Flies are adapted for aerial movement and typically have short and streamlined bodies. The first tagma of the fly, the head, consists of ocelli, antennae, compound eyes, and the mouthparts (the labrum, labium, mandible and maxilla make up the mouthparts). The second tagma, the thorax, bears the wings and contains the flight muscles on the second segment, which is greatly enlarged; the first and third segments have been reduced to collar-like structures. The third segment of the thorax bears the halteres, which help to balance the insect during flight. A further adaptation for flight is the reduction in number of the neural ganglia, and concentration of nerve tissue in the thorax, a feature that is most extreme in the highly derived Muscomorpha infraorder.

Reproduction and development

The genitalia of female flies are rotated to a varying degree from the position found in other insects. In some flies, this is a temporary rotation during mating, but in others, it is a permanent torsion of the organs that occurs during the pupal stage. This torsion may lead to the anus being located below the genitals, or, in the case of 360° torsion, to the sperm duct being wrapped around the gut, despite the external organs being in their usual position. When flies mate, the male initially flies on top of the female, facing in the same direction, but then turns round to face in the opposite direction. This forces the male to lie on his back for his genitalia to remain engaged with those of the female, or the torsion of the male genitals allows the male to mate while remaining upright. This leads to flies having more reproduction abilities than most insects, and at a much quicker rate. Flies occur in great populations due to their ability to mate effectively and in a short period of time during the mating season.

The female lays her eggs as close to the food source as possible, and development is rapid, allowing the larvae to consume as much food as possible in a short period of time before transforming into adults. The eggs hatch immediately after being laid, or the flies are ovoviviparous, with the larvae hatching inside the mother.

Larval flies have no true legs. Some Dipteran larvae, such as species of Simuliidae, Tabanidae, and Vermileonidae, have prolegs adapted to such functions as holding onto a substrate in flowing water, holding onto host tissues, or holding prey. Roughly speaking, there is some anatomical distinction between the larvae of the Nematocera and the Brachycera (see Classification section, below); especially in the Brachycera, there is little demarcation between the thorax and abdomen, though the demarcation may be very visible in many Nematocera, such as mosquitoes (see image, both here and in the mosquitoes article); in the Brachycera, the head of the larva is not clearly distinguishable from the rest of the body, and there are few, if any, sclerites. Informally, such Brachyceran larvae are called maggots, but the term is nontechnical and often applied indifferently to fly larvae or insect larvae in general. The eyes and antennae of Brachyceran larvae are reduced or absent, and the abdomen also lacks appendages such as cerci. This lack of features is an adaptation to food such as carrion, decaying detritus, or host tissues surrounding endoparasites. Nematoceran larvae generally have visible eyes and antennae, though usually small and of limited function.

The pupae take various forms, and in some cases develop inside a silk cocoon. Despite the myth that flies only live a day (which may have either arisen from confusion with the mayfly or the fact that a fly inside a house may starve to death in a few days), most adult dipterans can live about one month.