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Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) are
aquatic insect Aquatic insects or water insects live some portion of their life cycle in the water. They feed in the same ways as other insects. Some ''diving'' insects, such as predatory diving beetles, can hunt for food underwater where land-living insects c ...
s belonging to the
order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
Ephemeroptera. This order is part of an ancient group of insects termed the Palaeoptera, which also contains dragonflies and
damselflies Damselflies are flying insects of the suborder Zygoptera in the order Odonata. They are similar to dragonflies, which constitute the other odonatan suborder, Anisoptera, but are smaller and have slimmer bodies. Most species fold the wings along ...
. Over 3,000 species of mayfly are known worldwide, grouped into over 400
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
in 42 families. Mayflies have 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. Their immature stages are aquatic
fresh water Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does include ...
forms (called "naiads" or " nymphs"), whose presence indicates a clean, unpolluted and highly oxygenated aquatic environment. They are unique among insect orders in having a fully winged terrestrial preadult stage, the subimago, which moults into a sexually mature adult, the imago. Mayflies "hatch" (emerge as adults) from spring to autumn, not necessarily in May, in enormous numbers. Some hatches attract tourists.
Fly fishermen Fly fishing is an angling method that uses a light-weight lure—called an artificial fly—to catch fish. The fly is cast using a fly rod, reel, and specialized weighted line. The light weight requires casting techniques significantly diff ...
make use of mayfly hatches by choosing artificial fishing flies that resemble them. One of the most famous English mayflies is '' Rhithrogena germanica'', the fisherman's "March brown mayfly". The brief lives of mayfly adults have been noted by naturalists and encyclopaedists since Aristotle and Pliny the Elder in classical antiquity. The German engraver
Albrecht Dürer Albrecht Dürer (; ; hu, Ajtósi Adalbert; 21 May 1471 – 6 April 1528),Müller, Peter O. (1993) ''Substantiv-Derivation in Den Schriften Albrecht Dürers'', Walter de Gruyter. . sometimes spelled in English as Durer (without an umlaut) or Due ...
included a mayfly in his 1495 engraving '' The Holy Family with the Mayfly'' to suggest a link between heaven and earth. The English poet
George Crabbe George Crabbe ( ; 24 December 1754 – 3 February 1832) was an English poet, surgeon and clergyman. He is best known for his early use of the realistic narrative form and his descriptions of middle and working-class life and people. In the 177 ...
compared the brief life of a daily newspaper with that of a mayfly in the satirical poem "The Newspaper" (1785), both being known as "ephemera".


Description


Nymph

Immature mayflies are aquatic and are referred to as nymphs or naiads. In contrast to their short lives as adults, they may live for several years in the water. They have an elongated, cylindrical or somewhat flattened body that passes through a number of instars (stages), molting and increasing in size each time. When ready to emerge from the water, nymphs vary in length, depending on species, from . The
head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mouth, each of which aid in various sensory functions such as sight, hearing, smell, and taste. Some very simple animals may ...
has a tough outer covering of sclerotin, often with various hard ridges and projections; it points either forwards or downwards, with the mouth at the front. There are two large compound eyes, three ocelli (simple eyes) and a pair of antennae of variable lengths, set between or in front of the eyes. The mouthparts are designed for chewing and consist of a flap-like labrum, a pair of strong mandibles, a pair of
maxillae The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The t ...
, a membranous hypopharynx and a labium. The thorax consists of three segments – the hindmost two, the mesothorax and metathorax, being fused. Each segment bears a pair of legs which usually terminate in a single claw. The legs are robust and often clad in bristles, hairs or spines. Wing pads develop on the mesothorax, and in some species, hindwing 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 (expanded part of the prothorax) or the developing wing pads. 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 slender thread-like projections, consisting of a pair of cerci, with or without a third central
caudal filament Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insec ...
.


Subimago

The final moult of the nymph is not to the full adult form, but to a winged stage called a subimago that physically resembles the adult, but which is usually sexually immature and duller in colour. The subimago, or dun, often has partially cloudy wings fringed with minute hairs known as microtrichia; its eyes, legs and genitalia are not fully developed. Females of some mayflies (subfamily Palingeniinae) do not moult from a subimago state into an adult stage and are sexually mature while appearing like a subimago with microtrichia on the wing membrance. Oligoneuriine mayflies form another exception in retaining microtrichia on their wings but not on their bodies. Subimagos are generally poor fliers, have shorter appendages, and typically lack the colour patterns used to attract mates. In males of ''
Ephoron leukon ''Ephoron leukon'', the white fly, is a species of pale burrower mayfly in the family Polymitarcyidae Polymitarcyidae is a family of pale burrower mayflies in the order Ephemeroptera. There are about 10 genera and more than 90 described speci ...
'', the subimagos have forelegs that are short and compressed, with accordion like folds, and expands to more than double its length after moulting. 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.


Imago

Adult mayflies, or imagos, are relatively primitive in structure, exhibiting traits that were probably present in the first flying insects. These include long tails and wings that do not fold flat over the abdomen. Mayflies are delicate-looking insects with one or two pairs of membranous, triangular wings, which are extensively covered with
veins Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated b ...
. At rest, the wings are held upright, like those of a butterfly. The hind wings are much smaller than the forewings and may be vestigial or absent. The second segment of the thorax, which bears the forewings, is enlarged to hold 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 the males of some families, there are two large cylindrical "turban" eyes (also known as ''turbanate'' or ''turbinate'' eyes) that face upwards in addition to the lateral eyes. They are capable of detecting ultraviolet light and are thought to be used during courtship to detect females flying above them. 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. Like Entognatha, Archaeognatha and Zygentoma, the spiracles on the abdomen don't have closing muscles. Uniquely among insects, mayflies possess paired genitalia, with the male having two
aedeagi An aedeagus (plural aedeagi) is a reproductive organ of male arthropods through which they secrete sperm from the testes during copulation (zoology), copulation with a female. It can be thought of as the insect equivalent of a mammal's penis, t ...
(penis-like organs) and the female two gonopores (sexual openings).


Biology


Reproduction and life cycle

Mayflies are hemimetabolous (they have "incomplete
metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...
"). 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 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 An artificial fly or fly lure is a type of fishing lure, usually used in the sport of fly fishing (although they may also be used in other forms of angling). In general, artificial flies are an imitation of aquatic insects that are natural food ...
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 Palingeniidae is a family of mayflies, members of which are known as spiny-headed burrowing mayflies. These are generally quite large mayflies with more than four longitudinal cross-veins on their wings. Males have short, wide pronotums and the ...
have sexually mature subimagos and no true adult form at all. Often, all the individuals in a population mature at once (a hatch), and for a day or two in the spring or autumn, mayflies are extremely abundant, 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 ''Baetis intercalaris'' is a species of small minnow mayfly in the family Baetidae Baetidae is a family of mayfly, mayflies with about 1000 described species in 110 genera distributed worldwide. These are among the smallest of mayflies, adult ...
'', 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. The primary function of the adult is reproduction; adults do not feed and have only vestigial
mouthparts Mouthparts may refer to: * The parts of a mouth ** Arthropod mouthparts *** Insect mouthparts {{disambig ...
, while their digestive systems are filled with air. ''
Dolania americana ''Dolania'' is a monotypic genus of mayfly in the family Behningiidae containing the single species ''Dolania americana''. It is found in the southeastern United States, as far south as Florida, and is generally uncommon. The adult insects emer ...
'' has the shortest adult 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 above water with clear open sky above it, and perform a nuptial or 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. Males may spend the night in vegetation and return to their dance the following day. Although they do not feed, some briefly touch the surface to drink a little water before flying off. 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. 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. Around half of all mayfly species whose reproductive biology has been described are parthenogenetic (able to asexually reproduce), including both partially and exclusively parthogenetic populations and species. 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. For example, the female Tisza mayfly, the largest European species with a length of , flies up to upstream before depositing eggs on the water surface. These sink to the bottom and hatch after 45 days, the nymphs
burrow An Eastern chipmunk at the entrance of its burrow A burrow is a hole or tunnel excavated into the ground by an animal to construct a space suitable for habitation or temporary refuge, or as a byproduct of locomotion. Burrows provide a form of sh ...
ing 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
exuvia In biology, exuviae are the remains of an exoskeleton and related structures that are left after ecdysozoans (including insects, crustaceans and arachnids) have moulted. The exuviae of an animal can be important to biologists as they can often b ...
e (cast skin) and then flies upwards, and in some, the nymph climbs out of the water before transforming.


Ecology

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 Weather radar, also called weather surveillance radar (WSR) and Doppler weather radar, is a type of radar used to locate precipitation (meteorology), precipitation, calculate its motion, and estimate its type (rain, snow, hail etc.). Modern wea ...
by the shoreline 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 ''Ephemera vulgata'' is a species of mayfly in the genus ''Ephemera''. This mayfly breeds in stationary water in slow rivers and in ponds, the nymphs developing in the mud. Description ''Ephemera vulgata'' can be told in both adult and subimago ...
'' burrows, the filamentous gills act as true accessory respiratory organs and are used in gaseous exchange. In most species, the nymphs are
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...
s or
detritivore Detritivores (also known as detrivores, detritophages, detritus feeders, or detritus eaters) are heterotrophs that obtain nutrients by consuming detritus (decomposing plant and animal parts as well as feces). There are many kinds of invertebrates, ...
s, feeding on
algae Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
,
diatom A diatom (Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising sev ...
s or
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts commun ...
, but in a few species, they are
predators Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
of chironomid and other small insect larvae and nymphs. Nymphs of ''Povilla'' burrow into submerged wood and can be a problem for boat owners in Asia. 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 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 Plecoptera is an order of insects, commonly known as stoneflies. Some 3,500 species are described worldwide, with new species still being discovered. Stoneflies are found worldwide, except Antarctica. Stoneflies are believed to be one of the mo ...
, caddisfly, alderfly and
dragonfly A dragonfly is a flying insect belonging to the infraorder Anisoptera below the order Odonata. About 3,000 extant species of true dragonfly are known. Most are tropical, with fewer species in temperate regions. Loss of wetland habitat threate ...
larvae feed on bottom-dwelling mayfly nymphs, as do aquatic beetles, leeches, crayfish and
amphibian Amphibians are tetrapod, four-limbed and ectothermic vertebrates of the Class (biology), class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terres ...
s. 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, such as Rhamphomyia longicauda. Mayfly nymphs may serve as
hosts A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places *Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ( ...
for parasites such as
nematode The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-Parasitism, parasitic nematodes also known as eelworms. They are a diverse animal phylum inhab ...
s and trematodes. Some of these affect the nymphs' behaviour in such a way that they become more likely to be predated. Other nematodes turn adult male mayflies into quasi-females which haunt the edges of streams, enabling the parasites to break their way out into the aqueous environment they need to complete their life cycles. The nymphs can also serve as
intermediate hosts In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...
for the horsehair worm ''
Paragordius varius ''Paragordius varius'' is a parasite species in the horsehair worm group (Nematomorpha). They cycle between terrestrial and aquatic habitats and are most commonly known for their ability to manipulate their definitive host to jump into a pool of ...
'', which causes its definitive host, a
grasshopper Grasshoppers are a group of insects belonging to the suborder Caelifera. They are among what is possibly the most ancient living group of chewing herbivorous insects, dating back to the early Triassic around 250 million years ago. Grasshopp ...
, to jump into water and drown.


Effects on ecosystem functioning

Mayflies are involved in both primary production and
bioturbation Bioturbation is defined as the reworking of soils and sediments by animals or plants. It includes burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a pr ...
. A study in laboratory simulated streams revealed that the mayfly genus ''
Centroptilum ''Centroptilum'' is a genus of mayflies of the family Baetidae Baetidae is a family of mayflies with about 1000 described species in 110 genera distributed worldwide. These are among the smallest of mayflies, adults rarely exceeding 10 mm ...
'' 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. Once burrowing to the bottom of the lake, mayfly nymphs begin to billow their respiratory gills. 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.


Distribution

Mayflies are distributed all over the world in clean freshwater habitats, though absent from Antarctica. They tend to be absent from oceanic islands or represented by one or two species that have dispersed from nearby mainland. Female mayflies may be dispersed by wind, and eggs may be transferred by adhesion to the legs of waterbirds. The greatest generic diversity is found in the Neotropical realm, while the Holarctic has a smaller number of genera but a high degree of speciation. Some thirteen families are restricted to a single bioregion. The main families have some general habitat preferences: the Baetidae favour warm water; the Heptageniidae live under stones and prefer fast-flowing water; and the relatively large Ephemeridae make burrows in sandy lake or river beds.


Conservation

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 Sewage (or domestic sewage, domestic wastewater, municipal wastewater) is a type of wastewater that is produced by a community of people. It is typically transported through a sewer system. Sewage consists of wastewater discharged from residenc ...
,
pesticide Pesticides are substances that are meant to control pests. This includes herbicide, insecticide, nematicide, molluscicide, piscicide, avicide, rodenticide, bactericide, insect repellent, animal repellent, microbicide, fungicide, and lampri ...
s and
industrial effluent Industrial wastewater treatment describes the processes used for treating wastewater that is produced by industries as an undesirable by-product. After treatment, the treated industrial wastewater (or effluent) may be reused or released to a sa ...
. In general, mayflies are particularly sensitive to
acidification Acidification may refer to: * Ocean acidification, decrease in the pH of the Earth's oceans * Freshwater acidification, atmospheric depositions and soil leaching of SOx and NOx * Soil acidification, buildup of hydrogen cations, which reduces the ...
, but tolerances vary, and certain species are exceptionally tolerant to heavy metal contamination and to low pH levels.
Ephemerellidae Ephemerellidae are known as the spiny crawler mayflies. They are a family of the order Ephemeroptera. There are eight genera consisting of a total 90 species (Merritt & Cummins). They are distributed throughout North America as well as the UK. T ...
are among the most tolerant groups and
Siphlonuridae Siphlonuridae, also known as the primitive minnow mayfly is a family of insects belonging to the order Ephemeroptera. Taxonomy The family is divided into the following genera: * Genus: '' Edmundsius'' * Genus: '' Parameletus'' * Genus: '' Sip ...
and
Caenidae Caenidae, or the small squaregill mayflies, is a family of insects consisting of 5 genera comprising 26 individual species. They are found throughout the world in lotic, depositional environments, and they are sprawlers. Caenids occur in quiet an ...
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. Adult female mayflies find water by detecting the
polarization Polarization or polarisation may refer to: Mathematics *Polarization of an Abelian variety, in the mathematics of complex manifolds *Polarization of an algebraic form, a technique for expressing a homogeneous polynomial in a simpler fashion by ...
of reflected light. 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. "Modest levels" of pollution in rivers in England are sufficient to kill 80% of mayfly eggs, which are as vulnerable to pollutants as other life-cycle stages; numbers of the
blue-winged olive mayfly Blue-winged Olive flies is a collective term used by anglers in fly fishing to identify a broad array of mayflies having olive, olive-brown bodies and bluish wings in their adult form. Sometimes referred to as BWO , a wide array of artificial fl ...
(''Baetis'') have fallen dramatically, almost to none in some rivers. 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 ''Pentagenia robusta'', the robust burrowing mayfly, is a recently extinct species of mayfly in the family Ephemeridae. It was endemic to the United States, found in the states Indiana, Kentucky, and Ohio Ohio () is a state in the Midwester ...
'' was originally collected from the
Ohio River The Ohio River is a long river in the United States. It is located at the boundary of the Midwestern and Southern United States, flowing southwesterly from western Pennsylvania to its mouth on the Mississippi River at the southern tip of Illino ...
near Cincinnati, but this species has not been seen since its original collection in the 1800s. ''
Ephemera compar Most likely, ''Ephemera compar'' is an extinct species of burrowing mayfly in the family Ephemeridae. It was found in North America. Ephemera compar is known from a single specimen, collected from the "foothills of Colorado" in 1873, but despit ...
'' 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 ''Tasmanophlebia lacuscoerulei'' is a species of mayfly in the family Siphlonuridae. It is endemic to New South Wales in Australia. It is known commonly as the large Blue Lake mayfly. This mayfly has a limited distribution in an area of about 80 ...
'', the large blue lake mayfly, which is a native of
Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...
and is listed as endangered because its alpine habitat is vulnerable to climate change.


Taxonomy and phylogeny

As of 2012, over 3,000 species of mayfly in 42 families and over 400 genera are known worldwide, including about 630 species in
North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Car ...
. 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 Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...
. The name Ephemeroptera is from the Greek ἐφήμερος, ' "short-lived" (literally "lasting a day", cf. English " ephemeral"), and πτερόν, ', " wing", referring to the brief lifespan of adults. The English common name is for the insect's emergence in or around the month of May in the UK. The name shadfly is from the Atlantic fish the
shad The Alosinae, or the shads,Alosinae
, which runs up American East Coast rivers at the same time as many mayflies emerge. 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 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 Coxoplectoptera or "chimera wings" is an extinct Order (biology), order of stem-group Mayfly, mayflies containing one family, Mickoleitiidae. Two adult and more than 20 nymphal fossils of ''Mickoleitia'' have been scientifically described from Mes ...
(order
Coxoplectoptera Coxoplectoptera or "chimera wings" is an extinct order of stem-group mayflies containing one family, Mickoleitiidae. Two adult and more than 20 nymphal fossils of ''Mickoleitia'' have been scientifically described from Mesozoic outcrops, mainly f ...
) 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 ''Neoephemera'' is a genus of large squaregill mayflies in the family Neoephemeridae first described by McDunnough (1925). and containing approximately six described species in ''Neoephemera''. Species These six species belong to the genus ''N ...
'' 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 further 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 ''Ephemeroidea'' is a superfamily of mayflies. Members of this superfamily are found in most parts of the world with the exception of the Arctic, the Antarctic and Australia. The following families are recognised: * Behningiidae *Ephemeridae ...
, and families such as Ameletopsidae, were found not to be
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...
, through convergence among nymphal features. The following traditional classification is based on Peters and Campbell (1991), in ''Insects of Australia''. Suborder
Pannota Pannota is a suborder of mayflies. One of the differences between this suborder and its sister group Schistonota concerns the degree of fusion of the wing pads in the final-stage nymph; in Schistonota, the degree of fusion along the mesothorax ...
*Superfamily
Ephemerelloidea ''Ephemerelloidea'' is a superfamily of mayflies Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the Unite ...
**
Ephemerellidae Ephemerellidae are known as the spiny crawler mayflies. They are a family of the order Ephemeroptera. There are eight genera consisting of a total 90 species (Merritt & Cummins). They are distributed throughout North America as well as the UK. T ...
**
Leptohyphidae Leptohyphidae is a family of mayfly, mayflies with some 140 described species in 12 genera. References

* Mayflies Insect families {{Mayfly-stub ...
**
Tricorythidae Tricorythidae is a family of mayflies in the order Ephemeroptera. There are about six genera and at least 40 described species in Tricorythidae. Genera These six genera belong to the family Tricorythidae: * '' Madecassorythus'' Elouard & Oliari ...
*Superfamily
Caenoidea Caenoidea is a superfamily of mayflies in the suborder Pannota. Members of this superfamily can be distinguished from those of Ephemerelloidea by the fact that the gills of the nymphs are filamentous. The following families are recognised: * ...
**
Neoephemeridae Neoephemeridae is a family of large squaregill mayflies in the order Ephemeroptera. There are at least four genera and about 17 described species in Neoephemeridae. Genera These four genera belong to the family Neoephemeridae: * '' Leucorhoenan ...
**
Baetiscidae Baetiscidae is a family of mayflies. It contains a single extant genus, '' Baetisca'', native to North America with around 12 species. The family is noted for their spined armoured larvae, which live in flowing water pools and on the edges of str ...
**
Caenidae Caenidae, or the small squaregill mayflies, is a family of insects consisting of 5 genera comprising 26 individual species. They are found throughout the world in lotic, depositional environments, and they are sprawlers. Caenids occur in quiet an ...
**
Prosopistomatidae Prosopistomatidae is a family of mayflies. There is one extant genus, '' Prosopistoma,'' with several dozen species found across Afro-Eurasia and Oceania. They are noted for their unusual beetle-shaped larvae, which live beneath rocks and stones ...
Suborder
Schistonota Schistonota is a suborder of mayflies. One of the differences between this suborder and its sister group, Pannota, concerns the degree of fusion of the wing pads in the final-stage nymph; in Schistonota, the degree of fusion along the mesothora ...
*Superfamily
Baetoidea Baetoidea is a superfamily of mayflies, which probably includes the most primitive living species. The following families are recognised: * Ameletopsidae * Ametropodidae *Baetidae Baetidae is a family of mayflies with about 1000 described ...
**
Siphlonuridae Siphlonuridae, also known as the primitive minnow mayfly is a family of insects belonging to the order Ephemeroptera. Taxonomy The family is divided into the following genera: * Genus: '' Edmundsius'' * Genus: '' Parameletus'' * Genus: '' Sip ...
** Baetidae ** Oniscigastridae ** Ameletopsidae **
Ametropodidae Ametropodidae is a family of mayflies in the order Ephemeroptera. There are at least three genera and three described species in Ametropodidae. Genera These three genera belong to the family Ametropodidae: * '' Ametropus'' Albarda, 1878 * '' Br ...
*Superfamily
Heptagenioidea ''Heptagenioidea'' is a superfamily of mayflies. Members of this superfamily are found in most parts of the world apart from the Arctic and Antarctic, with Heptageniidae being the most widely distributed family. Heptagenioidea probably originat ...
**
Coloburiscidae Coloburiscidae is a family of mayflies in the order Ephemeroptera, with at least seven species in three genera native to Australia, New Zealand, and Chile. Genera These three genera belong to the family Coloburiscidae: * '' Coloburiscoides'' Le ...
**
Oligoneuriidae Oligoneuriidae is a family of mayflies with a pantropical distribution. They are also known as brushlegged mayflies due to the presence of two rows of setae used for filtration on the front legs of their nymphs. Nymphs also have tufts of gills at ...
** Isonychiidae **
Heptageniidae The Heptageniidae (synonym: Ecdyonuridae) are a family of mayflies with over 500 described species mainly distributed in the Holarctic, Oriental, and Afrotropical regions, and also present in the Central American Tropics and extreme northern Sou ...
*Superfamily
Leptophlebioidea Leptophlebiidae is a family belonging to the Ephemeropterans that are commonly known as the prong-gilled mayflies or leptophlebiids. It is the only family in the superfamily Leptophlebioidea. There are around 131 genera and 640 described specie ...
**
Leptophlebiidae Leptophlebiidae is a family belonging to the Ephemeropterans that are commonly known as the prong-gilled mayflies or leptophlebiids. It is the only family in the superfamily Leptophlebioidea. There are around 131 genera and 640 described specie ...
*Superfamily
Ephemeroidea ''Ephemeroidea'' is a superfamily of mayflies. Members of this superfamily are found in most parts of the world with the exception of the Arctic, the Antarctic and Australia. The following families are recognised: * Behningiidae *Ephemeridae ...
**
Behningiidae Behningiidae is a family of mayflies. It is a primitive family; the nymphs burrow in the sediment but lack tusks on their mandibles, and the forelegs are not modified for burrowing. The gills are ventral, and the ones on the first abdominal segme ...
**
Potamanthidae Potamanthidae is a family of mayflies Mayflies (also known as shadflies or fishflies in Canada and the upper Midwestern United States, as Canadian soldiers in the American Great Lakes region, and as up-winged flies in the United Kingdom) a ...
**
Euthyplociidae Euthyplociidae is a family of mayflies in the order Ephemeroptera. There are about 7 genera and more than 20 described species in Euthyplociidae. Genera These seven genera belong to the family Euthyplociidae: * '' Afroplocia'' Lestage, 1939 * ' ...
**
Polymitarcyidae Polymitarcyidae is a family of pale burrower mayflies in the order Ephemeroptera. There are about 10 genera and more than 90 described species in Polymitarcyidae. Genera These 10 genera belong to the family Polymitarcyidae: * '' Asthenopus'' Ea ...
**
Ephemeridae Ephemeridae is a family of mayflies with about 150 described species found throughout the world except Australia and Oceania. These are generally quite large mayflies (up to 35 mm) with either two or three very long tails. Many species have ...
**
Palingeniidae Palingeniidae is a family of mayflies, members of which are known as spiny-headed burrowing mayflies. These are generally quite large mayflies with more than four longitudinal cross-veins on their wings. Males have short, wide pronotums and the ...


Phylogeny

After


In human culture


In art

The
Dutch Golden Age The Dutch Golden Age ( nl, Gouden Eeuw ) was a period in the history of the Netherlands, roughly spanning the era from 1588 (the birth of the Dutch Republic) to 1672 (the Rampjaar, "Disaster Year"), in which Dutch trade, science, and Dutch art, ...
author Augerius Clutius ( Outgert Cluyt) illustrated some mayflies in his 1634 ''De Hemerobio'' ("On the Mayfly"), the earliest book written on the group. Maerten 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 Albrecht Dürer (; ; hu, Ajtósi Adalbert; 21 May 1471 – 6 April 1528),Müller, Peter O. (1993) ''Substantiv-Derivation in Den Schriften Albrecht Dürers'', Walter de Gruyter. . sometimes spelled in English as Durer (without an umlaut) or Due ...
included a mayfly in his engraving '' The Holy Family with the Mayfly''. The critics
Larry Silver Larry Silver (born 1934) is an American photographer. He was born in the Bronx. While a student at the High School of Industrial Art in Manhattan he met members of the Photo League, among them Lou Bernstein, W. Eugene Smith and Weegee. He won a ...
and
Pamela H. Smith Pamela H. Smith is a historian of science specializing in attitudes to nature in early modern Europe (1350-1700), with particular attention to craft knowledge and the role of craftspeople in the Scientific Revolution. She is the Seth Low Professor ...
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."


In literature

The Ancient Greek
biologist A biologist is a scientist who conducts research in biology. Biologists are interested in studying life on Earth, whether it is an individual cell, a multicellular organism, or a community of interacting populations. They usually specialize in ...
and philosopher Aristotle wrote in his '' History of Animals'' that The
Ancient Roman In modern historiography, ancient Rome refers to Roman civilisation from the founding of the city of Rome in the 8th century BC to the collapse of the Western Roman Empire in the 5th century AD. It encompasses the Roman Kingdom (753–509 BC ...
encyclopaedist Pliny the Elder described the mayfly as the "hemerobius" in his '' Natural History'': In his 1789 book '' The Natural History and Antiquities of Selborne'', Gilbert White described in the entry for "June 10th, 1771" how The mayfly has come to symbolise the transitoriness and brevity of life. The English poet
George Crabbe George Crabbe ( ; 24 December 1754 – 3 February 1832) was an English poet, surgeon and clergyman. He is best known for his early use of the realistic narrative form and his descriptions of middle and working-class life and people. In the 177 ...
, 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: 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". Another literary reference to mayflies is seen in ''
The Epic of Gilgamesh The ''Epic of Gilgamesh'' () is an epic poem from ancient Mesopotamia, and is regarded as the earliest surviving notable literature and the second oldest religious text, after the Pyramid Texts. The literary history of Gilgamesh begins with ...
'', one of the earliest surviving great works of literature. The briefness of Gilgamesh's life is compared to that of the adult mayfly. In Szeged, Hungary, mayflies are celebrated in a monument near the Belvárosi bridge, the work of local sculptor Pal Farkas, depicting the courtship dance of mayflies. The American playwright David Ives wrote a short comedic play, ''Time Flies'', in 2001, as to what two mayflies might discuss during their one day of existence.


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 Fly fishing is an angling method that uses a light-weight lure—called an artificial fly—to catch fish. The fly is cast using a fly rod, reel, and specialized weighted line. The light weight requires casting techniques significantly diffe ...
. 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 Izaak Walton (baptised 21 September 1593 – 15 December 1683) was an English writer. Best known as the author of ''The Compleat Angler'', he also wrote a number of short biographies including one of his friend John Donne. They have been colle ...
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 George Edward MacKenzie Skues, usually known as G. E. M. Skues (1858–1949), was a British lawyer, author and fly fisherman most noted for the invention of modern-day nymph fishing and the controversy it caused with the Chalk stream dry fly d ...
with his 1910 book ''
Minor Tactics of the Chalk Stream ''Minor Tactics of the Chalk Stream and Kindred Studies'' is a fly fishing book written by G. E. M. Skues published in London in 1910. ''Minor Tactics'' was Skues's first book and set the stage for his ascendancy as the inventor of nymph fishi ...
''. 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
angler Angler may refer to: * A fisherman who uses the fishing technique of angling * ''Angler'' (video game) * The angler, ''Lophius piscatorius'', a monkfish * More generally, any anglerfish in the order Lophiiformes * '' Angler: The Cheney Vice Presi ...
s to catch trout for over 500 years. Some English public houses beside trout streams such as the River Test in Hampshire are named "The Mayfly".


As a spectacle

The hatch of the giant mayfly ''Palingenia longicauda'' on the Tisza and
Maros River Maros is a town in the South Sulawesi province of Indonesia close to the provincial capital of Makassar. It is the capital of the Maros Regency. Maros is the location of the Indonesian Cereals Research Institute, a branch of the Indonesian ...
s in Hungary and Serbia, known as "Tisza blooming", is a tourist attraction. The 2014 hatch of the large black-brown mayfly ''
Hexagenia bilineata ''Hexagenia bilineata'' is a species of mayfly in the family Ephemeridae. It is native to North America where it is found in the Upper Mississippi Valley. Sometimes adults of this mayfly are so abundant as to cause a nuisance because of their en ...
'' on the Mississippi River in the US was imaged on weather radar; the swarm flew up to 760 m (2,500 feet) above the ground near
La Crosse, Wisconsin La Crosse is a city in the U.S. state of Wisconsin and the county seat of La Crosse County. Positioned alongside the Mississippi River, La Crosse is the largest city on Wisconsin's western border. La Crosse's population as of the 2020 census w ...
, creating a radar signature that resembled a "significant rain storm", and the mass of dead insects covering roads, cars and buildings caused a "slimy mess". During the weekend of 13–14 June 2015, a large swarm of mayflies caused several vehicular accidents on the
Columbia–Wrightsville Bridge The Columbia–Wrightsville Bridge, officially the Veterans Memorial Bridge, spans the Susquehanna River between Columbia and Wrightsville, Pennsylvania, and carries Pennsylvania Route 462 and BicyclePA Route S. Built originally as the Lanc ...
, carrying Pennsylvania Route 462 across the
Susquehanna River The Susquehanna River (; Lenape: Siskëwahane) is a major river located in the Mid-Atlantic region of the United States, overlapping between the lower Northeast and the Upland South. At long, it is the longest river on the East Coast of the ...
between
Columbia Columbia may refer to: * Columbia (personification), the historical female national personification of the United States, and a poetic name for America Places North America Natural features * Columbia Plateau, a geologic and geographic region in ...
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.


As food

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 Lake Victoria is one of the African Great Lakes. With a surface area of approximately , Lake Victoria is Africa's largest lake by area, the world's largest tropical lake, and the world's second-largest fresh water lake by surface area after ...
, ''Povilla'' mayflies are collected, dried and preserved for use in food preparations.


As a name for ships and aircraft

"Mayfly" was the crew's nickname for His Majesty's Airship No. 1, an aerial scout airship built by Vickers but wrecked by strong winds in 1911 before her trial flights. Two vessels of the Royal Navy were named : a torpedo boat launched in January 1907, and a ''Fly''-class river gunboat constructed in sections at Yarrow in 1915. The
Seddon Mayfly The Seddon Mayfly was a tandem biplane of unusual construction. It was designed by Royal Navy Lieutenant John W. Seddon and A. G. Hackett and built by Accles & Pollock. When built it was the largest aeroplane in the world, but it failed to f ...
, which was constructed in 1908, was an aircraft that was unsuccessful in early flight. The first aircraft designed by a woman, Lillian Bland, was titled the
Bland Mayfly The Bland Mayfly was an early aircraft constructed in 1910 by Lilian Bland, Lilian E. Bland in Carnmoney in Ireland. It is credited as the first aeroplane to be designed and constructed by a woman. Background Lillian E. Bland was a sports j ...
.


Other human uses

In pre-1950s France, "chute de manne" was obtained by pressing mayflies into cakes and using them as bird food and fishbait. From an economic standpoint, mayflies also provide fisheries with an excellent diet for fish. Mayflies could find uses in the biomedical, pharmaceutical, and cosmetic industries. Their exoskeleton contains
chitin Chitin ( C8 H13 O5 N)n ( ) is a long-chain polymer of ''N''-acetylglucosamine, an amide derivative of glucose. Chitin is probably the second most abundant polysaccharide in nature (behind only cellulose); an estimated 1 billion tons of chit ...
, which has applications in these industries. Research on genome expression in the mayfly '' Cloeon dipterum,'' has provided ideas on the evolution of the insect wing and giving support to the so-called gill theory which suggests that the ancestral insect wing may have evolved from larval gills of aquatic insects like mayflies. 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 2003 titled "Mayfly". The shoes were designed with a wing venation pattern like the mayfly and were also said to have a finite lifetime. The telecommunication company Vodafone featured mayflies in a 2006 branding campaign, telling consumers to "make the most of now".


Notes


References


External links

* *
Info about Ephemeroptera
on Tree of Life
Mayfly Central
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