Meganisoptera

Meganisoptera Temporal range: Pennsylvanian-Lopingian PreЄ Є O S D C P T J K Pg N
Illustration of a Meganeura species
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Superorder:	Odonatoptera
Order:	†Meganisoptera Martynov, 1932
Families
Meganeuridae Paralogidae
Synonyms
Protodonata C. Brongniart, 1893

Meganisoptera is an extinct order of very large to gigantic insects often called griffenflies. The order was formerly named Protodonata for their similar appearance and relation to dragonflies. They range in Palaeozoic (Late Carboniferous to Late Permian) insects, Most were only slightly larger than modern dragonflies and the order includes the largest known insect species such as the late Carboniferous Meganeura monyi, Megatypus, and the even larger early Permian Meganeuropsis permiana, with wingspans of up to 71 centimetres (2.33 ft).^[1]

The fore and hind wings are similar in venation (a primitive feature) except for the larger anal (rearwards) area in the hind wing. The fore wing is usually more slender and slightly longer than the hind wing. Unlike the true dragonflies, they had no pterostigma, and a somewhat simpler pattern of veins in the wings.

Most specimens are known from wing fragments only; with only a few as complete wings, and even fewer (of the family Meganeuridae) with body impressions. These show a globose head with large dentate mandibles, strong spiny legs, a large thorax, and long and slender dragonfly-like abdomen. Like true dragonflies, they were presumably predators.

A few nymphs are also known, and show similar mouthparts to those of modern dragonfly nymphs, suggesting that they were also active aquatic predators.^[2]

Although sometimes included under the dragonflies, the Protodonata lack certain distinctive wing features that characterise the Odonata. Grimaldi and Engel, pointing out that the colloquial term "giant dragonfly" is therefore misleading, suggest "griffinfly" instead.

Size

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 puts an upper limit on body size, which prehistoric insects seem to have well exceeded. It was originally proposed (Harlé & Harlé, 1911) 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.^[3] If this theory is correct, these insects would have been susceptible to falling oxygen levels and certainly could not survive in our modern atmosphere. Other research indicates that insects really do breathe, with "rapid cycles of tracheal compression and expansion".^[4] 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^[5].

A general problem with all oxygen related explanations of the giant dragonflies is the circumstance that very large Meganeuridae with a wing span of 45 cm also occurred in the Upper Permian of Lodève in France, when the oxygen content of the atmosphere was already much lower than in the Carboniferous and Lower Permian.^[6]

Bechly (2004) 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.^[7]

Cited references

1. ^ Grimaldi & Engel 2005 p.175
2. ^ Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. p. 321. ISBN 0-19-510033-6. 
3. ^ Gauthier Chapelle and Lloyd S. Peck (May 1999). "Polar gigantism dictated by oxygen availability". Nature 399 (6732): 114–115. doi:10.1038/20099. http://www.nature.com/nature/journal/v399/n6732/abs/399114b0.html. "Oxygen supply may also have led to insect gigantism in the Carboniferous period, because atmospheric oxygen was 30-35% (ref. 7). The demise of these insects when oxygen content fell indicates that large species may be susceptible to such change. Giant amphipods may therefore be among the first species to disappear if global temperatures are increased or global oxygen levels decline. Being close to the critical MPS limit may be seen as a specialization that makes giant species more prone to extinction over geological time." 
4. ^ Westneat MW, Betz O, Blob RW, Fezzaa K, Cooper WJ, Lee WK. (January 2003). "Tracheal respiration in insects visualized with synchrotron x-ray imaging". Science 299 (5606): 558–560. doi:10.1126/science.1078008. PMID 12543973. "Insects are known to exchange respiratory gases in their system of tracheal tubes by using either diffusion or changes in internal pressure that are produced through body motion or hemolymph circulation. However, the inability to see inside living insects has limited our understanding of their respiration mechanisms. We used a synchrotron beam to obtain x-ray videos of living, breathing insects. Beetles, crickets, and ants exhibited rapid cycles of tracheal compression and expansion in the head and thorax. Body movements and hemolymph circulation cannot account for these cycles; therefore, our observations demonstrate a previously unknown mechanism of respiration in insects analogous to the inflation and deflation of vertebrate lungs." 
5. ^ Robert Dudley (April 1998). "Atmospheric oxygen, giant Paleozoic insects and the evolution of aerial locomotion performance". The Journal of Experimental Biology 201 (Pt8): 1043–1050. PMID 9510518. "Uniformitarian approaches to the evolution of terrestrial locomotor physiology and animal flight performance have generally presupposed the constancy of atmospheric composition. Recent geophysical data as well as theoretical models suggest that, to the contrary, both oxygen and carbon dioxide concentrations have changed dramatically during defining periods of metazoan evolution. Hyperoxia in the late Paleozoic atmosphere may have physiologically enhanced the initial evolution of tetrapod locomotor energetics; a concurrently hyperdense atmosphere would have augmented aerodynamic force production in early flying insects. Multiple historical origins of vertebrate flight also correlate temporally with geological periods of increased oxygen concentration and atmospheric density. Arthropod as well as amphibian gigantism appear to have been facilitated by a hyperoxic Carboniferous atmosphere and were subsequently eliminated by a late Permian transition to hypoxia. For extant organisms, the transient, chronic and ontogenetic effects of exposure to hyperoxic gas mixtures are poorly understood relative to contemporary understanding of the physiology of oxygen deprivation. Experimentally, the biomechanical and physiological effects of hyperoxia on animal flight performance can be decoupled through the use of gas mixtures that vary in density and oxygen concentration. Such manipulations permit both paleophysiological simulation of ancestral locomotor performance and an analysis of maximal flight capacity in extant forms." 
6. ^ Nel A.N., Fleck G., Garrouste R. and Gand, G. (2008): The Odonatoptera of the Late Permian Lodève Basin (Insecta). Journal of Iberian Geology 34(1): 115-122 PDF
7. ^ Bechly G. (2004): Evolution and systematics. pp. 7-16 in: Hutchins M., Evans A.V., Garrison R.W. and Schlager N. (eds): Grzimek's Animal Life Encyclopedia. 2nd Edition. Volume 3, Insects. 472 pp. Gale Group, Farmington Hills, MI PDF

General references

• Carpenter, F. M. 1992. Superclass Hexapoda. Volume 3 of Part R, Arthropoda 4; Treatise on Invertebrate Paleontology, Boulder, Colorado, Geological Society of America.
• Grimaldi, David and Engel, Michael S. (2005-05-16). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 
• Tasch, Paul, 1973, 1980 Paleobiology of the Invertebrates, John Wiley and Sons, p. 617
• André Nel, Günther Fleck, Romain Garrouste, Georges Gand, Jean Lapeyrie, Seth M Bybee, and Jakub Prokop (2009): Revision of Permo-Carboniferous griffenflies (Insecta: Odonatoptera: Meganisoptera) based upon new species and redescription of selected poorly known taxa from Eurasia. Palaeontographica Abteilung A, 289(4-6): 89–121.

External links

• Phylogenetic Systematics of Odonata by Günter Bechly, 2005