Nymphs

In contrast to the adults, mayfly nymphs show considerable diversity in habit and appearance. Differences do not always follow taxonomic lines, and convergent and parallel evolution seems to be common (Fig. 2).

Growth and Development

Mayflies have a large number of postembryonic molts. Estimates of the number of instars vary between 10 and 50; most are in the range 15 to 25. The number of instars for a particular species does not seem to be constant, but probably varies within certain limits. Environmental conditions, such as food quality and temperature, may affect instar number. Because of its simplicity, by far the most common measure of development and growth in mayflies has been body length, although head width and other body dimensions also have been used. However, growth of the various body parts is not always isometric. Many authors have also used body weight, and the length—weight relationship is usually well expressed by a power function.

Nymphal growth rates are influenced by several environmental factors, although the major growth regulator is mean temperature, the scale of diurnal fluctuations, or total degrees-days. Other factors, such as food and current velocity, may exert a modifying influence on growth rates. No true diapausing nymphal stage has been reported in the Ephemeroptera, although growth rates often are very low during the winter.

Respiration

The gills of mayflies are very diverse in form, ranging from a single plate in Ameletus to fibrillar tufts in Hexagenia. Respiratory tufts are sometimes developed on other parts of the body besides the abdomen, such as those at the base of the coxa in Isonychia and Dactylobaetis. In several families the second abdominal gill has developed into an operculate (lidlike) gill cover for the remaining gills, and in certain Heptageniidae the gills are markedly expanded so that they together form an adhesion disc. In many of the Siphlonuridae, the gills are used as swimming paddles, which has been put forward as their original function. In respiring, the gills may function either as respiratory organs or as ventilatory organs for other respiratory exchange surfaces.

High rates of oxygen consumption are often reported in association with emergence and gonad maturation. High water temperatures at that time may mean that low oxygen concentrations can be critical. Many burrowing Ephemeridae and pond-dwelling Baetidae are able to survive moderately low oxygen concentrations, especially for short periods. However, so far only one species, the European Cloeon dipterum, has been shown to survive long-term anoxia.

Population Movements

During the final stages of nymphal life there is a movement to and a concentration in the shallower areas of lakes and rivers. In running waters, springtime mass movements of mayfly nymphs along the banks of the main river and into slower flowing tributary streams or into areas flooded by spring snowmelt have been observed. In running water, mayfly nymphs may move down into the substratum in response to spates or as part of a daily rhythm. Generally, however, mayflies do not extend far down into the substratum (i.e., the hyporheic zone).

Mayflies, especially Baetidae, are a major component of invertebrate drift in running waters. Their drift shows a strong diel periodicity, with a peak during the hours of darkness. Drift rates are not constant for a particular species, and the larger size classes are usually more in evidence. Other factors that have been shown to influence mayfly drift include changes in current velocity and discharge, increased sediment loading, temperature changes, oxygen conditions, density, food availability, and predators.

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