Temporal range: Late Carboniferous-Recent
|Rhithrogena germanica, the fly fisherman's "March brown mayfly"|
Hyatt & Arms, 1891
Mayflies or shadflies are aquatic insects belonging to the order Ephemeroptera (from the Greek εφήμερος, ephemeros = "short-lived" (literally "lasting a day", cf. English "ephemeral"), and πτερόν, pteron = "wing", referring to the brief lifespan of adults). They are in an ancient group of insects termed the Palaeoptera, which also contains the dragonflies and damselflies.
Mayflies are relatively primitive insects and exhibit a number of ancestral traits that were probably present in the first flying insects, such as long tails and wings that do not fold flat over the abdomen. They are aquatic insects whose immature stage (called "naiad" or "nymph") lives in fresh water. They are unique among insects in having a fully winged terrestrial adult stage, the subimago, that moults into a sexually mature adult. Over 3,000 species in 42 families and over 400 genera are known worldwide, including about 630 species in North America.
Mayflies "hatch" (emerge as adults) in spring, not necessarily in May, in enormous numbers. Some hatches attract tourists. Fly fishermen make use of mayfly hatches by choosing artificial fishing flies that resemble the species in question. One of the most famous English mayflies is Rhithrogena germanica, the fisherman's "March brown mayfly".
Immature mayflies are aquatic, and are known as nymphs or naiads. They have an elongated, cylindrical or somewhat flattened body that passes through a number of instars, moulting and increasing in size each time. When ready to emerge from the water, nymphs vary in length, depending on species, from 3 to 30 mm (0.12 to 1.18 in). The head is heavily sclerotinised, often with various ridges and projections, and is either forward pointing or downward pointing, in each case with a terminal mouth. It has two large compound eyes, three ocelli (simple eyes) and a pair of slender antennae. The mouthparts are designed for chewing and consist of a flap-like labrum, a pair of strong mandibles, a pair of maxillae, a membranous hypopharynx and a labium.
The thorax consists of three segments, the hindermost two, the mesothorax and metathorax being fused. Each segment bears a pair of legs which usually terminate in a single claw, though in a few species this is missing. The legs are robust and often clad in bristles, hairs or spines. Wing pads develop on the mesothorax, and in some species, hind wing pads develop on the metathorax.
The abdomen consists of ten segments, some of which may be obscured by a large pair of operculate gills, a thoracic shield or the developing wingpads. In most taxa up to seven pairs of gills arise from the top or sides of the abdomen, but in some species they are under the abdomen, and in a very few species the gills are instead located on the coxae of the legs, or the bases of the maxillae. The abdomen terminates in a pair of, or three, slender thread-like projections.
The final moult of the naiad is not to the full adult form, but to a winged subimago that physically resembles the adult, but which is usually sexually immature and duller in colour. It often has partially cloudy wings that are fringed with minute hairs, and the eyes, legs and genitalia are not fully developed. The subimagos are generally poor fliers, and typically lack the colour patterns used to attract mates. After a period, usually lasting one or two days but in some species only a few minutes, the subimago moults to the full adult form, making mayflies the only insects where a winged form undergoes a further moult.
Adult mayflies, or imagos, are delicate-looking insects with one or two pairs of membranous, triangular wings, with extensive venation. At rest, these are held upright, like those of a butterfly. The hindwings are much smaller than the forewings, and may be vestigial or absent. The second segment of the thorax, which bears the forewings, is enlarged, holding the main flight muscles. Adults have short, flexible antennae, large compound eyes, three ocelli and non-functional mouthparts. In most species, the males' eyes are large and the front legs unusually long, for use in locating and grasping females during the mid-air mating. In some species, all the legs are functionless, apart from the front pair in males. The abdomen is long and roughly cylindrical, with ten segments and two or three long cerci (tail-like appendages) at the tip. Uniquely among insects, mayflies possess paired genitalia, with the male having two aedeagi (penis-like organs) and the female two gonopores.
Reproduction and life cycle
Mayflies are hemimetabolous (they have "incomplete metamorphosis"). They are unique among insects in that they moult one more time after acquiring functional wings; this last-but-one winged (alate) instar usually lives a very short time, often a matter of hours, and is known as a subimago, or to fly fishermen as a dun. 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 fall, mayflies are everywhere, dancing around each other in large groups, or resting on every available surface. In many species the emergence is synchronised with dawn or dusk, and light intensity seems to be an important cue for emergence, but other factors may also be involved. 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 lifespan of an adult mayfly is very short, varying with the species. Its primary function at this stage is reproduction and it does not feed; its mouthparts are vestigial, and its digestive system is filled with air. Dolania americana has the shortest lifespan of any mayfly: the adult females of the species live for less than five minutes.
Male adults may patrol individually, but most congregate in swarms a few metres (yards) above water (or the ground away from water) and perform a nuptial dance. Each insect has a characteristic up-and-down pattern of movement; strong wing beats propel it upwards and forwards with the body sloping down posteriorly; a cessation of wing movement causes it to fall passively under gravity 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 the pair may remain in tandem and flutter to the ground. Males may spend the night in vegetation and return to the nuptial dance the following day. Although they do not feed, some have been observed landing briefly on the surface of the water before flying off. They are believed to imbibe moisture through their mouth to prevent desiccation.
Females typically lay somewhere between four hundred and three thousand eggs. The eggs are dropped onto the surface of the water, or the female deposits them by dipping the tip of the abdomen into the water during flight or while standing next to the water. The eggs then sink to the bottom where they hatch. In a few species, the female submerges and deposits the eggs among plants or in crevices underwater. The incubation time is variable, depending at least in part on temperature, and may be anything from a few days to nearly a year. The larval growth rate is also temperature-dependent, as is the number of moults. 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, and there is a tendency in these circumstances, for the eggs and nymphs to get washed downstream. To counteract this, females may fly upriver before depositing their eggs in the water. For example, the female Tisza mayfly, the largest European species with a length of 10 cm (4 in), flies up to 3 kilometres (1.9 mi) upstream before depositing eggs on the water surface. 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. In other species, the nymph rises to the surface, bursts out of its skin, remains quiescent for a minute or two resting on the exuviae and then flies upwards, and in some, the nymph climbs out of the water before transforming.
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 along the shores of Lake Erie in 2003. In the nymphs of most mayfly species, the paddle-like gills do not function as respiratory surfaces because sufficient oxygen is absorbed through the integument, instead serving to create a respiratory current. 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. Some are able to shift from one feeding group to another as they grow, thus enabling them to utilise a variety of food resources. 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. Along with caddisfly larvae and gastropod molluscs, the grazing of mayfly nymphs has a significant impact on the primary producers, the plants and algae, on the bed of streams and rivers.
The nymphs are eaten by a wide range of predators and form an important part of the aquatic food chain. Fish are probably 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 larvae feed on bottom-dwelling mayfly nymphs, and amphibians also take their toll. Besides the direct mortality caused by these predators, the behaviour of their potential prey is also affected, with the nymphs' growth rate being slowed by the need to hide rather than feed. The nymphs are highly susceptible to pollution and can be useful in the biomonitoring of water bodies. Once they have emerged, large numbers are preyed on by birds, bats and by other insects.
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. Fish that feed on mayfly nymphs that have bioaccumulated heavy metals are themselves at risk.
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 1800s. Ephemera compar is known from a single specimen, collected from the "foothills of Colorado" in 1873, but despite intensive surveys of the Colorado mayflies reported in 1984, 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.
Taxonomy and phylogeny
Mayflies are an ancient group of winged (pterygote) insects. Putative fossil stem group representatives (e.g. Syntonopteroidea like Lithoneura lameerrei) are already known from the late Carboniferous. The largest mayfly of all times may have been Bojophlebia prokopi from the Upper Carboniferous of Moravia with a wing span of 45 cm (18 in).
From the Permian, numerous stem group representatives of mayflies are known, which are often lumped into a separate taxon Permoplectoptera (e.g. including Protereisma permianum in the Protereismatidae, and Misthodotidae). The larvae of Permoplectoptera still had 9 pairs of abdominal gills, and the adults still had long hind wings. 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). The modern Genus Neoephemera is represented in the fossil record by the Ypresian species N. antiqua from Washington State.
Grimaldi and Engel, reviewing the phylogeny in 2005, 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. The phylogeny of the Ephemeroptera was first studied using molecular analysis by Ogden and Whiting in 2005. They recovered the Baetidae as sister to the other clades. Mayfly phylogeny was studied using morphological and molecular analyses by Ogden and others in 2009. They found that the Asian genus Siphluriscus was sister to all other mayflies. 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 (1991), in Insects of Australia.
In human culture
In art and literature
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."[b]
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.
The Dutch Golden Age author Augerius Clutius (Outgert Cluyt) illustrated some mayflies in his 1634 De Hemerobio ("On the Mayfly"), the earliest book written on the group. Maarten de Vos similarly illustrated a mayfly in his 1587 depiction of the fifth day of creation, amongst an assortment of fish and water birds.
In 1495 Albrecht Dürer included a mayfly in his engraving The Holy Family with the Mayfly. The critics Larry Silver and Pamela H. Smith argue that the image provides "an explicit link between heaven and earth .. to suggest a cosmic resonance between sacred and profane, celestial and terrestrial, macrocosm and microcosm."
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. The English poet George Crabbe, known to have been interested in insects, compared the brief life of a newspaper with that of mayflies, both being known as "Ephemera", things that live for a day:
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.—George Crabbe, "The Newspaper", 1785
The theme of brief life is echoed in the artist Douglas Florian's 1998 poem, "The Mayfly". The American Poet Laureate Richard Wilbur's 2005 poem "Mayflies" includes the lines "I saw from unseen pools a mist of flies, In their quadrillions rise, And animate a ragged patch of glow, With sudden glittering".
In fly fishing
Mayflies are the primary source of models for artificial flies, hooks tied with coloured materials such as threads and feathers, used in fly fishing. These are based on different life-cycle stages of mayflies. 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. In 1983, Patrick McCafferty recorded that artificial flies had been based on 36 genera of North American mayfly, from a total of 63 western species and 103 eastern/central species. 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 1653 book The Compleat Angler; for example, he names the "Green-drake" for use as a natural fly, and "duns" (mayfly subimagos) as artificial flies. These include for example the "Great Dun" and the "Great Blue Dun" in February; the "Whitish Dun" in March; the "Whirling Dun" and the "Yellow Dun" in April; the "Green-drake", the "Little Yellow May-Fly" and the "Grey-Drake" in May; and the "Black-Blue Dun" in July. Nymph or "wet fly" fishing was restored to popularity on the chalk streams of England by G.E.M. Skues with his 1910 book Minor Tactics of the Chalk Stream . In the book, Skues discusses the use of duns to catch trout. The March brown is "probably the most famous of all British mayflies", having been copied by anglers to catch trout for over 500 years.
As a spectacle
The 2014 hatch of the large black-brown mayfly Hexagenia bilineata on the Mississippi River was imaged on weather radar; the swarm flew up to 2,500 feet high around La Crosse, Wisconsin, creating a radar signature that resembled a "significant rain storm" and covering cars and buildings with a "slimy mess".
As a name for vessels
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It has been advanced as an argument against the use of the wet fly, that duns and the other small insects which drift down upon the surface of a stream are never seen by the fish under water, and that a wet fly is therefore an unnatural object, especially if winged. "Never " is a big word, and I venture to think the case is overstated. I have watched an eddy with little swirling whirlpools in it for an hour together, and again and again I have seen little groups of flies caught in one or other of the whirls, sucked under and thrown scatterwise through the water
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|Wikispecies has information related to: Ephemeroptera|
|Wikisource has the text of the 1911 Encyclopædia Britannica article May-fly.|
|Wikimedia Commons has media related to Ephemeroptera.|
- Tree of Life info for Ephemeroptera
- Mayfly Central hosted by Purdue University
- Bibliography of Ephemeroptera