Controversy has prevailed as to how insects of the Carboniferous period were able to grow so large. The way
oxygen is
diffused through the insect's body via its
tracheal breathing system (see
Respiratory system of insects) puts an upper limit on body size, which prehistoric insects seem to have well exceeded. It was originally proposed in that
Meganeura was only able to fly because the atmosphere at that time contained more oxygen than the present 20%. This theory was dismissed by fellow scientists, but has found approval more recently through further study into the relationship between
gigantism and oxygen availability. If this theory is correct, these insects would have been susceptible to falling oxygen levels and certainly could not survive in the modern atmosphere. Other research indicates that insects really do breathe, with "rapid cycles of tracheal compression and expansion". Recent analysis of the flight energetics of modern insects and birds suggests that both the oxygen levels and air density provide a bound on size. A general problem with all oxygen related explanations of giant griffenflies is the circumstance that very large Meganeuridae (
Arctotypus sp.) with a wingspan of 45 cm also occurred in the
Late Permian of
Lodève in France, when the oxygen content of the atmosphere was already much lower than in the
Carboniferous and
Early Permian. suggested that the lack of aerial vertebrate predators allowed
pterygote insects to evolve to maximum sizes during the Carboniferous and Permian periods, maybe accelerated by an "
evolutionary arms race" for increase in body size between plant-feeding
Palaeodictyoptera and
meganeurids as their predators. ==Families and genera==