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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.
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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).
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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.
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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.
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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.
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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.
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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.
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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).
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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.
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which are generally considered as corresponding structures to insect wings, articulated within the dorsal tergite plates. This cannot be seen in modern mayfly larvae, because their abdominal tergites and sternites are fused to rings, without any traces left even in embryonic development. If larval gills and wings are corresponding ("serial homologous") structures and thus share the same evolutionary origin, the new results from
Coxoplectoptera demonstrate that also wings are of tergal origin, as proposed by the classical paranotal-hypothesis. Staniczek, Bechly & Godunko (2011) therefore suggested a new hypothesis that could reconcile the apparently conflicting evidence from
2374:(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
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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.
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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.
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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.
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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.
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1431:. 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.
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566:. 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 remigiumturns 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.
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839:) 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 (
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1357:– 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
2406:, 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:
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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.
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91:
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36:
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3068:, 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.
895:, 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
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707:(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,
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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.
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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.
725:, 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.
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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.
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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.
3443:", 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.
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2680:(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:
3254:) 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.
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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.
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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.
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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
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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
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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
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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
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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
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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
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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
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The axillary region is region containing the axillary sclerites 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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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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
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variation displayed a mutation transforming normal wings to what was interpreted as a triple-jointed leg arrangement with some additional appendages but lacking the tarsus, where the wing's costal surface normally would be. This mutation was reinterpreted as strong evidence for a dorsal exite and
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and enable the insect to land more softly. The theory suggests that these lobes gradually grew larger and in a later stage developed a joint with the thorax. Even later would appear the muscles to move these crude wings. This model implies a progressive increase in the effectiveness of the wings,
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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
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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
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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
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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
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The front and rear wings at rest: The front wing covers most of the hindwing, with only the joint projects in the form of a quarter circle forward with a central white spot under the forewing. On the right hand side of the forewing is opened to the right (blue arrow), which from this perspective
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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
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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
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of the insect wing has been universally considered as conclusive evidence in favour of the leg-exite-hypothesis, which proposes that insect wings are derived from mobile leg appendages (exites). However, the larvae of Coxoplectoptera show that the abdominal gills of mayflies and their ancestors,
1043:(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.
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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
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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
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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
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The front wing is open to the left (blue arrow) with the right side of the forewing removed; the hindwing is half open. With greater resolution, the multiple folding is shown, resembling a fan which is parallel to the lines b and c. The arrow points to the e point where the fan is closed again,
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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
800:
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.
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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
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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
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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
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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
951:(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 (
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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
2439:
The same wing, half folded: The two joints of the cross-folding form an obtuse angle. The right is already in the wings folded in three layers. With greater resolution, the third arch of the wing margin in the first and second is visible. To the left of the fifth arch appears in the
2534:), 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
328:, 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
2526:). 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
793:) 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.
277:
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
352:
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
1154:
gills, which started their way as exits of the respiratory system and over time were modified into locomotive purposes, eventually developed into wings. The tracheal gills are equipped with little winglets that perpetually vibrate and have their own tiny straight
1223:), and only later in evolution became mobile, articulated appendages through secondary recruiting of leg genes. More recent fossil analysis of Paleozoic nymph wing pads provides additional support for the fusion of the paranota elements and arthopodan leg genes.
2660:. 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
1396:
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
2815:(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:
1118:
that is believed to have assisted stabilization while hopping or falling. In favor of this hypothesis is the tendency of most insects, when startled while climbing on branches, to escape by dropping to the ground. Such lobes would have served as
3345:
The main fold line of the forewing seen halfway up as a bright horizontal line. The wing part that is behind this line is turned back down. The narrow strip at the front edge of the wing is in front of the first strong wire folded forward and
140:, 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
2742:– third and fourth longitudinal vein, the strongest vein on the wing, with branches, R1-R4, reach the wing margin, the media anterior (MA) are also reach the wing margin. IR2 and IR3 are intercalary veins behind R2 and R3 respectively.
234:
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
768:
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.,
226:
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
578:
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
1161:: This hypothesis stems from the adaptation of endites and exites, appendages on the respective inner and outer aspects of the primitive arthropod limb. It was advanced by Trueman based on a study by Goldschmidt in 1945 on
1019:
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.
1172:
endite fusion, rather than a leg, with the appendages fitting in much better with this hypothesis. The innervation, articulation and musculature required for the evolution of wings are already present in podomeres.
4876:
Riek EF Kukalova-Peck J (1984). "A new interpretation of dragonfly wing venation based on early Upper Carboniferous fossils from Argentina (Insecta: Odonatoida and basic character states in Pterygote wings.)".
1136:. Still, lack of substantial fossil evidence of the development of the wing joints and muscles poses a major difficulty to the theory, as does the seemingly spontaneous development of articulation and venation.
4469:
Dinges, G.F., Chockley, A.S., Bockemühl, T., Ito, K., Blanke, A. and Büschges, A., 2021. Location and arrangement of campaniform sensilla in Drosophila melanogaster. Journal of Comparative Neurology, 529(4),
3800:
Almudi, Isabel; Vizueta, Joel; Wyatt, Christopher D. R.; de Mendoza, Alex; Marlétaz, Ferdinand; Firbas, Panos N.; Feuda, Roberto; Masiero, Giulio; Medina, Patricia; Alcaina-Caro, Ana; Cruz, Fernando (2020).
4460:
Valmalette, J.C., Raad, H., Qiu, N., Ohara, S., Capovilla, M. and Robichon, A., 2015. Nano-architecture of gustatory chemosensory bristles and trachea in Drosophila wings. Scientific reports, 5(1), pp.1-11.
1416:
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
2643:
There is the enlargement of the humeral area of the hindwing which is overlapped with the forewing. The humeral vein strengthened the hindwing overlapped area so that the two wings coupling better.
2367:
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.
1405:
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 (
1243:. 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.
1142:: This theory, first proposed in 1870 by Carl Gegenbaur, suggested that a possible origin for insect wings might have been the movable abdominal gills found in many aquatic insects, such as on
3360:
So in rest position, the outer lining forms the tough outer edge of the wing, which protects the sides of the abdomen as a shock absorber. The rear wing is covered largely by the forewing.
4488:
Wolf, H., 1993. The locust tegula: significance for flight rhythm generation, wing movement control and aerodynamic force production. Journal of Experimental Biology, 182(1), pp.229-253.
677:
2603:– radius divides into branches beyond the middle of the wing up to five branches in Papilionidae. On forewing, the last R is stalked in all butterflies except Hesperiidae is separated.
1427:. 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
979:
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
1389:. 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
148:). 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
211:. 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
2514:
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 (
4511:
605:
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:
3463:
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 (
2791:) 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
2571:
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,
1353:
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
2597:– at the leading wing marginal, fused or very close for most of the length, in hindwing fused and well developed in the humeral area, subcosta never branches in butterfly.
5527:
5074:
4031:
3917:
3786:
3723:
3689:
3660:
219:
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.
5654:
988:, estimated at 396–407 million years old, it possessed dicondylic mandibles, a feature associated with winged insects, although it is later considered as possible
1361:
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.
3866:
Bruce, Heather, N.H. Patel. (2020). "Knockout of crustacean leg patterning genes suggests that insect wings and body walls evolved from ancient leg segments".
2343:
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,
2554:
2370:
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
1186:
and the leg-exite-hypothesis have been considered as incompatible alternative explanations, which have both been supported by a set of evidences from the
500:
fold line); forming variable and unsatisfactory boundaries. Wing foldings can be very complicated, with transverse folding occurring in the hindwings of
5027:"Coxoplectoptera, a new fossil order of Palaeoptera (Arthropoda: Insecta), with comments on the phylogeny of the stem group of mayflies (Ephemeroptera)"
6010:
4970:
V. A. Blagoderov; E. D. Lukashevich; M. B. Mostovski (2002). "Order Diptera Linné, 1758. The true flies". In A. P. Rasnitsyn; D. L. J. Quicke (eds.).
3225:- unbranched, CuP is reduced in Diptera. Some species CuA and 1A are separated, some species meets when reaching the wing margin, some species fused.
5026:
1007:) were the dominant aerial predator and probably dominated terrestrial insect predation as well. True Odonata appeared in the Permian and all are
3044:(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
5088:
Prokop, Jakub; Pecharová, Martina; Nel, André; Hörnschemeyer, Thomas; Krzemińska, Ewa; Krzemiński, Wiesław; Engel, Michael S. (January 2017).
558:, which is responsible for most of the flight, powered by the thoracic muscles. The posterior portion of the remigium is sometimes called the
257:; or that insect wings arose from the fusion of pre-existing endite and exite structures each with pre-existing articulation and tracheation.
6015:
5329:
4736:
4666:
4525:
4517:
1423:
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
5489:"Wing morphology in featherwing beetles (Coleoptera: Ptiliidae): Features associated with miniaturization and functional scaling analysis"
4243:
4479:
Field, L.H. and Matheson, T., 1998. Chordotonal organs of insects. In Advances in insect physiology (Vol. 27, pp. 1-228). Academic Press.
1182:
provided important new clues to the disputed question of the evolutionary origin of insect wings. Before the larvae fossil discovery the
5647:
5620:
4123:
332:(grasshoppers and crickets). Large numbers of cross-veins are present in some insects, and they may form a reticulum as in the wings of
3478:
that evolved from the forewings instead of the hindwings. This means that only their hindwings are functional at flying, as opposed to
1015:, and are different from other wings in every way. Their prototypes may have had the beginnings of many modern attributes even by late
4497:
Zhang, N. and Simpson, J.H., 2022. A pair of commissural command neurons induces Drosophila wing grooming. Iscience, 25(2), p.103792.
5599:
5453:
4985:
4226:
4172:
4097:
4067:
1286:(below). Some insects evolved other wing features that are not advantageous for flight, but play a role in something else, such as
4381:"The Flight Muscles of Insects-Their Anatomy and Histology; with Some Observations on the Structure of Striated Muscle in General"
508:, and in some insects the anal area can be folded like a fan. There are about four different fields found on the insect wings:
5640:
5469:
3933:"Insights into insect wing origin provided by functional analysis of vestigial in the red flour beetle, Tribolium castaneum"
5752:
3423:– Row of hooks on the leading edge of hindwing engage the hind margin of the forewing, strongly couple the wings in flight.
2926:– veins behind the cubitus, unbranched, two in forewing, many in hindwing 1A-7A in one group and the rest in another group.
3535:
Crampton, G. (1916). "The Phylogenetic Origin and the Nature of the Wings of Insects According to the Paranotal Theory".
4531:
579:
559:
353:
318:
5774:
2856:– fifth longitudinal vein, on forewing and hindwing dividing near the wing base into branched CuA, and unbranched CuP.
1227:
Suggestions have been made that wings may have evolved initially for sailing on the surface of water as seen in some
781:", which lock onto the forewing, keeping them held together (hamulate coupling). In some other insect species (e.g.,
3429:– Some species, including Vespidae, the forewing are longitudinally folded along the 'line of wing folding' at rest.
933:
715:(dragonflies), use direct muscles that are responsible for developing the needed power for the up and down strokes.
5866:
5702:
4977:
5015:
Trueman JWH (1990), Comment: evolution of insect wings: a limb exite plus endite model Canadian Journal of Zoology
4604:
1231:. An alternative idea is that it derives from directed aerial gliding descent—a preflight phenomena found in some
191:. In some cases, wings are produced only at particular times in the life cycle, such as in the dispersal phase of
3990:
Prokop, Jakub, Pecharová, M., Nel, A., Hörnschemeyer, T., Krzemińska, E., Krzemiński, W., & Engel, M (2017).
3165:, which help the fly to sense its orientation and movement, as well as to improve balance by acting similar to
1261:. One noteworthy trait is wing twist. Most insect wings are twisted, as are helicopter blades, with a higher
861:
There are two basic aerodynamic models of insect flight. Most insects use a method that creates a spiralling
6041:
3500:
1283:
1163:
918:
896:
236:
121:
4863:
Grzimek HC Bernhard (1975) Grzimek's Animal Life Encyclopedia Vol 22 Insects. Van Nostrand Reinhold Co. NY.
2425:
The hindwing, spread: by folding lines, it is divided into five fields that are completed each to the rear.
939:
6046:
6005:
5949:
5944:
5929:
5488:
5057:
3097:– in forewing Sc and R fused together to the node. Radial sector (Rs) arises near the node and unbranches.
1216:
1191:
876:
Many insects can hover by beating their wings rapidly, requiring sideways stabilization as well as lift.
6020:
5068:
4025:
3911:
3780:
3654:
1195:
972:
223:
appendages of legs, and lateral (paranotal) and posterolateral projections of the thorax to name a few.
5129:
Adrian L. R. Thomas; R. Åke Norberg (1996). "Skimming the surface — the origin of flight in insects?".
3609:
Averof, Michalis, and S. M. Cohen. (1997). "Evolutionary origin of insect wings from ancestral gills".
5487:
Polilov, Alexey A.; Reshetnikova, Natalia I.; Petrov, Pyotr N.; Farisenkov, Sergey E. (January 2019).
3370:
adults, including sawflies, wasps, bees, and non-worker ants, all have two pairs of membranous wings.
1454:
5959:
5871:
5500:
4763:
4574:
4392:
3944:
3875:
3814:
3618:
1737:
Insects that undergo incomplete metamorphosis (and thus have externally visible wing buds as nymphs)
1424:
1151:
892:
749:
5362:
2510:
In most species of beetles, the front pair of wings are modified and sclerotised (hardened) to form
1025:
5954:
5881:
2648:
984:
856:
496:
5615:
5292:
5246:
5227:
1057:
1029:
265:
5969:
5934:
5687:
5395:
5049:
4787:
4637:
4559:
4410:
4244:"A New Method of Studying the Wing Veins of the Mayflies and Some Results Therefrom (Ephemerida)"
3899:
3768:
3717:
3683:
3642:
3544:
1407:
813:
753:
4430:"Chemosensory sensilla of the Drosophila wing express a candidate ionotropic pheromone receptor"
1412:
563:
231:
arthropods with a direct crustacean ancestor and shared genetic mechanisms of limb development.
4254:
3161:), there is only one pair of functional wings, with the posterior pair of wings are reduced to
1270:. Thus, the wing's basic shape already is capable of generating a small amount of lift at zero
809:
492:
5985:
5964:
5924:
5804:
5722:
5595:
5568:
5550:
5449:
5387:
5325:
5195:
5146:
5111:
4981:
4925:
4846:
4828:
4779:
4732:
4662:
4629:
4521:
4361:
4222:
4168:
4093:
4063:
4013:
3972:
3891:
3848:
3830:
3760:
3634:
3588:
3145:
Also notice there are the ambient veins and peripheral membranes on the margin of both wings.
3045:
2006:
1246:
1008:
683:
555:
5528:"General Biology and Current Management Approaches of Soft Scale Pests (Hemiptera: Coccidae)"
5443:
4724:
4127:
4087:
2454:
The same wing, folded completely: The five fields are aligned (The elytra have been removed).
1239:
with two sets of wings, direct flight muscles, and no ability to fold their wings over their
5995:
5769:
5558:
5542:
5508:
5377:
5185:
5177:
5138:
5101:
5041:
4952:
4917:
4886:
4836:
4818:
4771:
4754:
4619:
4582:
4441:
4400:
4353:
4271:
4003:
3962:
3952:
3883:
3838:
3822:
3750:
3626:
3578:
2993:
2804:
2657:
2543:
1598:
975:. Very early Blattopterans (during the Carboniferous) had a very large discoid pronotum and
880:
869:
865:
745:
149:
4586:
4341:
3803:"Genomic adaptations to aquatic and aerial life in mayflies and the origin of insect wings"
3482:
which have functional forewings and halteres for hindwings. Also the hindwings in males of
3173:, the very hindmost portion of the wings are modified into somewhat thickened flaps called
2750:– fifth longitudinal vein, cubitus posterior (CuP) is unbranched and reach the wing margin.
5939:
5843:
5826:
5697:
3257:
2701:
2689:
1271:
1262:
1179:
1143:
910:
734:
594:
250:
196:
157:
55:
4164:
1369:
While the development of wings in insects is clearly defined in those who are members of
5504:
5221:
4767:
4578:
4428:
He, Zhe; Luo, Yichen; Shang, Xueying; Sun, Jennifer S.; Carlson, John R. (21 May 2019).
4396:
4385:
Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences
4323:
3948:
3879:
3818:
3622:
2721:– at the leading edge of the wing, strong and marginal, extends to the apex of the wing.
1176:
Paranota plus leg gene recruitment hypothesis (also known as the dual origin hypothesis)
313:
6000:
5891:
5814:
5761:
5732:
5692:
5667:
5563:
5318:
5190:
5165:
4841:
4806:
4057:
3967:
3932:
3843:
3802:
3084:– at the leading wing marginal, in forewing extends to the node and lies close to Sc+R.
3001:
2873:
2651:, from which feature the order 'Lepidoptera' derives its names, the word "lepteron" in
2576:
2432:
2383:
2375:
1957:
Includes all currently living orders of flying insects except mayflies and dragonflies
1862:
1382:
1374:
1254:
1250:
757:
337:
3071:
687:
and their structural relationship to the tube-like heart (also in green). Red depicts
6035:
5821:
5737:
5727:
5142:
3903:
2652:
2531:
2527:
2519:
2189:
1702:
1448:
1398:
1386:
1370:
1183:
1111:
1110:: This hypothesis suggests that the insect's wings developed from paranotal lobes, a
1052:
1016:
964:
943:. 49.5 Million Years old; "Boot Hill", Klondike Mountain Formation, Washington, USA.
821:
708:
176:
117:
5399:
4905:
4641:
4295:
3772:
3008:) have rigid leathery forewings that are not flapped while flying, sometimes called
2522:, but tend to cover the hind part of the body and protect the second pair of wings (
1235:, a wingless sister taxa to the winged insects. The earliest fliers were similar to
484:
5886:
5053:
4791:
4752:
Michael S. Engel; David A. Grimaldi (2004). "New light shed on the oldest insect".
4555:
3646:
3471:
3417:– only two anal veins 1A and 2A are present, 2A is not distinctive in some species.
3233:– only two anal veins 1A and 2A are present, 2A is not distinctive in some species.
3141:– veins behind the cubitus, 1A and 2A fused in the forewing, CuP and 2A are folded.
2765:
is formed where the second main vein meets the leading edge of the wing. The black
2647:
The wings, head parts of thorax and abdomen of Lepidoptera are covered with minute
2418:
2379:
2251:
2231:
2209:
2123:
2077:
2052:
1722:
1682:
1444:
1212:
960:
862:
840:
348:
325:
228:
172:
3338:
3180:
2402:), the only functional wings are the hindwings. The hindwings are longer than the
90:
5473:
4656:
4446:
4429:
4157:
3300:– fifth longitudinal vein, with dichotomous branches occupy large part of tegmen.
5848:
5789:
5784:
5779:
3739:"Evolution of insect wings and development – new details from Palaeozoic nymphs"
3710:
On the Wing: Insects, Pterosaurs, Birds, Bats and the Evolution of Animal Flight
3367:
3170:
3057:
2989:
2788:
2784:
2766:
2539:
2447:
2271:
2032:
1902:
1842:
1802:
1762:
1618:
1125:
1033:
1000:
956:
790:
786:
774:
770:
692:
688:
113:
35:
3937:
Proceedings of the National Academy of Sciences of the United States of America
3887:
3826:
3353:
5799:
5742:
5712:
5707:
5512:
5382:
5106:
5089:
4008:
3991:
3583:
3566:
2985:
2902:– third longitudinal vein, branched to Rs in hindwing, unbranched in forewing.
2812:
2775:
2661:
2640:
2567:
2563:
2371:
2356:
2149:
2103:
1986:
1922:
1658:
1578:
1558:
1532:
1524:
1491:
1258:
1232:
976:
968:
505:
501:
341:
329:
307:
168:
167:
The wings are present in only one sex (often the male) in some groups such as
145:
129:
125:
79:
5554:
5045:
4971:
4832:
3834:
2184:
In class, unlike Apterygota, including winged and wingless secondary systems
5903:
5876:
3957:
3495:
3464:
3460:
3251:
3174:
3166:
3061:
3041:
3005:
2910:– fourth longitudinal vein, branched in basal part as Media posterior (MP).
2848:– fourth longitudinal vein, branched in basal part as Media posterior (MP).
2779:
2572:
2352:
2348:
2344:
2311:
2291:
2169:
1882:
1782:
1742:
1428:
1354:
1275:
1236:
1120:
1021:
989:
952:
782:
738:
208:
97:
45:
5572:
5391:
5199:
5181:
5150:
5115:
4850:
4783:
4633:
4405:
4380:
4365:
4126:. Department of Entomology, North Carolina State University. Archived from
4017:
3976:
3895:
3852:
3764:
3592:
2778:
settle with the wings held out sideways or slightly downward, however most
756:. These sensors provide the nervous system with both external and internal
5546:
4929:
4921:
3638:
3439:
The forward margin of the hindwing bears a number of hooked bristles, or "
2964:
2948:
2886:– at the leading marginal of the hindwing, unbranched, absent in forewing.
2864:– veins behind the cubitus, unbranched, two in forewing, many in hindwing.
5908:
5831:
5809:
5717:
4357:
3676:
Form and Function of Insect Wings: The Evolution of Biological Structures
3483:
3475:
3401:– M is unbranched, in forewing M is fused with Rs for part of its length.
2997:
2535:
2364:
2360:
1942:
1822:
1417:
1402:
1228:
1220:
1207:
1199:
1040:
1012:
980:
948:
922:
844:
741:
291:
279:
212:
188:
61:
5632:
4956:
4775:
3548:
2956:
appears narrower than it is with the rear wing still folded completely.
2351:
rest with their wings spread out horizontally, while groups such as the
17:
5836:
5090:"Paleozoic Nymphal Wing Pads Support Dual Model of Insect Wing Origins"
4823:
4414:
3992:"Paleozoic Nymphal Wing Pads Support Dual Model of Insect Wing Origins"
3511:
3479:
3162:
3154:
3020:
2981:
2677:
2665:
2511:
1638:
1358:
1267:
1240:
1129:
1004:
996:
903:
836:
778:
721:
712:
704:
333:
287:
243:
184:
141:
4624:
3755:
3738:
582:
is the outer margin of the wing, between apex and hind or anal angle.
405:– fifth longitudinal vein, one to three branches reach the wing margin
395:– fourth longitudinal vein, one to four branches reach the wing margin
5794:
3630:
3515:
3440:
3065:
3048:
is hardened, while the posterior part is membranous. They are called
3010:
2562:
The two pairs of wings are found on the middle and third segment, or
2403:
2399:
1385:
do not have a pupal stage, therefore they must have a different wing
1291:
1287:
1279:
1187:
1147:
1133:
1115:
914:
899:
832:
777:) is a row of small hooks on the forward margin of the hindwing, or "
385:– third longitudinal vein, one to five branches reach the wing margin
254:
239:
204:
110:
4890:
3467:) have hairy wings with the front and hindwings clothed with setae.
324:
In some very small insects, the venation may be greatly reduced. In
164:
attach to and deform the thorax, causing the wings to move as well.
4943:
Matsuda R (1970). "Morphology and evolution of the insect thorax".
2840:– third longitudinal vein, branched to Rs in forewing and hindwing.
2824:– at the leading marginal of the forewing and hindwing, unbranched.
2558:
Transition of scales color on a butterfly wing (30x magnification).
375:– second longitudinal vein (behind the costa), typically unbranched
3456:
3256:
3179:
3070:
2872:
2803:
2792:
2553:
1390:
932:
808:
676:
336:(dragonflies and damselflies) and at the base of the forewings of
283:
264:
200:
192:
153:
72:
4062:(4th ed.). Cambridge, New York: Cambridge University Press.
1011:. Their prototypes are the oldest winged fossils, go back to the
5990:
5858:
5671:
3505:
3329:
2639:– The hindwing of most butterflies has the humeral vein, except
1378:
1203:
1032:
having forewings with unusual venation, possibly diverging from
906:
246:
5636:
4342:"Diffusion In Insect Wing Muscle, The Most Active Tissue Known"
4329:. Mechanical and Aerospace Engineering, University of Virginia.
2729:– second longitudinal vein, it is unbranched, joins C at nodus.
1381:
stage of the insects life cycle. However, insects that undergo
1219:: wings first originated as stiff outgrowths of tergal plates (
5679:
5663:
3158:
1300:
Evolution of the ways the wings at rest to the body to create
306:"Cross vein" redirects here. For the Japanese metal band, see
290:, and, since the cavities of the veins are connected with the
180:
5166:"Gliding hexapods and the origins of insect aerial behaviour"
2471:– at the leading wing marginal, fused for most of the length.
5421:
4807:"The presumed oldest flying insect: more likely a myriapod?"
554:
Most veins and crossveins occur in the anterior area of the
5625:
1420:, the wing begins to expand and form its definitive shape.
873:
are increased, at the price of wear and tear on the wings.
831:
Two groups of relatively large insects, the Ephemeroptera (
5445:
Termites: evolution, sociality, symbioses, ecology, ecolab
4731:(2 (illustrated) ed.). Academic Press. p. 1132.
2477:– divided into two branches beyond the middle of the wing.
2783:
horizontally at rest, and in one anisopteran genus (e.g.
681:
The diamond-shaped alary muscles (green) of the mosquito
5592:
Borror and DeLong's introduction to the study of insects
5420:
Trueman, John W. H.; Richard J. Rowe (16 October 2009).
5025:
Staniczek, A.H.; Bechly, G. & Godunko, R.J. (2011).
3284:– third longitudinal vein, with many pectinate branches.
1347:
Subjects folding (such as the rear wing of the earwigs)
282:
under a vein. Within each of the major veins there is a
5226:. Ithaca, NY: The Comstack Publishing Company. p.
187:. Rarely, the female is winged but the male not, as in
5621:
North-Carolina State University course on insect wings
2518:
meaning "wing" in Greek). The elytra are not used for
1274:. Most insects control their wings by adjusting tilt,
760:
feedback necessary for effective flight and grooming.
1087:
Hypothetical insect with wings from annex of the legs
1039:
A single large wing from a species of Diptera in the
5526:
Camacho, Ernesto Robayo; Chong, Juang-Horng (2015).
4513:
Numbers of living species in Australia and the World
3737:
Haug, Joachim, C. Haug., and R. J. Garwood. (2016).
1075:
Hypothetical insect with wings from the back (Notum)
868:. Some very small insects use the fling and clap or
5978:
5917:
5857:
5760:
5678:
5495:. Special Issue: Miniaturization in Panarthropoda.
3278:– second longitudinal vein, it is relatively short.
2530:(family Carabidae) and some "true weevils" (family
2506:– veins behind the cubitus, separated by anal fold.
1377:; in these species, the wing develops while in the
1249:has played an enormous role in refining the wings,
5590:Triplehorn, Charles A.; Johnson Norman F. (2005).
5472:. Texas AgriLife Extension Service. Archived from
5317:
4906:"Dragonfly flight III lift and power requirements"
4324:"Insect Flight Mechanisms: Anatomy and Kinematics"
4212:
4210:
4208:
4206:
4204:
4156:
3308:– is unbranched, curved and reach the wing margin.
3292:– fourth longitudinal vein, reach the wing margin.
2069:Part of Neoptera, mostly with piercing mouthparts
562:; the two other posterior fields are the anal and
132:), and the two pairs are often referred to as the
4718:
4716:
4714:
4712:
4710:
4202:
4200:
4198:
4196:
4194:
4192:
4190:
4188:
4186:
4184:
2047:Mayflies, dragonflies, and several fossil orders
1757:Hemiptera (true bugs, leafhoppers, aphids, etc.)
1342:Cross fold (such as the rear wing of the beetle)
1257:, and anything else that affects aerodynamics or
195:. Wing structure and colouration often vary with
82:'s hardened forewings raised, hindwings unfolding
4081:
4079:
1278:, and flapping frequency of the wings with tiny
5343:
5341:
4598:
4596:
4051:
4049:
4047:
4045:
4043:
4041:
1080:Hypothetical insect with wings from the Pleurum
816:in flight; it uses the direct flight mechanism.
657:the epimeron inserted into the axillary plate.
5320:The Lepidoptera: Form, function, and diversity
4805:Haug, Carolin; Haug, Joachim T. (2017-05-30).
4727:. In Resh, Vincent H.; Cardé, Ring T. (eds.).
2579:Modifications in the wing's venation include:
5648:
4092:(2 ed.). Springer-Verlag New York, LLC.
3537:Journal of the New York Entomological Society
1053:Insect flight § Evolution and adaptation
447:– run between adjacent branches of the radius
8:
5241:
5239:
5237:
5073:: CS1 maint: multiple names: authors list (
4030:: CS1 maint: multiple names: authors list (
3916:: CS1 maint: multiple names: authors list (
3785:: CS1 maint: multiple names: authors list (
3722:: CS1 maint: multiple names: authors list (
3688:: CS1 maint: multiple names: authors list (
3659:: CS1 maint: multiple names: authors list (
2633:– either one vein A, or two veins 1A+2A, 3A.
4703:. New York, NY: Cambridge University Press.
4699:Grimaldi, David; Engel, Michael S. (2005).
4694:
4692:
4690:
4688:
4686:
4684:
4682:
4680:
4678:
3712:. Oxford University Press. pp. 74–101.
3409:– unbranched, CuP is absent in Hymenoptera.
3201:– became the leading wing vein, unbranched.
1461:
1452:
1024:, or true bugs had appeared in the form of
703:In more derived orders of insects, such as
491:Wing areas are delimited and subdivided by
5655:
5641:
5633:
5616:Brisbane University course on insect wings
5286:
5284:
5282:
5280:
5278:
5276:
5274:
5272:
5270:
5268:
4550:
4548:
4505:
4503:
2027:Primitive palaeozoic paleopterous insects
1296:
344:(katydids and grasshoppers respectively).
175:, or are selectively lost in "workers" of
6011:Tradeoffs for locomotion in air and water
5562:
5381:
5189:
5164:Yanoviak SP, Kaspari M, Dudley R (2009).
5105:
4840:
4822:
4623:
4445:
4404:
4317:
4315:
4272:"EXPLANATION OF TERMS USED IN ENTOMOLOGY"
4150:
4148:
4146:
4144:
4007:
3966:
3956:
3842:
3754:
3582:
3486:are reduced to halteres (or are absent).
2144:Neoptera with hemimetabolous development
1401:, a cluster of cells grow underneath the
902:; that they are modifications of movable
242:; that they are modifications of movable
120:. They are found on the second and third
5215:
5213:
5211:
5209:
4117:
4115:
4113:
4111:
4109:
3324:
2938:
2408:
1473:
1056:
593:
483:
312:
5442:Abe T., Bignell D.E; Higashi M (2000).
4155:Gullan, P. J.; Cranston, P. S. (2004).
3567:"Insect Evolution: The Origin of Wings"
3527:
3056:). They are only found in the suborder
2894:– second longitudinal vein, unbranched.
2832:– second longitudinal vein, unbranched.
317:Venation of insect wings, based on the
5422:"Odonata. Dragonflies and damselflies"
5066:
4587:10.1146/annurev.fluid.36.050802.121940
4023:
3909:
3778:
3715:
3681:
3652:
3328:An example of Longitudinal folding in
2918:– fifth longitudinal vein, unbranched.
2758:– unbranched veins behind the cubitus.
2412:Cross folding in the wings of beetles
1337:fan-fold (e.g., front wings of wasps)
1329:wings not foldable (e.g., stoneflies)
5594:(7th ed.). Thomson Brooks/Cole.
5535:Journal of Integrated Pest Management
5493:Arthropod Structure & Development
5003:Grundzüge der vergleichenden Anatomie
4871:
4869:
4518:Australian Biological Resources Study
4159:The insects: an outline of entomology
2980:Other orders such as the Dermaptera (
2001:Grasshoppers, katydids, and crickets
1814:Insects with wings of thin membranes
1304:
764:Coupling, folding, and other features
415:– unbranched veins behind the cubitus
294:, hemolymph can flow into the wings.
7:
4163:. UK: Blackwell Publishing. p.
3703:
3701:
3699:
3604:
3602:
3560:
3558:
1937:Lacewings, owlflies, antlions, etc.
1102:Exit (outer attachments of the legs)
437:– run between the costa and subcosta
4612:The Journal of Experimental Biology
4605:"The aerodynamics of insect flight"
4059:The Insects: Structure and function
2575:, has the wings modified to act as
1633:Cockroaches, mantises and termites
1483:Translation of the Scientific name
883:flight, without the use of thrust.
495:along which the wing can fold, and
467:– run between the media and cubitus
5034:Insect Systematics & Evolution
4661:. Academic Press. pp. 78–79.
4217:Snodgrass, R. E. (December 1993).
3270:– at the leading edge of the wing.
2617:– the basal section has been lost.
2072:True bugs, lice, barklice, thrips
1315:over the back against one another
457:– run between the radius and media
25:
5363:"Structural color in Lepidoptera"
5131:Trends in Ecology & Evolution
4904:Wakeling JM Ellington CP (1997).
4122:Meyer, John R. (5 January 2007).
2491:– branches, long and strong vein.
1691:, inside; πτερύγιον, small wing)
1447:of insect orders is based on the
1310:spread laterally (large bubbles)
1150:. According to this theory these
5751:
4567:Annual Review of Fluid Mechanics
4086:Gilliott, Cedric (August 1995).
3931:Clark-Hatchel, Courtney (2013).
3352:
3337:
2963:
2947:
2700:
2688:
2446:
2431:
2417:
89:
71:
54:
34:
27:Body part used by insects to fly
4910:Journal of Experimental Biology
4658:Physics in Biology and Medicine
4219:Principles of Insect Morphology
3674:Grodnitsky, Dmitry, L. (1999).
3241:– well defined in most species.
1674:Interior living winged insects
5448:. Kluwer academic publishers.
4573:(1). Annual Reviews: 183–210.
4379:Tiegs, O. W. (February 1954).
3868:Nature Ecology & Evolution
3435:– is present for some species.
1607:, transparent or translucent)
1068:Hypothetical wingless ancestor
917:; or that they developed from
365:– the leading edge of the wing
1:
4322:Knospe, Carl R. (Fall 1998).
3708:Alexander, David, E. (2015).
2769:is carried near the wing tip.
1062:Diagram of different theories
937:Holotype wing of the extinct
41:
5297:Brisbane Insects and Spiders
5143:10.1016/0169-5347(96)30022-0
4447:10.1371/journal.pbio.2006619
3125:– branches to CuA1 and CuA2.
2625:– CuP section has been lost.
2622:Cubitus anterior (CuA1-CuA2)
2483:– basal connection is lost.
1411:). In the second instar the
109:are adult outgrowths of the
5291:Chew, Peter (May 9, 2009).
5220:H. Comstock, Henry (1918).
3649:– via Web of Science.
3595:– via Web of Science.
3379:– not found in Hymenoptera.
3316:– veins behind the cubitus.
2972:having been folded by 180°.
2774:identification. Almost all
2658:three-dimensional structure
2588:– not found in Butterflies.
2542:, in which the females are
1714:Short lived winged insects
1630:Wings with netted venation
837:dragonflies and damselflies
737:are found on insect wings:
6063:
5867:Flying and gliding animals
5703:Fin and flipper locomotion
5253:. Cislunar Aerospace. 1997
4978:Kluwer Academic Publishers
4560:"Dissecting Insect Flight"
4340:Weis-Fogh, T (July 1963).
3888:10.1038/s41559-020-01349-0
3827:10.1038/s41467-020-16284-8
3177:which cover the halteres.
2577:tympanal or hearing organs
2398:In species of Coleoptera (
2243:Rotating or twisted wings
1050:
854:
819:
305:
5749:
5513:10.1016/j.asd.2019.01.003
5383:10.1016/j.cub.2006.07.040
5247:"Insect Wings in General"
5107:10.1016/j.cub.2016.11.021
4723:Powell, Jerry A. (2009).
4655:Davidovits, Paul (2008).
4124:"External Anatomy: WINGS"
4009:10.1016/j.cub.2016.11.021
3584:10.1016/j.cub.2016.12.014
3455:Species of Thysanoptera (
3327:
2942:Unfolding of earwig wing
2941:
2931:stick insect wing veins.
2630:Anal veins (A, 1A+2A, 3A)
2411:
2246:twisted-winged parasites
1981:Division of the Neoptera
1453:
1333:
1328:
1321:
1299:
5046:10.1163/187631211X578406
5001:Gegenbaur, Carl (1870).
4701:Evolution of the Insects
4603:Sane, Sanjay P. (2003).
3510:Insect inspired robots:
2546:throughout their lives.
2468:Subcosta posterior (ScP)
1857:Butterflies & Moths
1817:bees, wasps, ants, etc.
1383:incomplete metamorphosis
1317:(damselflies, mayflies)
1202:. The expression of leg
5347:Scoble (1995). Section
5251:Aerodynamics of Insects
4916:(Pt 3): 583–600 (589).
4729:Encyclopedia of Insects
4510:Chapman, A. D. (2006).
4300:www.merriam-webster.com
3958:10.1073/pnas.1304332110
3501:Comstock-Needham system
3305:Cubitus posterior (CuP)
3193:– not found in Diptera.
3130:Cubitus posterior (CuP)
3117:– branches to M1 to M4.
1917:Dobsonflies, fishflies
1697:Holometabolous insects
1610:With transparent wings
1305:wings do not fold back
1178:: The fossil larvae of
1164:Drosophila melanogaster
1159:Endite-exite hypothesis
963:, primitive stem-group
412:Anal veins (A1, A2, A3)
354:Comstock–Needham system
319:Comstock–Needham system
215:, it is the forewings.
96:Outstretched wing of a
6006:Terrestrial locomotion
5950:Evolution of cetaceans
5945:Origin of avian flight
5930:Evolution of tetrapods
5324:. Oxford Univ. Press.
5182:10.1098/rsbl.2009.0029
4945:Mem. Entomol. Soc. Can
4406:10.1098/rstb.1955.0001
4296:"Definition of TERMEN"
4221:. Cornell Univ Press.
4056:Chapman, R.F. (1998).
3297:Cubitus anterior (CuA)
3261:
3250:Species of Blattodea (
3184:
3122:Cubitus anterior (CuA)
3075:
2877:
2808:
2559:
2496:Cubitus anterior (CuA)
1978:New with little veins
1462:
1375:complete metamorphosis
1307:(recent Archaeoptera)
1217:developmental genetics
1192:comparative morphology
1103:
944:
817:
696:
598:
488:
321:
269:
203:, migratory phases of
44:and wing posture of a
6021:Undulatory locomotion
5970:Homologous structures
4922:10.1242/jeb.200.3.583
3807:Nature Communications
3565:Ross, Andrew (2017).
3260:
3222:Cubitus anterior(CuA)
3183:
3074:
2876:
2807:
2668:, and to seek mates.
2656:within or due to its
2557:
2481:Radius posterior (RP)
1196:developmental biology
1060:
973:Palaeodictyopteroidea
940:Cimbrophlebia brooksi
936:
843:) for the infraclass
812:
680:
597:
487:
316:
268:
5965:Analogous structures
5960:Convergent evolution
5626:Insect wing drawings
5361:Vukusic, P. (2006).
5316:Scoble, MJ. (1992).
5223:The Wings of Insects
4358:10.1242/jeb.41.2.229
3427:Line of wing folding
3393:– branched to R1-R5.
3217:– branched to M1-M4.
3209:– branched to R1-R5.
2488:Media posterior (MP)
2475:Radius anterior (RA)
1711:about one day long)
1613:diaphanopteroideans
1184:paranotal-hypothesis
1108:Paranotal hypothesis
921:protrusions used as
909:as found on aquatic
893:Carboniferous Period
750:campaniform sensilla
249:as found on aquatic
6016:Rotating locomotion
5955:Comparative anatomy
5547:10.1093/jipm/pmv016
5505:2019ArtSD..48...56P
4957:10.4039/entm10276fv
4776:10.1038/nature02291
4768:2004Natur.427..627E
4579:2005AnRFM..37..183W
4397:1955RSPTB.238..221T
4242:Spieth, HT (1932).
3949:2013PNAS..11016951C
3943:(42): 16951–16956.
3880:2020NatEE...4.1703B
3819:2020NatCo..11.2631A
3623:1997Natur.385..627A
3060:; the wings of the
2907:Media anterior (MA)
2845:Media anterior (MA)
2503:Anal veins (AA, AP)
2164:Barklice, booklice
2095:Lice without wings
1754:Halfwinged insects
1140:Epicoxal hypothesis
1132:and finally active
985:Rhyniognatha hirsti
857:Insect aerodynamics
835:) and the Odonata (
5935:Evolution of birds
5688:Aquatic locomotion
4973:History of Insects
4824:10.7717/peerj.3402
4251:Entomological News
3743:Biological Reviews
3262:
3185:
3076:
2878:
2809:
2607:Radius sector (Rs)
2595:+ Radius 1 (Sc+R1)
2560:
2240:, to turn around)
2141:Many veined wings
1897:Snake flies, etc.
1104:
945:
879:A few insects use
818:
814:Australian emperor
754:chordotonal organs
697:
599:
523:Anal area (vannus)
489:
322:
270:
6029:
6028:
5986:Animal locomotion
5925:Evolution of fish
5805:facultative biped
5331:978-1-4020-6242-1
4762:(6975): 627–630.
4738:978-0-12-374144-8
4668:978-0-12-369411-9
4625:10.1242/jeb.00663
4618:(23): 4191–4208.
4527:978-0-642-56850-2
4520:. pp. 60pp.
4276:www.gutenberg.org
3874:(12): 1703–1712.
3756:10.1111/brv.12159
3678:. pp. 82–83.
3617:(6617): 627–630.
3365:
3364:
3361:
3347:
3004:) and Blattodea (
2978:
2977:
2973:
2957:
2459:
2458:
2455:
2441:
2426:
2382:while minimizing
2330:
2329:
2024:Old veined wings
2021:meaning network)
2007:Palaeodictyoptera
1774:Different winged
1587:, skin, leather)
1467:), as the suffix
1351:
1350:
1247:Natural selection
1073:Paranotal theory:
955:), including the
947:Fossils from the
827:Flight mechanisms
733:Several types of
689:pericardial cells
684:Anopheles gambiae
199:, such as in the
16:(Redirected from
6054:
5996:Robot locomotion
5770:Limb development
5755:
5728:Lobe-finned fish
5657:
5650:
5643:
5634:
5605:
5577:
5576:
5566:
5532:
5523:
5517:
5516:
5484:
5478:
5477:
5466:
5460:
5459:
5439:
5433:
5432:
5430:
5429:
5417:
5411:
5410:
5408:
5406:
5385:
5367:
5358:
5352:
5345:
5336:
5335:
5323:
5313:
5307:
5306:
5304:
5303:
5288:
5263:
5262:
5260:
5258:
5243:
5232:
5231:
5217:
5204:
5203:
5193:
5161:
5155:
5154:
5126:
5120:
5119:
5109:
5085:
5079:
5078:
5072:
5064:
5062:
5056:. Archived from
5031:
5022:
5016:
5013:
5007:
5006:
4998:
4992:
4991:
4967:
4961:
4960:
4940:
4934:
4933:
4901:
4895:
4894:
4885:(6): 1150–1160.
4873:
4864:
4861:
4855:
4854:
4844:
4826:
4802:
4796:
4795:
4749:
4743:
4742:
4720:
4705:
4704:
4696:
4673:
4672:
4652:
4646:
4645:
4627:
4609:
4600:
4591:
4590:
4564:
4552:
4543:
4542:
4540:
4539:
4530:. Archived from
4507:
4498:
4495:
4489:
4486:
4480:
4477:
4471:
4467:
4461:
4458:
4452:
4451:
4449:
4425:
4419:
4418:
4408:
4391:(656): 221–348.
4376:
4370:
4369:
4337:
4331:
4330:
4328:
4319:
4310:
4309:
4307:
4306:
4292:
4286:
4285:
4283:
4282:
4268:
4262:
4261:
4259:
4253:. Archived from
4248:
4239:
4233:
4232:
4214:
4179:
4178:
4162:
4152:
4139:
4138:
4136:
4135:
4119:
4104:
4103:
4083:
4074:
4073:
4053:
4036:
4035:
4029:
4021:
4011:
3987:
3981:
3980:
3970:
3960:
3928:
3922:
3921:
3915:
3907:
3863:
3857:
3856:
3846:
3797:
3791:
3790:
3784:
3776:
3758:
3734:
3728:
3727:
3721:
3713:
3705:
3694:
3693:
3687:
3679:
3671:
3665:
3664:
3658:
3650:
3631:10.1038/385627a0
3606:
3597:
3596:
3586:
3577:(3): R113–R115.
3562:
3553:
3552:
3532:
3415:
3407:
3399:
3385:
3377:
3359:
3356:
3344:
3341:
3325:
3314:
3306:
3298:
3290:
3276:
3268:
3239:
3238:Discal Cell (dc)
3231:
3223:
3215:
3207:
3199:
3191:
3139:
3131:
3123:
3115:
3107:Radius posterior
3094:
3090:
3082:
2970:
2967:
2954:
2951:
2939:
2924:
2916:
2908:
2900:
2892:
2884:
2862:
2854:
2846:
2838:
2830:
2822:
2756:
2748:
2739:
2735:
2727:
2719:
2704:
2692:
2631:
2623:
2615:
2594:
2586:
2504:
2497:
2489:
2469:
2465:
2453:
2450:
2438:
2435:
2424:
2421:
2409:
2300:meaning strong)
2223:Wingless siphon
1599:Diaphanopterodea
1480:linguistic root
1477:Scientific Name
1474:
1465:
1458:
1457:
1373:, which undergo
1297:
1114:found in insect
891:Sometime in the
650:
646:
633:
626:
619:
612:
542:
533:
524:
515:
465:
464:M-Cu cross-veins
455:
445:
435:
434:C-Sc cross-veins
413:
403:
393:
383:
373:
363:
326:chalcidoid wasps
207:and polymorphic
93:
75:
64:hovering to mate
58:
38:
21:
6062:
6061:
6057:
6056:
6055:
6053:
6052:
6051:
6032:
6031:
6030:
6025:
5974:
5940:Origin of birds
5913:
5853:
5775:Limb morphology
5756:
5747:
5733:Ray-finned fish
5698:Fish locomotion
5674:
5661:
5612:
5602:
5589:
5586:
5581:
5580:
5530:
5525:
5524:
5520:
5486:
5485:
5481:
5468:
5467:
5463:
5456:
5441:
5440:
5436:
5427:
5425:
5419:
5418:
5414:
5404:
5402:
5370:Current Biology
5365:
5360:
5359:
5355:
5351:, (pp 63 – 66).
5346:
5339:
5332:
5315:
5314:
5310:
5301:
5299:
5290:
5289:
5266:
5256:
5254:
5245:
5244:
5235:
5219:
5218:
5207:
5170:Biology Letters
5163:
5162:
5158:
5128:
5127:
5123:
5094:Current Biology
5087:
5086:
5082:
5065:
5060:
5029:
5024:
5023:
5019:
5014:
5010:
5005:. W. Engelmann.
5000:
4999:
4995:
4988:
4969:
4968:
4964:
4942:
4941:
4937:
4903:
4902:
4898:
4891:10.1139/z84-166
4875:
4874:
4867:
4862:
4858:
4804:
4803:
4799:
4751:
4750:
4746:
4739:
4722:
4721:
4708:
4698:
4697:
4676:
4669:
4654:
4653:
4649:
4607:
4602:
4601:
4594:
4562:
4554:
4553:
4546:
4537:
4535:
4528:
4509:
4508:
4501:
4496:
4492:
4487:
4483:
4478:
4474:
4468:
4464:
4459:
4455:
4440:(5): e2006619.
4427:
4426:
4422:
4378:
4377:
4373:
4339:
4338:
4334:
4326:
4321:
4320:
4313:
4304:
4302:
4294:
4293:
4289:
4280:
4278:
4270:
4269:
4265:
4257:
4246:
4241:
4240:
4236:
4229:
4216:
4215:
4182:
4175:
4154:
4153:
4142:
4133:
4131:
4130:on 16 July 2011
4121:
4120:
4107:
4100:
4085:
4084:
4077:
4070:
4055:
4054:
4039:
4022:
3996:Current Biology
3989:
3988:
3984:
3930:
3929:
3925:
3908:
3865:
3864:
3860:
3799:
3798:
3794:
3777:
3736:
3735:
3731:
3714:
3707:
3706:
3697:
3680:
3673:
3672:
3668:
3651:
3608:
3607:
3600:
3571:Current Biology
3564:
3563:
3556:
3534:
3533:
3529:
3524:
3492:
3449:
3413:
3405:
3397:
3383:
3375:
3323:
3312:
3304:
3296:
3288:
3274:
3266:
3248:
3237:
3229:
3221:
3213:
3205:
3197:
3189:
3151:
3137:
3129:
3121:
3113:
3101:Radius anterior
3092:
3088:
3080:
3038:
2937:
2922:
2914:
2906:
2898:
2890:
2882:
2871:
2860:
2852:
2844:
2836:
2828:
2820:
2802:
2755:Anal veins (A1)
2754:
2746:
2737:
2733:
2725:
2717:
2713:
2712:
2711:
2710:
2709:
2708:
2705:
2697:
2696:
2693:
2674:
2629:
2621:
2613:
2592:
2584:
2552:
2502:
2495:
2487:
2467:
2463:
2396:
2340:
2335:
2219:
2181:Winged insects
2133:
2087:
2066:
2016:
1998:Straight wings
1970:
1734:External wings
1667:, inside; βίος
1570:Hardened wings
1542:
1527:, now obsolete
1451:word for wing,
1441:
1367:
1323:
1316:
1306:
1272:angle of attack
1263:angle of attack
1255:sensory systems
1180:Coxoplectoptera
1098:
1093:
1088:
1086:
1085:Epicoxal theory
1081:
1076:
1074:
1069:
1064:
1055:
1049:
965:Ephemeropterans
931:
889:
859:
853:
829:
824:
807:
766:
735:sensory neurons
731:
711:(mayflies) and
675:
648:
644:
643:Median plates (
638:Fourth Axillary
631:
625:Second Axillary
624:
617:
610:
592:
540:
531:
522:
513:
482:
463:
454:R-M cross-veins
453:
443:
433:
411:
401:
391:
381:
371:
361:
311:
304:
275:
263:
104:
103:
102:
101:
100:
94:
85:
84:
83:
76:
67:
66:
65:
59:
50:
49:
48:
39:
28:
23:
22:
15:
12:
11:
5:
6060:
6058:
6050:
6049:
6044:
6042:Insect anatomy
6034:
6033:
6027:
6026:
6024:
6023:
6018:
6013:
6008:
6003:
5998:
5993:
5988:
5982:
5980:
5976:
5975:
5973:
5972:
5967:
5962:
5957:
5952:
5947:
5942:
5937:
5932:
5927:
5921:
5919:
5915:
5914:
5912:
5911:
5906:
5904:Pterosaur wing
5901:
5896:
5895:
5894:
5889:
5884:
5874:
5869:
5863:
5861:
5855:
5854:
5852:
5851:
5846:
5841:
5840:
5839:
5829:
5824:
5819:
5818:
5817:
5812:
5807:
5802:
5797:
5792:
5787:
5782:
5772:
5766:
5764:
5758:
5757:
5750:
5748:
5746:
5745:
5740:
5735:
5730:
5725:
5720:
5715:
5710:
5705:
5700:
5695:
5693:Cephalopod fin
5690:
5684:
5682:
5676:
5675:
5662:
5660:
5659:
5652:
5645:
5637:
5629:
5628:
5623:
5618:
5611:
5610:External links
5608:
5607:
5606:
5600:
5585:
5582:
5579:
5578:
5518:
5479:
5476:on 2011-04-13.
5461:
5454:
5434:
5412:
5376:(16): R621–3.
5353:
5337:
5330:
5308:
5293:"Insect Wings"
5264:
5233:
5205:
5156:
5137:(5): 187–188.
5121:
5100:(2): 263–269.
5080:
5063:on 2011-10-03.
5040:(2): 101–138.
5017:
5008:
4993:
4986:
4962:
4935:
4896:
4865:
4856:
4797:
4744:
4737:
4706:
4674:
4667:
4647:
4592:
4544:
4526:
4499:
4490:
4481:
4472:
4462:
4453:
4420:
4371:
4352:(2): 229–256.
4332:
4311:
4287:
4263:
4260:on 2011-09-30.
4234:
4227:
4180:
4173:
4140:
4105:
4098:
4075:
4068:
4037:
4002:(2): 263–269.
3982:
3923:
3858:
3792:
3729:
3695:
3666:
3598:
3554:
3526:
3525:
3523:
3520:
3519:
3518:
3508:
3503:
3498:
3491:
3488:
3448:
3447:Other families
3445:
3437:
3436:
3430:
3424:
3418:
3414:Anal veins (A)
3410:
3402:
3394:
3388:
3380:
3363:
3362:
3357:
3349:
3348:
3342:
3334:
3333:
3322:
3319:
3318:
3317:
3313:Anal veins (A)
3309:
3301:
3293:
3285:
3279:
3271:
3247:
3244:
3243:
3242:
3234:
3230:Anal veins (A)
3226:
3218:
3210:
3202:
3194:
3150:
3147:
3143:
3142:
3138:Anal veins (A)
3134:
3126:
3118:
3110:
3104:
3098:
3085:
3064:, such as the
3037:
3034:
3002:praying mantis
2976:
2975:
2968:
2960:
2959:
2952:
2944:
2943:
2936:
2933:
2928:
2927:
2923:Anal veins (A)
2919:
2911:
2903:
2895:
2887:
2870:
2867:
2866:
2865:
2861:Anal veins (A)
2857:
2849:
2841:
2833:
2825:
2801:
2798:
2771:
2770:
2759:
2751:
2743:
2730:
2722:
2706:
2699:
2698:
2694:
2687:
2686:
2685:
2684:
2683:
2682:
2673:
2670:
2645:
2644:
2634:
2626:
2618:
2610:
2609:– in hindwing.
2604:
2601:Radius (R2-R5)
2598:
2589:
2551:
2548:
2538:of the family
2528:ground beetles
2508:
2507:
2499:
2492:
2484:
2478:
2472:
2457:
2456:
2451:
2443:
2442:
2436:
2428:
2427:
2422:
2414:
2413:
2395:
2392:
2339:
2336:
2334:
2331:
2328:
2327:
2324:
2321:
2320:meaning pair)
2314:
2308:
2307:
2304:
2301:
2294:
2288:
2287:
2284:
2281:
2274:
2268:
2267:
2264:
2263:Fringe winged
2261:
2254:
2248:
2247:
2244:
2241:
2234:
2228:
2227:
2224:
2221:
2220:ἀ- or without
2212:
2206:
2205:
2202:
2199:
2192:
2186:
2185:
2182:
2179:
2172:
2166:
2165:
2162:
2161:Rubbing wings
2159:
2152:
2146:
2145:
2142:
2139:
2126:
2120:
2119:
2116:
2113:
2106:
2100:
2099:
2096:
2093:
2080:
2074:
2073:
2070:
2067:
2055:
2049:
2048:
2045:
2042:
2035:
2029:
2028:
2025:
2022:
2009:
2003:
2002:
1999:
1996:
1989:
1983:
1982:
1979:
1976:
1963:
1962:Oligoneoptera
1959:
1958:
1955:
1952:
1951:, new, young)
1945:
1939:
1938:
1935:
1932:
1925:
1919:
1918:
1915:
1912:
1905:
1899:
1898:
1895:
1892:
1885:
1879:
1878:
1875:
1872:
1865:
1863:Lonchopteridae
1859:
1858:
1855:
1852:
1845:
1839:
1838:
1835:
1832:
1825:
1819:
1818:
1815:
1812:
1805:
1799:
1798:
1795:
1792:
1785:
1779:
1778:
1775:
1772:
1765:
1759:
1758:
1755:
1752:
1745:
1739:
1738:
1735:
1732:
1725:
1719:
1718:
1715:
1712:
1705:
1699:
1698:
1695:
1692:
1685:
1679:
1678:
1675:
1672:
1661:
1655:
1654:
1651:
1648:
1641:
1635:
1634:
1631:
1628:
1621:
1615:
1614:
1611:
1608:
1601:
1595:
1594:
1591:
1590:Leather wings
1588:
1581:
1575:
1574:
1571:
1568:
1561:
1555:
1554:
1551:
1548:
1535:
1529:
1528:
1522:
1519:
1512:
1508:
1507:
1504:
1503:Unequal wings
1501:
1494:
1488:
1487:
1484:
1481:
1478:
1440:
1437:
1366:
1363:
1349:
1348:
1344:
1343:
1339:
1338:
1335:
1331:
1330:
1326:
1325:
1319:
1318:
1312:
1311:
1308:
1302:
1301:
1225:
1224:
1173:
1156:
1137:
1124:starting with
1048:
1045:
1026:Arctiniscytina
930:
927:
888:
885:
855:Main article:
852:
849:
828:
825:
820:Main article:
806:
803:
765:
762:
758:proprioceptive
746:mechanosensory
730:
727:
674:
671:
654:
653:
640:
635:
632:Third Axillary
628:
621:
618:First Axillary
614:
611:Humeral plates
591:
588:
552:
551:
545:
544:
536:
535:
527:
526:
518:
517:
481:
478:
469:
468:
459:
458:
449:
448:
439:
438:
417:
416:
407:
406:
397:
396:
387:
386:
377:
376:
367:
366:
338:Tettigonioidea
303:
300:
274:
271:
262:
259:
229:pan-crustacean
177:social insects
156:level in many
124:segments (the
118:insects to fly
95:
88:
87:
86:
77:
70:
69:
68:
60:
53:
52:
51:
42:Original veins
40:
33:
32:
31:
30:
29:
26:
24:
14:
13:
10:
9:
6:
4:
3:
2:
6059:
6048:
6047:Animal flight
6045:
6043:
6040:
6039:
6037:
6022:
6019:
6017:
6014:
6012:
6009:
6007:
6004:
6002:
5999:
5997:
5994:
5992:
5989:
5987:
5984:
5983:
5981:
5977:
5971:
5968:
5966:
5963:
5961:
5958:
5956:
5953:
5951:
5948:
5946:
5943:
5941:
5938:
5936:
5933:
5931:
5928:
5926:
5923:
5922:
5920:
5916:
5910:
5907:
5905:
5902:
5900:
5897:
5893:
5890:
5888:
5885:
5883:
5880:
5879:
5878:
5875:
5873:
5870:
5868:
5865:
5864:
5862:
5860:
5856:
5850:
5847:
5845:
5842:
5838:
5835:
5834:
5833:
5830:
5828:
5825:
5823:
5820:
5816:
5813:
5811:
5808:
5806:
5803:
5801:
5798:
5796:
5793:
5791:
5788:
5786:
5783:
5781:
5778:
5777:
5776:
5773:
5771:
5768:
5767:
5765:
5763:
5759:
5754:
5744:
5741:
5739:
5738:Pectoral fins
5736:
5734:
5731:
5729:
5726:
5724:
5721:
5719:
5716:
5714:
5711:
5709:
5706:
5704:
5701:
5699:
5696:
5694:
5691:
5689:
5686:
5685:
5683:
5681:
5677:
5673:
5669:
5665:
5658:
5653:
5651:
5646:
5644:
5639:
5638:
5635:
5631:
5627:
5624:
5622:
5619:
5617:
5614:
5613:
5609:
5603:
5601:0-03-096835-6
5597:
5593:
5588:
5587:
5583:
5574:
5570:
5565:
5560:
5556:
5552:
5548:
5544:
5540:
5536:
5529:
5522:
5519:
5514:
5510:
5506:
5502:
5498:
5494:
5490:
5483:
5480:
5475:
5471:
5465:
5462:
5457:
5455:0-7923-6361-2
5451:
5447:
5446:
5438:
5435:
5423:
5416:
5413:
5401:
5397:
5393:
5389:
5384:
5379:
5375:
5371:
5364:
5357:
5354:
5350:
5344:
5342:
5338:
5333:
5327:
5322:
5321:
5312:
5309:
5298:
5294:
5287:
5285:
5283:
5281:
5279:
5277:
5275:
5273:
5271:
5269:
5265:
5252:
5248:
5242:
5240:
5238:
5234:
5229:
5225:
5224:
5216:
5214:
5212:
5210:
5206:
5201:
5197:
5192:
5187:
5183:
5179:
5175:
5171:
5167:
5160:
5157:
5152:
5148:
5144:
5140:
5136:
5132:
5125:
5122:
5117:
5113:
5108:
5103:
5099:
5095:
5091:
5084:
5081:
5076:
5070:
5059:
5055:
5051:
5047:
5043:
5039:
5035:
5028:
5021:
5018:
5012:
5009:
5004:
4997:
4994:
4989:
4987:1-4020-0026-X
4983:
4979:
4975:
4974:
4966:
4963:
4958:
4954:
4951:(76): 1–431.
4950:
4946:
4939:
4936:
4931:
4927:
4923:
4919:
4915:
4911:
4907:
4900:
4897:
4892:
4888:
4884:
4880:
4872:
4870:
4866:
4860:
4857:
4852:
4848:
4843:
4838:
4834:
4830:
4825:
4820:
4816:
4812:
4808:
4801:
4798:
4793:
4789:
4785:
4781:
4777:
4773:
4769:
4765:
4761:
4757:
4756:
4748:
4745:
4740:
4734:
4730:
4726:
4719:
4717:
4715:
4713:
4711:
4707:
4702:
4695:
4693:
4691:
4689:
4687:
4685:
4683:
4681:
4679:
4675:
4670:
4664:
4660:
4659:
4651:
4648:
4643:
4639:
4635:
4631:
4626:
4621:
4617:
4613:
4606:
4599:
4597:
4593:
4588:
4584:
4580:
4576:
4572:
4568:
4561:
4557:
4556:Wang, Z. Jane
4551:
4549:
4545:
4534:on 2009-05-19
4533:
4529:
4523:
4519:
4515:
4514:
4506:
4504:
4500:
4494:
4491:
4485:
4482:
4476:
4473:
4466:
4463:
4457:
4454:
4448:
4443:
4439:
4435:
4431:
4424:
4421:
4416:
4412:
4407:
4402:
4398:
4394:
4390:
4386:
4382:
4375:
4372:
4367:
4363:
4359:
4355:
4351:
4347:
4343:
4336:
4333:
4325:
4318:
4316:
4312:
4301:
4297:
4291:
4288:
4277:
4273:
4267:
4264:
4256:
4252:
4245:
4238:
4235:
4230:
4228:0-8014-8125-2
4224:
4220:
4213:
4211:
4209:
4207:
4205:
4203:
4201:
4199:
4197:
4195:
4193:
4191:
4189:
4187:
4185:
4181:
4176:
4174:1-4051-1113-5
4170:
4166:
4161:
4160:
4151:
4149:
4147:
4145:
4141:
4129:
4125:
4118:
4116:
4114:
4112:
4110:
4106:
4101:
4099:0-306-44967-6
4095:
4091:
4090:
4082:
4080:
4076:
4071:
4069:0-521-57048-4
4065:
4061:
4060:
4052:
4050:
4048:
4046:
4044:
4042:
4038:
4033:
4027:
4019:
4015:
4010:
4005:
4001:
3997:
3993:
3986:
3983:
3978:
3974:
3969:
3964:
3959:
3954:
3950:
3946:
3942:
3938:
3934:
3927:
3924:
3919:
3913:
3905:
3901:
3897:
3893:
3889:
3885:
3881:
3877:
3873:
3869:
3862:
3859:
3854:
3850:
3845:
3840:
3836:
3832:
3828:
3824:
3820:
3816:
3812:
3808:
3804:
3796:
3793:
3788:
3782:
3774:
3770:
3766:
3762:
3757:
3752:
3748:
3744:
3740:
3733:
3730:
3725:
3719:
3711:
3704:
3702:
3700:
3696:
3691:
3685:
3677:
3670:
3667:
3662:
3656:
3648:
3644:
3640:
3636:
3632:
3628:
3624:
3620:
3616:
3612:
3605:
3603:
3599:
3594:
3590:
3585:
3580:
3576:
3572:
3568:
3561:
3559:
3555:
3550:
3546:
3542:
3538:
3531:
3528:
3521:
3517:
3513:
3509:
3507:
3504:
3502:
3499:
3497:
3494:
3493:
3489:
3487:
3485:
3481:
3477:
3473:
3468:
3466:
3462:
3458:
3453:
3446:
3444:
3442:
3434:
3431:
3428:
3425:
3422:
3421:Wing-coupling
3419:
3416:
3411:
3408:
3406:Cubitus (CuA)
3403:
3400:
3395:
3392:
3389:
3387:– unbranched.
3386:
3384:Subcosta (Sc)
3381:
3378:
3373:
3372:
3371:
3369:
3358:
3355:
3351:
3350:
3343:
3340:
3336:
3335:
3331:
3326:
3320:
3315:
3310:
3307:
3302:
3299:
3294:
3291:
3286:
3283:
3280:
3277:
3275:Subcosta (Sc)
3272:
3269:
3264:
3263:
3259:
3255:
3253:
3245:
3240:
3235:
3232:
3227:
3224:
3219:
3216:
3211:
3208:
3203:
3200:
3198:Subcosta (Sc)
3195:
3192:
3187:
3186:
3182:
3178:
3176:
3172:
3168:
3164:
3160:
3156:
3148:
3146:
3140:
3135:
3133:– unbranches.
3132:
3127:
3124:
3119:
3116:
3111:
3108:
3105:
3102:
3099:
3096:
3086:
3083:
3078:
3077:
3073:
3069:
3067:
3063:
3059:
3055:
3051:
3047:
3043:
3035:
3033:
3031:
3030:pseudoelytron
3027:
3023:
3022:
3017:
3013:
3012:
3007:
3003:
2999:
2995:
2991:
2987:
2983:
2969:
2966:
2962:
2961:
2953:
2950:
2946:
2945:
2940:
2934:
2932:
2925:
2920:
2917:
2912:
2909:
2904:
2901:
2896:
2893:
2891:Subcosta (Sc)
2888:
2885:
2880:
2879:
2875:
2868:
2863:
2858:
2855:
2850:
2847:
2842:
2839:
2834:
2831:
2829:Subcosta (Sc)
2826:
2823:
2818:
2817:
2816:
2814:
2806:
2799:
2797:
2794:
2790:
2786:
2781:
2777:
2768:
2764:
2760:
2757:
2752:
2749:
2744:
2741:
2731:
2728:
2726:Subcosta (Sc)
2723:
2720:
2715:
2714:
2703:
2691:
2681:
2679:
2671:
2669:
2667:
2663:
2659:
2654:
2653:Ancient Greek
2650:
2642:
2638:
2635:
2632:
2627:
2624:
2619:
2616:
2614:Media (M1-M3)
2611:
2608:
2605:
2602:
2599:
2596:
2593:Subcosta (Sc)
2590:
2587:
2582:
2581:
2580:
2578:
2574:
2569:
2565:
2556:
2549:
2547:
2545:
2541:
2537:
2533:
2532:Curculionidae
2529:
2525:
2521:
2517:
2513:
2505:
2500:
2498:
2493:
2490:
2485:
2482:
2479:
2476:
2473:
2470:
2461:
2460:
2452:
2449:
2445:
2444:
2437:
2434:
2430:
2429:
2423:
2420:
2416:
2415:
2410:
2407:
2405:
2401:
2393:
2391:
2387:
2385:
2381:
2377:
2373:
2368:
2366:
2362:
2358:
2354:
2350:
2346:
2337:
2332:
2325:
2323:Paired wings
2322:
2319:
2315:
2313:
2310:
2309:
2305:
2303:Strong wings
2302:
2299:
2295:
2293:
2290:
2289:
2285:
2283:Haired wings
2282:
2279:
2275:
2273:
2270:
2269:
2265:
2262:
2259:
2255:
2253:
2250:
2249:
2245:
2242:
2239:
2235:
2233:
2230:
2229:
2225:
2222:
2217:
2213:
2211:
2208:
2207:
2203:
2201:Needle wings
2200:
2197:
2193:
2191:
2190:Raphidioptera
2188:
2187:
2183:
2180:
2177:
2173:
2171:
2168:
2167:
2163:
2160:
2157:
2153:
2151:
2148:
2147:
2143:
2140:
2137:
2131:
2127:
2125:
2122:
2121:
2117:
2115:Folded wings
2114:
2111:
2107:
2105:
2102:
2101:
2097:
2094:
2091:
2085:
2081:
2079:
2076:
2075:
2071:
2068:
2064:
2060:
2056:
2054:
2051:
2050:
2046:
2043:
2040:
2036:
2034:
2031:
2030:
2026:
2023:
2020:
2014:
2010:
2008:
2005:
2004:
2000:
1997:
1994:
1990:
1988:
1985:
1984:
1980:
1977:
1974:
1968:
1964:
1961:
1960:
1956:
1953:
1950:
1946:
1944:
1941:
1940:
1936:
1933:
1930:
1926:
1924:
1921:
1920:
1916:
1913:
1910:
1906:
1904:
1901:
1900:
1896:
1893:
1890:
1886:
1884:
1881:
1880:
1876:
1873:
1870:
1866:
1864:
1861:
1860:
1856:
1854:Scaled wings
1853:
1850:
1846:
1844:
1841:
1840:
1836:
1833:
1830:
1826:
1824:
1821:
1820:
1816:
1813:
1810:
1806:
1804:
1801:
1800:
1797:now obsolete
1796:
1793:
1790:
1786:
1784:
1781:
1780:
1776:
1773:
1771:, different)
1770:
1766:
1764:
1761:
1760:
1756:
1753:
1750:
1746:
1744:
1741:
1740:
1736:
1733:
1730:
1726:
1724:
1721:
1720:
1716:
1713:
1710:
1706:
1704:
1703:Ephemeroptera
1701:
1700:
1696:
1694:Inside wings
1693:
1690:
1686:
1684:
1683:Endopterygota
1681:
1680:
1676:
1673:
1670:
1666:
1662:
1660:
1657:
1656:
1652:
1649:
1646:
1642:
1640:
1637:
1636:
1632:
1629:
1626:
1622:
1620:
1617:
1616:
1612:
1609:
1606:
1602:
1600:
1597:
1596:
1592:
1589:
1586:
1582:
1580:
1577:
1576:
1572:
1569:
1566:
1562:
1560:
1557:
1556:
1553:Apterygotans
1552:
1549:
1546:
1540:
1536:
1534:
1531:
1530:
1526:
1523:
1520:
1517:
1513:
1510:
1509:
1505:
1502:
1499:
1495:
1493:
1490:
1489:
1486:English Name
1485:
1482:
1479:
1476:
1475:
1472:
1470:
1466:
1464:
1456:
1450:
1449:Ancient Greek
1446:
1438:
1436:
1432:
1430:
1426:
1421:
1419:
1414:
1410:
1409:
1404:
1400:
1399:embryogenesis
1394:
1392:
1388:
1387:morphogenesis
1384:
1380:
1376:
1372:
1371:Endopterygota
1365:Morphogenesis
1364:
1362:
1360:
1356:
1346:
1345:
1341:
1340:
1336:
1332:
1327:
1320:
1314:
1313:
1309:
1303:
1298:
1295:
1293:
1289:
1285:
1281:
1277:
1273:
1269:
1264:
1260:
1256:
1252:
1248:
1244:
1242:
1238:
1234:
1230:
1222:
1218:
1214:
1209:
1205:
1201:
1197:
1193:
1189:
1185:
1181:
1177:
1174:
1170:
1167:, in which a
1166:
1165:
1160:
1157:
1153:
1149:
1145:
1141:
1138:
1135:
1131:
1127:
1122:
1117:
1113:
1112:preadaptation
1109:
1106:
1105:
1101:
1096:
1091:
1084:
1079:
1072:
1067:
1063:
1059:
1054:
1046:
1044:
1042:
1037:
1035:
1031:
1027:
1023:
1018:
1017:Carboniferous
1014:
1010:
1006:
1002:
998:
993:
991:
987:
986:
982:
978:
974:
970:
966:
962:
958:
954:
950:
942:
941:
935:
928:
926:
924:
920:
916:
912:
908:
905:
901:
898:
894:
886:
884:
882:
877:
874:
871:
867:
864:
858:
850:
848:
846:
842:
838:
834:
826:
823:
822:Insect flight
815:
811:
804:
802:
798:
794:
792:
788:
784:
780:
776:
772:
763:
761:
759:
755:
751:
747:
743:
740:
736:
728:
726:
724:
723:
716:
714:
710:
709:Ephemeroptera
706:
701:
694:
690:
686:
685:
679:
672:
670:
666:
662:
658:
652:
641:
639:
636:
634:
629:
627:
622:
620:
615:
613:
608:
607:
606:
603:
596:
589:
587:
583:
581:
575:
571:
567:
565:
561:
557:
550:
547:
546:
543:
541:Axillary area
538:
537:
534:
529:
528:
525:
520:
519:
516:
511:
510:
509:
507:
503:
498:
497:flexion-lines
494:
486:
479:
477:
473:
466:
461:
460:
456:
451:
450:
446:
444:R cross-veins
441:
440:
436:
431:
430:
429:
425:
421:
414:
409:
408:
404:
399:
398:
394:
389:
388:
384:
379:
378:
374:
372:Subcosta (Sc)
369:
368:
364:
359:
358:
357:
355:
350:
345:
343:
339:
335:
331:
327:
320:
315:
309:
301:
299:
295:
293:
289:
285:
281:
272:
267:
260:
258:
256:
252:
248:
245:
241:
238:
232:
230:
224:
220:
216:
214:
210:
206:
202:
198:
194:
190:
186:
182:
178:
174:
170:
165:
161:
159:
155:
151:
147:
143:
139:
135:
131:
127:
123:
119:
115:
112:
108:
99:
92:
81:
74:
63:
57:
47:
43:
37:
19:
5898:
5630:
5591:
5538:
5534:
5521:
5496:
5492:
5482:
5474:the original
5464:
5444:
5437:
5426:. Retrieved
5424:. Tolweb.org
5415:
5403:. Retrieved
5373:
5369:
5356:
5348:
5319:
5311:
5300:. Retrieved
5296:
5255:. Retrieved
5250:
5222:
5176:(4): 510–2.
5173:
5169:
5159:
5134:
5130:
5124:
5097:
5093:
5083:
5069:cite journal
5058:the original
5037:
5033:
5020:
5011:
5002:
4996:
4972:
4965:
4948:
4944:
4938:
4913:
4909:
4899:
4882:
4879:Can. J. Zool
4878:
4859:
4814:
4810:
4800:
4759:
4753:
4747:
4728:
4725:"Coleoptera"
4700:
4657:
4650:
4615:
4611:
4570:
4566:
4536:. Retrieved
4532:the original
4516:. Canberra:
4512:
4493:
4484:
4475:
4465:
4456:
4437:
4434:PLOS Biology
4433:
4423:
4388:
4384:
4374:
4349:
4345:
4335:
4303:. Retrieved
4299:
4290:
4279:. Retrieved
4275:
4266:
4255:the original
4250:
4237:
4218:
4158:
4132:. Retrieved
4128:the original
4088:
4058:
4026:cite journal
3999:
3995:
3985:
3940:
3936:
3926:
3912:cite journal
3871:
3867:
3861:
3810:
3806:
3795:
3781:cite journal
3749:(1): 53–69.
3746:
3742:
3732:
3709:
3675:
3669:
3655:cite journal
3614:
3610:
3574:
3570:
3540:
3536:
3530:
3472:Strepsiptera
3469:
3454:
3450:
3438:
3432:
3426:
3420:
3412:
3404:
3396:
3390:
3382:
3374:
3366:
3311:
3303:
3295:
3287:
3281:
3273:
3265:
3249:
3236:
3228:
3220:
3212:
3204:
3196:
3188:
3152:
3144:
3136:
3128:
3120:
3112:
3106:
3100:
3087:
3079:
3053:
3049:
3039:
3029:
3025:
3019:
3015:
3009:
2990:grasshoppers
2979:
2929:
2921:
2915:Cubitus (Cu)
2913:
2905:
2897:
2889:
2881:
2859:
2853:Cubitus (Cu)
2851:
2843:
2835:
2827:
2819:
2810:
2772:
2762:
2753:
2747:Cubitus (Cu)
2745:
2732:
2724:
2716:
2675:
2646:
2637:Humeral vein
2636:
2628:
2620:
2612:
2606:
2600:
2591:
2583:
2561:
2523:
2515:
2509:
2501:
2494:
2486:
2480:
2474:
2462:
2397:
2388:
2369:
2341:
2326:Damselflies
2317:
2306:Zorapterans
2297:
2286:Caddisflies
2277:
2257:
2252:Thysanoptera
2237:
2232:Strepsiptera
2215:
2210:Siphonaptera
2195:
2175:
2155:
2135:
2129:
2124:Polyneoptera
2109:
2098:Animal lice
2089:
2083:
2078:Phthiraptera
2062:
2058:
2053:Paraneoptera
2038:
2018:
2012:
1995:, straight)
1992:
1972:
1966:
1948:
1934:Veined wing
1928:
1914:Large wings
1908:
1888:
1877:Lance flies
1874:Lance wings
1868:
1848:
1834:Same winged
1828:
1811:, membrane)
1808:
1794:Same winged
1788:
1768:
1748:
1731:, external)
1728:
1723:Exopterygota
1708:
1688:
1677:Webspinners
1668:
1664:
1644:
1643:Δύο- (dyo-,
1624:
1604:
1584:
1564:
1544:
1538:
1525:Apterygotans
1515:
1497:
1468:
1460:
1445:nomenclature
1443:Most of the
1442:
1439:Nomenclature
1433:
1422:
1406:
1395:
1368:
1352:
1245:
1226:
1213:paleontology
1175:
1168:
1162:
1158:
1139:
1107:
1099:
1094:
1092:Notum (back)
1089:
1082:
1077:
1070:
1065:
1061:
1038:
1030:Paraknightia
994:
983:
961:Caloneurodea
946:
938:
890:
878:
875:
863:leading edge
860:
851:Aerodynamics
841:synapomorphy
830:
799:
795:
767:
732:
720:
717:
702:
698:
682:
667:
663:
659:
655:
642:
637:
630:
623:
616:
609:
604:
600:
584:
576:
572:
568:
553:
548:
539:
530:
521:
512:
490:
474:
470:
462:
452:
442:
432:
426:
422:
418:
410:
402:Cubitus (Cu)
400:
390:
380:
370:
360:
349:archedictyon
346:
323:
296:
276:
233:
225:
221:
217:
213:vespid wasps
173:Strepsiptera
166:
162:
160:of insects.
137:
133:
116:that enable
107:Insect wings
106:
105:
5899:Insect wing
5849:Webbed foot
5790:unguligrade
5785:plantigrade
5780:digitigrade
5405:11 November
4470:pp.905-925.
3813:(1): 2631.
3543:(1): 1–39.
3433:Pterostigma
3368:Hymenoptera
3332:(Vespidae)
3321:Hymenoptera
3252:cockroaches
3171:Calyptratae
3058:Heteroptera
3006:cockroaches
2869:Phasmatodea
2811:Species of
2789:Corduliidae
2785:Cordulephya
2767:pterostigma
2676:Species of
2550:Lepidoptera
2540:Phengodidae
2353:caddisflies
2349:damselflies
2345:dragonflies
2333:Adaptations
2272:Trichoptera
2260:, fringes)
2204:Snakeflies
2174:Πτερύγιον (
2118:Stoneflies
2033:Palaeoptera
1903:Megaloptera
1894:Long wings
1843:Lepidoptera
1803:Hymenoptera
1791:, similar)
1763:Heteroptera
1627:, network)
1619:Dictyoptera
1541:small wing)
1537:πτερύγιον (
1324:(Neoptera)
1237:dragonflies
1126:parachuting
1034:Blattoptera
1001:dragonflies
995:During the
957:Blattoptera
791:Trichoptera
789:, and some
787:Lepidoptera
775:Trichoptera
771:Hymenoptera
693:cell nuclei
280:sclerotized
209:butterflies
169:velvet ants
142:dragonflies
114:exoskeleton
6036:Categories
5827:Cephalopod
5743:Pelvic fin
5713:Dorsal fin
5708:Caudal fin
5584:References
5428:2011-03-21
5302:2011-03-21
4538:2012-06-18
4346:J Exp Biol
4305:2024-01-08
4281:2024-01-08
4134:2011-03-21
4089:Entomology
3474:also have
3391:Radius (R)
3282:Radius (R)
3206:Radius (R)
3167:gyroscopes
3159:true flies
3050:hemelytron
3046:stink bugs
2986:Orthoptera
2935:Dermaptera
2899:Radius (R)
2837:Radius (R)
2813:Orthoptera
2800:Orthoptera
2776:Anisoptera
2662:camouflage
2641:Lycaenidae
2568:metathorax
2564:mesothorax
2536:glow-worms
2394:Coleoptera
2376:propulsion
2372:Sphingidae
2361:alderflies
2357:stoneflies
2198:, needle)
2158:, to rub)
2150:Psocoptera
2104:Plecoptera
2044:Old wings
1987:Orthoptera
1954:New wings
1923:Neuroptera
1891:, length)
1777:True bugs
1707:ἐφήμερος (
1659:Embioptera
1650:Two wings
1603:Διαφανής (
1579:Dermaptera
1567:, sheath)
1559:Coleoptera
1533:Apterygota
1506:Dragonfly
1492:Anisoptera
1413:histoblast
1292:protection
1259:kinematics
1233:apterygote
1229:stoneflies
1121:parachutes
1051:See also:
1047:Hypotheses
1009:amphibious
977:coriaceous
969:Orthoptera
748:bristles,
564:jugal ares
532:Jugal area
506:Coleoptera
502:Dermaptera
493:fold-lines
382:Radius (R)
342:Acridoidea
330:Orthoptera
308:Cross Vein
261:Morphology
130:metathorax
126:mesothorax
80:cockchafer
62:Hoverflies
5918:Evolution
5877:Bird wing
5822:Arthropod
5815:quadruped
5555:2155-7470
5541:(1): 17.
5499:: 56–70.
5470:"Termite"
5257:March 28,
4833:2167-8359
4817:: e3402.
3904:227253368
3835:2041-1723
3718:cite book
3684:cite book
3496:Appendage
3465:caddisfly
3461:Ptiliidae
3398:Media (M)
3376:Costa (C)
3289:Media (M)
3267:Costa (C)
3246:Blattodea
3214:Media (M)
3190:Costa (C)
3175:calypters
3114:Media (M)
3081:Costa (C)
3062:Homoptera
3054:hemelytra
3042:Hemiptera
3036:Hemiptera
2883:Costa (C)
2821:Costa (C)
2780:Zygoptera
2718:Costa (C)
2707:Damselfly
2695:Dragonfly
2585:Costa (C)
2573:Noctuidae
2544:larviform
2464:Costa (C)
2365:lacewings
2347:and some
2338:Variation
2312:Zygoptera
2292:Zoraptera
2276:Τρίχωμα (
2256:Θύσανοι (
2236:Στρέψις (
2176:pterygion
2170:Pterygota
2108:Πλέκειν (
2037:Παλαιός (
2017:δίκτυον (
2011:Παλαιός (
1965:ὀλίγον- (
1911:, large)
1907:Μεγαλο- (
1883:Mecoptera
1871:, lance)
1851:, scale)
1837:Termites
1831:, equal)
1807:ὑμένιον (
1783:Homoptera
1743:Hemiptera
1717:Mayflies
1709:ephemeros
1623:Δίκτυον (
1605:diaphanes
1550:Wingless
1539:pterygion
1521:Wingless
1429:tracheole
1355:predators
1276:stiffness
1022:Hemiptera
953:Pterygota
923:radiators
904:abdominal
887:Evolution
870:Weis-Fogh
783:Mecoptera
739:gustatory
392:Media (M)
362:Costa (C)
244:abdominal
189:fig wasps
146:lacewings
138:hindwings
134:forewings
98:bumblebee
46:dragonfly
5909:Wingspan
5892:feathers
5887:skeleton
5872:Bat wing
5832:Tetrapod
5718:Fish fin
5573:26823990
5400:52828850
5392:16920604
5200:19324632
5151:21237803
5116:28089512
4851:28584727
4784:14961119
4642:17453426
4634:14581590
4558:(2005).
4366:14187297
4018:28089512
3977:24085843
3896:33262517
3853:32457347
3773:21031689
3765:25400084
3593:28171756
3549:25003692
3490:See also
3484:Coccidae
3476:halteres
3163:halteres
3089:Subcosta
2998:Mantodea
2994:crickets
2316:ζεῦγος (
2280:, hair)
2278:trichoma
2258:thysanoi
2238:strepsis
2178:, wing)
2112:, fold)
2086:, lice)
2061:) νέος (
2013:palaios-
1975:or new)
1943:Neoptera
1931:, vein)
1927:νεῦρον (
1823:Isoptera
1809:hymenion
1767:ἑτερο- (
1751:, half)
1671:, life)
1593:Earwigs
1573:Beetles
1563:Κολεός (
1496:ἀνισο- (
1418:eclosion
1403:ectoderm
1359:odonates
1334:Folding
1322:Folding
1241:abdomens
1221:paranota
1208:ontogeny
1200:genetics
1190:record,
1155:muscles.
1152:tracheal
1148:mayflies
1041:Triassic
1013:Devonian
990:myriapod
981:Devonian
949:Devonian
919:thoracic
915:mayflies
897:thoracic
845:Neoptera
833:mayflies
742:bristles
556:remigium
514:Remigium
302:Venation
292:hemocoel
273:Internal
255:mayflies
237:thoracic
185:termites
179:such as
152:or even
122:thoracic
18:Forewing
5979:Related
5837:dactyly
5723:Flipper
5564:4725186
5501:Bibcode
5191:2781901
5054:4986911
4930:9318294
4842:5452959
4792:4431205
4764:Bibcode
4575:Bibcode
4415:3739600
4393:Bibcode
3968:3801059
3945:Bibcode
3876:Bibcode
3844:7250882
3815:Bibcode
3647:4257270
3639:9024659
3619:Bibcode
3512:RoboBee
3480:Diptera
3155:Diptera
3153:In the
3149:Diptera
3093:Radius
3021:elytron
3016:tegmina
2982:earwigs
2678:Odonata
2672:Odonata
2666:mimicry
2440:fourth.
2400:beetles
2296:Ζωρός (
2266:Thrips
2218:, tube)
2214:Σίφων (
2196:rhaphis
2194:ῥαφίς (
2132:, many
2128:Πολύς (
2110:plekein
2084:phtheir
2082:Φθείρ (
2065:, new)
2057:Παρα- (
2041:, old)
2039:Palaios
2019:diktyon
1969:, few)
1967:oligon-
1909:megalo-
1887:μῆκος (
1867:Λόγχη (
1847:Λεπίς (
1769:hetero-
1687:ἐντός (
1639:Diptera
1625:diktyon
1583:Δέρμα (
1547:), not
1518:), not
1511:Aptera
1425:trachea
1282:in the
1280:muscles
1268:airfoil
1251:control
1206:in the
1130:gliding
1128:, then
1116:fossils
1097:Pleurum
1005:Odonata
997:Permian
929:Fossils
881:gliding
729:Sensors
722:Rutilia
713:Odonata
705:Diptera
691:, blue
673:Muscles
334:Odonata
288:trachea
205:locusts
6001:Samara
5810:triped
5795:uniped
5598:
5571:
5561:
5553:
5452:
5398:
5390:
5349:Scales
5328:
5198:
5188:
5149:
5114:
5052:
4984:
4928:
4849:
4839:
4831:
4790:
4782:
4755:Nature
4735:
4665:
4640:
4632:
4524:
4413:
4364:
4225:
4171:
4096:
4066:
4016:
3975:
3965:
3902:
3894:
3851:
3841:
3833:
3771:
3763:
3645:
3637:
3611:Nature
3591:
3547:
3516:DelFly
3457:thrips
3441:hamuli
3095:(Sc+R)
3066:cicada
3028:), or
3026:elytra
3011:tegmen
2734:Radius
2649:scales
2520:flight
2516:pteron
2512:elytra
2404:elytra
2363:, and
2318:zeugos
2226:Fleas
2216:siphon
2156:psocho
2154:Ψώχω (
2092:, not
2015:, old)
1993:ortho-
1991:ὀρθο (
1971:νέος (
1947:νέος (
1929:neuron
1869:lonche
1827:ἶσον (
1787:ὅμο- (
1747:ἡμι- (
1653:Flies
1565:koleos
1498:aniso-
1469:-ptera
1463:pteron
1455:πτερόν
1408:Pieris
1288:mating
1284:thorax
1188:fossil
1144:naiads
1134:flight
999:, the
911:naiads
866:vortex
805:Flight
779:hamuli
752:, and
590:Joints
580:termen
560:clavus
480:Fields
286:and a
251:naiads
201:aphids
197:morphs
193:aphids
158:orders
150:family
111:insect
5859:Wings
5844:Digit
5800:biped
5762:Limbs
5672:wings
5668:limbs
5531:(PDF)
5396:S2CID
5366:(PDF)
5061:(PDF)
5050:S2CID
5030:(PDF)
4811:PeerJ
4788:S2CID
4638:S2CID
4608:(PDF)
4563:(PDF)
4411:JSTOR
4327:(PDF)
4258:(PDF)
4247:(PDF)
3900:S2CID
3769:S2CID
3643:S2CID
3545:JSTOR
3522:Notes
3470:Male
3346:down.
3330:wasps
3169:. In
3052:(pl.
3024:(pl.
3014:(pl.
2793:nodus
2763:nodus
2740:(R+M)
2738:Media
2298:zōros
2138:new)
2130:polys
2059:Para-
1889:mekos
1849:lepis
1789:homo-
1749:hemi-
1727:ἔξω (
1689:entos
1663:ἐν- (
1585:derma
1391:nymph
1379:pupal
1204:genes
907:gills
900:terga
549:Alula
284:nerve
247:gills
240:terga
154:genus
5991:Gait
5882:keel
5680:Fins
5670:and
5664:Fins
5596:ISBN
5569:PMID
5551:ISSN
5450:ISBN
5407:2010
5388:PMID
5326:ISBN
5259:2011
5196:PMID
5147:PMID
5112:PMID
5075:link
4982:ISBN
4926:PMID
4847:PMID
4829:ISSN
4780:PMID
4733:ISBN
4663:ISBN
4630:PMID
4522:ISBN
4362:PMID
4223:ISBN
4169:ISBN
4094:ISBN
4064:ISBN
4032:link
4014:PMID
3973:PMID
3918:link
3892:PMID
3849:PMID
3831:ISSN
3787:link
3761:PMID
3724:link
3690:link
3661:link
3635:PMID
3589:PMID
3506:Wing
3109:(RP)
3103:(RA)
2736:and
2566:and
2524:alae
2384:drag
2380:lift
2378:and
2136:neos
2134:νέος
2063:neos
1973:neos
1949:neos
1829:ison
1669:bios
1543:ἀ- (
1514:ἀ- (
1253:and
1215:and
1198:and
1028:and
971:and
773:and
504:and
347:The
340:and
183:and
181:ants
171:and
144:and
136:and
128:and
5559:PMC
5543:doi
5509:doi
5378:doi
5228:114
5186:PMC
5178:doi
5139:doi
5102:doi
5042:doi
4953:doi
4949:102
4918:doi
4914:200
4887:doi
4837:PMC
4819:doi
4772:doi
4760:427
4620:doi
4616:206
4583:doi
4442:doi
4401:doi
4389:238
4354:doi
4004:doi
3963:PMC
3953:doi
3941:110
3884:doi
3839:PMC
3823:doi
3751:doi
3627:doi
3615:385
3579:doi
3459:),
3040:In
3018:),
2996:),
2984:),
2088:ἀ,
1729:exo
1645:two
1290:or
1169:pod
1146:of
913:of
253:of
6038::
5666:,
5567:.
5557:.
5549:.
5537:.
5533:.
5507:.
5497:48
5491:.
5394:.
5386:.
5374:16
5372:.
5368:.
5340:^
5295:.
5267:^
5249:.
5236:^
5208:^
5194:.
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5145:.
5135:11
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5067:{{
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