Mayfly
Mayflies are aquatic insects belonging to the order Ephemeroptera. This order is part of an ancient group of insects termed the Palaeoptera, which also contains dragonflies and damselflies. Over 3,000 species of mayfly are known worldwide, grouped into over 400 genera 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 forms, 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" from spring to autumn, 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 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 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 compared the brief life of a daily newspaper with that of a mayfly in the satirical poem "The Newspaper", 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, moulting and increasing in size each time. When ready to emerge from the water, nymphs vary in length, depending on species, from. The head 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 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, 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 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.
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 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 membrane. 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, 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. 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 that face upwards in addition to the lateral eyes that resemble those of females. The lateral eyes are apposition compound eyes but the turbinate eyes are thought by some to be superposition compound eyes while others suggest they may appear like superposition eyes but are based on more complex wave-guide like systems meant to detect ultraviolet light. They 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 at the tip. Like Entognatha, Archaeognatha and Zygentoma, the spiracles on the abdomen do not have closing muscles. Uniquely among insects, mayflies possess paired genitalia, with the male having two aedeagi and the female two gonopores.Biology
Reproduction and life cycle
Mayflies are hemimetabolous. They are unique among insects in that they moult one more time after acquiring functional wings; this last-but-one winged 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 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 individuals in a population mature at once, 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, 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, while their digestive systems are filled with air. Dolania americana 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. The sperm transfer is direct as in Neoptera, contrasting with the indirect insemination of Odonata.
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, including both partially and exclusively parthenogenetic 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 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.