Mayflies at the subimago stage are a favourite food of many fish, and many fishing flies are modelled to resemble them. The subimago stage does not survive for long, rarely for more than 24 hours. In some species, it may last for just a few minutes, while the mayflies in the family Palingeniidae have sexually mature subimagos and no true adult form at all. Often, all the mayflies in a population mature at once a hatch , and for a day or two in the spring or autumn, mayflies are everywhere, dancing around each other in large groups, or resting on every available surface. Baetis intercalaris , for example, usually emerges just after sunset in July and August, but in one year, a large hatch was observed at midday in June.
The soft-bodied subimagos are very attractive to predators. Synchronous emergence is probably an adaptive strategy that reduces the individual's risk of being eaten. The primary function of the adult is reproduction; adults do not feed, and have only vestigial unusable mouthparts , while their digestive systems are filled with air. Male adults may patrol individually, but most congregate in swarms a few metres above water with clear open sky above it, and perform a nuptial courtship dance.
Each insect has a characteristic up-and-down pattern of movement; strong wingbeats propel it upwards and forwards with the tail sloping down; when it stops moving its wings, it falls passively with the abdomen tilted upwards. Females fly into these swarms, and mating takes place in the air. A rising male clasps the thorax of a female from below using his front legs bent upwards, and inseminates her.
Copulation may last just a few seconds, but occasionally a pair remains in tandem and flutters to the ground. Although they do not feed, some briefly touch the surface to drink a little water before flying off.
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Females typically lay between four hundred and three thousand eggs. The eggs are often dropped onto the surface of the water; sometimes the female deposits them by dipping the tip of her abdomen into the water during flight, releasing a small batch of eggs each time, or deposits them in bulk while standing next to the water.
In a few species, the female submerges and places the eggs among plants or in crevices underwater, but in general, they sink to the bottom. The incubation time is variable, depending at least in part on temperature, and may be anything from a few days to nearly a year. Eggs can go into a quiet dormant phase or diapause. At anywhere between ten and fifty, these post-embryonic moults are more numerous in mayflies than in most other insect orders.
The nymphal stage of mayflies may last from several months to several years, depending on species and environmental conditions. Many species breed in moving water, where there is a tendency for the eggs and nymphs to get washed downstream. To counteract this, females may fly upriver before depositing their eggs. These sink to the bottom and hatch after 45 days, the nymphs burrowing their way into the sediment where they spend two or three years before hatching into subimagos.
When ready to emerge, several different strategies are used. In some species, the transformation of the nymph occurs underwater and the subimago swims to the surface and launches itself into the air. Nymphs live primarily in streams under rocks, in decaying vegetation, or in sediments. Few species live in lakes, but they are among the most prolific. For example, the emergence of one species of Hexagenia was recorded on Doppler weather radar by the shoreline of Lake Erie in However, in low-oxygen environments such as the mud at the bottom of ponds in which Ephemera vulgata burrows, the filamentous gills act as true accessory respiratory organs and are used in gaseous exchange.
In most species, the nymphs are herbivores or detritivores , feeding on algae , diatoms or detritus , but in a few species, they are predators of chironomid and other small insect larvae and nymphs. They process a great quantity of organic matter as nymphs and transfer a lot of phosphates and nitrates to terrestrial environments when they emerge from the water, thus helping to remove pollutants from aqueous systems. The nymphs are eaten by a wide range of predators and form an important part of the aquatic food chain. Fish are among the main predators, picking nymphs off the bottom or ingesting them in the water column, and feeding on emerging nymphs and adults on the water surface.
Carnivorous stonefly , caddisfly , alderfly and dragonfly larvae feed on bottom-dwelling mayfly nymphs, as do aquatic beetles, leeches, crayfish and amphibians. Mayfly nymphs may serve as hosts for parasites such as nematodes and trematodes. Some of these affect the nymphs' behaviour in such a way that they become more likely to be predated. Mayflies are involved in both primary production and bioturbation. A study in laboratory simulated streams revealed that the mayfly genus Centroptilum increased the export of periphyton ,  thus indirectly affecting primary production positively, which is an essential process for ecosystems.
The mayfly can also reallocate and alter the nutrient availability in aquatic habitats through the process of bioturbation. By burrowing in the bottom of lakes and redistributing nutrients, mayflies indirectly regulate phytoplankton and epibenthic primary production. This motion creates current that carries food particles through the burrow and allows the nymph to filter feed. Other mayfly nymphs possess elaborate filter feeding mechanisms like that of the genus Isonychia.
The nymph have forelegs that contain long bristle-like structures that have two rows of hairs. Interlocking hairs form the filter by which the insect traps food particles. The action of filter feeding has a small impact on water purification but an even larger impact on the convergence of small particulate matter into matter of a more complex form that goes on to benefit consumers later in the food chain.
Mayflies are distributed all over the world in clean freshwater habitats,  though absent from Antarctica. Female mayflies may be dispersed by wind, and eggs may be transferred by adhesion to the legs of waterbirds. Some thirteen families are restricted to a single bioregion. The nymph is the dominant life history stage of the mayfly. Different insect species vary in their tolerance to water pollution, but in general, the larval stages of mayflies, stoneflies Plecoptera and caddis flies Trichoptera are susceptible to a number of pollutants including sewage , pesticides and industrial effluent.
In general, mayflies are particularly sensitive to acidification , but tolerances vary, and certain species are exceptionally tolerant to heavy metal contamination and to low pH levels. Ephemerellidae are among the most tolerant groups and Siphlonuridae and Caenidae the least. The adverse effects on the insects of pollution may be either lethal or sub-lethal, in the latter case resulting in altered enzyme function, poor growth, changed behaviour or lack of reproductive success.
As important parts of the food chain, pollution can cause knock-on effects to other organisms; a dearth of herbivorous nymphs can cause overgrowth of algae, and a scarcity of predacious nymphs can result in an over-abundance of their prey species. They are easily fooled by other polished surfaces which can act as traps for swarming mayflies.
The threat to mayflies applies also to their eggs. The major pollutants thought to be responsible are fine sediment and phosphate from agriculture and sewage. The status of many species of mayflies is unknown because they are known from only the original collection data. Four North American species are believed to be extinct. Among these, Pentagenia robusta was originally collected from the Ohio River near Cincinnati , but this species has not been seen since its original collection in the s. Ephemera compar is known from a single specimen, collected from the "foothills of Colorado" in , but despite intensive surveys of the Colorado mayflies reported in , it has not been rediscovered.
The International Union for Conservation of Nature IUCN red list of threatened species includes one mayfly: Tasmanophlebi lacuscoerulei , the large blue lake mayfly, which is a native of Australia and is listed as endangered because its alpine habitat is vulnerable to climate change. Over 3, species of mayfly in 42 families and over genera are known worldwide,  including about species in North America. Putative fossil stem group representatives e. Syntonopteroidea-like Lithoneura lameerrei are already known from the late Carboniferous.
The English common name is for the insect's emergence in or around the month of May in the UK. From the Permian , numerous stem group representatives of mayflies are known, which are often lumped into a separate taxon Permoplectoptera e. The larvae of Permoplectoptera still had 9 pairs of abdominal gills, and the adults still had long hindwings. Maybe the fossil family Cretereismatidae from the Lower Cretaceous Crato Formation of Brazil also belongs as the last offshoot to Permoplectoptera.
The Crato outcrops otherwise yielded fossil specimens of modern mayfly families or the extinct but modern family Hexagenitidae. However, from the same locality the strange larvae and adults of the extinct family Mickoleitiidae order Coxoplectoptera have been described,  which represents the fossil sister group of modern mayflies, even though they had very peculiar adaptations such as raptorial forelegs.
The oldest mayfly inclusion in amber is Cretoneta zherichini Leptophlebiidae from the Lower Cretaceous of Siberia. In the much younger Baltic amber numerous inclusions of several modern families of mayflies have been found Ephemeridae, Potamanthidae, Leptophlebiidae, Ametropodidae, Siphlonuridae, Isonychiidae, Heptageniidae, and Ephemerellidae. Grimaldi and Engel, reviewing the phylogeny in , commented that many cladistic studies had been made with no stability in Ephemeroptera suborders and infraorders; the traditional division into Schistonota and Pannota was wrong because Pannota is derived from the Schistonota.
They recovered the Baetidae as sister to the other clades. They found that the Asian genus Siphluriscus was sister to all other mayflies.
Like a Fish Out of Water
Some existing lineages such as Ephemeroidea , and families such as Ameletopsidae, were found not to be monophyletic , through convergence among nymphal features. The following traditional classification is based on Peters and Campbell , in Insects of Australia. Bloodless and many footed animals, whether furnished with wings or feet, move with more than four points of motion; as, for instance, the dayfly ephemeron moves with four feet and four wings: and, I may observe in passing, this creature is exceptional not only in regard to the duration of its existence, whence it receives its name, but also because though a quadruped it has wings also.
The River Bug on the Black Sea at midsummer brings down some thin membranes that look like berries out of which burst a four-legged caterpillar in the manner of the creature mentioned above, but it does not live beyond one day, owing to which it is called the hemerobius. Maerten de Vos similarly illustrated a mayfly in his depiction of the fifth day of creation, amongst an assortment of fish and water birds.
Smith argue that the image provides "an explicit link between heaven and earth.. Myriads of May-flies appear for the first time on the Alresford stream. The air was crowded with them, and the surface of the water covered. Large trouts sucked them in as they lay struggling on the surface of the stream, unable to rise till their wings were dried Their motions are very peculiar, up and down for so many yards almost in a perpendicular line.
The mayfly has come to symbolise the transitoriness and brevity of life. In shoals the hours their constant numbers bring Like insects waking to th' advancing spring; Which take their rise from grubs obscene that lie In shallow pools, or thence ascend the sky: Such are these base ephemeras, so born To die before the next revolving morn. The theme of brief life is echoed in the artist Douglas Florian's poem, "The Mayfly". Another literary reference to mayflies is seen in The Epic of Gilgamesh , one of the earliest surviving great works of literature.
The briefness of Gilgamesh's life is compared to that of the adult mayfly. The American playwright David Ives wrote a short comedic play, Time Flies , in , as to what two mayflies might discuss during their one day of existence. Mayflies are the primary source of models for artificial flies, hooks tied with coloured materials such as threads and feathers, used in fly fishing.
For example, the flies known as "emergers" in North America are designed by fly fishermen to resemble subimago mayflies, and are intended to lure freshwater trout. A large number of these species have common names among fly fishermen, who need to develop a substantial knowledge of mayfly "habitat, distribution, seasonality, morphology and behavior" in order to match precisely the look and movements of the insects that the local trout are expecting. Izaak Walton describes the use of mayflies for catching trout in his book The Compleat Angler ; for example, he names the "Green-drake" for use as a natural fly, and "duns" mayfly subimagos as artificial flies.
Skues with his book Minor Tactics of the Chalk Stream. In the book, Skues discusses the use of duns to catch trout. The hatch of the giant mayfly Palingenia longicauda on the Tisza and Maros Rivers in Hungary and Serbia , known as "Tisza blooming", is a tourist attraction. During the weekend of June 13—14, , a large swarm of mayflies caused several vehicular accidents on the Columbia—Wrightsville Bridge , carrying Pennsylvania Route across the Susquehanna River between Columbia and Wrightsville, Pennsylvania. The bridge had to be closed to traffic twice during that period due to impaired visibility and obstructions posed by piles of dead insects.
Mayflies are consumed in several cultures and are estimated to contain the most raw protein content of any edible insect by dry weight. In Malawi , kungu, a paste of mayflies Caenis kungu and mosquitoes is made into a cake for eating. Adult mayflies are collected and eaten in many parts of China and Japan. Near Lake Victoria , Povilla mayflies are collected, dried and preserved for use in food preparations.
Two vessels of the Royal Navy were named HMS Mayfly : a torpedo boat launched in January ,  and a Fly -class river gunboat constructed in sections at Yarrow in The Seddon Mayfly , which was constructed in , was an aircraft that was unsuccessful in early flight.
The first aircraft designed by a woman, Lillian Bland , was titled the Bland Mayfly. In pres France, "chute de manne" was obtained by pressing mayflies into cakes and using them as bird food and fishbait. Their exoskeleton contains chitin , which has applications in these industries. Mayfly larvae do not survive in polluted aquatic habitats and, thus, have been chosen as bioindicators, markers of water quality in ecological assessments. In marketing, Nike produced a line of running shoes in titled "Mayfly".
The shoes were designed with a wing venation pattern like the mayfly and were also said to have a finite lifetime. From Wikipedia, the free encyclopedia. For other uses, see Mayfly disambiguation. Aquatic insects of the order Ephemeroptera.
Aristotle also describes the mayfly in History of Animals , b. Introduction to Insect Biology and Diversity 2nd ed. Oxford University Press. Retrieved 22 May The Language of Fly-Fishing. Patrick Pensoft Publishers. Systematic Biology. Aquatic Entomology. Eric Advances in Ephemeroptera Biology. Book of Insect Records. University of Florida. Archived from the original on The nuptial flight".
Journal of the New York Entomological Society. Quaestiones Entomologicae. Cambridge University Press.
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The Mayflies of Florida. University Press of Florida. Pennsylvania Sea Grant. Archived from the original PDF on 27 September Retrieved 30 May Journal of Experimental Biology. Aquatic Insects. In Thorp; Rogers eds. Thorp and Covich's Freshwater Invertebrates 4th ed. Academic Press. Richard; Lamberti, Gary A. Methods in Stream Ecology. Christopher 6 September Poulin, R. Ephemeroptera ". Journal of Parasitology. Journal of the North American Benthological Society.
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