1911 Encyclopædia Britannica/May-fly
MAY-FLY. The Mayflies belong to the Ephemeridae, a remarkable family of winged insects, included by Linnaeus in his order Neuroptera, which derive their scientific name from ἑφήμερος, in allusion to their very short lives. In some species it is possible that they have scarcely more than one day's existence, but others are far longer lived, though the extreme limit is probably rarely more than a week. The family has very sharply defined characters, which separate its members at once from all other neuropterous (or pseudo-neuropterous) groups.
These insects are universally aquatic in their preparatory states. The eggs are dropped into the water by the female in large masses, resembling, in some species, bunches of grapes in miniature. Probably several months elapse before the young larvae are excluded. The sub-aquatic condition lasts a considerable time: in Cloeon, a genus of small and delicate species, Sir J. Lubbock (Lord Avebury) proved it to extend over more than six months; but in larger and more robust genera (e.g. Palingenia) there appears reason to believe that the greater part of three years is occupied in preparatory conditions.
The larva is elongate and campodeiform. The head is rather large, and is furnished at first with five simple eyes of nearly equal size; but as it increases in size the homologues of the facetted eyes of the imago become larger, whereas those equivalent to the ocelli remain small. The antennae are long and thread-like, composed at first of few joints, but the number of these latter apparently increases at each moult. The mouth parts are well developed, consisting of an upper lip, powerful mandibles, maxillae with three-jointed palpi, and a deeply quadrifid labium or lower lip with three-jointed labial palpi. Distinct and conspicuous maxillulae are associated with the tongue or hypopharynx. There are three distinct and large thoracic segments, whereof the prothorax is narrower than the others; the legs are much shorter and stouter than in the winged insect, with monomerous tarsi terminated by a single claw. The abdomen consists of ten segments, the tenth furnished with long and slender multi-articulate tails, which appear to be only two in number at first, but an intermediate one gradually develops itself (though this latter is often lost in the winged insect). Respiration is effected by means of external gills placed along both sides of the dorsum of the abdomen and hinder segments of the thorax. These vary in form: in some species they are entire plates, in others they are cut up into numerous divisions, in all cases traversed by numerous tracheal ramifications. According to the researches of Lubbock and of E. Joly, the very young larvae have no breathing organs, and respiration is effected through the skin. Lubbock traced at least twenty moults in Cloeon; at about the tenth rudiments of the wing-cases began to appear. These gradually become larger, and when so the creature may be said to have entered its “nymph” stage; but there is no condition analogous to the pupa-stage of insects with complete metamorphoses.
There may be said to be three or four different modes of life in these larvae: some are fossorial, and form tubes in the mud or clay in which they live; others are found on or beneath stones; while others again swim and crawl freely among water plants. It is probable that some are carnivorous, either attacking other larvae or subsisting on more minute forms of animal life; but others perhaps feed more exclusively on vegetable matters of a low type, such as diatoms.
The most aberrant type of larva is that of the genus Prosopistoma, which was originally described as an entomostracous crustacean on account of the presence of a large carapace overlapping the greater part of the body. The dorsal skeletal elements of the thorax and of the anterior six abdominal segments unite with the wing-cases to form a large respiratory chamber, containing five pairs of tracheal gills, with lateral slits for the inflow and a posterior orifice for the outflow of water. Species of this genus occur in Europe, Africa and Madagascar.
When the aquatic insect has reached its full growth it emerges from the water or seeks its surface; the thorax splits down the back and the winged form appears. But this is not yet perfect, although it has all the form of a perfect insect and is capable of flight; it is what is variously termed a “pseud-imago,” “sub-imago” or “pro-imago.” Contrary to the habits of all other insects, there yet remains a pellicle that has to be shed, covering every part of the body. This final moult is effected soon after the insect's appearance in the winged form; the creature seeks a temporary resting-place, the pellicle splits down the back, and the now perfect insect comes forth, often differing very greatly in colours and markings from the condition in which it was only a few moments before. If the observer takes up a suitable position near water, his coat is often seen to be covered with the cast sub-imaginal skins of these insects, which had chosen him as a convenient object upon which to undergo their final change. In some few genera of very low type it appears probable that, at any rate in the female, this final change is never effected and that the creature dies a sub-imago.
The winged insect differs considerably in form from its sub-aquatic condition. The head is smaller, often occupied almost entirely above in the male by the very large eyes, which in some species are curiously double in that sex, one portion being pillared, and forming what is termed a “turban,” the mouth parts are aborted, for the creature is now incapable of taking nutriment either solid or fluid; the antennae are mere short bristles, consisting of two rather large basal joints and a multi-articulate thread. The prothorax is much narrowed, whereas the other segments (especially the mesothorax) are greatly enlarged; the legs long and slender, the anterior pair often very much longer in the male than in the female; the tarsi four- or five-jointed; but in some genera (e.g. Oligoneuria and allies) the legs are aborted, and the creatures are driven helplessly about by the wind. The wings are carried erect: the anterior pair large, with numerous longitudinal nervures, and usually abundant transverse reticulation; the posterior pair very much smaller, often lanceolate, and frequently wanting absolutely. The abdomen consists of ten segments; at the end are either two or three long multi-articulate tails; in the male the ninth joint bears forcipated appendages; in the female the oviducts terminate at the junction of the seventh and eighth ventral segments. The independent opening of the, genital ducts and the absence of an ectodermal vagina and ejaculatory duct are remarkable archaic features of these insects, as has been pointed out by J. A. Palmén. The sexual act takes place in the air, and is of very short duration, but is apparently repeated several times, at any rate in some cases.
Ephemeridae are found all over the world, even up to high northern latitudes. F. J. Pictet, A. E. Eaton and others have given us valuable works or monographs on the family; but the subject still remains little understood, partly owing to the great difficulty of preserving such delicate insects; and it appears probable they can only be satisfactorily investigated as moist preparations. The number of described species is less than 200, spread over many genera.
From the earliest times attention has been drawn to the enormous abundance of species of the family in certain localities. Johann Anton Scopoli, writing in the 18th century, speaks of them as so abundant in one place in Carniola that in June twenty cartloads were carried away for manure! Polymitarcys virgo, which, though not found in England, occurs in many parts of Europe (and is common at Paris), emerges from the water soon after sunset, and continues for several hours in such myriads as to resemble snow showers, putting out lights, and causing inconvenience to man, and annoyance to horses by entering their nostrils. In other parts of the world they have been recorded in multitudes that obscured passers-by on the other side of the street. And similar records might be multiplied almost to any extent. In Britain, although they are often very abundant, we have scarcely anything analogous.
Fish, as is well known, devour them greedily, and enjoy a veritable feast during the short period in which any particular species appears. By anglers the common English species of Ephemera (vulgata and danica, but more especially the latter, which is more abundant) is known as the “may-fly,” but the terms “green drake” and “bastard drake” are applied to conditions of the same species. Useful information on this point will be found in Ronalds's Fly-Fisher's Entomology, edited by Westwood.
Ephemeridae belong to a very ancient type of insects, and fossil imprints of allied forms occur even in the Devonian and Carboniferous formations.
There is much to be said in favour of the view entertained by some entomologists that the structural and developmental characteristics of may-flies are sufficiently peculiar to warrant the formation for them of a special order of insects, for which the names Agnatha, Plectoptera and Ephemeroptera have been proposed. (See Hexapoda, Neuroptera.)
Bibliography. — Of especial value to students of these insects are A. E. Eaton's monograph (Trans. Linn. Soc. (2) iii. 1883-1885) and A. Vayssière's “Recherches sur l'organisation des larves” (Ann. Nat. Zool. (6) xiii. 1882 (7) ix. 1890). J. A. Palmén's memoirs Zur Morphologie des Tracheensystems (Leipzig, 1877) and Über paarige Ausführungsgänge der Geschlechtsorgane bei Insekten (Helsingfors, 1884), contain important observations on may-flies. See also L. C. Miall, Nat. Hist. Aquatic Insects (London, 1895); J. G. Needham and others (New York State Museum, Bull. 86, 1905).