Insect wing
Body part used by insects to fly
Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwings, respectively, though a few insects lack hindwings, even rudiments. The wings are strengthened by a number of longitudinal veins, which often have cross-connections that form closed "cells" in the membrane (extreme examples include the dragonflies and lacewings). The patterns resulting from the fusion and cross-connection of the wing veins are often diagnostic for different evolutionary lineages and can be used for identification to the family or even genus level in many orders of insects.
Physically, some insects move their flight muscles directly, others indirectly. In insects with direct flight, the wing muscles directly attach to the wing base, so that a small downward movement of the wing base lifts the wing itself upward. Those insects with indirect flight have muscles that attach to and deform the thorax, causing the wings to move as well.
The wings are present in only one sex (often the male) in some groups such as velvet ants and Strepsiptera, or are selectively lost in "workers" of social insects such as ants and termites. Rarely, the female is winged but the male not, as in fig wasps. In some cases, wings are produced only at particular times in the life cycle, such as in the dispersal phase of aphids. Wing structure and colouration often vary with morphs, such as in the aphids, migratory phases of locusts and polymorphic butterflies. At rest, the wings may be held flat, or folded a number of times along specific patterns; most typically, it is the hindwings which are folded, but in a few groups such as the vespid wasps, it is the forewings.
The evolutionary origin of the insect wing is debated. During the 19th century, the question of insect wing evolution originally rested on two main positions. One position postulated insect wings evolved from pre-existing structures, while the second proposed insect wings were entirely novel formations. The “novel” hypothesis suggested that insect wings did not form from pre-existing ancestral appendages but rather as outgrowths from the insect body wall.
Long since, research on insect wing origins has built on the “pre-existing structures” position that was originally proposed in the 19th century. Recent literature has pointed to several ancestral structures as being important to the origin of insect wings. Among these include: gills, respiratory appendages of legs, and lateral (paranotal) and posterolateral projections of the thorax to name a few.
According to more current literature, possible candidates include gill-like structures, the paranotal lobe, and the crustacean tergal plate. The latter is based on recent insect genetic research which indicates that insects are pan-crustacean arthropods with a direct crustacean ancestor and shared genetic mechanisms of limb development.
Other theories of the origin of insect wings are the paranotal lobe theory, the gill theory and the dual theory of insect wing evolution. These theories postulate that wings either developed from paranotal lobes, extensions of the thoracic terga; that they are modifications of movable abdominal gills as found on aquatic naiads of mayflies; or that insect wings arose from the fusion of pre-existing endite and exite structures each with pre-existing articulation and tracheation.
Morphology.
Internal.
Each of the wings consists of a thin membrane supported by a system of veins. The membrane is formed by two layers of integument closely apposed, while the veins are formed where the two layers remain separate; sometimes the lower cuticle is thicker and more heavily sclerotized under a vein. Within each of the major veins there is a nerve and a trachea, and, since the cavities of the veins are connected with the hemocoel, hemolymph can flow into the wings.
As the wing develops, the dorsal and ventral integumental layers become closely apposed over most of their area forming the wing membrane. The remaining areas form channels, the future veins, in which the nerves and tracheae may occur. The cuticle surrounding the veins becomes thickened and more heavily sclerotized to provide strength and rigidity to the wing. Two types of hair may occur on the wings: microtrichia, which are small and irregularly scattered, and macrotrichia, which are larger, socketed, and may be restricted to veins. The scales of Lepidoptera and Trichoptera are highly modified macrotrichia.
Venation.
In some very small insects, the venation may be greatly reduced. In chalcidoid wasps, for instance, only the subcosta and part of the radius are present. Conversely, an increase in venation may occur by the branching of existing veins to produce accessory veins or by the development of additional, intercalary veins between the original ones, as in the wings of Orthoptera (grasshoppers and crickets). Large numbers of cross-veins are present in some insects, and they may form a reticulum as in the wings of Odonata (dragonflies and damselflies) and at the base of the forewings of Tettigonioidea and Acridoidea (katydids and grasshoppers respectively).
The archedictyon is the name given to a hypothetical scheme of wing venation proposed for the very first winged insect. It is based on a combination of speculation and fossil data. Since all winged insects are believed to have evolved from a common ancestor, the archedictyon represents the "template" that has been modified (and streamlined) by natural selection for 200 million years. According to current dogma, the archedictyon contained 6–8 longitudinal veins. These veins (and their branches) are named according to a system devised by John Comstock and George Needham—the Comstock–Needham system:
Costa (C) – the leading edge of the wing
Subcosta (Sc) – second longitudinal vein (behind the costa), typically unbranched
Radius (R) – third longitudinal vein, one to five branches reach the wing margin
Media (M) – fourth longitudinal vein, one to four branches reach the wing margin
Cubitus (Cu) – fifth longitudinal vein, one to three branches reach the wing margin
Anal veins (A1, A2, A3) – unbranched veins behind the cubitus
The costa (C) is the leading marginal vein on most insects. Sometimes, there is a small vein above the costa called the precosta, although in almost all extant insects, the precosta is fused with the costa. The costa rarely ever branches because it is at the leading edge, which is associated at its base with the humeral plate. The trachea of the costal vein is perhaps a branch of the subcostal trachea. Located after the costa is the third vein, the subcosta, which branches into two separate veins: the anterior and posterior. The base of the subcosta is associated with the distal end of the neck of the first axillary (see section below). The fourth vein is the radius (R), which is branched into five separate veins. The radius is generally the strongest vein of the wing. Toward the middle of the wing, it forks into a first undivided branch (R1) and a second branch, called the radial sector (Ra), which subdivides dichotomously into four distal branches (R2, R3, R4, R5). Basally, the radius is flexibly united with the anterior end of the second axillary (2Ax).
The fifth vein of the wing is the media. In the archetype pattern (A), the media forks into two main branches: a media anterior (MA), which divides into two distal branches (MA1, MA2), and a median sector, or media posterior (MP), which has four terminal branches (M1, M2, M3, M4). In most modern insects the media anterior has been lost, and the usual "media" is the four-branched media posterior with the common basal stem. In the Ephemerida, according to present interpretations of the wing venation, both branches of the media are retained, while in Odonata the persisting media is the primitive anterior branch. The stem of the media is often united with the radius, but when it occurs as a distinct vein its base is associated with the distal median plate (m') or is continuously sclerotized with the latter. The cubitus, the sixth vein of the wing, is primarily two-branched. The primary forking of the takes place near the base of the wing, forming the two principal branches (Cu1, Cu2). The anterior branch may break up into a number of secondary branches, but commonly it forks into two distal branches. The second branch of the cubitus (Cu2) in Hymenoptera, Trichoptera, and Lepidoptera was mistaken by Comstock and Needham for the first anal. Proximally the main stem of the cubitus is associated with the distal median plate (m') of the wing base.
Postcubitus (Pcu) is the first anal of the Comstock–Needham system. The postcubitus, however, has the status of an independent wing vein and should be recognized as such. In nymphal wings, its trachea arises between the cubital trachea and the group of vannal tracheae. In the mature wings of more generalized insect the Postcubitus is always associated proximally with the cubitus and is never intimately connected with the flexor sclerite (3Ax) of the wing base. In Neuroptera, Mecoptera, and Trichoptera the postcubitus may be more closely associated with the vannal veins, but its base is always free from the latter. The postcubitus is usually unbranched; it is primitively two branched. The vannal veins (lV to nV) are the anal veins that are immediately associated with the third axillary, and which are directly affected by the movement of this sclerite that brings about the flexion of the wings. In number the vannal veins vary. from 1 to 12, according to the expansion of the vannal area of the wing. The vannal tracheae usually arise from a common tracheal stem in nymphal insects, and the veins are regarded as branches of a single anal vein. Distally the vannal veins are either simple or branched. Jugal Veins (J) of the jugal lobe of the wing is often occupied by a network of irregular veins, or it may be entirely membranous; but sometimes it contains one or two distinct small veins, the first jugal vein, or vena arcuata, and the second jugal vein, or vena cardinalis (2J).
C-Sc cross-veins – run between the costa and subcosta
R cross-veins – run between adjacent branches of the radius
R-M cross-veins – run between the radius and media
M-Cu cross-veins – run between the media and cubitus
All the veins of the wing are subject to secondary forking and to union by cross-veins. In some orders of insects the cross-veins are so numerous that the whole venational pattern becomes a close network of branching veins and cross-veins. Ordinarily, however, there is a definite number of cross-veins having specific locations. The more constant cross-veins are the humeral cross-vein (h) between costa and subcosta, the radial cross-vein (r) between R and the first fork of Rs, the sectorial cross-vein (s) between the two forks of R8, the median cross-vein (m–m) between M2 and M3, and the mediocubital cross-vein (m-cu) between media and cubitus.
The veins of insect wings are characterized by a convex-concave placement, such as those seen in mayflies (i.e., concave is "down" and convex is "up") which alternate regularly and by its triadic type of branching; whenever a vein forks there is always an interpolated vein of the opposite position between the two branches. A concave vein will fork into two concave veins (with the interpolated vein being convex) and the regular alteration of the veins is preserved. The veins of the wing appear to fall into an undulating pattern according to whether they have a tendency to fold up or down when the wing is relaxed. The basal shafts of the veins are convex, but each vein forks distally into an anterior convex branch and a posterior concave branch. Thus the costa and subcosta are regarded as convex and concave branches of a primary first vein, Rs is the concave branch of the radius, posterior media the concave branch of the media, Cu1 and Cu2 are respectively convex and concave, while the primitive Postcubitus and the first vannal have each an anterior convex branch and a posterior concave branch. The convex or concave nature of the veins has been used as evidence in determining the identities of the persisting distal branches of the veins of modern insects, but it has not been demonstrated to be consistent for all wings.
Fields.
Wing areas are delimited and subdivided by fold-lines along which the wing can fold, and flexion-lines along which the wing can flex during flight. The fundamental distinction between the flexion-lines and the fold-lines is often blurred, as fold-lines may permit some flexibility or vice versa. Two constants that are found in nearly all insect wings are the claval (a flexion-line) and jugal folds (or fold line); forming variable and unsatisfactory boundaries. Wing foldings can be very complicated, with transverse folding occurring in the hindwings of Dermaptera and Coleoptera, and in some insects the anal area can be folded like a fan.
There are about four different fields found on the insect wings:
Remigium
Anal area (vannus)
Jugal area
Axillary area
Alula
Most veins and crossveins occur in the anterior area of the remigium, which is responsible for most of the flight, powered by the thoracic muscles. The posterior portion of the remigium is sometimes called the clavus; the two other posterior fields are the anal and jugal areas. When the vannal fold has the usual position anterior to the group of anal veins, the remigium contains the costal, subcostal, radial, medial, cubital, and postcubital veins. In the flexed wing the remigium turns posteriorly on the flexible basal connection of the radius with the second axillary, and the base of the mediocubital field is folded medially on the axillary region along the plica basalis (bf) between the median plates (m, m') of the wing base.
The "vannus" is bordered by the vannal fold, which typically occurs between the postcubitus and the first vannal vein. In Orthoptera it usually has this position. In the forewing of Blattidae, however, the only fold in this part of the wing lies immediately before the postcubitus. In Plecoptera the vannal fold is posterior to the postcubitus, but proximally it crosses the base of the first vannal vein. In the cicada the vannal fold lies immediately behind the first vannal vein (lV). These small variations in the actual position of the vannal fold, however, do not affect the unity of action of the vannal veins, controlled by the flexor sclerite (3Ax), in the flexion of the wing. In the hindwings of most Orthoptera a secondary vena dividens forms a rib in the vannal fold. The vannus is usually triangular in shape, and its veins typically spread out from the third axillary like the ribs of a fan. Some of the vannal veins may be branched, and secondary veins may alternate with the primary veins. The vannal region is usually best developed in the hindwing, in which it may be enlarged to form a sustaining surface, as in Plecoptera and Orthoptera. The great fanlike expansions of the hindwings of Acrididae are clearly the vannal regions, since their veins are all supported on the third axillary sclerites on the wing bases, though Martynov (1925) ascribes most of the fan areas in Acrididae to the jugal regions of the wings. The true jugum of the acridid wing is represented only by the small membrane (Ju) mesad of the last vannal vein. The jugum is more highly developed in some other Polyneoptera, as in the Mantidae. In most of the higher insects with narrow wings the vannus becomes reduced, and the vannal fold is lost, but even in such cases the flexed wing may bend along a line between the postcubitus and the first vannal vein.
The "jugal region", or neala, is a region of the wing that is usually a small membranous area proximal to the base of the vannus strengthened by a few small, irregular veinlike thickenings; but when well developed it is a distinct section of the wing and may contain one or two jugal veins. When the jugal area of the forewing is developed as a free lobe, it projects beneath the humeral angle of the hindwing and thus serves to yoke the two wings together. In the Jugatae group of Lepidoptera it bears a long finger-like lobe. The jugal region was termed the neala ("new wing") because it is evidently a secondary and recently developed part of the wing.
The "axillary region" is the region containing the axillary sclerites and has in general the form of a scalene triangle. The base of the triangle (a-b) is the hinge of the wing with the body; the apex (c) is the distal end of the third axillary sclerite; the longer side is anterior to the apex. The point d on the anterior side of the triangle marks the articulation of the radial vein with the second axillary sclerite. The line between d and c is the plica basalis (bf), or fold of the wing at the base of the mediocubital field. The is the outer margin of the wing, between apex and hind or anal angle.
At the posterior angle of the wing base in some Diptera there is a pair of membranous lobes (squamae, or calypteres) known as the alula. The alula is well developed in the house fly. The outer squama (c) arises from the wing base behind the third axillary sclerite (3Ax) and evidently represents the jugal lobe of other insects (A, D); the larger inner squama (d) arises from the posterior scutellar margin of the tergum of the wing-bearing segment and forms a protective, hoodlike canopy over the haltere. In the flexed wing the outer squama of the alula is turned upside down above the inner squama, the latter not being affected by the movement of the wing. In many Diptera a deep incision of the anal area of the wing membrane behind the single vannal vein sets off a proximal alar lobe distal to the outer squama of the alula.
Joints.
The various movements of the wings, especially in insects that flex the wings horizontally over the back when at rest, demand a more complicated articular structure at the wing base than a mere hinge of the wing with the body. Each wing is attached to the body by a membranous basal area, but the articular membrane contains a number of small articular sclerites, collectively known as the pteralia. The pteralia include an anterior humeral plate at the base of the costal vein, a group of axillaries (Ax) associated with the subcostal, radial, and vannal veins, and two less definite median plates (m, m') at the base of the mediocubital area. The axillaries are specifically developed only in the wing-flexing insects, where they constitute the flexor mechanism of the wing operated by the flexor muscle arising on the pleuron. Characteristic of the wing base is also a small lobe on the anterior margin of the articular area proximal to the humeral plate, which, in the forewing of some insects, is developed into a large, flat, scale-like flap, the tegula, overlapping the base of the wing. Posteriorly the articular membrane often forms an ample lobe between the wing and the body, and its margin is generally thickened and corrugated, giving the appearance of a ligament, the so-called axillary cord, continuous mesally with the posterior marginal scutellar fold of the tergal plate bearing the wing.
The articular sclerites, or pteralia, of the wing base of the wing-flexing insects and their relations to the body and the wing veins, shown diagrammatically, are as follows:
 Humeral plates
 First Axillary
 Second Axillary
 Third Axillary
 Fourth Axillary
 Median plates (m, m')
The humeral plate is usually a small sclerite on the anterior margin of the wing base, movable and articulated with the base of the costal vein. Odonata have their humeral plate greatly enlarged, with two muscles arising from the episternum inserted into the Humeral plates and two from the edge of the epimeron inserted into the axillary plate.
The first axillary sclerite (lAx) is the anterior hinge plate of the wing base. Its anterior part is supported on the anterior notal wing process of the tergum (ANP); its posterior part articulates with the tergal margin. The anterior end of the sclerite is generally produced as a slender arm, the apex of which (e) is always associated with the base of the subcostal vein (Sc), though it is not united with the latter. The body of the sclerite articulates laterally with the second axillary. The second axillary sclerite (2Ax) is more variable in form than the first axillary, but its mechanical relations are no less definite. It is obliquely hinged to the outer margin of the body of the first axillary, and the radial vein (R) is always flexibly attached to its anterior end (d). The second axillary presents both a dorsal and a ventral sclerotization in the wing base; its ventral surface rests upon the fulcral wing process of the pleuron. The second axillary, therefore, is the pivotal sclerite of the wing base, and it specifically manipulates the radial vein.
The third axillary sclerite (3Ax) lies in the posterior part of the articular region of the wing. Its form is highly variable and often irregular, but the third axillary is the sclerite on which is inserted the flexor muscle of the wing (D). Mesally it articulates anteriorly (f) with the posterior end of the second axillary, and posteriorly (b) with the posterior wing process of the tergum (PNP), or with a small fourth axillary when the latter is present. Distally the third axillary is prolonged in a process which is always associated with the bases of the group of veins in the anal region of the wing here termed the vannal veins (V). The third axillary, therefore, is usually the posterior hinge plate of the wing base and is the active sclerite of the flexor mechanism, which directly manipulates the vannal veins. The contraction of the flexor muscle (D) revolves the third axillary on its mesal articulations (b, f) and thereby lifts its distal arm; this movement produces the flexion of the wing. The Fourth Axillary sclerite is not a constant element of the wing base. When present it is usually a small plate intervening between the third axillary and the posterior notal wing process and is probably a detached piece of the latter.
The median plates (m, m') are also sclerites that are not so definitely differentiated as specific plates as are the three principal axillaries, but nevertheless they are important elements of the flexor apparatus. They lie in the median area of the wing base distal to the second and third axillaries and are separated from each other by an oblique line (bf) which forms a prominent convex fold during flexion of the wing. The proximal plate (m) is usually attached to the distal arm of the third axillary and perhaps should be regarded as a part of the latter. The distal plate (m') is less constantly present as a distinct sclerite and may be represented by a general sclerotization of the base of the mediocubital field of the wing. When the veins of this region are distinct at their bases, they are associated with the outer median plate.
Muscles.
The muscles that control flight in insects can take up to 10% to 30% of the total body mass. The muscles that control flight vary with the two types of flight found in insects: indirect and direct. Insects that use first, indirect, have the muscles attach to the tergum instead of the wings, as the name suggests. As the muscles contract, the thoracic box becomes distorted, transferring the energy to the wing. There are two "bundles" of muscles, those that span parallel to the tergum, the dorsolongitudinals, and those that are attached to the tegum and extend to the sternum, the dorsoventrals. In direct muscle, the connection is directly from the pleuron (thoracic wall) to individual sclerites located at the base of the wing. The subalar and basilar muscles have ligament attachments to the subalar and basilar sclerites. Here resilin, a highly elastic material, forms the ligaments connecting flight muscles to the wing apparatus.
In more derived orders of insects, such as Diptera (flies) and Hymenoptera (wasp), the indirect muscles occupy the greatest volume of the pterothorax and function as the primary source of power for the wingstroke. Contraction of the dorsolongitudinal muscles causes the severe arching of the notum which depresses the wing while contraction of the dorsoventral muscles causes opposite motion of notum. The most primitive extant flying insects, Ephemeroptera (mayflies) and Odonata (dragonflies), use direct muscles that are responsible for developing the needed power for the up and down strokes.
Insect wing muscle is a strictly aerobic tissue. Per unit protein it consumes fuel and oxygen at rates taking place in a very concentrated and highly organized tissue so that the steady-state rates per unit volume represent an absolute record in biology. The fuel and oxygen rich blood is carried to the muscles through diffusion occurring in large amounts, in order to maintain the high level of energy used during flight. Many wing muscles are large and may be as large as 10 mm in length and 2 mm in width. Moreover, in some Diptera the fibres are of giant dimensions. For instance, in the very active "Rutilia", the cross-section is 1800 μm long and more than 500 μm wide. The transport of fuel and oxygen from the surroundings to the sites of consumption and the reverse transport of carbon dioxide therefore represent a challenge to the biologist both in relation to transport in the liquid phase and in the intricate system of air tubes, i.e. in the tracheal system.
Sensors.
Several types of sensory neurons are found on insect wings: gustatory bristles, mechanosensory bristles, campaniform sensilla, and chordotonal organs. These sensors provide the nervous system with both external and internal proprioceptive feedback necessary for effective flight and grooming.
Coupling, folding, and other features.
In many insect species, the forewing and hindwing can be coupled together, which improves the aerodynamic efficiency of flight by joining the forewing and hindwing into one bigger wing. The most common coupling mechanism (e.g., Hymenoptera and Trichoptera) is a row of small hooks on the forward margin of the hindwing, or "hamuli", which lock onto the forewing, keeping them held together (hamulate coupling). In some other insect species (e.g., Mecoptera, Lepidoptera, and some Trichoptera) the jugal lobe of the forewing covers a portion of the hindwing (jugal coupling), or the margins of the forewing and hindwing overlap broadly (amplexiform coupling), or the hindwing bristles, or frenulum, hook under the retaining structure or retinaculum on the forewing.
When at rest, the wings are held over the back in most insects, which may involve longitudinal folding of the wing membrane and sometimes also transverse folding. Folding may sometimes occur along the flexion lines. Though fold lines may be transverse, as in the hindwings of beetles and earwigs, they are normally radial to the base of the wing, allowing adjacent sections of a wing to be folded over or under each other. The commonest fold line is the jugal fold, situated just behind the third anal vein, although, most Neoptera have a jugal fold just behind vein 3A on the forewings. It is sometimes also present on the hindwings. Where the anal area of the hindwing is large, as in Orthoptera and Blattodea, the whole of this part may be folded under the anterior part of the wing along a vannal fold a little posterior to the claval furrow. In addition, in Orthoptera and Blattodea, the anal area is folded like a fan along the veins, the anal veins being convex, at the crests of the folds, and the accessory veins concave. Whereas the claval furrow and jugal fold are probably homologous in different species, the vannal fold varies in position in different taxa. Folding is produced by a muscle arising on the pleuron and inserted into the third axillary sclerite in such a way that, when it contracts, the sclerite pivots about its points of articulation with the posterior notal process and the second axillary sclerite.
As a result, the distal arm of the third axillary sclerite rotates upwards and inwards, so that finally its position is completely reversed. The anal veins are articulated with this sclerite in such a way that when it moves they are carried with it and become flexed over the back of the insect. Activity of the same muscle in flight affects the power output of the wing and so it is also important in flight control. In orthopteroid insects, the elasticity of the cuticle causes the vannal area of the wing to fold along the veins. Consequently, energy is expended in unfolding this region when the wings are moved to the flight position. In general, wing extension probably results from the contraction of muscles attached to the basilar sclerite or, in some insects, to the subalar sclerite.
Flight.
Flight mechanisms.
Two groups of relatively large insects, the Ephemeroptera (mayflies) and the Odonata (dragonflies and damselflies) have the flight muscles attached directly to their wings; the wings can beat no faster than the rate at which nerves can send impulses to command the muscles to beat. All other living winged insects fly using a different mechanism, involving indirect flight muscles which cause the thorax to vibrate; the wings can beat faster than the rate at which the muscles receive nerve impulses. This mechanism evolved once, and is the defining feature (synapomorphy) for the infraclass Neoptera.
Aerodynamics.
There are two basic aerodynamic models of insect flight. Most insects use a method that creates a spiralling leading edge vortex. Some very small insects use the fling and clap or Weis-Fogh mechanism in which the wings clap together above the insect's body and then fling apart. As they fling open, the air gets sucked in and creates a vortex over each wing. This bound vortex then moves across the wing and, in the clap, acts as the starting vortex for the other wing. Circulation and lift are increased, at the price of wear and tear on the wings.
Many insects can hover by beating their wings rapidly, requiring sideways stabilization as well as lift.
A few insects use gliding flight, without the use of thrust.
Evolution.
Sometime in the Carboniferous Period, some 350 million years ago, when there were only two major land masses, insects began flying. How and why insect wings developed, however, is not well understood, largely due to the scarcity of appropriate fossils from the period of their development in the Lower Carboniferous. Three main theories on the origins of insect flight are that wings developed from paranotal lobes, extensions of the thoracic terga; that they are modifications of movable abdominal gills as found on aquatic naiads of mayflies; or that they developed from thoracic protrusions used as radiators.
Fossils.
Fossils from the Devonian (400 million years ago) are all wingless, but by the Carboniferous (320 million years ago), more than 10 different genera of insects had fully functional wings. There is little preservation of transitional forms between the two periods. The earliest winged insects are from this time period (Pterygota), including the Blattoptera, Caloneurodea, primitive stem-group Ephemeropterans, Orthoptera and Palaeodictyopteroidea. Very early Blattopterans (during the Carboniferous) had a very large discoid pronotum and coriaceous forewings with a distinct CuP vein (an unbranched wing vein, lying near the claval fold and reaching the wing posterior margin). Even though the oldest possible insect fossil is the Devonian "Rhyniognatha hirsti", estimated at 396–407 million years old, it possessed dicondylic mandibles, a feature associated with winged insects, although it is later considered as possible myriapod.
During the Permian, the dragonflies (Odonata) were the dominant aerial predator and probably dominated terrestrial insect predation as well. True Odonata appeared in the Permian and all are amphibious. Their prototypes are the oldest winged fossils, go back to the Devonian, and are different from other wings in every way. Their prototypes may have had the beginnings of many modern attributes even by late Carboniferous and it is possible that they even captured small vertebrates, for some species had a wing span of 71 cm. The earliest beetle-like species during the Permian had pointed, leather like forewings with cells and pits. Hemiptera, or true bugs had appeared in the form of Arctiniscytina and Paraknightia having forewings with unusual venation, possibly diverging from Blattoptera.
A single large wing from a species of Diptera in the Triassic (10 mm instead of usual 2–6 mm) was found in Australia (Mt. Crosby).This family Tilliardipteridae, despite the numerous 'tipuloid' features, should be included in Psychodomorpha sensu Hennig on account of loss of the convex distal 1A reaching wing margin and formation of the anal loop.
Hypotheses.
Suggestions have been made that wings may have evolved initially for sailing on the surface of water as seen in some stoneflies. An alternative idea is that it derives from directed aerial gliding descent—a preflight phenomena found in some apterygote, a wingless sister taxa to the winged insects. The earliest fliers were similar to dragonflies with two sets of wings, direct flight muscles, and no ability to fold their wings over their abdomens. Most insects today, which evolved from those first fliers, have simplified to either one pair of wings or two pairs functioning as a single pair and using a system of indirect flight muscles.
Natural selection has played an enormous role in refining the wings, control and sensory systems, and anything else that affects aerodynamics or kinematics. One noteworthy trait is wing twist. Most insect wings are twisted, as are helicopter blades, with a higher angle of attack at the base. The twist generally is between 10 and 20 degrees. In addition to this twist, the wing surfaces are not necessarily flat or featureless; most larger insects have wing membranes distorted and angled between the veins in such a way that the cross-section of the wings approximates an airfoil. Thus, the wing's basic shape already is capable of generating a small amount of lift at zero angle of attack. Most insects control their wings by adjusting tilt, stiffness, and flapping frequency of the wings with tiny muscles in the thorax (below). Some insects evolved other wing features that are not advantageous for flight, but play a role in something else, such as mating or protection.
Some insects, occupying the biological niches that they do, need to be incredibly maneuverable. They must find their food in tight spaces and be capable of escaping larger predators – or they may themselves be predators, and need to capture prey. Their maneuverability, from an aerodynamic viewpoint, is provided by high lift and thrust forces. Typical insect fliers can attain lift forces up to three times their weight and horizontal thrust forces up to five times their weight. There are two substantially different insect flight mechanisms, and each has its own advantages and disadvantages – just because odonates have a more primitive flight mechanism does not mean they are less able fliers; they are, in certain ways, more agile than anything that has evolved afterward.
Morphogenesis.
While the development of wings in insects is clearly defined in those who are members of Endopterygota, which undergo complete metamorphosis; in these species, the wing develops while in the pupal stage of the insects life cycle. However, insects that undergo incomplete metamorphosis do not have a pupal stage, therefore they must have a different wing morphogenesis. Insects such as those that are hemimetabolic have wings that start out as buds, which are found underneath the exoskeleton, and do not become exposed until the last instar of the nymph.
The first indication of the wing buds is of a thickening of the hypodermis, which can be observed in insect species as early the embryo, and in the earliest stages of the life cycle. During the development of morphological features while in the embryo, or embryogenesis, a cluster of cells grow underneath the ectoderm which later in development, after the lateral ectoderm has grown dorsally to form wind imaginal disc. An example of wing bud development in the larvae, can be seen in those of White butterflies ("Pieris"). In the second instar the histoblast become more prominent, which now form a pocket-like structure. As of the third and fourth instars, the histoblast become more elongated. This greatly extended and evaginated, or protruding, part is what becomes the wing. By the close of the last instar, or fifth, the wing is pushed out of the wing-pocket, although continues to lie under the old larval cuticle while in its prepupal stage. It is not until the butterfly is in its pupal stage that the wing-bud becomes exposed, and shortly after eclosion, the wing begins to expand and form its definitive shape.
The development of tracheation of the wings begin before the wing histoblast form, as it is important to note that they develop near a large trachea. During the fourth instar, cells from the epithelium of this trachea become greatly enlarged extend into the cavity of the wing bud, with each cell having developed a closely coiled tracheole. Each trachcole is of unicellular origin, and is at first intracellular in position; while tracheae are of multicellular origin and the lumen of each is intercellular in position. The development of tracheoles, each coiled within a single cell of the epithelium of a trachea, and the subsequent opening of communication between the tracheoles and the lumen of the trachea, and the uncoiling and stretching out of the tracheoles, so that they reach all parts of the wing.
In the earlier stages of its development, the wing-bud is not provided with special organs of respiration such as tracheation, as it resembles in this respect the other portions of the hypodermis of which it is still a part. The histoblast is developed near a large trachea, a cross-section of which is shown in, which represents sections of these parts of the first, second, third and fourth instars respectively. At the same time the tracheoles uncoil, and extend in bundles in the forming vein-cavities of the wing-bud. At the molt that marks the beginning of the pupal stadium stage, they become functional. At the same time, the larval tracheoles degenerate; their function having been replaced by the wing tracheae.
Nomenclature.
Most of the nomenclature of insect orders is based on the Ancient Greek word for wing, (""), as the suffix "-ptera".
Adaptations.
Variation.
Insect wings are fundamental in identifying and classifying species as there is no other set of structures in studying insects more significant. Each order and insect family has distinctive wing shapes and features. In many cases, even species may be distinguished from each other by differences of color and pattern. For example, just by position one can identify species, albeit to a much lesser extent. Though most insects fold their wings when at rest, dragonflies and some damselflies rest with their wings spread out horizontally, while groups such as the caddisflies, stoneflies, alderflies, and lacewings hold their wings sloped roof-like over their backs. A few moths wrap their wings around their bodies, while many flies and most butterflies close their wings together straight upward over the back.
Many times the shape of the wings correlates with the type of insect flight. The best-flying insects tend to have long, slender wings. In many species of Sphingidae (sphinx moths), the forewings are large and sharply pointed, forming with the small hindwings a triangle that is suggestive of the wings of fast, modern airplanes. Another, possibly more important correlation, is that of the size and power of the muscles to the speed and power of flight. In the powerfully flying insects, the wings are most adapted for the stresses and aerodynamics of flight. The veins are thicker, stronger, and closer together toward the front edge (or "leading edge") and thinner yet flexible toward the rear edge (or "trailing edge"). This makes the insect wing an excellently constructed airfoil, capable of exerting both propulsion and lift while minimizing drag.
Variation of the wing beat may also occur, not just amongst different species, but even among individuals at different times. In general, the frequency is dependent upon the ratio between the power of the wing muscles and the resistance of the load. Large-winged, light-bodied butterflies may have a wing beat frequency of 4–20 per second whereas small-winged, heavy-bodied flies and bees beat their wings more than 100 times a second and mosquitoes can beat up to 988–1046 times a second. The same goes for flight; though it is generally difficult to estimate the speed of insects in flight, most insects can probably fly faster in nature than they do in controlled experiments.
Coleoptera.
In species of Coleoptera (beetles), the only functional wings are the hindwings. The hindwings are longer than the elytra, folded longitudinally and transversely under the elytra. The wing is rotated forwards on its base into flight position. This action spread the wing and unfolded longitudinally and transversely. There is the spring mechanism in the wing structure, sometimes with the help of abdomen movement, to keep the wing in folded position. The beetle wing venation is reduced and modified due to the folding structure, which include:
In most species of beetles, the front pair of wings are modified and sclerotised (hardened) to form elytra and they protect the delicate hindwings which are folded beneath. The elytra are connected to the pterathorax; being called as such because it is where the wings are connected ("pteron" meaning "wing" in Greek). The elytra are not used for flight, but tend to cover the hind part of the body and protect the second pair of wings ("alae"). The elytra must be raised in order to move the hind flight wings. A beetle's flight wings are crossed with veins and are folded after landing, often along these veins, and are stored below the elytra. In some beetles, the ability to fly has been lost. These include some ground beetles (family Carabidae) and some "true weevils" (family Curculionidae), but also some desert and cave-dwelling species of other families. Many of these species have the two elytra fused together, forming a solid shield over the abdomen. In a few families, both the ability to fly and the elytra have been lost, with the best known example being the glow-worms of the family Phengodidae, in which the females are larviform throughout their lives.
Lepidoptera.
The two pairs of wings are found on the middle and third segment, or mesothorax and metathorax respectively. In the more recent genera, the wings of the second segment are much more pronounced, however some more primitive forms have similarly sized wings of both segments. The wings are covered in scales arranged like shingles, forming the extraordinary variety seen in color. The mesothorax is evolved to have more powerful muscles to propel moth or butterfly through the air, with the wing of said segment having a stronger vein structure. The largest superfamily, Noctuidae, has the wings modified to act as tympanal or hearing organs Modifications in the wing's venation include:
The wings, head parts of thorax and abdomen of Lepidoptera are covered with minute scales, from which feature the order 'Lepidoptera' derives its names, the word "lepteron" in Ancient Greek meaning 'scale'. Most scales are lamellar, or blade-like and attached with a pedicel, while other forms may be hair-like or specialized as secondary sexual characteristics. The lumen or surface of the lamella, has a complex structure. It gives color either due to the pigmentary colors contained within or due to its three-dimensional structure. Scales provide a number of functions, which include insulation, thermoregulation, aiding gliding flight, amongst others, the most important of which is the large diversity of vivid or indistinct patterns they provide which help the organism protect itself by camouflage, mimicry, and to seek mates.
Odonata.
Species of Odonata (Damselflies and dragonflies) both have two pairs of wings which are about equal in size and shape and are clear in color. There are five, if the R+M is counted as 1, main vein stems on dragonfly and damselfly wings, and wing veins are fused at their bases and the wings cannot be folded over the body at rest, which also include:
The main veins and the crossveins form the wing venation pattern. The venation patterns are different in different species. There may be very numerous crossveins or rather few. The Australian Flatwing Damselfly's wings are one of the few veins patterns. The venation pattern is useful for species identification. Almost all Anisoptera settle with the wings held out sideways or slightly downward, however most Zygoptera settle with the wings held together, dorsal surfaces apposed. The thorax of Zygoptera is so oblique that when held in this way the wings fit neatly along the top of the abdomen. They do not appear to be held straight up as in butterflies or mayflies. In a few zygopteran families the wings are held horizontally at rest, and in one anisopteran genus (e.g. Cordulephya, Corduliidae) the wings are held in the typical damselfly resting position. Adult species possess two pairs of equal or subequal wings. There appear to be only five main vein stems. A nodus is formed where the second main vein (subcosta) meets the leading edge of the wing. In most families a conspicuous pterostigma is carried near the wing tip. Identification as Odonata can be based on the venation. The only likely confusion is with some lacewings (order Neuroptera) which have many crossveins in the wings. Until the early years of the 20th century Odonata were often regarded as being related to lacewings and were given the ordinal name Paraneuroptera, but any resemblance between these two orders is entirely superficial. In Anisoptera the hindwing is broader than the forewing and in both wings a crossvein divides the discoidal cell into a Triangle and Supertriangle.
Orthoptera.
Species of Orthoptera (grasshoppers and crickets) have forewings that are tough opaque tegmina, narrow which are normally covering the hindwings and abdomen at rest. The hindwings are board membranous and folded in fan-like manner, which include the following venation:
Phasmatodea.
Stick insect have forewings that are tough, opaque tegmina, short and covering only the base part of the hindwings at rest. Hindwings from costa to Cubitus are tough and opaque like the forewings. The large anal area are membranous and folded in fan-like manner. There are no or very few branching in stick insect wing veins.
Dermaptera.
Other orders such as the Dermaptera (earwigs), Orthoptera (grasshoppers, crickets), Mantodea (praying mantis) and Blattodea (cockroaches) have rigid leathery forewings that are not flapped while flying, sometimes called "tegmen" (pl. "tegmina"), "elytron" (pl. "elytra"), or "pseudoelytron".
Hemiptera.
In Hemiptera (true bugs), the forewings may be hardened, though to a lesser extent than in the beetles. For example, the anterior part of the front wings of stink bugs is hardened, while the posterior part is membranous. They are called "hemelytron" (pl. "hemelytra"). They are only found in the suborder Heteroptera; the wings of the Homoptera, such as the cicada, are typically entirely membranous. Both forewings and hindwings of Cicada are membranous. Most species are glass-like although some are opaque. Cicadas are not good fliers and most fly only a few seconds. When flying, forewing and hindwing are hooked together by a grooved coupling along the hindwing costa and forewing margin. Most species have a basic venation as shown in the following picture.
Also notice there are the ambient veins and peripheral membranes on the margin of both wings.
Diptera.
In the Diptera (true flies), there is only one pair of functional wings, with the posterior pair of wings are reduced to halteres, which help the fly to sense its orientation and movement, as well as to improve balance by acting similar to gyroscopes. In Calyptratae, the very hindmost portion of the wings are modified into somewhat thickened flaps called calypters which cover the halteres.
Blattodea.
Species of Blattodea (cockroaches) have a forewing, are also known as tegmen, that is more or less sclerotized. It is used in flight as well as a form of protection of the membranous hindwings. The veins of hindwing are about the same as front wing but with large anal lobe folded at rest between CuP and 1A. The anal lobe usually folded in a fan-like manner.
Hymenoptera.
Hymenoptera adults, including sawflies, wasps, bees, and non-worker ants, all have two pairs of membranous wings.
The forward margin of the hindwing bears a number of hooked bristles, or "hamuli", which lock onto the forewing, keeping them held together. The smaller species may have only two or three hamuli on each side, but the largest wasps may have a considerable number, keeping the wings gripped together especially tightly. Hymenopteran wings have relatively few veins compared with many other insects, especially in the smaller species.
Other families.
Termites are relatively poor fliers and are readily blown downwind in wind speeds of less than 2 km/h, shedding their wings soon after landing at an acceptable site, where they mate and attempt to form a nest in damp timber or earth. Wings of most termites have three heavy veins along the basal part of the front edge of the forewing and the crossveins near the wing tip are angled, making trapezoidal cells. Although subterranean termite wings have just two major veins along the front edge of the forewing and the cross veins towards the wingtip are perpendicular to these veins, making square and rectangular cells.
Species of Thysanoptera (thrips), Ptiliidae and other flying microinsects have slender front and hindwings with long fringes of hair, called fringed wings, also referred to as ptiloptery. While species of Trichoptera (caddisfly) have hairy wings with the front and hindwings clothed with setae.
Male Strepsiptera also have halteres that evolved from the forewings instead of the hindwings. This means that only their hindwings are functional at flying, as opposed to Diptera which have functional forewings and halteres for hindwings. Also the hindwings in males of Coccidae are reduced to halteres (or are absent).
Notes.
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Elytron
Hardened insect forewing
An elytron (; from grc " "ἔλυτρον" (élutron)" 'sheath, cover'; pl.: elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometimes alternatively spelled as "hemielytra"), and in most species only the basal half is thickened while the apex is membranous, but when they are entirely thickened the condition is referred to as "coleopteroid". An elytron is sometimes also referred to as a shard.
Description.
The elytra primarily serve as protective wing-cases for the hindwings underneath, which are used for flying. To fly, a beetle typically opens the elytra and then extends the hindwings, flying while still holding the elytra open, though many beetles in the families Scarabaeidae and Buprestidae can fly with the elytra closed (e.g., most Cetoniinae; ).
In a number of groups, the elytra are reduced to various degrees, (e.g., the beetle families Staphylinidae and Ripiphoridae), or secondarily lost altogether, as in various Elateroidea lineages with wingless females.
In some flightless groups, the elytra are present but fused together, and the hindwings are absent (e.g., some ground beetles (Carabidae), scarab beetles, and weevils).
References.
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Elytron (annelid anatomy)
In annelids, elytra (; from Greek ἔλυτρον "sheath, cover"; sg.: elytron ) are shield-like scales that are attached dorsally, one pair on each of a number of alternating segments and entirely or partly cover the dorsum. Elytra are modified dorsal cirri, and their number, size, location, and ornamentation are important taxonomic characters. The basal part of the elytra is known as the elytrophore; if (as is often the case) elytra are lost their presence is indicated by the elytrophore which is still present and visible.
Annelids possessing elytra are also known as "scale worms". Possession of elytra is characteristic of the annelid suborder Aphroditiformia.
References.
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Platypria echidna
Species of beetle
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Platypria (Platypria) echidna, is a species of leaf beetle found India, Sri Lanka, Myanmar and Vietnam.
Description.
Antenna thin, which is extending beyond scutellum over pronotum. There are six spines on each side of the anterior lateral lobe of the elytra. Elytra is covered with white pubescence. Elytral punctures are large and sub-quadrate. The anterior and posterior-lateral lobes on the elytra are reddish.
Biology.
Adult female lays a single egg covered with secretions that becomes an ootheca. Larva is a leaf miner that feed and live in mesophyll tissues. It is found in wide varieties of host plants including: "Mallotus", "Desmodium gangeticum", "Erythrina subumbrans", "Erythrina lithosperma", "Erythrina variegata", "Pueraria tuberosa", "Ziziphus mauritiana", "Ziziphus nummularia", "Ziziphus oenoplia", "Ziziphus rugosa", and "Ziziphus xylopyrus".
References.
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Insect morphology
Description of the physical form of insects
Insect morphology is the study and description of the physical form of insects. The terminology used to describe insects is similar to that used for other arthropods due to their shared evolutionary history. Three physical features separate insects from other arthropods: they have a body divided into three regions (called tagmata) (head, thorax, and abdomen), three pairs of legs, and mouthparts located "outside" of the head capsule. This position of the mouthparts divides them from their closest relatives, the non-insect hexapods, which include Protura, Diplura, and Collembola.
There is enormous variation in body structure amongst insect species. Individuals can range from 0.3 mm (fairyflies) to 30 cm across (great owlet moth); have no eyes or many; well-developed wings or none; and legs modified for running, jumping, swimming, or even digging. These modifications allow insects to occupy almost every ecological niche except the deep ocean. This article describes the basic insect body and some variations of the different body parts; in the process, it defines many of the technical terms used to describe insect bodies.
Anatomy summary.
Insects, like all arthropods, have no interior skeleton; instead, they have an exoskeleton, a hard outer layer made mostly of chitin that protects and supports the body. The insect body is divided into three parts: the head, thorax, and abdomen. The head is specialized for sensory input and food intake; the thorax, which is the anchor point for the legs and wings (if present), is specialized for locomotion; and the abdomen is for digestion, respiration, excretion, and reproduction. Although the general function of the three body regions is the same across all insect species, there are major differences in basic structure, with wings, legs, antennae, and mouthparts being variable from group to group.
External.
Exoskeleton.
The insect's outer skeleton, the cuticle, consists of two layers; the epicuticle, which is a thin, waxy, water-resistant outer layer that lacks chitin, and the layer under it is called the procuticle. This is chitinous and much thicker than the epicuticle and has two layers, the outer is the exocuticle while the inner is the endocuticle. The tough and flexible endocuticle is built from numerous layers of fibrous chitin and proteins, crisscrossing each other in a sandwich pattern, while the exocuticle is rigid and sclerotized. The exocuticle is greatly reduced in many soft-bodied insects, especially the larval stages (e.g., caterpillars). Chemically, chitin is a long-chain polymer of a N-acetylglucosamine, a derivative of glucose. In its unmodified form, chitin is translucent, pliable, and resilient. In arthropods, however, it is often modified, becoming embedded in a hardened proteinaceous matrix, which forms much of the exoskeleton. In its pure form, it is leathery, but when encrusted in calcium carbonate, it becomes much harder. The difference between the unmodified and modified forms is evident when comparing the body wall of a caterpillar (unmodified) to a beetle (modified).
From the embryonic stages, a layer of columnar or cuboidal epithelial cells gives rise to the external cuticle and an internal basement membrane. The majority of insect material is inside of the endocuticle. The cuticle provides muscular support and acts as a protective shield as the insect develops. However, since it cannot grow, the external sclerotized part of the cuticle is periodically shed in a process called "molting". As the time for molting approaches, most of the exocuticle material is reabsorbed. In molting, the old cuticle separates from the epidermis (apolysis). Enzymatic molting fluid is then released between the old cuticle and epidermis, which separates the exocuticle by digesting the endocuticle and sequestering its material for the new cuticle. When the new cuticle has formed sufficiently, the epicuticle and reduced exocuticle are shed in ecdysis.
The four principal regions of an insect body segment are the tergum or dorsal, sternum or ventral, and the two pleura or laterals. Hardened plates in the exoskeleton are called sclerites, which are subdivisions of the major regions – tergites, sternites, and pleurites, for respective regions tergum, sternum, and pleuron.
Head.
The head in most insects is enclosed in a hard, heavily sclerotized, exoskeletal head capsule. The main exception is in those species whose larvae are not fully sclerotized, mainly some holometabola; but even most unsclerotized or weakly sclerotized larvae tend to have well-sclerotized head capsules, for example, the larvae of Coleoptera and Hymenoptera. The larvae of Cyclorrhapha however, tend to have hardly any head capsule at all.
The head capsule bears most of the sensory organs, including the antennae, ocelli, and compound eyes, along with the mouthparts. In the adult insect, the head capsule appears unsegmented, though embryological studies show it to consist of six segments that bear the paired head appendages, including the mouthparts, each pair on a specific segment. Each such pair occupies one segment, though not all segments in modern insects bear any visible appendages.
Of all the insect orders, Orthoptera displays the greatest variety of features found in the heads of insects, including the sutures and sclerites. Here, the vertex, or the apex (dorsal region), is situated between the compound eyes of insects with hypognathous and opisthognathous heads. In prognathous insects, the vertex is not found between the compound eyes, but rather where the ocelli are normally found. This is because the primary axis of the head is rotated 90° to become parallel to the primary axis of the body. In some species, this region is modified and assumes a different name.
The ecdysial suture is made of the coronal, frontal, and epicranial sutures plus the ecdysial and cleavage lines, which vary among different species of insects. The ecdysial suture is longitudinally placed on the vertex, separating the epicranial halves of the head to the left and right sides. Depending on the insect, the suture may come in different shapes: like either a Y, U or V. Those diverging lines that make up the ecdysial suture are called the frontal or frontogenal sutures. Not all species of insects have frontal sutures, but in those that do, the sutures split open during ecdysis, which provides an opening for the new instar to emerge from the integument.
The frons is that part of the head capsule that lies ventrad or anteriad of the vertex. The frons varies in size relative to the insect, and in many species, the definition of its borders is arbitrary, even in some insect taxa that have well-defined head capsules. In most species, though, the frons is bordered at its anterior by the frontoclypeal or epistomal sulcus above the clypeus. Laterally it is limited by the fronto-genal sulcus, if present, and the boundary with the vertex, by the ecdysial cleavage line, if it is visible. If there is a median ocellus, it generally is on the frons, though in some insects such as many Hymenoptera, all three ocelli appear on the vertex. A more formal definition is that it is the sclerite from which the pharyngeal dilator muscles arise, but in many contexts that too, is not helpful. In the anatomy of some taxa, such as many Cicadomorpha, the front of the head is fairly clearly distinguished and tends to be broad and sub-vertical; that median area commonly is taken to be the frons.
The clypeus is a sclerite between the face and labrum, which is dorsally separated from the frons by the frontoclypeal suture in primitive insects. The clypeogenal suture laterally demarcates the clypeus, with the clypeus ventrally separated from the labrum by the clypeolabral suture. The clypeus differs in shape and size, such as species of Lepidoptera with a large clypeus with elongated mouthparts. The cheek or gena forms the sclerotized area on each side of the head below the compound eyes extending to the gular suture. Like many parts making up the insect's head, the gena varies among species, with its boundaries difficult to establish. In dragonflies and damselflies, it is between the compound eyes, clypeus, and mouthparts. The postgena is the area immediately posteriad, or posterior or lower on the gena of pterygote insects, and forms the lateral and ventral parts of the occipital arch. The occipital arch is a narrow band forming the posterior edge of the head capsule arching dorsally over the foramen. The subgenal area is usually narrow, located above the mouthparts; this area also includes the hypostoma and pleurostoma. The vertex extends anteriorly above the bases of the antennae as a prominent, pointed, concave rostrum. The posterior wall of the head capsule is penetrated by a large aperture, the foramen. Through it passes the organ systems, such as the nerve cord, esophagus, salivary ducts, and musculature, connecting the head with the thorax.
On the posterior aspect of the head are the occiput, postgena, occipital foramen, posterior tentorial pit, gula, postgenal bridge, hypostomal suture and bridge, and the mandibles, labium, and maxilla. The occipital suture is well-founded in species of Orthoptera, but not so much in other orders. Where found, the occipital suture is the arched, horseshoe-shaped groove on the back of the head ending at the posterior of each mandible. The postoccipital suture is a landmark on the posterior surface of the head, and is typically near the occipital foremen. In pterygotes, the postocciput forms the extreme posterior, often U-shaped, which forms the rim of the head extending to the postoccipital suture. In pterygotes, such as those of Orthoptera, the occipital foramen and the mouth are not separated. The three types of occipital closures, or points under the occipital foramen that separate the two lower halves of the postgena, are the hypostomal bridge, the postgenal bridge, and the gula. The hypostomal bridge is usually found in insects with hypognathous orientation. The postgenal bridge is found in the adults of species of higher Diptera and aculeate Hymenoptera, while the gula is found on some Coleoptera, Neuroptera, and Isoptera, which typically display prognathous-oriented mouthparts.
Compound eyes and ocelli.
Most insects have one pair of large, prominent compound eyes composed of units called ommatidia (ommatidium, singular), up to 30,000 in a single compound eye of, for example, large dragonflies. This type of eye gives less resolution than eyes found in vertebrates, but it gives an acute perception of movement and usually possesses UV- and green sensitivity, and may have additional sensitivity peaks in other regions of the visual spectrum. Often an ability to detect the E-vector of polarized light exists in polarization of light. There can also be an additional two or three ocelli, which help detect low light or small changes in light intensity. The image perceived is a combination of inputs from the numerous ommatidia, located on a convex surface, thus pointing in slightly different directions. Compared with simple eyes, compound eyes possess very large view angles and better acuity than the insect's dorsal ocelli, but some stemmatal (= larval eyes), for example, those of sawfly larvae (Tenthredinidae) with an acuity of 4 degrees and very high polarization sensitivity, match the performance of compound eyes.
Because the individual lenses are so small, the effects of diffraction impose a limit on the possible resolution that can be obtained (assuming they do not function as phased arrays). This can only be countered by increasing lens size and number. To see with a resolution comparable to our simple eyes, humans would require compound eyes that would each reach the size of their heads. Compound eyes fall into two groups: apposition eyes, which form multiple inverted images, and superposition eyes, which form a single erect image. Compound eyes grow at their margins with the addition of new ommatidia.
Antennae.
Antennae, sometimes called "feelers", are flexible appendages located on the insect's head which are used for sensing the environment. Insects "can" feel with their antennae because of the fine hairs (setae) that cover them. However, touch is not the only thing that antennae can detect; numerous tiny sensory structures on the antennae allow insects to sense smells, temperature, humidity, pressure, and even potentially sense themselves in space. Some insects, including bees and some groups of flies, can also detect sound with their antennae.
The number of segments in an antenna varies amongst insects, with higher flies having 3-6 segments, while adult cockroaches can have over 140. The general shape of the antennae is also quite variable, but the first segment (the one attached to the head) is always called the scape, and the second segment is called the pedicel. The remaining antennal segments or flagellomeres are called the flagellum.
General insect antenna types are shown below:
Mouthparts.
The insect mouthparts consist of the maxilla, labium, and in some species, the mandibles. The labrum is a simple, fused sclerite, often called the upper lip, and moves longitudinally. It is hinged to the clypeus. The mandibles (jaws) are a highly sclerotized pair of structures that move at right angles to the body, used for biting, chewing, and severing food. The maxillae are paired structures that can also move at right angles to the body and possess segmented palps. The labium (lower lip) is the fused structure that moves longitudinally and has a pair of segmented palps.
The mouthparts and rest of the head can be articulated in at least three different positions: prognathous, opisthognathous, and hypognathous. In species with prognathous articulation, the head is vertically aligned with the body, such as species of Formicidae; while in a hypognathous type, the head is aligned horizontally adjacent to the body. An opisthognathous head is positioned diagonally, such as in species of Blattodea and some Coleoptera. The mouthparts vary greatly between insects of different orders, but the two main functional groups are mandibulate and haustellate. Haustellate mouthparts are used for sucking liquids and can be further classified by the presence of stylets, which include piercing-sucking, sponging, and siphoning. The stylets are needle-like projections used to penetrate plant and animal tissues. The stylets and the feeding tube form the modified mandibles, maxilla, and hypopharynx.
Mandibular mouthparts are found in species of Odonata, adult Neuroptera, Coleoptera, Hymenoptera, Blattodea, Orthoptera, and Lepidoptera. However, most adult Lepidoptera have siphoning mouthparts, while their larvae (commonly called caterpillars) have mandibles.
Mandibulate.
The labrum is a broad lobe forming the roof of the preoral cavity, suspended from the clypeus in front of the mouth and forming the upper lip. On its inner side, it is membranous and may be produced into a median lobe, the epipharynx, bearing some sensilla. The labrum is raised away