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Because of the structural variety in Diptera, especially among larvae, it is difficult to generalize about morphology. Despite this variety, flies share a number of features. Except for certain forms (e.g., cave-dwelling species), adult flies usually possess large compound eyes. In some species, eyes meet or almost meet dorsally (holoptic); in other groups, eyes are widely separated (dichoptic). Further modifications include eyes that are divided into distinct dorsal and ventral components, a feature found in many Simuliidae, Blephariceridae, and other groups. These modifications are among many that might be related to swarming behavior. The

FIGURES 1-6 Adult head of (1) Tipulidae, (2) Blephariceridae, (3) Asilidae, (4) Empididae, (5) Tachinidae, (6) Syrphidae. (Photographs by G. Courtney.)

regions of a fly head include the vertex, a dorsomedial area above and posterior to the eyes; the frons, an area extending from the vertex to the antennal insertions; and the face, which extends from the antennal insertions to the clypeus, a region intimately associated with the mouthparts. All of these areas can bear a variety of setae, the number and position of which often are useful in identification.

Nearly all flies have well-developed antennae, with the flagellum being the most varied component. In nemato-cerous families, the antennae are usually composed of many segments and are filiform, plumose, or pectinate (Figs. 1—2), whereas brachycerous flies typically have the first flagellomere enlarged and the remaining flagellomeres stylate or aristate (Figs. 3—6). The mouthparts of adult flies also vary between groups, ranging from vestigial forms (e.g., Deuterophlebidae, Oestridae) to those that are well developed. The latter include two general types: (1) piercing and sucking, as seen in simuliids, culicids, and asilids, and (2) lapping and sucking, as seen in tipulids and most brachycerous groups. Typically, the proboscis comprises the unpaired labrum— epipharynx, labium, and hypopharynx and the paired mandibles and maxillae. In most groups, the base of each maxilla bears a distinct palpus and the apex of the labium is modified into a labellum, which consists of membranous lobes derived from the labial palpi.

Perhaps the most distinct feature of the adult fly is the single pair of wings (hence, the ordinal name, Diptera, meaning "two wings"). A related characteristic is the highly modified thorax, with a reduced prothorax and metathorax, and a greatly enlarged mesothorax. The latter includes several prominent dorsal and lateral sclerites and, internally, houses much of the wing musculature. Wing venation varies greatly throughout the Diptera and can be extremely important for identification. The metathoracic wings are modified into distinct club-shaped halteres, which are thought to play an important role as balancing organs. Interestingly, halteres are distinct in some groups that are otherwise wingless (e.g., Hippoboscidae). The legs of an adult fly are typical of most insects, each with a coxa, trochanter, femur, tibia, and, in nearly all groups, a tarsus comprising five tarsomeres. Beyond this basic arrangement, there is considerable diversity of leg structure in Diptera, with this diversity often providing useful taxonomic information.

The adult abdomen also shows considerable variety. In basic structure, the abdomen consists ofll segments, the last 2 or 3 of which are highly modified for reproduction. Most abdominal segments consist of a dorsal and ventral sclerite, connected laterally by a pleural membrane of varying width. There is a general trend toward a shortening of the abdomen in Diptera (cf. Tipulidae and Muscidae). The terminalia of Diptera are complex, highly variable, and of considerable use in taxonomic and phylogenetic studies. Details of terminalic structure are beyond the scope of this article; however, the structural variety of Diptera terminalia and the controversy about interpreting their homologies can be found in some of the general references listed at the end.

The dipteran pupa also varies considerably in form. Some fly pupae look like a cross between the worm-like larva and the adult, whereas others are relatively featureless and seed-like in appearance. The former are typical of the Nematocera and are described as obtect, or having the appendages fused to the body (Figs. 7-10). For instance, a crane fly (Tipulidae) pupa has identifiable head, thoracic, and abdominal segments, but the antennal sheaths, legs, and wing pads adhere to the pupal body (Fig. 9). Nematocerous pupae are frequently leathery to the touch. The exterior of the nematoceran pupa may be adorned with spines, gill-like respiratory devices, or locomotory paddles (Figs. 7-10). The Brachycera and Cyclorrhapha form the pupal stage in a different, more concealed manner. Families of the so-called higher Diptera form pupae that are described as coarctate, which literally means "compacted" or "contracted" (Figs. 11-15). These taxa (e.g., Syrphidae, Drosophilidae, Muscidae) form a puparium that is composed of the hardened skin of the last larval instar (Fig. 14). This relatively tough, desiccation-resistant structure houses and protects the pupa; the adult

FIGURES 7-15 Pupa of (7) Ptychopteridae, (8) Simuliidae, (9) Tipulidae, (10) Chironomidae, showing anal division below, (11) Tabanidae, (12) Empididae, (13) Syrphidae, (14) Muscidae, (15) Ephydridae. (All illustrations modified, with permission, from Merritt and Cummins, 1996.)

also forms within the puparium. The enclosed adult must break through the puparial skin and does so by extruding a balloon-like structure from the frons called the ptilinum. The ptilinum is used to break the cephalic cap, a lid-like structure positioned anteriorly on the puparium, thus liberating the teneral (or newly emerged) adult. Very few external features are noticeable on the puparium, although careful examination will reveal the spiracles through which atmospheric air is obtained by the pupa.

Diptera larvae can be distinguished from the larvae of most other insects by the lack of jointed thoracic legs. In other features, larval dipterans show tremendous structural variety. This variation is exemplified by cranial structure. Larvae of most nematocerous flies are eucephalic, i.e., characterized by a complete, fully exposed, and heavily sclerotized head capsule (Figs. 17-19 and 24). Larval tipulids are special among nematocerous flies, as the head capsule often is fully retracted into the thorax (Fig. 16) and the posterior cranial margin may possess small to extensive longitudinal incisions (Fig. 23). In contrast to the condition in nematoceran larvae, the cranial sclerites of brachyceran larvae are greatly reduced or absent. The hemicephalic head capsule of many orthorrhaphous Brachycera consists of slender arms and rods that are partly retracted into the thorax (Figs. 25-26). The culmination of cranial reduction is in the acephalic head of larval Cyclorrhapha, in which the external portions of the head are membranous, and much of the head is retracted into the

Pics Eucephalus Larvae

FIGURES 16-27 Larva of (16) Tipulidae, (17) Ceratopogonidae, (18) Chironomidae, (19) Simuliidae, (20) Tabanidae, (21) Syrphidae, (22) Ephydridae. Larval head capsule of (23) Tipulidae, (24) Chironomidae. Cranial sclerites and mouth parts of (25) Tabanidae, (26) Dolichopodidae. (27) Cephalopharyngeal skeleton of Sciomyzidae. (All illustrations modified, with permission, from Merritt and Cummins, 1996.)

FIGURES 16-27 Larva of (16) Tipulidae, (17) Ceratopogonidae, (18) Chironomidae, (19) Simuliidae, (20) Tabanidae, (21) Syrphidae, (22) Ephydridae. Larval head capsule of (23) Tipulidae, (24) Chironomidae. Cranial sclerites and mouth parts of (25) Tabanidae, (26) Dolichopodidae. (27) Cephalopharyngeal skeleton of Sciomyzidae. (All illustrations modified, with permission, from Merritt and Cummins, 1996.)

thorax (Fig. 27). The internal portion, or cephalopharyngeal skeleton, is thought to comprise the remnants of internal cranial sclerites (tentorium) and various mouthparts. Although referred to as "acephalic," the primary difference between the head of a cyclorrhaphan larva and that of a nematoceran larva is that most of the constituent segments are withdrawn into the thorax and thus externally hidden (Fig. 22). Cranial modifications are accompanied by general changes in the shape and rotation of mandibles and other mouthparts. The mandible of larval nematocerans typically consists of a stout, toothed structure that moves in a horizontal or oblique plane and operates as a biting and chewing organ. The brachyceran larval mandible usually is more claw-like, has fewer teeth along the inner surface, moves in a vertical plane, and operates as a piercing or slashing organ.

In most Diptera larvae, the thorax and abdomen are soft, flexible, and only occasionally provided with sclerotized plates. The thorax usually consists of three distinct segments and the abdomen usually eight or nine segments (Figs. 17-19). Body form varies almost as much as does cranial diversity and ecological habits. In many nematoceran groups (e.g., most Chironomidae, Tipulidae, and Simuliidae), the body is sub-cylindrical (Figs. 16, 18, and 19). Other groups are predominantly fusiform (e.g., Cecidomyiidae) or elongated and serpentine (e.g., Ceratopogonidae) (Fig. 17). The latter body form is common in groups inhabiting soil and interstitial aquatic habitats. The larvae of some groups (e.g., Culicidae) are unusual in that the thoracic segments are indistinctly differentiated and form a single large segment that is wider than the rest of the body (Fig. 48). The typical body shape of a cyclorrhaphan larva is that of a maggot (i.e., pointed at the anterior end, with the thoracic segments approaching the maximum body diameter). The variation in body form is particularly impressive in families whose larvae feed on a variety of substrates (e.g., Syrphidae). Cyclorrhaphan larvae can be dorsoventrally flattened, a feature often associated with the presence of segmental or branched body protuberances. The syrphid genus Microdon has one of the most unusual larvae, being ventrally flattened, dorsally dome-shaped, and sluglike in overall appearance. Larvae with parasitoid and parasitic life styles (e.g., Pipunculidae, Oestridae) are often extremely stout or pear-shaped, their body form being closely adapted to that of the host.

Despite the absence of jointed thoracic legs, locomotion is highly diverse in fly larvae, reflecting the group's diversity in habitat and habits. Locomotory appendages operate through a combination of turgor pressure and muscle action and include creeping welts, prolegs, and other specialized structures (e.g., suctorial discs). Creeping welts are transverse, swollen areas (ridges) that bear one to several modified setae or spines; creeping welts are characteristic of several groups, including many crane flies, dance flies, and deer and horse flies (Fig. 20). Among orthorrhaphous groups, ventral creeping welts are common in the larvae of Rhagionidae and Empididae. Cyclorrhaphan larvae typically use creeping welts as anchoring devices, with welts usually comprising bands of small spines on abdominal segments. The distribution and morphology of creeping welts vary considerably between families, species, instars, and segments. Prolegs usually are paired, round, elongate, fleshy, retractile processes that bear apical spines or crochets; prolegs come in a diversity of shapes, sizes, and positions and are typical of Chironomidae, Deuterophlebidae, Simuliidae, Rhagionidae, and various members of other groups (Figs. 16, 18, 19, 21, and 22). Other specialized structures used for locomotion or attachment include friction pads and suctorial discs. Several genera of Psychodidae possess friction pads, which are areas of modified cuticle on the ventral surface of the thorax or abdomen. Functionally similar structures may occur in certain Ephydridae, particularly in groups inhabiting waterfalls and thin films of flowing water. Suctorial disks are true suction devices on the ventral body surface of larval net-winged midges and are an obvious adaptation to life in torrential streams.

Larval Diptera show a variety of respiratory adaptations, many a reflection of life in fluid or semifluid habitats. The basic respiratory system comprises an internal system of tracheae and the external spiracles. Respiration may be directly from the atmosphere, from plant tissues, or from oxygenated fluids. The presence of hemoglobin in the blood of some midges can assist the absorption of oxygen. Many aquatic larvae, particularly those from well-oxygenated streams, are apneustic (lack spiracles) and absorb oxygen directly through the skin. Some families (e.g., Psychodidae) possess spiracles on the prothorax and last abdominal segment, whereas others (e.g., Culicidae and most cyclorrhaphans) have spiracles on only the last segment. In several groups (e.g., many Ephydridae and Syrphidae), the spiracles are at the end of a retractile respiratory siphon (Figs. 21 and 22).

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