Misplaced Pages

#839160

125-418: (traditional, but see below ): Coccinellidae ( / ˌ k ɒ k s ɪ ˈ n ɛ l ɪ d iː / ) is a widespread family of small beetles . They are commonly known as ladybugs in North America and ladybirds in the United Kingdom; "lady" refers to mother Mary . Entomologists use the names ladybird beetles or lady beetles to avoid confusion with true bugs . The more than 6,000 described species have

250-580: A global distribution and are found in a variety of habitats. They are oval beetles with a domed back and flat underside. Many of the species have conspicuous aposematic (warning) colours and patterns, such as red with black spots, that warn potential predators that they taste bad. Most coccinellid species are carnivorous predators, preying on insects such as aphids and scale insects . Other species are known to consume non-animal matter, including plants and fungi. They are promiscuous breeders, reproducing in spring and summer in temperate regions and during

375-554: A lack of widespread consensus within the scientific community for extended periods. The continual publication of new data and diverse opinions plays a crucial role in facilitating adjustments and ultimately reaching a consensus over time. The naming of families is codified by various international bodies using the following suffixes: The taxonomic term familia was first used by French botanist Pierre Magnol in his Prodromus historiae generalis plantarum, in quo familiae plantarum per tabulas disponuntur (1689) where he called

500-444: A large trachea, a cross-section of which is shown in, which represents sections of these parts of the first, second, third and fourth instars respectively. At the same time the tracheoles uncoil, and extend in bundles in the forming vein-cavities of the wing-bud. At the molt that marks the beginning of the pupal stadium stage, they become functional. At the same time, the larval tracheoles degenerate; their function having been replaced by

625-431: A membranous basal area, but the articular membrane contains a number of small articular sclerites, collectively known as the pteralia. The pteralia include an anterior humeral plate at the base of the costal vein, a group of axillaries (Ax) associated with the subcostal, radial, and vannal veins, and two less definite median plates (m, m') at the base of the mediocubital area. The axillaries are specifically developed only in

750-462: 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. While the development of wings in insects is clearly defined in those who are members of Endopterygota , which undergo complete metamorphosis ; in these species, the wing develops while in the pupal stage of the insects life cycle. However, insects that undergo incomplete metamorphosis do not have

875-415: 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

1000-408: A pupal stage, therefore they must have a different wing morphogenesis . Insects such as those that are hemimetabolic have wings that start out as buds, which are found underneath the exoskeleton, and do not become exposed until the last instar of the nymph . The first indication of the wing buds is of a thickening of the hypodermis, which can be observed in insect species as early the embryo, and in

1125-630: A relatively small effect on aphid populations; at others they cause significant seasonal declines. Family (biology) Family ( Latin : familia , pl. : familiae ) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy . It is classified between order and genus . A family may be divided into subfamilies , which are intermediate ranks between the ranks of family and genus. The official family names are Latin in origin; however, popular names are often used: for example, walnut trees and hickory trees belong to

1250-690: A role in something else, such as mating or protection . Some insects, occupying the biological niches that they do, need to be incredibly maneuverable. They must find their food in tight spaces and be capable of escaping larger predators – or they may themselves be predators, and need to capture prey. Their maneuverability, from an aerodynamic viewpoint, is provided by high lift and thrust forces. Typical insect fliers can attain lift forces up to three times their weight and horizontal thrust forces up to five times their weight. There are two substantially different insect flight mechanisms, and each has its own advantages and disadvantages – just because odonates have

1375-502: A sample of their toxic and bitter body fluid. Predator-deterring poisons are particularly important for the immobile pupa. Access to food can affect the concentration of both pigments and toxins. The similarity of coccinellid patterning in red and orange with black markings has led to suggestions that they and some species of chrysomelids form Müllerian mimicry rings particularly to defend them from birds. Despite their chemical defenses, coccinellids are preyed on by some clerid beetles in

SECTION 10

#1732790208840

1500-474: 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). All the veins of the wing are subject to secondary forking and to union by cross-veins. In some orders of insects

1625-404: A vortex over each wing. This bound vortex then moves across the wing and, in the clap, acts as the starting vortex for the other wing. Circulation and lift are increased, at the price of wear and tear on the wings. Many insects can hover by beating their wings rapidly, requiring sideways stabilization as well as lift. A few insects use gliding flight, without the use of thrust. Sometime in

1750-456: A wingless sister taxa to the winged insects. The earliest fliers were similar to dragonflies with two sets of wings, direct flight muscles, and no ability to fold their wings over their abdomens . Most insects today, which evolved from those first fliers, have simplified to either one pair of wings or two pairs functioning as a single pair and using a system of indirect flight muscles. Natural selection has played an enormous role in refining

1875-526: Is a strictly aerobic tissue. Per unit protein it consumes fuel and oxygen at rates taking place in a very concentrated and highly organized tissue so that the steady-state rates per unit volume represent an absolute record in biology. The fuel and oxygen rich blood is carried to the muscles through diffusion occurring in large amounts, in order to maintain the high level of energy used during flight. Many wing muscles are large and may be as large as 10 mm in length and 2 mm in width. Moreover, in some Diptera

2000-411: Is at the leading edge, which is associated at its base with the humeral plate. The trachea of the costal vein is perhaps a branch of the subcostal trachea. Located after the costa is the third vein, the subcosta, which branches into two separate veins: the anterior and posterior. The base of the subcosta is associated with the distal end of the neck of the first axillary (see section below). The fourth vein

2125-402: Is broad and convex, and can cover the back of the head. Being beetles, they have hardened, non-overlapping forewings , known as elytra , which cover up the more fragile hindwings when the insects are not in flight. Their legs are relatively short, with a tarsal formula of 4-4-4 (may appear 3-3-3 because the third segment of each tarsus is reduced). The tarsus (end of leg) has two claws at

2250-571: Is derived from the Latin word coccineus meaning ' scarlet ' . The common English name ladybird originated in Britain where the insects became known as "Our Lady's birds". Mary ("Our Lady") was often depicted wearing a red cloak in early art, and the seven spots of the species Coccinella septempunctata (the most common in Europe) were said to represent her seven joys and seven sorrows . In

2375-821: Is further aided by creases in the membrane. These beetles may migrate long distances to hibernation and breeding sites, and areas with more food. They appear to be drawn to recognisable landmarks. The more crowded an area is, the more individuals leave, but will remain if there are enough prey species to feed on. "Trivial flights" refer to flying while foraging or when finding a place to lay eggs. One study of species in Britain found that coccinellids can fly as far as 120 km (75 mi). They flew at speeds of 30 km/h (19 mph) and could reach altitudes close to 1,100 m (3,600 ft). In temperate climates, coccinellids typically breed from late spring to early summer. In warmer temperate regions, reproduction may occur in spring, fall and winter; tropical species reproduce during

2500-515: Is limited to adult male spiders which are actively searching for females and exposed – unlike the females and young, which remain sheltered in burrows. Coccinellidae are found on every continent except Antarctica. Asian and African species are less studied than others. Coccinellids can be found in a variety of habitats, both on the ground and in the trees. They may specialise using certain plants. Some species can live in extreme environments such as high mountains, arid deserts and cold regions. Several of

2625-429: Is not until the butterfly is in its pupal stage that the wing-bud becomes exposed, and shortly after eclosion , the wing begins to expand and form its definitive shape. The development of tracheation of the wings begin before the wing histoblast form, as it is important to note that they develop near a large trachea . During the fourth instar, cells from the epithelium of this trachea become greatly enlarged extend into

SECTION 20

#1732790208840

2750-438: Is the anterior hinge plate of the wing base. Its anterior part is supported on the anterior notal wing process of the tergum (ANP); its posterior part articulates with the tergal margin. The anterior end of the sclerite is generally produced as a slender arm, the apex of which (e) is always associated with the base of the subcostal vein (Sc), though it is not united with the latter. The body of the sclerite articulates laterally with

2875-416: Is the defining feature ( synapomorphy ) for the infraclass Neoptera . There are two basic aerodynamic models of insect flight. Most insects use a method that creates a spiralling leading edge vortex . Some very small insects use the fling and clap or Weis-Fogh mechanism in which the wings clap together above the insect's body and then fling apart. As they fling open, the air gets sucked in and creates

3000-545: Is the hindwings which are folded, but in a few groups such as the vespid wasps , it is the forewings. The evolutionary origin of the insect wing is debated. During the 19th century, the question of insect wing evolution originally rested on two main positions. One position postulated insect wings evolved from pre-existing structures, while the second proposed insect wings were entirely novel formations. The “novel” hypothesis suggested that insect wings did not form from pre-existing ancestral appendages but rather as outgrowths from

3125-404: Is the most basal diet of Coccinellidae. Aphid-eating evolved three separate times and leaf-eating evolved twice, one of which evolved from a clade that contains both aphid-eating and pollen -eating. The fungi-eating also evolved from aphid-eating. Coccinellids mostly fly during the day. Springy, cylindrical veins in the hindwings stiffen when in flight and bend when folding. Folding of the wings

3250-399: Is the name given to a hypothetical scheme of wing venation proposed for the very first winged insect. It is based on a combination of speculation and fossil data. Since all winged insects are believed to have evolved from a common ancestor, the archedictyon represents the "template" that has been modified (and streamlined) by natural selection for 200 million years. According to current dogma,

3375-432: Is the radius (R), which is branched into five separate veins. The radius is generally the strongest vein of the wing. Toward the middle of the wing, it forks into a first undivided branch (R1) and a second branch, called the radial sector (Ra), which subdivides dichotomously into four distal branches (R2, R3, R4, R5). Basally, the radius is flexibly united with the anterior end of the second axillary (2Ax). The fifth vein of

3500-421: Is usually a small plate intervening between the third axillary and the posterior notal wing process and is probably a detached piece of the latter. The median plates (m, m') are also sclerites that are not so definitely differentiated as specific plates as are the three principal axillaries, but nevertheless they are important elements of the flexor apparatus. They lie in the median area of the wing base distal to

3625-536: The Genera Plantarum of George Bentham and Joseph Dalton Hooker this word ordo was used for what now is given the rank of family. Families serve as valuable units for evolutionary, paleontological, and genetic studies due to their relatively greater stability compared to lower taxonomic levels like genera and species. Hindwing Physically, some insects move their flight muscles directly, others indirectly. In insects with direct flight,

3750-561: The Carboniferous Period , some 350 million years ago, when there were only two major land masses, insects began flying. How and why insect wings developed, however, is not well understood, largely due to the scarcity of appropriate fossils from the period of their development in the Lower Carboniferous. Three main theories on the origins of insect flight are that wings developed from paranotal lobes, extensions of

3875-582: The Cretaceous , the oldest fossils of the group are known from the Oise amber of France, dating to the Early Eocene ( Ypresian ) around 53 million years ago, which belong to the extant genera Rhyzobius and Nephus . The greatest number of fossils comes from the younger Eocene Baltic amber , including members of the extant genera Serangium and Rhyzobius as well as extinct genera belonging to

Coccinellidae - Misplaced Pages Continue

4000-406: The hemocoel , hemolymph can flow into the wings. As the wing develops, the dorsal and ventral integumental layers become closely apposed over most of their area forming the wing membrane. The remaining areas form channels, the future veins, in which the nerves and tracheae may occur. The cuticle surrounding the veins becomes thickened and more heavily sclerotized to provide strength and rigidity to

4125-410: The histoblast become more prominent, which now form a pocket-like structure. As of the third and fourth instars, the histoblast become more elongated. This greatly extended and evaginated, or protruding, part is what becomes the wing. By the close of the last instar, or fifth, the wing is pushed out of the wing-pocket, although continues to lie under the old larval cuticle while in its prepupal stage. It

4250-461: The monophyly (single ancestry) of most of these subfamilies. The monophyly of Coccinellinae has the most support. A 2021 genetic study sampling many species, identified three subfamilies, Microweiseinae (with three tribes), Coccinellinae (26 tribes) and a newly identified group, the Monocoryninae (one tribe). All three subfamilies were strongly supported, but the study noted that although

4375-567: The thoracic terga ; that they are modifications of movable abdominal gills as found on aquatic naiads of mayflies ; or that they developed from thoracic protrusions used as radiators . Fossils from the Devonian (400 million years ago) are all wingless, but by the Carboniferous (320 million years ago), more than 10 different genera of insects had fully functional wings. There is little preservation of transitional forms between

4500-1082: The wet season in tropical regions. Many predatory species lay their eggs near colonies of prey, providing their larvae with a food source. Like most insects, they develop from larva to pupa to adult. Temperate species hibernate and diapause during the winter; tropical species are dormant during the dry season . Coccinellids migrate between dormancy and breeding sites. Species that prey on agricultural pests are considered beneficial insects . Several species have been introduced outside their range as biological control agents, with varying degrees of success. Some species are pests themselves and attack agricultural crops, or can infest people's homes, particularly in winter. Invasive species like Harmonia axyridis can pose an ecological threat to native coccinellid species. Other threats to coccinellids include climate change and habitat destruction . These insects have played roles in folklore , religion and poetry, and are particularly popular in nursery rhymes . The name Coccinellidae , created by Pierre André Latreille in 1807,

4625-469: The wet season . Mating is promiscuous . In some species, females appear to be selective in their partners, preferring males of a certain size and colour. Males produce sperm packets each with 14,000 sperm, and insert three of them into the female, even though she can only hold 18,000 sperm. This is likely a form of sperm competition . Like other insects, coccinellids develop from egg, to larva, to pupa and finally adult. Eggs tend to be bright yellow, and

4750-530: 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

4875-463: The Odonata ( dragonflies and damselflies ) have the flight muscles attached directly to their wings; the wings can beat no faster than the rate at which nerves can send impulses to command the muscles to beat. All other living winged insects fly using a different mechanism, involving indirect flight muscles which cause the thorax to vibrate; the wings can beat faster than the rate at which the muscles receive nerve impulses. This mechanism evolved once, and

5000-471: The Permian and all are amphibious . Their prototypes are the oldest winged fossils, go back to the Devonian , and are different from other wings in every way. Their prototypes may have had the beginnings of many modern attributes even by late Carboniferous and it is possible that they even captured small vertebrates, for some species had a wing span of 71 cm. The earliest beetle-like species during

5125-534: The Permian had pointed, leather like forewings with cells and pits. Hemiptera , or true bugs had appeared in the form of Arctiniscytina and Paraknightia having forewings with unusual venation, possibly diverging from Blattoptera . A single large wing from a species of Diptera in the Triassic (10 mm instead of usual 2–6 mm) was found in Australia (Mt. Crosby).This family Tilliardipteridae, despite

Coccinellidae - Misplaced Pages Continue

5250-467: 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

5375-900: The United States, the name was popularly adapted to ladybug . Entomologists prefer the names ladybird beetles or lady beetles to avoid confusion with true bugs . Names in some other countries may be similar; for example, in Germany they are known as Marienkäfer meaning ' Marybeetle ' or ' ladybeetle ' . Coccinellids range in size from 0.8 to 18 mm (0.03–0.7 in). They are sexually dimorphic ; adult females tend to be slightly larger than males. They are generally oval with domed backs and flattened undersides. They have large compound eyes and clubbed antennae with seven to eleven segments. The powerful mandibles (equivalent to jaws) typically have pairs of "teeth" which face each other. The coccinellid prothorax (front of thorax)

5500-461: The abdominal segments, in particular, each having six divided into pairs, and one to three segmented antennae. Their colouration varies from grey, blue-grey, grey-brown or brown and spotted with white, yellow, red or orange. They tend to brighten as they get closer to adulthood. Over 6,000 living species of Coccinellidae have been described. They are sparsely preserved in the fossil record. Although molecular clock estimates have placed their origin in

5625-414: The aerodynamic efficiency of flight by joining the forewing and hindwing into one bigger wing. The most common coupling mechanism (e.g., Hymenoptera and Trichoptera ) is a row of small hooks on the forward margin of the hindwing, or " hamuli ", which lock onto the forewing, keeping them held together (hamulate coupling). In some other insect species (e.g., Mecoptera , Lepidoptera , and some Trichoptera )

5750-417: The anal area can be folded like a fan. There are about four different fields found on the insect wings: Most veins and crossveins occur in the anterior area of the remigium , which is responsible for most of the flight, powered by the thoracic muscles. The posterior portion of the remigium is sometimes called the clavus ; the two other posterior fields are the anal and jugal ares . When the vannal fold has

5875-402: 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. 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

6000-410: 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

6125-503: The archedictyon contained 6–8 longitudinal veins. These veins (and their branches) are named according to a system devised by John Comstock and George Needham—the Comstock–Needham system : 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

6250-408: The articular membrane often forms an ample lobe between the wing and the body, and its margin is generally thickened and corrugated, giving the appearance of a ligament, the so-called axillary cord, continuous mesally with the posterior marginal scutellar fold of the tergal plate bearing the wing. The articular sclerites, or pteralia, of the wing base of the wing-flexing insects and their relations to

6375-433: 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 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

SECTION 50

#1732790208840

6500-417: 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. 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

6625-429: The base of the wing. The subalar and basilar muscles have ligament attachments to the subalar and basilar sclerites. Here resilin, a highly elastic material, forms the ligaments connecting flight muscles to the wing apparatus. In more derived orders of insects, such as Diptera (flies) and Hymenoptera (wasp), the indirect muscles occupy the greatest volume of the pterothorax and function as the primary source of power for

6750-509: The body and the wing veins, shown diagrammatically, are as follows: The humeral plate is usually a small sclerite on the anterior margin of the wing base, movable and articulated with the base of the costal vein. Odonata have their humeral plate greatly enlarged, with two muscles arising from the episternum inserted into the Humeral plates and two from the edge of the epimeron inserted into the axillary plate. The first axillary sclerite (lAx)

6875-481: The branching of existing veins to produce accessory veins or by the development of additional, intercalary veins between the original ones, as in the wings of Orthoptera (grasshoppers and crickets). Large numbers of cross-veins are present in some insects, and they may form a reticulum as in the wings of Odonata (dragonflies and damselflies) and at the base of the forewings of Tettigonioidea and Acridoidea (katydids and grasshoppers respectively). The archedictyon

7000-556: The brood of the ant Wasmannia auropunctata . Cannibalism has been recorded in several species; which includes larvae eating eggs or other larvae, and adults feeding on individuals of any life stage. Some coccinellids are mostly non-predatory, such as some species in the genera Epilachna and Henosepilachna . The majority of predatory species may also supplement their diet with other sources of food both in their larval and adult stages. Non-animal matter consumed include leaves, pollen, nectar , sap , fungi, and honeydew . Members of

7125-416: The cavity of the wing bud, with each cell having developed a closely coiled tracheole . Each trachcole is of unicellular origin, and is at first intracellular in position; while tracheae are of multicellular origin and the lumen of each is intercellular in position. The development of tracheoles, each coiled within a single cell of the epithelium of a trachea, and the subsequent opening of communication between

7250-454: 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

7375-408: The claval furrow and jugal fold are probably homologous in different species, the vannal fold varies in position in different taxa. Folding is produced by a muscle arising on the pleuron and inserted into the third axillary sclerite in such a way that, when it contracts, the sclerite pivots about its points of articulation with the posterior notal process and the second axillary sclerite. As a result,

7500-403: The cross-veins are so numerous that the whole venational pattern becomes a close network of branching veins and cross-veins. Ordinarily, however, there is a definite number of cross-veins having specific locations. The more constant cross-veins are the humeral cross-vein (h) between costa and subcosta, the radial cross-vein (r) between R and the first fork of Rs, the sectorial cross-vein (s) between

7625-429: The distal arm of the third axillary sclerite rotates upwards and inwards, so that finally its position is completely reversed. The anal veins are articulated with this sclerite in such a way that when it moves they are carried with it and become flexed over the back of the insect. Activity of the same muscle in flight affects the power output of the wing and so it is also important in flight control. In orthopteroid insects,

SECTION 60

#1732790208840

7750-407: The earliest stages of the life cycle. During the development of morphological features while in the embryo, or embryogenesis , a cluster of cells grow underneath the ectoderm which later in development, after the lateral ectoderm has grown dorsally to form wind imaginal disc. An example of wing bud development in the larvae, can be seen in those of White butterflies ( Pieris ). In the second instar

7875-492: 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. Two groups of relatively large insects, the Ephemeroptera ( mayflies ) and

8000-542: The family Juglandaceae , but that family is commonly referred to as the "walnut family". The delineation of what constitutes a family— or whether a described family should be acknowledged— is established and decided upon by active taxonomists . There are not strict regulations for outlining or acknowledging a family, yet in the realm of plants, these classifications often rely on both the vegetative and reproductive characteristics of plant species. Taxonomists frequently hold varying perspectives on these descriptions, leading to

8125-540: The family as a rank intermediate between order and genus was introduced by Pierre André Latreille in his Précis des caractères génériques des insectes, disposés dans un ordre naturel (1796). He used families (some of them were not named) in some but not in all his orders of "insects" (which then included all arthropods ). In nineteenth-century works such as the Prodromus of Augustin Pyramus de Candolle and

8250-431: The female is winged but the male not, as in fig wasps . In some cases, wings are produced only at particular times in the life cycle, such as in the dispersal phase of aphids . Wing structure and colouration often vary with morphs , such as in the aphids , migratory phases of locusts and polymorphic butterflies . At rest, the wings may be held flat, or folded a number of times along specific patterns; most typically, it

8375-406: The females lay them close together, standing upright and near where they can access food. The number of eggs in a cluster can vary depending on the species; it is typically in the double digits but some species can lay over a thousand eggs in their lifetime. After hatching, the larvae will begin eating, including the other eggs in their clutch. Certain species lay extra infertile trophic eggs with

8500-402: The fertile eggs, providing a backup food source for the larvae when they hatch. The ratio of infertile to fertile eggs increases with scarcity of food at the time of egg laying. Larvae typically have four instar stages with three moults between them. The larva eventually transitions into a pupa; which involves the development of a hunch, the fusion of the legs to the body, and the attachment of

8625-428: The fibres are of giant dimensions. For instance, in the very active Rutilia , the cross-section is 1800 μm long and more than 500 μm wide. The transport of fuel and oxygen from the surroundings to the sites of consumption and the reverse transport of carbon dioxide therefore represent a challenge to the biologist both in relation to transport in the liquid phase and in the intricate system of air tubes, i.e. in

8750-423: The flexion lines. Though fold lines may be transverse, as in the hindwings of beetles and earwigs, they are normally radial to the base of the wing, allowing adjacent sections of a wing to be folded over or under each other. The commonest fold line is the jugal fold, situated just behind the third anal vein, although, most Neoptera have a jugal fold just behind vein 3A on the forewings. It is sometimes also present on

8875-401: The fold-lines is often blurred, as fold-lines may permit some flexibility or vice versa. Two constants that are found in nearly all insect wings are the claval (a flexion-line) and jugal folds (or fold line); forming variable and unsatisfactory boundaries. Wing foldings can be very complicated, with transverse folding occurring in the hindwings of Dermaptera and Coleoptera , and in some insects

9000-451: The fulcral wing process of the pleuron. The second axillary, therefore, is the pivotal sclerite of the wing base, and it specifically manipulates the radial vein. The third axillary sclerite (3Ax) lies in the posterior part of the articular region of the wing. Its form is highly variable and often irregular, but the third axillary is the sclerite on which is inserted the flexor muscle of the wing (D). Mesally it articulates anteriorly (f) with

9125-425: The genus Enoclerus , several species of which are brightly coloured in red and black, and which possibly sequester the toxins of the prey to defend themselves against other predators. As an anti-predator defense, spiders of the genus Eresus , known as ladybird spiders, have evolved to replicate the patterns of coccinellids. This is a form of Batesian mimicry , as the spiders lack the chemicals. This resemblance

9250-477: 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

9375-401: The insect body wall. Long since, research on insect wing origins has built on the “pre-existing structures” position that was originally proposed in the 19th century. Recent literature has pointed to several ancestral structures as being important to the origin of insect wings. Among these include: gills, respiratory appendages of legs, and lateral (paranotal) and posterolateral projections of

9500-493: The insects, including citizen science and education programs, habitat preservation and restoration, prevention of the spread of invasive species and a global monitoring program. Coccinellids have been valued in biological pest control , as they prey on agricultural pests such as aphids and scale insects. Their importance in controlling pests was noted as far back as 1814 in England. Their efficiency can vary: sometimes they have

9625-458: The jugal area of the forewing is developed as a free lobe, it projects beneath the humeral angle of the hindwing and thus serves to yoke the two wings together. In the Jugatae group of Lepidoptera it bears a long finger-like lobe. The jugal region was termed the neala ("new wing") because it is evidently a secondary and recently developed part of the wing. The axillary region is region containing

9750-399: The jugal lobe of other insects (A, D); the larger inner squama (d) arises from the posterior scutellar margin of the tergum of the wing-bearing segment and forms a protective, hoodlike canopy over the haltere. In the flexed wing the outer squama of the alula is turned upside down above the inner squama, the latter not being affected by the movement of the wing. In many Diptera a deep incision of

9875-472: The jugal lobe of the forewing covers a portion of the hindwing (jugal coupling), or the margins of the forewing and hindwing overlap broadly (amplexiform coupling), or the hindwing bristles, or frenulum, hook under the retaining structure or retinaculum on the forewing. When at rest, the wings are held over the back in most insects, which may involve longitudinal folding of the wing membrane and sometimes also transverse folding. Folding may sometimes occur along

10000-411: The larva stage lasts around three weeks and the pupa lasts seven to ten days. Adult coccinellids develop much of their final colouration within hours, but may not fully darken for weeks or months. The lifespan of an adult reaches up to a year. In temperate areas, coccinellids may hibernate or enter diapause during the winter. Individuals during this period gather in clumps, large or small depending on

10125-426: The larvae of moths and other beetles, as well as mites . Since much of their prey are agricultural pests, coccinellids are considered to be beneficial insects . A 2009 metastudy by Hodek and Honěk found that aphid-eaters constituted around 68 percent of species that live in temperate areas but only 20 percent of species worldwide. Around 36 percent of total species mostly feed on scale insects. Larvae and adults eat

10250-431: The lighter colours, and melanins create darker colours. Other parts of the body also vary in colouration. These colour patterns typically serve as warning colouration , but some can act as camouflage , attract mates or even regulate heat. Several individual species may display polymorphism and even change colour between seasons. Coccinellid larvae are elongated with square heads. They are covered in hairs or setae ,

10375-474: The main stem of the cubitus is associated with the distal median plate (m') of the wing base. Postcubitus (Pcu) is the first anal of the Comstock–Needham system. The postcubitus, however, has the status of an independent wing vein and should be recognized as such. In nymphal wings, its trachea arises between the cubital trachea and the group of vannal tracheae. In the mature wings of more generalized insect

10500-539: The most famous species have wide ranges, but others are more endemic and possibly threatened. Threats to coccinellids include climate change , agriculture, urbanisation, and invasive species . Coccinellid biodiversity will likely be affected by the rising of both average temperatures and heat fluctuations. Climate change may lead to smaller larvae, as well as increase energy and metabolic needs and interspecific predation. Agriculture and urbanisation threatens these insects though habitat destruction and homogenisation and

10625-504: 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. Suggestions have been made that wings may have evolved initially for sailing on the surface of water as seen in some stoneflies . An alternative idea is that it derives from directed aerial gliding descent—a preflight phenomena found in some apterygote ,

10750-449: The opposite position between the two branches. A concave vein will fork into two concave veins (with the interpolated vein being convex) and the regular alteration of the veins is preserved. The veins of the wing appear to fall into an undulating pattern according to whether they have a tendency to fold up or down when the wing is relaxed. The basal shafts of the veins are convex, but each vein forks distally into an anterior convex branch and

10875-458: The paranotal lobe theory, the gill theory and the dual theory of insect wing evolution. These theories postulate that wings either developed from paranotal lobes, extensions of the thoracic terga ; that they are modifications of movable abdominal gills as found on aquatic naiads of mayflies ; or that insect wings arose from the fusion of pre-existing endite and exite structures each with pre-existing articulation and tracheation. Each of

11000-399: The posterior end of the second axillary, and posteriorly (b) with the posterior wing process of the tergum (PNP), or with a small fourth axillary when the latter is present. Distally the third axillary is prolonged in a process which is always associated with the bases of the group of veins in the anal region of the wing here termed the vannal veins (V). The third axillary, therefore, is usually

11125-413: 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

11250-479: The posterior to the surface. Pupae may be uncovered, partially covered or fully covered by larval skin depending on the species. The pupa is mostly immobile, but the head can move in response to irritation. When the adult emerges, it has its hindwings, while the elytron starts out softer and lighter in colour, with no patterns. The length of each development stage varies based on climate and between species. For Adalia bipunctata , eggs hatch after four to eight days,

11375-969: The same foods, unlike in other insect groups. Ladybird species vary in dietary specificity . An example of a specialist species is those of the genus Stethorus , which feed on spider mites . Aphid-eaters tend to be generalist; they have a high voracity and can multiply quickly in response to outbreaks, and switch to other prey when the ephemeral aphids become scarce. Predators of scale insects tend to be less voracious and are slower breeders and developers; matching their prey. Under pressure from coccinellid predation, aphid species have evolved to become more toxic, forcing coccinellids to develop immunities. Coccinellid predators of aphids need to defend themselves against ants that tend and defend aphids for their honeydew, and coccinellid eggs laid near aphids are disposed of. Some species including Coccinella magnifica and Diomus have adapted to grow within ant nests as larvae, and some like Diomus thoracicus are predators of

11500-418: The second and third axillaries and are separated from each other by an oblique line (bf) which forms a prominent convex fold during flexion of the wing. The proximal plate (m) is usually attached to the distal arm of the third axillary and perhaps should be regarded as a part of the latter. The distal plate (m') is less constantly present as a distinct sclerite and may be represented by a general sclerotization of

11625-432: 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

11750-575: The seventy-six groups of plants he recognised in his tables families ( familiae ). The concept of rank at that time was not yet settled, and in the preface to the Prodromus Magnol spoke of uniting his families into larger genera , which is far from how the term is used today. In his work Philosophia Botanica published in 1751, Carl Linnaeus employed the term familia to categorize significant plant groups such as trees , herbs , ferns , palms , and so on. Notably, he restricted

11875-628: The species. Overwintering insects can be found both in lowland areas, aggregating under dead vegetation, and at the tops of hills, hibernating under rocks and on grass tussocks . In areas with particularly hot summers, the insects experience summer dormancy or aestivation ; in the tropics, coccinellids enter dormancy during the dry season . Coccinellids act both as predators, prey and parasitic hosts in food webs . The majority of coccinellids are carnivorous and predatory , typically preying on Sternorrhyncha insects like aphids, scale insects, whiteflies , psyllids and adelgids . Some species feed on

12000-453: The tergum instead of the wings, as the name suggests. As the muscles contract, the thoracic box becomes distorted, transferring the energy to the wing. There are two "bundles" of muscles, those that span parallel to the tergum, the dorsolongitudinals, and those that are attached to the tegum and extend to the sternum, the dorsoventrals. In direct muscle, the connection is directly from the pleuron (thoracic wall) to individual sclerites located at

12125-483: 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

12250-538: 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

12375-425: The thorax to name a few. According to more current literature, possible candidates include gill-like structures, the paranotal lobe, and the crustacean tergal plate. The latter is based on recent insect genetic research which indicates that insects are pan-crustacean arthropods with a direct crustacean ancestor and shared genetic mechanisms of limb development. Other theories of the origin of insect wings are

12500-443: The tip. As adults, these beetles differ from their closest relatives with the following morphological characteristics: Coccinellids are often distinctively coloured and patterned. The elytron may be light with dark spots or dark with light spots. Light areas are typically yellow, red, orange or brown, and the spots vary in size and shape and numbers. Some species have striped or checkered patterns . The pigment carotene creates

12625-428: The tracheal system. Several types of sensory neurons are found on insect wings: gustatory bristles , mechanosensory bristles, campaniform sensilla , and chordotonal organs . These sensors provide the nervous system with both external and internal proprioceptive feedback necessary for effective flight and grooming. In many insect species, the forewing and hindwing can be coupled together, which improves

12750-403: The tracheoles and the lumen of the trachea, and the uncoiling and stretching out of the tracheoles, so that they reach all parts of the wing. In the earlier stages of its development, the wing-bud is not provided with special organs of respiration such as tracheation, as it resembles in this respect the other portions of the hypodermis of which it is still a part. The histoblast is developed near

12875-741: The tribe Halyziini of the subfamily Coccinellinae are obligate fungus feeders. Coccinellids of any lifestage are preyed on by predators such as birds, spiders, ants and lacewings . They are also hosts for parasites, including some flies , ticks , mites, hymenopterans and nematodes , and pathogens, including bacteria, fungi and protozoa . Wolbachia bacteria infects eggs and kills male zygotes. The promiscuity of Coccinellids has led to their being affected by sexually transmitted infections . The bright warning colouration of many coccinellids discourage potential predators , warning of their toxicity . A 2015 study of five ladybird species found that their colouration honestly signalled their toxicity, implying

13000-1196: The tribes Microweiseini ( Baltosidis ) and Sticholotidini ( Electrolotis ). The Coccinellidae are within the superfamily Coccinelloidea , which in turn is part of the infraorder Cucujiformia , a group containing most of the plant-eating beetles. The ladybirds form the majority of the species in the Coccinelloidea; many of the rest are fungus-feeding beetles or scavengers . Lymexyloidea [REDACTED] Tenebrionoidea [REDACTED] Cleroidea [REDACTED] Chrysomeloidea [REDACTED] Curculionoidea [REDACTED] Cucujoidea [REDACTED] Bothrideridae and allies [REDACTED] Latridiidae [REDACTED] Akalyptoischiidae [REDACTED] Alexiidae [REDACTED] Corylophidae and allies [REDACTED] Endomychidae [REDACTED] Coccinellidae [REDACTED] Coccinellidae have historically been divided into up seven subfamilies ( Chilocorinae , Coccidulinae , Coccinellinae , Epilachninae , Microweiseinae , Scymninae and Sticholotidinae ) and 35 tribes based on morphology. However, genetics studies have called into question

13125-775: The tribes are mostly monophyletic, their relationships are only weakly supported. The study suggests that the crown group appeared some 143 Mya in the Early Cretaceous , and that the group diversified rapidly during the Late Cretaceous, perhaps because the growth in diversity of angiosperm plants then encouraged the radiation of insects of the clade Sternorrhyncha such as aphids , on which ladybirds could feed. Microweiseinae [REDACTED] Monocoryninae Stethorini [REDACTED] Coccinellini [REDACTED] other tribes [REDACTED] An earlier 2009 study concluded that consumption of scale insects

13250-413: The two forks of R8, the median cross-vein (m–m) between M2 and M3, and the mediocubital cross-vein (m-cu) between media and cubitus. The veins of insect wings are characterized by a convex-concave placement, such as those seen in mayflies (i.e., concave is "down" and convex is "up") which alternate regularly and by its triadic type of branching; whenever a vein forks there is always an interpolated vein of

13375-538: The two periods. The earliest winged insects are from this time period ( Pterygota ), including the Blattoptera , Caloneurodea , primitive stem-group Ephemeropterans , Orthoptera and Palaeodictyopteroidea . Very early Blattopterans (during the Carboniferous) had a very large discoid pronotum and coriaceous forewings with a distinct CuP vein (an unbranched wing vein, lying near the claval fold and reaching

13500-558: The use of pesticides . Invasive threats include other coccinellids, particularly C. septempunctata in North America and H. axyridis globally. These invaders outcompete the native species as well as eat their eggs. As of 2022, the IUCN Red List does not list the conservation status for any coccinellid, though there is an IUCN SSC Ladybird Specialist Group. Conservationists have suggested several measures for protecting

13625-497: The use of this term solely within the book's morphological section, where he delved into discussions regarding the vegetative and generative aspects of plants. Subsequently, in French botanical publications, from Michel Adanson 's Familles naturelles des plantes (1763) and until the end of the 19th century, the word famille was used as a French equivalent of the Latin ordo (or ordo naturalis ). In zoology ,

13750-417: 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

13875-433: The vannal fold is lost, but even in such cases the flexed wing may bend along a line between the postcubitus and the first vannal vein. The Jugal Region, or Neala, is a region of the wing that is usually a small membranous area proximal to the base of the vannus strengthened by a few small, irregular veinlike thickenings; but when well developed it is a distinct section of the wing and may contain one or two jugal veins. When

14000-410: 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. Wing areas are delimited and subdivided by fold-lines along which the wing can fold, and flexion-lines along which the wing can flex during flight. The fundamental distinction between the flexion-lines and

14125-432: The veins in such a way that the cross-section of the wings approximates an airfoil . Thus, the wing's basic shape already is capable of generating a small amount of lift at zero angle of attack . Most insects control their wings by adjusting tilt, stiffness , and flapping frequency of the wings with tiny muscles in the thorax (below). Some insects evolved other wing features that are not advantageous for flight, but play

14250-463: The warning is genuine. Species with more contrast with the background environment tended to be more toxic. Coccinellid haemolymph (blood) contains toxic alkaloids , azamacrolides and polyamines , as well as foul-smelling pyrazines . Coccinellids can produce at least 50 types of alkaloids. When disturbed, ladybirds further defend themselves with reflex bleeding , exuding drops from their tibio-femoral (knee) joints, effectively presenting predators with

14375-431: The wing at the base of the mediocubital field. The termen is the outer margin of the wing, between apex and hind or anal angle. At the posterior angle of the wing base in some Diptera there is a pair of membranous lobes (squamae, or calypteres) known as the alula. The alula is well developed in the house fly. The outer squama (c) arises from the wing base behind the third axillary sclerite (3Ax) and evidently represents

14500-424: The wing base. The vannus is bordered by the vannal fold, which typically occurs between the postcubitus and the first vannal vein. In Orthoptera it usually has this position. In the forewing of Blattidae, however, the only fold in this part of the wing lies immediately before the postcubitus. In Plecoptera the vannal fold is posterior to the postcubitus, but proximally it crosses the base of the first vannal vein. In

14625-410: The wing is the media. In the archetype pattern (A), the media forks into two main branches: a media anterior (MA), which divides into two distal branches (MA1, MA2), and a median sector, or media posterior (MP), which has four terminal branches (M1, M2, M3, M4). In most modern insects the media anterior has been lost, and the usual "media" is the four-branched media posterior with the common basal stem. In

14750-462: The wing muscles directly attach to the wing base, so that a small downward movement of the wing base lifts the wing itself upward. Those insects with indirect flight have muscles that attach to and deform the thorax, causing the wings to move as well. The wings are present in only one sex (often the male) in some groups such as velvet ants and Strepsiptera , or are selectively lost in "workers" of social insects such as ants and termites . Rarely,

14875-593: The wing posterior margin). Even though the oldest possible insect fossil is the Devonian Rhyniognatha hirsti , estimated at 396–407 million years old, it possessed dicondylic mandibles, a feature associated with winged insects, although it is later considered as possible myriapod . During the Permian , the dragonflies ( Odonata ) were the dominant aerial predator and probably dominated terrestrial insect predation as well. True Odonata appeared in

15000-479: 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

15125-411: 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

15250-470: 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. In some very small insects, the venation may be greatly reduced. In chalcidoid wasps , for instance, only the subcosta and part of the radius are present. Conversely, an increase in venation may occur by

15375-409: The wings consists of a thin membrane supported by a system of veins. The membrane is formed by two layers of integument closely apposed, while the veins are formed where the two layers remain separate; sometimes the lower cuticle is thicker and more heavily sclerotized under a vein. Within each of the major veins there is a nerve and a trachea , and, since the cavities of the veins are connected with

15500-447: The wings, control and sensory systems , and anything else that affects aerodynamics or kinematics . One noteworthy trait is wing twist. Most insect wings are twisted, as are helicopter blades, with a higher angle of attack at the base. The twist generally is between 10 and 20 degrees. In addition to this twist, the wing surfaces are not necessarily flat or featureless; most larger insects have wing membranes distorted and angled between

15625-425: The wingstroke. Contraction of the dorsolongitudinal muscles causes the severe arching of the notum which depresses the wing while contraction of the dorsoventral muscles causes opposite motion of notum. The most primitive extant flying insects, Ephemeroptera (mayflies) and Odonata (dragonflies), use direct muscles that are responsible for developing the needed power for the up and down strokes. Insect wing muscle

#839160