Dipterans

aka: True Flies

The order Diptera belongs to the Phylum Arthropoda, Subphylum Labiata, Superclass Hexapoda, and Class Insecta. Among all insect orders, the order is number two in total number of species, ranked only behind beetles (Coleoptera), with about 125,000 described species (and an estimated 1,000,000 total species). They include house flies, blow flies, mosquitoes, gnats, black flies, midges, crane flies, horseflies, fruit flies, and others. Arkansas is home to many of these species.

The first true fossil dipterans are known from the Middle Triassic Period (about 240 million years ago [mya]), and became more widespread during the Middle to Late Triassic. Although modern flowering plants did not appear until the Cretaceous (around 140 mya), the original dipterans may have had a different source of sustenance other than nectar feeding, perhaps honeydew produced by sap-sucking bugs (hemipterans). Several new species of lower dipterans emerged in the Triassic (220 mya). During the Jurassic (180 mya), many lower brachycerans emerged. In addition, a third radiation took place among the Schizophora (consisting mainly of the Acalyptratae and Calyptratae) at the start of the Paleogene (66 mya). Evolutionarily, the basal clades in the Diptera include the mountain midges (Deuterophlebiidae) and the perplexing Nymphomyiidae (a family of tiny, slender, delicate flies with larvae found in northeastern North America, Japan, the Himalayas, and eastern Russia). Three episodes of evolutionary radiation are thought to have occurred, based on the fossil record.

Morphologically, the first tagma of the fly is a mobile head that supports a pair of large compound eyes, the antennae, and the mouthparts (the labrum, labium, mandible, and maxilla), which are characteristically suctoral. Dipterans also have, depending on the species involved, mouthparts designed for stabbing, piercing, and sucking (as in the black flies, mosquitoes, robber flies, and dance flies), or for lapping and sucking as in several other groups. The antennae are variable but well-developed, being thread-like, feathery, or comb-like in the different families.

The second tagma, the mesothorax, bears one pair of wings and contains the greatly enlarged flight muscles on the second segment; the first and third segments have been reduced to collar-like structures, and the third segment (metathorax) includes a single pair of halteres (mechanosensory organs), which help to balance the fly during flight. Flies are well adapted for aerial movement and typically have short and streamlined bodies. The wings and halteres function as high-speed sensors for rotational movement and allow them to perform advanced aerobatics. Interestingly, some species, such as the Hippoboscidae (louse flies), have lost wings secondarily.

The third tagma is the abdomen, which consists of eleven segments in primitive groups and ten segments in more derived groups, the tenth and eleventh segments being fused. The hindmost two or three segments are modified for reproduction. It shows considerable variability among members of the order. Each segment is made up of a dorsal and a ventral sclerite, connected by an elastic membrane. In some females, the sclerites are rolled into a flexible, telescopic ovipositor.

Each of the fly’s six legs possesses the typical insect structure of a coxa, trochanter, femur, tibia, and tarsus, with the tarsus, in most instances, being subdivided into five tarsomeres. At the tip of the limb is a pair of claws, and between these are pulvilli (cushion-like foot pads) that provide adhesion and permit them to cling to smooth surfaces.

Dipterans are often abundant, and are found in almost all terrestrial habitats in the world except for Antarctica. There are estimated to be a total of about 19,000 species in Europe, 22,000 in the Nearctic region, 20,000 in the Afrotropical region, 23,000 in the Oriental region, and 19,000 in the Australasian region. While most species have restricted overall distributions, a few, like the common housefly, Musca domestica (Muscidae), are cosmopolitan and found in most environs. On the other hand, larval shore flies (Ephydridae) and some chironomids (non-biting midges and lake flies) survive in extreme environments such as glaciers, geysers, hot springs, saline pools, septic tanks, and sulfur pools; some even live in crude oil, like the petroleum fly (Helaeomyia petrolei) of California. The largest fly in the world is the mydas fly, Gauromydas heros (Mydidae), of Brazil, Bolivia, and Paraguay, with a length up to 7 cm (2.8 in.), whereas the smallest is Euryplatea nanaknihali (Phoridae) of Thailand, at only 0.4 mm (0.016 in.).

The greatest diversity of gall-forming insects is found among the flies, principally in the family Cecidomyiidae (gall midges). Many flies (most importantly in the leaf-miner family Agromyzidae) lay their eggs in the mesophyll tissue of leaves, with larvae feeding between the surfaces, forming blisters and mines. Some families (like the dark-winged fungus gnats, Sciaridae) are fungus feeders (mycophagous). These also include the cave-dwelling fungus gnats (Mycetophilidae) whose larvae are the only dipterans possessing the ability for bioluminescence. Some plants are pollinated by fungus-feeding flies that visit fungus-infected flowers.

The larvae of the laboratory fly, Megaselia scalaris (Phoridae), are almost omnivorous and consume such odd substances as paint and shoe polish. Tachinid flies, Exorista mella, are considered generalists and parasitoids of a variety of hosts. Well known for their mimicry are adult flower or hoverflies (Syrphidae), whose larvae adopt diverse lifestyles including being inquiline scavengers inside the nests of social insects.

Diptera is one of the major insect orders and of considerable human and ecological importance. Interestingly, flies represent a very large group of pollinators, second only to the Hymenoptera (bees, wasps, and relatives). In colder and damper environments, dipterans are significantly more important as pollinators. Compared to bees, they require fewer nutrients, as they do not need to provision their young. Many flowers that possess low nectar and those that have evolved trap pollination are dependent on flies. Fruit flies (Drosophila) have been used for decades as important model organisms in biological, genetic, and medical research. Blowfly larvae (Calliphoridae), also known as gentles, and other dipteran larvae, known more generally as maggots, are used as fishing bait and serve as food for carnivores.

Houseflies, blowflies, and fungus gnats (Mycetophilidae) are scavengers and aid in decomposition. Robber flies (Asilidae), tachinids (Tachinidae), and dagger flies and balloon flies (Empididae) are predators and parasitoids of other insects, helping to control a variety of pests. Many dipterans such as bee flies (Bombyliidae) and hoverflies (Syrphidae) are pollinators of crop plants.

Dipterans, most famously the mosquitoes (Culicidae), are vectors of many serious diseases affecting humans and other animals, including encephalitis, filariasis, dengue fever, heartworm, malaria, arboviruses, West Nile fever, Zika virus, and yellow fever. Houseflies (Muscidae), for example, are commensal with all peoples of the world and are capable of spreading food-borne illnesses. Muscid and stable flies can be annoying, especially in some parts of the world where they can occur in large populations, buzzing and settling on the skin or eyes to bite or seek fluids. Some, such as some larger flies like tsetse flies (Glossina) and screwworms (Cochliomya), also cause significant economic harm to cattle.

Other important parasitic dipterans include those in the several other families: Simulidae (black flies, turkey gnats, and buffalo gnats); Psychodidae (subfamily Phlebotominae) or sand flies, important as vectors of Leishmania; Ceratopogonidae, the biting midges (no-see-ums or punkies); Tabanidae, deer flies (Chrysops spp.), which transmit the eye worm (Loa loa); Hippoboscidae, louse flies or sheep ked (Melophagus ovinus); drain flies (Psychodidae); Sarcophagidae, flesh flies; Oestridae, nasal bots, pharyngeal bots, head maggots, rodent or skin bots; Gastrophilidae, stomach bots; Nycteribiidae, bat spider flies; and Streblidae, bat flies. In addition, some leaf-miner flies (Agromyzidae), fruit flies (Tephritidae and Drosophilidae), and gall midges (Cecidomyiidae) are pests of agricultural crops.

In many dipteran groups, swarming is a behavioral feature of adult life, with clouds of flies gathering in certain locations; these insects are mostly males, and the swarm may serve the purpose of making their location more visible to females.

In general, flies undergo complete metamorphosis with four distinct life stages: egg, larva, pupa, and adult. The eggs are laid on the larval food source, and the larvae (with a long life) lack true limbs and develop as maggots often inside their food source. Many larvae are aquatic, and others are found in moist places such as carrion, fruit, fungi, vegetable matter, and, in the case of parasitic species, inside their hosts. Larval dipterans lack true legs and move by muscular contraction (peristaltic waves) throughout the body. The larvae of most species of flies have a reduced head capsule, and all that remains are the mandibles and some associated sclerites, which are collectively referred to as the cephalopharyngeal skeleton. The pupae take various forms that are formed in a tough desiccation-resistant capsule (puparium), from which the adult emerges as it inflates a balloon-like structure on its head, and forces its way out when ready. Dipteran pupae have non-functional mandibles and may have the appendages free from the body, or glued to the body. For the most part, flies have a short life span as adults. The adult stage functions mainly to mate and lay eggs.

In some groups, larval dipterans complete their development without feeding, and in others the adults do not feed at all. However, flies that do feed, especially those that feed on blood, possess special sensory structures that can detect infrared emissions, which they use to home in on their hosts. Indeed, many blood-sucking flies can detect the raised concentration of carbon dioxide that occurs near large animals. Larvae can be herbivores, scavengers, decomposers, predators, or parasites, with the consumption of decaying organic matter being one of the most prevalent feeding behaviors. Fruit or detritus are consumed along with the associated micro-organisms, a sieve-like filter in the pharynx being utilized to concentrate the particles, while flesh-eating larvae have mouth-hooks to help shred their food. The larvae of some groups feed on or in the living tissues of plants and fungi, and some of these are serious pests of agricultural crops. Some aquatic larvae consume algal films that form under water on rocks and plants. Many of the parasitoid larvae grow inside and eventually kill other arthropods, while parasitic larvae may attack vertebrate hosts.

In terms of food habits, similar to other insects, flies possess chemoreceptors that detect smell and taste, and mechanoreceptors that respond to touch. The third segments of the antennae and the maxillary palps bear the main olfactory receptors, while the gustatory (taste) receptors are in the labium, pharynx, feet, wing margins, and female genitalia, enabling flies to taste their food by walking on it. The taste receptors in females at the tip of the abdomen receive information on the suitability of a site for ovipositing. Some tachinid flies (subfamily Ormiinae) that are parasitoids of bush crickets (Tettigonidae) have sound receptors to help them locate their singing orthopteran hosts. Whereas many dipteran larvae are aquatic or live in enclosed terrestrial locations, the majority of adults are found above ground and are capable of flight. Predominantly, they feed on nectar or plant or animal exudates, such as honeydew, for which their lapping mouthparts are adapted. The flies that feed on vertebrate blood have sharp stylets that pierce the skin, the insects inserting anticoagulant saliva and absorbing the blood that flows; in this process, certain diseases can be transmitted.

Flies are predated on by other animals at all stages of their development. The eggs and larvae are often parasitized by other insects and eaten by many organisms, some of which specialize in feeding on flies but most of which consume them as part of a mixed diet. Birds, bats, frogs and toads, lizards, dragonflies, and spiders are among the prime predators of flies.

Many flies have evolved mimetic resemblances that aid their protection. Batesian mimicry is widespread, as many hoverflies resemble bees, wasps, and ants. Some species of tephritid fruit flies resemble jumping spiders, and this mimicry apparently serves to protect them from some predators. Some species of hoverflies are myrmecophilous; their young live and grow within the nests of ants. They are protected from the ants by imitating chemical scents produced by members of the ant colony. Large bombyliid bee flies, such as Bombylius major, are furry, short-bodied, round, and distinctly bee-like as they light on flowers for nectar; they are likely also Batesian mimics of bees.

Caves provide an excellent habitat for some flies. In Arkansas, several adult dipteran species have been reported from Ozark caves. The mycetophilids, Exechiopsis umbratica and Rymosia sp., and several culicids are among the most common cave dipterans in Ozarkian caves. Only one true troglobitic dipteran is found in Ozark caves, the sphaerocerid, Spelobia tenebrarum. Other subterranean-inhabiting dipterans (scuttle flies) in the state include Conicera slayi, found in caves as well as mines, and Megaselia taylori, found in caves in Arkansas and Missouri.

There is no comprehensive list of the dipterans of Arkansas; most of the research on dipterans of state has been conducted on various species of mosquitoes. There are about sixty species of mosquitoes that occur in Arkansas and Missouri. A checklist of 131 species of robber flies from Arkansas is available. There are at least ten genera and eighty species of horseflies and deerflies (Tabanidae) from Arkansas. A single endemic fly is known from the state: the robber fly (Atomosia arkansensis) was reported from Grandview Prairie Wildlife Management Area in Hempstead County.

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Chris T. McAllister
Eastern Oklahoma State College

Last Updated: 09/28/2021

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