at least with its long bill and neck while floating on the surface, and they may have reached even greater depths by plunge-diving into the water from the air like some modern long-winged seabirds. In 1994, Bennett noted that the head, neck, and shoulders of Pteranodon were as heavily built as diving birds, and suggested that they could dive by folding back their wings like the modern gannet. Crest function Pteranodon was notable for its skull crest, though the function of this crest has been a subject of debate. Most explanations have focused on the blade-like, backward pointed crest of male P. longiceps, however, and ignored the wide range of variation across age and sex. The fact that the crests vary so much rules out most practical functions other than for use in mating displays. Therefore, display was probably the main function of the crest, and any other functions were secondary. Scientific interpretations of the crest's function began in 1910, when George Francis Eaton proposed two possibilities: an aerodynamic counterbalance and a muscle attachment point. He suggested that the crest might have anchored large, long jaw muscles, but admitted that this function alone could not explain the large size of some crests. Bennett (1992) agreed with Eaton's own assessment that the crest was too large and variable to have been a muscle attachment site. Eaton had suggested that a secondary function of the crest might have been as a counterbalance against the long beak, reducing the need for heavy neck muscles to control the orientation of the head. Wind tunnel tests showed that the crest did function as an effective counterbalance to a degree, but Bennett noted that, again, the hypothesis focuses only on the long crests of male P. longiceps, not on the larger crests of P. sternbergi and very small crests that existed among the females. Bennett found that the crests of females had no counterbalancing effect, and that the crests of male P. sternbergi would, by themselves, have a negative effect on the balance of the head. In fact, side to side movement of the crests would have required more, not less, neck musculature to control balance. In 1943, Dominik von Kripp suggested that the crest may have served as a rudder, an idea embraced by several later researchers. One researcher, Ross S. Stein, even suggested that the crest may have supported a membrane of skin connecting