sankar Chatterjee and engineer R. Jack Templin said that smaller pterosaurs may have been able to skim-feed. They doubted that this was possible for larger ones, due to their lesser manoeuvrability and flying capability while resisting water. Chatterjee and Templin noted that skimmers have blunter beaks than pterosaurs like Thalassodromeus, to direct water from the jaw while skimming. In 2007, biophysicist Stuart Humphries and colleagues questioned whether any pterosaurs would have commonly fed by skimming and said that such conclusions had been based on anatomical comparisons rather than biomechanical data. The drag experienced by bird bills and pterosaur jaws was hydrodynamically and aerodynamically tested by creating model bills of the black skimmer, Thalassodromeus, and the (presumably) non-skimming Tupuxuara and towing them along a water-filled trough at varying speeds. The researchers found that skimming used more energy for skimmers than previously thought, and would have been impossible for a pterosaur weighing more than due to the metabolic power required. They found that even smaller pterosaurs, like Rhamphorhynchus, were not adapted for skimming. The aluminium rigging of the Thalassodromeus model was destroyed during the experiment, due to the high and unstable forces exerted on it while skimming at high speed, casting further doubt on this feeding method. The authors used the jaw tip of T. oberlii to model the performance of Thalassodromeus, since it was assigned to T. sethi at the time. Unwin and Martill suggested in 2007 that thalassodromids may have foraged similarly to storks, as had been suggested for azhdarchids. Witton said in 2013 that although skim-feeding had been suggested for many pterosaur groups, the idea was criticised in recent years; pterosaurs lacked virtually all adaptations for skim-feeding, making it unlikely that they fed this way. Thalassodromeus (unlike skimmers) did not have a particularly wide or robust skull or especially large jaw-muscle attachment sites, and its mandible was comparatively short and stubby. Witton agreed with Unwin and Martill that thalassodromids, with their equal limb proportions and elongated jaws, were suited to roaming terrestrially and feeding opportunistically; their shorter, more flexible necks indicated a different manner of feeding than azhdarchids, which had longer, stiffer necks. He suggested that thalassodromids may have had more generalised feeding habits, and azhdarchids may have been