Mecoptera

Mecoptera Temporal range: Permian - Recent 299–0 Ma PreЄ Є O S D C P T J K Pg N
Panorpa communis, male
Scientific classification
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Superorder:	Panorpida
(unranked):	Antliophora
Order:	Mecoptera Hyatt & Arms, 1891
Families
Apteropanorpidae Bittacidae (hangingflies) Boreidae (snow scorpionflies) Choristidae †Dinopanorpidae Eomeropidae Meropeidae (earwigflies) Nannochoristidae Panorpidae (common scorpionflies) Panorpodidae (short-faced scorpionflies)

Mecoptera (from the Greek: meco- = "long", -ptera = "wings") are an order of insects with about 550 species in nine families worldwide. Mecoptera are sometimes called scorpionflies after their largest family, Panorpidae, in which the males have enlarged genitals that look similar to the stinger of a scorpion. The Bittacidae, or hangingflies, are a prominent family of elongate insects known for their elaborate mating rituals, in which females choose mates based on the quality of gift prey offered by various males.

While modern mecoptera are overwhelmingly predators or consumers of dead organisms, early ones might have played an important role before the evolution of other insects in pollinating extinct gymnosperms.^[1][2]

Anatomy and biology

Mecoptera are small to medium insects with slender, elongated, bodies. They have relatively simple mouthparts, with long mandibles and fleshy palps, which resemble those of the more primitive true flies. Like many other insects, they possess compound eyes on the side of the head, and three ocelli on the top. Most Mecoptera feed on vegetation in moist environments; in hotter climates, they may therefore be active only for short periods of the year.^[3]

The wings are narrow in shape, with numerous cross-veins, and somewhat resemble those of primitive insects such as mayflies. A few genera, however, have reduced wings, or have lost them altogether. The abdomen is cylindrical, and typically curves upwards in the male, superficially resembling the tail of a scorpion.

The female lays the eggs in close contact with moisture, and the eggs typically absorb water and increase in size after deposition. In species that live in hot conditions, the eggs may not hatch for several months, the larvae only emerging when the dry season has finished. More typically, however, they hatch after a relatively short period of time.^[3]

The larvae are usually quite caterpillar-like, with short, clawed, true legs, and a number of abdominal prolegs. They have a sclerotised head with compound eyes and mandibulate mouthparts. The tenth abdominal segment bears either a suction disc, or, less commonly, a pair of hooks. They generally eat vegetation or scavenge for dead insects, although some predatory larvae are known.^[3]

The larva crawls into the soil or decaying wood to pupate, and does not spin a cocoon. The pupae are exarate, meaning that the limbs are free of the body, and are able to move their mandibles, but are otherwise entirely non-motile. In drier environments, they may spend several months in diapause, before emerging as adults once the conditions are more suitable.^[3]

Evolution

DNA evidence indicates that fleas, which are traditionally considered a separate order (Order Siphonaptera), are instead highly specialized Mecoptera^[4]. Grouped together with the fleas, Mecoptera would have about 3000 known species.

Mecoptera have special importance in evolution of Insecta. Two of the most important insect orders, Lepidoptera and Diptera, along with Trichoptera, probably evolved from ancestors belonging to, or strictly related to, the Mecoptera. This is apparent from anatomical and biochemical similarities, but, moreover, transitional fossils, such as Permotanyderus and Choristotanyderus, have been discovered that lie between the Mecoptera and Diptera.^[3]

The group was once much more widespread and diverse than at present, having as many as four suborders during the Mesozoic.^[3]

First pollinators

It has been proposed that extinct mecoptera species were important plant pollinators.^[1][2] Early non-angiosperm gymnosperm seed plants during the late Middle Jurassic to mid–Early Cretaceous period have been believed to be mainly wind-pollinated. However examination of fossil mecoptera show that they had siphon feeding apparatus that could fertilize early gymnosperms by feeding on their nectar and pollen. The lack of iron enrichment in their fossilized proboscis rules out a use in blood drinking. One question over this suggestion is that so far pollen has not been found associated with these feeding parts which is surprising for the amber-encased insects which should have preserved pollen but "further fossils may provide this information".^[1]

11 species have been identified belonged to three families, Mesopsychidae, Aneuretopsychidae, and Pseudopolycentropodidae for which "the encompassing name Aneuretopsychina is available".^[2] Their length ranges from 3 mm in Parapolycentropus burmiticus to 28 mm in Lichnomesopsyche gloriae.^[2] The proboscis could be as long as 10 mm. Pollen transfer has been suggested to occur by body surface transport on mouthpart and head surfaces like that in bee flies and hover flies—however no such associated pollen has been found even though the insects were preserved in amber.^[2] It is thought that they pollinated such plants as Caytoniaceae, Cheirolepidiaceae, Czekanowskiaceae, Pentoxylaceae, and Gnetales as these have ovulate organs that are either poorly suited for wind pollination or have structures that could support long-proboscid fluid feeding.^[2]

Mating

Gullen and Cranston ^[5] offer a startling description of the mating habits of the Bittacidae:

“

"Food items such as caterpillars, bugs, and flies are offered to be eaten during copulation. The female is first attracted by a pheromone emitted by one or more vesicles or pouches at the end of the male's abdomen. When the female is near, the vesicles are retracted. The female examines the offering while the male searches for her genitalia with his own. If the gift is rejected, the female flies away. If the gift is accepted, the genitalia of the male couples with that of the female, who lowers herself until she is hanging upside down. She consumes the offering during copulation. The male supports the female by holding her legs or the prey. Field observations show that both sexes mate several times per day. Small or unacceptable offerings result in no or a very short copulation time. Duration depends on the size of the gift. It has been observed that prey 3 to 14 mm long will provide 1 to 17 minutes of copulation in Hylobittacus apicalis. Larger H. apicalis give prey the size of houseflies in return for 20 to 29 minutes of copulation. This results in a maximum sperm transfer, increased oviposition, and a refractory period."

”

• 

  Male genital (Panorpa communis)

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  Panorpa meridionalis, from Portugal

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  Bittacidae species from Australia

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  Panorpa communis female with prey

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  Panorpa communis male

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  Detail of head, male (Panorpa communis)

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  Dinokanaga andersoni Forewing

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  Male Panorpa communis

References

1. ^ ^{a b c} Ollerton J. Coulthard E. (2009). Evolution of Animal Pollination. Science, 326: 808-809. doi:10.1126/science.1181154
2. ^ ^{a b c d e f} Ren D, Labandeira CC, Santiago-Blay JA, Rasnitsyn A, Shih CK, Bashkuev A, Logan MA, Hotton CL, Dilcher D. (2009). Probable Pollination Mode Before Angiosperms: Eurasian, Long-Proboscid Scorpionflies. Science, 326 (5954), 840-847. doi:10.1126/science.1178338
3. ^ ^{a b c d e f} Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. pp. 488–491. ISBN 0-19-510033-6. 
4. ^ Whiting, Michael F. (2002). "Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera". Zoologica Scripta 31 (1): 93–104. doi:10.1046/j.0300-3256.2001.00095.x. http://www3.interscience.wiley.com/journal/118916123/abstract. 
5. ^ Gullan, P.J. and P.S. Cranston. The Insects: An Outline of Entomology, second edition. 2000: Blackwell Science, Ltd.

• Grimaldi, D. and Engel, M.S. (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 
• Archibald, S.B. (2005). "New Dinopanorpida (Insecta: Mecoptera) from the Eocene Okanogan Highlands (British Columbia, Canada and Washington State, USA)". Canadian Journal of Earth Sciences 4 (2): 119–136. doi:10.1139/e04-073. 

External links

• Mecoptera at the Tree of Life
• Mecoptera image gallery at myrmecos.net
• Video of Mecoptera from Austria
• Mecoptera in UK on BBC wildlife website (third image in)