Induced Power Scaling Alone Cannot Explain Griffenfly Gigantism.

Paleozoic skies were ruled by extinct odonatopteran insects called 'griffenflies', some with wingspans three times that of the largest extant dragonflies and ten times that of common extant dragonflies. Previous studies suggested that flight was possible for larger fliers because of higher atmospheric oxygen levels that would have increased air density. We use actuator disk theory to evaluate this hypothesis. Actuator disk theory gives similar estimates of induced power as has been estimated for micro-air vehicles based on insect flight. We calculate that for a given mass of griffenfly, and assuming isometry, a higher density atmosphere would only have reduced the induced power required to hover by 11%, which would have supported a flyer 3% larger in linear dimensions. Steady level forward flight would have further reduced induced power but could only account for a flier 5% larger in linear dimensions. Further accounting for the higher power available due to high oxygen air, and assuming isometry, we calculate that the largest flyer hovering would have been only 1.19 times longer than extant dragonflies. We also consider known allometry in dragonflies and estimated allometry in extinct griffenflies. But such allometry only increases flyer size to 1.22 times longer while hovering. We also consider profile and parasite power, but both would have been higher in denser air and thus would not have enhanced the flyability of larger griffenflies. The largest meganeurid griffenflies might have adjusted flight behaviors to reduce power required. Alternatively, the scaling of flight muscle power may have been sufficient to support the power demands of large griffenflies. In literature estimates, mass-specific power output scales as mass0.24 in extant dragonflies. We need only more conservatively assume that mass-specific muscle power scales with mass0, when combined with higher oxygen concentrations and induced power reductions in higher density air to explain griffenflies 3.4 times larger than extant odonates. Experimental measurement of flight muscle power scaling in odonates is necessary to test this hypothesis.

Palaeoatmosphere facilitates a gliding transition to powered flight in the Eocene bat, Onychonycteris finneyi.

The evolutionary transition to powered flight remains controversial in bats, the only flying mammals. We applied aerodynamic modeling to reconstruct flight in the oldest complete fossil bat, the archaic Onychonycteris finneyi from the early Eocene of North America. Results indicate that Onychonycteris was capable of both gliding and powered flight either in a standard normodense aerial medium or in the hyperdense atmosphere that we estimate for the Eocene from two independent palaeogeochemical proxies. Aerodynamic continuity across a morphological gradient is further demonstrated by modeled intermediate forms with increasing aspect ratio (AR) produced by digital elongation based on chiropteran developmental data. Here a gliding performance gradient emerged of decreasing sink rate with increasing AR that eventually allowed applying available muscle power to achieve level flight using flapping, which is greatly facilitated in hyperdense air. This gradient strongly supports a gliding (trees-down) transition to powered flight in bats.

The engineering of the giant dragonflies of the Permian: revised body mass, power, air supply, thermoregulation and the role of air density.

An engineering examination of allometric and analogical data on the flight of giant Permian insects (Protodonata, Meganeura or griffinflies) indicates that previous estimates of the body mass of these insects are too low and that the largest of these insects (wingspan of 70 cm or more) would have had a mass of 100-150 g, several times greater than previously thought. Here, the power needed to generate lift and fly at the speeds typical of modern large dragonflies is examined together with the metabolic rate and subsequent heat generated by the thoracic muscles. This evaluation agrees with previous work suggesting that the larger specimens would rapidly overheat in the high ambient temperatures assumed in the Permian. Various extant mechanisms of thermoregulation are modelled and quantified, including behaviour, radiation and the constraints on convective respiration and evaporation imposed by air flow through spiracles. However, the effects of these on cooling an overheated insect are found to be limited. Instead, an examination of the heat budget in the flight medium indicates that, at about 1.6 bar (160 kPa), thermoregulation supply enters into equilibrium and, even at high ambient temperatures, overheating can be avoided and enough oxygen sourced. This approach indicates how fossil biology can be used to examine past atmospheres.