Heavily loaded flight and limits to the maximum size of dragonflies (Anisoptera) and griffenflies (Meganisoptera)

An original hypothesis is presented that the maximum mass and size of living anisopteran dragonflies are constrained by a physiological performance limit: the wing muscle power required to permit reproductively successful males to carry heavier females in the so-called wheel position' in flight. It is proposed that the same limit cannot have applied to all fossil Odonatoptera. As the physiology of the giant Carboniferous griffenfly Namurotypus sippeli precludes flight in the wheel position, it did not need to carry any substantial load aside from exogenous aerial prey. Based on its thorax dimensions, it is argued that Namurotypus flew with a relatively low maximum specific muscle power output in comparison with living Anisoptera. The extinction of some families of large Mesozoic Odonatoptera may have been exacerbated by competition with smaller (stem-) Anisoptera that evolved higher specific power outputs and superior flight performance similar to living Anisoptera. To investigate the credibility of this flight-performance size-limit hypothesis and its consequences, an analysis of the scaling of the required flight power and available muscle power is presented using allometric relations. It is found that for living Anisoptera and fossil Odonatoptera, there are different limiting sizes, above which the required specific flight power would exceed the available muscle specific power. These limits are directly related to maximum load-carrying capacity and the atmospheric air density at the habitual altitude. It is suggested that the largest living species of Petaluridae, Petalura ingentissima, is close to the proposed Anisoptera size limit at current near-sea-level air density conditions.