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Synopsis Flying insects vary greatly in body size and wing proportions, significantly impacting their flight energetics. Generally, the larger the insect, the slower its flight wingbeat frequency. However, variation in frequency is also explained by differences in wing proportions, where larger-winged insects tend to have lower frequencies. These associations affect the energy required for flight. The correlated evolution of flight form and function can be further defined using a lineage of closely related bee species varying in body mass. The decline in flight wingbeat frequency with increasing size is paralleled by the flight mass-specific metabolic rate. The specific scaling exponents observed can be predicted from the wing area allometry, where a greater increase (hyperallometry) leads to a more pronounced effect on flight energetics, and hypoallometry can lead to no change in frequency and metabolic rate across species. The metabolic properties of the flight muscles also vary with body mass and wing proportions, as observed from the activity of glycolytic enzymes and the phospholipid compositions of muscle tissue, connecting morphological differences with muscle metabolic properties. The evolutionary scaling observed across species is recapitulated within species. The static allometry observed within the bumblebee Bombus impatiens, where the wing area is proportional and isometric, affects wingbeat frequency and metabolic rate, which is predicted to decrease with an increase in size. Intraspecific variation in flight muscle tissue properties is also related to flight metabolic rate. The role of developmental processes and phenotypic plasticity in explaining intraspecific differences is central to our understanding of flight energetics. These studies provide a framework where static allometry observed within species gives rise to evolutionary allometry, connecting the evolution of size, form, and function associated with insect flight.
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Induced Power Scaling Alone Cannot Explain Griffenfly Gigantism
Synopsis Paleozoic skies were ruled by extinct odonatopteran insects called “griffenflies,” some with wingspans 3 times that of the largest extant dragonflies and 10 times that of common extant dragonflies. Previous studies suggested that flight was possible for larger fliers because of higher atmospheric oxygen levels, which would have increased air density. We use actuator disk theory to evaluate this hypothesis. Actuator disk theory gives similar estimates of induced power as have 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.
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- Award ID(s):
- 2326924
- PAR ID:
- 10542455
- Publisher / Repository:
- Oxford University Press
- Date Published:
- Journal Name:
- Integrative And Comparative Biology
- Volume:
- 64
- Issue:
- 2
- ISSN:
- 1540-7063
- Format(s):
- Medium: X Size: p. 598-610
- Size(s):
- p. 598-610
- Sponsoring Org:
- National Science Foundation
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