/koh'lee op"teuhr euhn, kol'ee-/, adj.
1. belonging or pertaining to the order Coleoptera.
2. a beetle.
[1840-50; COLEOPTER(A) + -AN]

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  any member of the insect order Coleoptera, consisting of the beetles and weevils. It is the largest order of insects, representing about 40 percent of the known insect species. Among the over 350,000 species of Coleoptera are many of the largest and most conspicuous insects, some of which also have brilliant metallic colours, showy patterns, or striking form. Beetles can usually be recognized by their two pairs of wings; the front pair is modified into horny covers (elytra) that hide the rear pair and most of the abdomen and usually meet down the back in a straight line. Coleoptera occur in nearly all climates. They may be divided into four groups: the first three, the Archostemata, the Adephaga, and the Myxophaga, contain relatively few families; the majority of beetles are placed in the fourth group, the Polyphaga.

  Beetles attract attention for many different reasons, including their economic importance, size, abundance, appearance, and remarkable habits. Several groups of beetles (e.g., Lampyridae) are among the few terrestrial animals capable of producing light; members of several other families (e.g., Cerambycidae) can produce sound (stridulate). Most large beetles make a loud noise during flight, and many species, both large and small, are attracted to light at night. Some beetles (e.g., burying beetles of the family Silphidae and whirligig beetles of the family Gyrinidae) attract attention by their bizarre habits; others do so by their grotesque forms (e.g., Scarabaeidae). Many beetles have become adapted to an aquatic environment (e.g., Hydrophilidae); others (e.g., Thorictidae) live in association with ants and termites.

General features

Distribution and abundance
  Beetles are found in nearly all climates and latitudes, except in such extreme environments as those in Antarctica and at the highest altitudes. They are found on sub-Antarctic islands, close to the northern extremes in the Arctic, and on many mountain tops. Although many species occur in temperate environments, the number of species is greatest in the tropics; in general, individuals of a species are most abundant in temperate areas, and fewer individuals of more species are found in the tropics.

 Some species are solitary, others occur in aggregations. Predators like the ground beetles (ground beetle) (Carabidae) are more apt to be found alone, as are many long-horned wood-boring beetles (Cerambycidae (long-horned beetle)) and weevils (Curculionidae). Ladybugs (Coccinellidae (ladybird beetle)), leaf beetles (Chrysomelidae), pleasing fungus beetles (pleasing fungus beetle) (Erotylidae), darkling beetles (darkling beetle) (Tenebrionidae), checkered beetles (checkered beetle) (Cleridae), bess beetles (bess beetle) (Passalidae), sap beetles (sap beetle) (Nitidulidae), and some species of scarab beetles (scarab beetle) (Scarabaeoidea) are often found in aggregations of one or several different species.

      Most families contain both widely distributed species and some with very limited ranges. Wide distribution in this sense refers to a zoogeographical or faunal region, limited distribution to a single valley, plain, island, altitude zone, or vegetation type on a mountain.

Size range and diversity of structure
 Coleoptera vary greatly in size, from a fraction of a millimetre to more than 200 mm (almost 8 inches) in length (e.g., rhinoceros beetle, Xyloryctes satyrus) and up to 75 mm (2.95 inches) in width (e.g., goliath beetle, Goliathus goliathus).

      Diversity of structure among adult beetles is as great as range of size. The ground beetles (Carabidae) have a rather generalized (primitive) form—the flattened, oval body has a relatively even surface, with regular ridges or grooves; antennae and legs are of moderate length and slender. The underside of most water beetles (Hydrophilidae) is oval, smooth, and flattened, the antennae either short or very slender, and the forelegs short, the hindlegs long and fringed with hairs that are used as paddles. Rove beetles (rove beetle) (Staphylinidae) have very short elytra and a slender abdomen. Soldier beetles (soldier beetle) (Cantharidae), fireflies (Lampyridae (firefly)), and net-winged beetles (net-winged beetle) (Lycidae) have soft elytra.

      Click beetles (click beetle) (Elateridae) have a hingelike joint in the body region called the thorax that enables them to snap their bodies and jump high in the air; their relatives, the Buprestidae (metallic wood-boring beetle) (metallic wood borers), cannot jump but take flight very quickly. Cleridae (checkered beetles) are usually oblong or cylindrical, fairly active, and often brightly coloured. Nitidulidae (sap beetles) are short, flattened, and have slightly shortened elytra. Coccinellidae (ladybugs, ladybird beetles) are rounded, with a smooth, raised upper surface and a flat underside. The Endomychidae (handsome fungus beetles) often have enlarged, rounded elytra. The Erotylidae (pleasing fungus beetles) are usually slender, smooth, and shiny, as are the Languriidae.

      Among the stout or cylindrical lamellicorns (Scarabaeoidea) are a number of bizarre forms. The male rhinoceros beetles (Dynastinae) have one or more horns on the head and sometimes on part of the thorax. Many of the true scarabs (Scarabaeinae) and other dung-feeding groups of the lamellicorns also have horns, including some of the goliath beetles (Cetoniinae (flower chafer)). Male stag beetles (stag beetle) (Lucanidae) have greatly enlarged mandibles (jaws); some are as long as the rest of the body.

      The Chrysomelidae (leaf beetles (leaf beetle)) vary from simple eggshaped forms to slender, flat, or wedge-shaped ones, with wide elytra in the tortoise beetles (Cassidinae) and often numerous spines or tubercles in the leaf-mining leaf beetles (Hispinae). The Bruchidae (pea weevils) are short and stumpy, with short stout legs. The head, slightly elongated in front, is similar to that of some curculionid weevils. The Cerambycidae (long-horned wood-boring beetles), diverse in form and structure, usually have antennae that are longer than the body. Cerambycids may be slender and medium to large in size or very small.

      The extremely diverse Tenebrionidae (darkling beetles) are not always recognized as members of one family. Most arboreal (tree-dwelling) forms in the tropics or subtropics are slender and long legged. Some slender and egg-shaped forms have a metallic sheen; most of the ground-dwelling forms are black and robust. The large, tropical Trictenotomidae resemble some cerambycids (Prioninae) but are not related to them. The Alleculidae (comb-clawed beetles) resemble some of the slender tenebrionids but are usually more active. Lagriidae (lagriids) have a characteristic shape, usually widened behind, and sometimes a metallic sheen. Colydiidae (cylindrical bark beetles), hard-bodied and shiny or roughened, may be cylindrical and somewhat flattened.

      The Curculionidae (weevils (weevil)) range from slender to stout, elongated to egg-shaped; the bodies of some species contain many rounded projections (tubercles), those of others may be smooth or grooved. The mouth is located on the end of a snoutlike projection, which varies in shape from short and stout to long and slender and sometimes exceeds the length of the rest of the body. Some Anthribidae (fungus weevils (fungus weevil)), usually cylindrical in shape, have slender antennae that may be longer than the rest of the body; they are easily confused with the cerambycids. Brenthidae (primitive weevils) usually are long and slender with antennae projecting from the sides of the snout. Scolytidae (bark beetles (bark beetle), ambrosia beetles) do not have a distinct snout and are usually cylindrical in shape, as are Platypodidae.


As predators
      Predators such as Carabidae (ground beetles) and Staphylinidae (rove beetles) help to control the populations of many insects by feeding on caterpillars and other immature insects (larvae), many soft-bodied adult insects, and insect eggs. Most of the Coccinellidae (ladybugs, ladybird beetles) are highly beneficial to man; both larvae and adults feed on plant-sucking insects (Homoptera) such as aphids and scale insects. Only a few coccinellids (e.g., Epilachna) feed on plants.

As plant feeders
      Most of the beetles and weevils harmful to man are phytophagous (plant feeders). Of primary importance are the leaf beetles (Chrysomelidae) and the weevils and their relatives (Curculionoidea). Leaf-beetle larvae feed on leaves, stems, or roots of plants; and most adults chew leaves. Various species of weevil larvae or adults have been found feeding on almost every plant part; especially numerous are species that bore into trunks, stems, and seeds. Both larval and adult forms of Scolytidae (bark beetles) are serious pests; they feed beneath tree bark, harming vital areas of living trees (e.g., cambium, pine needles, leaf stalks). The Platypodidae have similar habits, but the tropical Brenthidae (primitive weevils) usually attack deadwood.

      The long-horned beetles (Cerambycidae) bore into stems, trunks, roots, and cones of living and dead trees and large semiwoody herbs; adults often feed on tender, new bark. Most pea-weevil larvae (Bruchidae) develop in dried seeds of leguminous plants (peas, beans). Buprestidae (metallic wood borers) have habits similar to those of cerambycids, and many kill trees or branches by boring in the cambium. The scarab beetles (Scarabaeidae) include many important pests of crop plants, lawns, and pastures. The larvae of many Melolonthinae (June beetles (June beetle), chafers (chafer)), for example, feed on grass roots. The Dynastinae (rhinoceros, unicorn (unicorn beetle), and elephant beetles) are often pests of palms, killing them by destroying the growing points. Lumber, furniture, and other items made from wood are sometimes severely damaged by several groups of beetles that bore in dry wood; e.g., powder-post beetles (Lyctidae), deathwatch beetles (Anobiidae (borer beetle)), and branch borers (Bostrychidae). Some anobiids also feed on various types of dried products; e.g., the cigarette beetle, Lasioderma serricorne, feeds on tobacco, various dried foods, and drugs.

As scavengers
      Many groups of beetles function as scavengers, breaking down materials such as dead logs, lumber used in houses (in which case they are pests), dead plant and animal matter, excrement, and other waste products. Coleopterans that function as scavengers include Scarabaeidae, Tenebrionidae (darkling beetles), Silphidae (carrion and burying beetles (carrion beetle)), and Dermestidae (dermestid or hide beetles (dermestid beetle)). Some dermestid species cause serious damage in museums by feeding on dried animal materials. The larvae of several species of small dermestids damage carpets, upholstery, and clothing. However, some dermestids are valuable as scavengers; some of the carrion-feeding species (e.g., Dermestes caninus) are used by zoologists to clean skeletons of animals.

In transmitting plant (plant disease) diseases
      Little is known concerning the role of beetles in transmitting plant diseases. Since beetles do not suck plant juices as do plant-sucking insects (Homoptera), there is less likelihood of disease transmission. Transmission of diseases may occur, however, if beetles carry fungal spores on or in the body; e.g., the fungus that causes Dutch elm disease is transmitted by the European elm bark beetle, Scolytus multistriatus.

Natural history

Reproduction and life cycle
General features
      Reproduction is almost always bisexual in beetles, although some species are always parthenogenetic (reproducing without fertilization), and consist of females only. In some species parthenogenesis can sometimes occur. The male reproductive organ is a hardened tubelike structure called the aedeagus. The aedeagus enters a structure in the tip of the abdomen of the female (bursa copulatrix), and the sperm are stored in a saclike structure in the female (spermatheca) until they are needed to fertilize eggs. The females of most species lay eggs. After the larva has hatched, it feeds until its skin (cuticle) becomes too small and splits; the larva crawls out of the old skin (exuviae), and a new skin forms and hardens. The process, called molting, is repeated, usually from three to five times, until the larva is mature. During a nonfeeding period (prepupal stage) the insect enters the pupal stage. The pupa, which forms beneath the final larval skin and emerges when it splits, resembles the adult, except that it is soft and pale; in addition, the appendages are curled or loosely attached to the body, and the wings are folded in flat bags called wing pads. The adult emerges when the thin skin of the pupa is shed; the wings stretch out to full size, and the new outer skeleton hardens and becomes coloured. No further growth of the hardened skeleton occurs; the abdomen of the gravid (pregnant) female may enlarge, by stretching of membranes between the abdominal segments. The four developmental stages of Coleoptera—egg, larva, pupa, adult—constitute complete metamorphosis. The length of each stage in the life cycle depends on several factors; e.g., climate, nature of habitat, available food.

      Eggs vary in form, may be laid singly or in groups, and usually are laid at a site that allows proper development of the larva—on a leaf of a host plant (leaf-eating species), in bark, or in tree trunks (wood-borers). Eggs also may be laid near roots, in flowers, in fruits, in tree injuries, on water plants, or under rocks.

      There are several types of coleopteran larvae. Carabid larvae have a tapering, flattened, smooth body, as do those of staphylinids (rove beetles) and silphids (carrion beetles); larvae of the Dytiscidae (diving beetles), although somewhat similar to those of carabids, have a lobed air-float at the end. Larvae of click beetles (Elateridae) are cylindrical or flat and slender and have a hard surface; some click beetle larvae, called wireworms, feed on newly planted seeds and roots of plant crops (e.g., maize, cotton, potatoes); others feed in deadwood or on wood-boring beetle larvae (Cerambycidae). Larvae of Buprestidae (metallic wood borers), which are soft-bodied and slender, bore under the bark of trees or burrow beneath the surface of leaves.

 Dermestid larvae, somewhat tapering and cylindrical, have whorls of short bristles and some longer ones and resemble hedgehogs or porcupines. Coccinellid larvae, flattened, broad in the middle and tapering at the back, sometimes have a few low projections (tubercles) bearing short hairs and are often strikingly coloured with red or yellow and black. Larvae of the plant-feeding epilachnines often are yellow with black bristles. Scarabaeid larvae are soft-bodied, thick, strongly C-shaped, somewhat flattened beneath and round above. Cetonine larvae, similar to those of geotrupids and lucanids, are often short, less C-shaped than most scarabaeids, hairy, active, and capable of locomotion on their backs through movement of body segments. Passalid larvae, white and slender, are found with their parents. Chrysomelid larvae are short, flattish or fat, and sometimes have lobes at the sides or appendages at the hind end. Cerambycid larvae are long and slender, with swellings at the sides of the segments; pale, almost hairless, and fairly soft, they either have minute legs or none at all, and the eyes are poorly developed. Weevil larvae, usually white and soft, are fatter in the abdominal region than at the head end. The head capsule may be hardened, brown in colour, and have strong mandibles.

      Some beetles undergo hypermetamorphosis, in which they have different larval types in different instars (the stages between molts). The early larval stages usually are active, and the later stages are parasitic on other organisms. The active, young larvae of most Meloidae (blister beetles (blister beetle)), called triungulins, for example, hatch from eggs laid on flowers, become attached to bees visiting the flowers, and thus are carried to a bee nest, where they become parasitic on bee larvae.

      Pupae of beetles usually have a form similar to that of the adult, except that the elytra are represented by pads on the exterior of the body; the colour, generally white, is sometimes pale brown or patterned. As the time for emergence of the adult approaches, the pupa may darken, especially the mandibles and eyes. After emerging from the pupal skin, the adult rapidly assumes its final adult form and coloration, although metallic colours may take some days to develop their final appearance.

      Larvae that bore in wood, cones, or seeds and those that live in the ground or in excrement chew or dig a cavity, or pupal cell. In some cases, the pupa lies on a cushion of frass (chewed or torn wood fibres) or other material; in others, it is enclosed in a cocoon of frass or other material (e.g., a smooth, white, hard covering similar to the shell of a bird's egg). Sometimes material is used only to seal off the open end of the tube, gallery, or cell for protection from ants and other predators. After the body of the adult has hardened, the adult breaks or dissolves the barrier and emerges. Many wood borer adults must chew through solid wood to emerge, although the larva usually chews close to the surface of the tree, under the bark, before pupating.

Typical life cycle
 The adult Aspidomorpha furcata, a tortoise beetle of South China, feeds on the leaves of the host plant Ipomoea (sweet potato), where the entire life cycle takes place. Eggs, laid in small groups, are cemented together in a thin paperlike egg capsule whose thin brown layers separate and camouflage them. The larva, which hatches in four to six days, burrows directly through the egg capsule and feeds on the leaf epidermis for about three days before it molts. Later it eats through the entire leaf. When the fifth instar larva is fully grown, it goes through a resting, prepupal period, before pupation occurs. During each molt, the old larval skin is pushed back and attached to processes at the hind end. The dried and shrunken skins plus extruded feces combine to camouflage the larva. When pupation takes place, the combination of exuviae (molted skins) and feces becomes attached to a paired process at the hind end of the pupa, thus camouflaging and shielding it. The last larval skin is used to attach the end of the abdomen of the pupa to the leaf surface. The pupa usually rests with its camouflage flat over its back, although it may erect the camouflage and turn it back to discourage an enemy.

      The pupal stage lasts four days or longer. The life cycle from egg to adult requires 21 to 27 days in mild weather, longer in winter; adults may live more than 230 days. Females lay from 63 to 228 egg cases, with an average of about 3 eggs per case. There may be several generations per year.

      In cooler temperate areas, life cycles may occupy much longer periods, even up to four years or more. In general, wood-boring beetles and root feeders have the longest life cycles, while leaf-feeding species have shorter ones. Several generations per year are possible with subtropical and tropical species.

      Ecology, the relationship of organisms to each other and to their environment, represents fundamental interactions in nature. The habits of some of the small families of beetles have not yet been established.

Beetles as prey
      In general, beetles are very well armoured insects and thus are reasonably protected against enemies; most, however, have parasites. Some tachinid flies, for example, lay their eggs on adult beetles, and the larvae feed inside their bodies. Beetle larvae also often have hymenopterous parasites; e.g., wasps. Wood-boring long-horned beetle larvae (Cerambycidae) are parasitized by wasps that lay their eggs directly on or in beetle larvae. Beetles are probably attacked by fewer predators than are many other insects; birds that often feed on various kinds of insects may not eat some kinds of beetles. Swifts and other birds, insectivorous mammals including bats, reptiles, frogs, and other insects may act as beetle predators. Some beetle predators feed particularly on beetle larvae, although many beetle larvae that feed on plants and in the ground probably are distasteful to birds and other predators.

Feeding habits and habitats
      Beetles are found in almost any habitat occupied by insects and feed on a variety of plant and animal materials. Many are predatory, some are scavengers, many are plant feeders (phytophagous), others feed on fungi, and a few are parasitic on other organisms. Beetles may live beneath the ground, in water, or as commensals in the nests of social insects such as ants and termites. Plant-feeding species may eat foliage, bore in wood or fruit, and attack roots or blossoms; any part of a plant may be a food source for some type of beetle. Many beetles eat stored plant or animal products, including various types of foods and clothing.

 Adult carabids are nocturnal hunting predators whose prey ranges from miscellaneous small insects to fairly large caterpillars and land snails. Many small carabids live in moss on tree trunks and feed on small insects hidden in the moss. The predatory larvae remain more hidden than the adults. Both larval and adult dytiscids (diving beetles) feed on various water insects, small frogs, and small fish. Gyrinids (whirligig beetles (whirligig beetle)) feed both on water insects and on other insects that fall into the water. Staphylinids (rove beetles) are usually predatory, both as larvae and as adults. Hydrophilids are water scavenger beetles (water scavenger beetle) and eat various dead organisms and live algae in freshwater. Silphids, which dig under small dead animals so that they settle into the ground, lay their eggs on the carrion, on which the larvae feed. Dermestids feed on dead animal skins, dead insects, birds' nests, other animal nests, and accumulations of debris. Ptinidae (spider beetles (spider beetle)) feed on dead insects and animal skins, as do certain Anobiidae (e.g., the cigarette beetle, which also feeds on tobacco and other dried products). Heterocerids and histerids prey on fly larvae or those of beetles living in excrement or in carrion. Some Elateridae (click beetles) are predatory as larvae, feeding on the larvae of wood-boring insects; others feed on roots of various crops. Most larvae of Cantharidae (soldier beetles), which prey on worms and larvae of other insects, occur under bark, in rotting wood, or in soil; the adults are usually found on flowers. The lampyrids (fireflies), which are often luminous both as larvae and as adults, are primarily predators upon snails. Clerid larvae are largely predators of wood-boring beetle larvae, although some of the adults are flower feeders.

 Among the lamellicorns, the true scarabs (Scarabaeinae (dung beetle)) and several related groups (e.g., Aphodiinae, Geotrupinae) are dung feeders. Some make balls of animal excrement (e.g., of cattle) and roll them to protected spots for burying. Eggs are laid on the buried balls, and the larvae feed on the excrement.

      The Melolonthinae (June beetles, chafers) are phytophagous, the larvae usually feeding on roots of grasses or other plants, the adults feeding on leaves. The larvae of Rutelinae (shining leaf chafers (shining leaf chafer)), which have similar habits, sometimes feed in humus or in rotten wood. The adults of Cetoniinae feed on pollen and tree juices; the larvae, often called white grubs, feed on organic matter in the soil and may damage plant roots. The Dynastinae (rhinoceros, unicorn, and elephant beetles) feed in rotting wood, decaying vegetation, or humus. Buprestidae bore into living and dead trees, generally feeding on the cambium layer. Some of the small buprestids are leaf miners; that is, they feed on leaf cells between the upper and lower surfaces of a leaf. Moderate numbers of larvae and adults of Passalidae (bess beetles (bess beetle)), which feed on rotting logs, generally are found together; apparently the adults protect and feed their young, an unusual habit among beetles. The lucanids feed in logs and stumps as larvae; the adults often feed on juices from damaged trees, particularly at openings of larval tunnels of Cerambycidae (long-horned wood-boring beetles).

      Tenebrionidae (darkling beetles) live in various habitats as scavengers or predators. Those in rotten wood and under bark may be predatory on other larvae; those in damp places may feed on rodent excrement, on other wastes, or on fungi; and those in stored products feed on grain, meal, and other staples. Lagriidae (lagriids) and Alleculidae (comb-clawed beetles) feed in rotten logs. Melandryidae (false darkling beetles) usually feed on fungi or in old wood. Pythids usually are scavengers in burrows of other beetles, including weevils. Rhipiphoridae (wedge-shaped beetles), which usually are parasites in wasps' nests and undergo hypermetamorphosis, are related to another group of insects, the Strepsiptera, which are mostly parasitic in the bodies of wasps, leafhoppers, grasshoppers, and other insects. Some larvae of Mordellidae (tumbling flower beetles (tumbling flower beetle)) may live in dead or dying deciduous wood or attack the heartwood of weak trees; others may be found in pith or herbaceous weeds. The adults frequent flowers and are good fliers. Meloidae (blister beetles) undergo hypermetamorphosis and usually live in bee nests, feeding on eggs and stored food. The larvae of Pyrochroidae (fire-coloured beetles) live under bark and in rotting wood. Most cerambycid larvae are wood borers; a few live in large herbs or in cones, and many feed in roots. The adults, which chew new bark or other plant parts, at times cut rings in branches, killing them so that the larvae can bore in. Some chrysomelid larvae eat leaves; many others are root feeders. Members of one group of chrysomelid larvae (Sagrinae) bore in stems of leguminous vines, causing gall-like swellings; others (Donaciinae) feed on and in the stems of freshwater plants, often below the water level, or hide (Hispinae) in unopened parts of monocot plants such as palms, grasses, bamboos, and gingers; still others (Alticinae) may feed on both roots and leaves or act as leaf miners. Bruchidae live primarily in the seeds of legumes; hence their common name, seed beetles (or pea weevils).

      Weevils also are diverse in habits. Many larvae bore into solid wood of living or dead trees and stumps; some feed on or in roots or in stems of semiwoody plants; and others feed in seeds, pods, grain, meal, fruits, nuts, and other parts of plants; some are hidden inside the plants. Most adults are able to fly; nonflying forms, however, are more abundant on small oceanic islands than are flying forms. Some species, which develop in rotting stumps, soil, or palm trunks, make a rough cocoon from frass. Brenthids develop in dead trunks and stumps; anthribids live in various parts of plants, often in seeds; and scolytids feed largely under the bark of living trees, although some bore in other parts of plants, seeds, cones, and needles.

Special ecological relationships
      Certain beetles have special ecological relationships. Beetles that live in nests of ants and in nests of termites have become modified in form because certain structures that are no longer needed have degenerated (e.g., wings, wing covers); some have almost lost the power of locomotion. Some species have evolved glands that produce secretions attractive to the host ants or termites.

      An association of a different type is known involving beetles in the mountains of New Guinea; called epizoic symbiosis, the association occurs on the backs of large leaf-feeding weevils found on Nothofagus and other trees in the moss forests. Various kinds of algae, fungi, lichens, liverworts, mosses, and diatoms develop on the backs of the weevils. Among them live protozoans, rotifers, nematodes, phytophagous mites (mite), and parasitic mites. The phytophagous mites, known only from this association, feed primarily in the fungal growth on the backs of the beetles. Plant spores, which may be carried from one weevil to another either by the mites during mating of the weevils or by air dispersal, are trapped with a sticky fluid that may be produced by the weevils for this purpose.

      Ambrosia beetles (Scolytidae) associate with fungi in the host tree. Certain adult scolytids and platypodids have specialized structures called mycetangia, which are used to carry the fungi when the beetles seek out new host trees.

Form and function

Adult features
 Many structural modifications are found among the beetles. So varied is the structure that it is difficult to make general statements; for example, a few beetles have no elytra, and some have no wings.

      As in all adult insects, the segmented body consists of three primary body regions: head, thorax, and abdomen. In beetles, however, two of the three thoracic segments (mesothorax and metathorax) are attached to the abdomen; the third (prothorax), isolated as the region between the head and trunk, is covered by a dorsal plate, the pronotum. The body covering (exoskeleton) varies from very horny and rigid to soft and flexible, but it usually consists of hard plates (sclerites) separated by flexible membranes.

      The antennae are usually 11-segmented but vary widely in form. The jaws (mandibles) may be relatively large, in some as long as the rest of the body, or almost completely absent; usually they are triangular in shape and suitable for biting or chewing. The paired maxillary and labial palps are usually small and are used for feeding or handling food, but in some beetles one or the other pair may be greatly enlarged. The compound eyes are usually prominent, but are sometimes reduced or absent, and occasionally divided. Simple eyes (ocelli) are rarely present. A neck is sometimes evident, but in many beetles the head is recessed into the prothorax or under the pronotum.

      The prothorax is generally very distinct; the mesothorax and metathorax are hidden under the elytra along with most of the abdomen. The pronotum may be four-sided very wide or very long, and sculptured with lateral spines or dorsal grooves and pits. The front legs emerge from cavities in the underside that may be confluent or separated by other parts. The mesothoracic spiracle (respiratory opening) is often visible just behind the base of the front legs. The mesothorax bears the elytra (wing covers) and the second pair of legs. The metathorax bears the flying wings (hindwings) and the third pair of legs.

      The legs are modified in various ways, for running, swimming, jumping, digging, or clasping. In some beetles, wings are not capable of producing flight, but in many others, they are powerful and sustain strong flight.

      The abdomen is composed of nine or ten segments, but often some of these are not externally visible. From five to eight segments can usually be seen, with short apical appendages evident in some beetles. Each abdominal segment has a pair of spiracles, the openings into the air-tube (tracheal) system. Predatory beetles generally have short digestive tracts. Differences in salivary glands occur, depending on the food source.

Larval features
      Coleoptera larvae differ in appearance from adults. This is characteristic of insects with complete metamorphosis (Endopterygota), in which the wings develop internally until they become apparent in the pupal stage. The differences in body form of the larvae are closely associated with larval habitats and modes of feeding. The numerous predatory types, for example, have slender or gradually tapered bodies, with large, slender, mandibles, and relatively long legs; thus, these larvae are adapted to be rapid runners that capture and hold prey with their mandibles. A mouthpart structure (epipharynx) may be adapted for imbibing body fluids that exude from wounds caused by the mandibles. Typical predatory larvae are found in the dytiscids (predacious water beetles) and carabids (ground beetles). The larvae of tiger beetles (tiger beetle) (Cicindelidae), although similar to the dytiscid type, live in holes in the ground or in branches and capture passing insects. The enlarged head of the tiger beetle larva fills the burrow opening, and its legs are modified for attachment and leverage. Haliplid, staphylinid, and gyrinid larvae are also similar to those of dytiscids, except that the gyrinids have a gill on the side of each abdominal segment, rather than at the tip of the abdomen, as do most of the others. Some predatory larvae are generally less tapered, sometimes less armoured, and have shorter appendages than do the dytiscids; e.g., larvae predatory on other larvae in deadwood or in the ground. Larvae such as those of the Histeridae (hister beetles (hister beetle)), which usually live in special environments such as dung and tunnels in wood, have short appendages but slender mandibles.

      Another larval type includes many of the scavengers; e.g., silphids, hydrophilids. These larvae have short legs and mandibles.

      Wood-feeding borer larvae (cerambycid and buprestid type) have soft white bodies that may be cylindrical or flattened. The thoracic region of buprestids is flat and broad, the head is dark and retracted in the thorax, and the mandibles are short and stout. Larvae of wood-boring beetles usually have yeast associated with the digestive tract that helps to convert wood to digestible compounds.

      Larvae that feed on leaves, stems, and roots (chrysomelid type) are short and oval shaped. Coccinellid larvae are similar in form except for longer legs, and prey on soft-bodied insects like aphids.

      The lamellicorn larval type is C-shaped, has a soft body, and a hard, dark, nonretractile head. These larvae usually are found in protected habitats, where they feed on roots, rotten wood, or excrement. Weevil larvae have a similar form, although the head may be a little smaller, the body less arched. Another larval type is that of the elaterids and many tenebrionids, which have very slender usually brown bodies, with a hard outer skeleton.

 Coleoptera protect themselves against enemies in various ways. Some closely resemble their surroundings; the upper surface of one African species (Petrognatha gigas), for example, resembles dead velvety moss, and its irregular antennae are very much like dried tendrils or twigs. Many weevils fall and feign death at the least alarm and, folding their limbs closely around the body, look like seeds or particles of soil, thus escaping observation.

      Certain beetles, especially those living in ants' nests, resemble ants, and the common wasp beetle of Europe, Clytus arietis, both in its movements and coloration, closely resembles a wasp.

      Some beetles obtain some measure of protection possibly from their repellent appearance or from their evil-smelling or distasteful secretions, either in the form of exudations of blood from definite parts of the body or as the product of special fetid glands. The so-called bombardier beetles of the Carabidae have the property of secreting an evil-smelling defensive fluid from the anal end of the body. In some cases, this fluid volatilizes explosively into a gas at high temperature when it comes into contact with the air; it acts as a repellent to other insects or enemies. A number of beetles secure protection by virtue of their agility; many ground beetles and tiger beetles run rapidly, and the latter also readily take flight. The flea beetles (flea beetle) (a group of the Chrysomelidae) have remarkable powers of leaping.

      Many beetles produce sound, usually by rubbing one part of the body (a scraper) against another part (the file). These stridulating organs are generally present in both sexes and probably serve for mutual sex calling. Some beetles have a filelike area on the head that is rasped by the front margin of the prothorax. Among the cerambycids, sound is produced either by rubbing the rear margin of the prothorax over a grooved area on the mesothorax or by rubbing the femurs of the hind legs against the margins of the elytra.

      Stridulation, however, is not confined to adult beetles; it occurs also in certain larvae. Some larvae of the Scarabaeoidea, for example, have a series of ridges, or tubercles, on the coxal segment of the middle pair of legs, and the hind legs are modified in various ways as rasping organs. In the larvae of some chafers (Melolonthinae), a ridged area on the mandible is rasped by a series of teeth on the maxillae. Stridulation in larvae is independent of sex and may be used to warn neighbouring larvae to avoid getting in each other's way.

Light production
      Some beetles emit a bright light, whose source is in special luminous organs that consist of an outer, light-producing layer and an inner reflector layer. The outer layer is supplied with oxygen by means of air tubes (tracheae), and the reflector layer contains crystals that apparently act as a background, scattering the light and preventing its dispersion internally. The light is produced during a reaction involving a compound (luciferin) and an enzyme-like substance (luciferase) in the outer layer of the luminous organ. Luminous beetles include the Lampyridae (fireflies), Phengodidae, Drilidae, and certain Elateridae (click beetles). A familiar example of a firefly is the common European glowworm (Lampyris noctiluca), whose wingless female emits a bright light near the hind end of the body; the winged male emits a much feebler light. Both sexes of the elaterid genera Pyrophorus and Photophorus are winged and luminous.

Evolution and paleontology
      Coleopterans are very ancient insects; they date from the Permian Period (about 225,000,000–280,000,000 years ago), after the appearance of gymnosperm plants. Although the beetles have a number of similarities to another ancient group of insects, the cockroaches (Blattaria), they probably evolved from ancestors of the present-day Neuroptera. This theory is based largely on the nature of the life cycle of beetles and on their larval structure. Although many beetle fossils are known, they consist mostly of isolated elytra, which reveal little about the history of the order. Complete fossil specimens are closely related to living forms. The evolution of elytra may have been associated with the habit of living under bark of trees, where protection for flying wings is required. Most of the insects that live under bark are beetles.


Distinguishing taxonomic features
      One distinctive feature of coleopterans is wing structure. Most beetles have two pairs of wings. The front pair, which may be thickened, leathery, or hard and brittle, are called elytra and usually serve only as protective covers. A few beetles have greatly reduced wings. Variations in the structure of the first abdominal segment is one criterion used to separate the various suborders of Coleoptera; the hind coxal leg segments (by which the legs are attached to the body) may divide the abdominal segment partially or completely. Sometimes the abdominal segments are fused, the articulations marked by form sutures.

      Variation in length, texture, and appearance of elytra, as well as the number of abdominal segments exposed by short elytra, are used to distinguish the various superfamilies. Characters associated with the size and shape of the coxae also are used as distinguishing features. Structure of antennae and legs are important considerations for taxonomic criteria, as are larval structure, head structure (including mandibles, or jaws), pattern of veins in wings, habitats, and behaviour.

Annotated classification
      About 135 families of beetles are known, of which 120 occur in the Western Hemisphere.

      There have been a number of different classifications of Coleoptera. Many were based on the suborders Adephaga and Polyphaga; the latter, which contains about 90 percent of the beetles, included a number of divisions (e.g., clavicorns, serricorns, lamellicorns, phytophagous beetles, and weevils). Sometimes these divisions are considered as superfamilies or series, and sometimes (particularly weevils and relatives) as suborders. The classification below is based on that of R.A. Crowson (1955); it includes four suborders.

Order Coleoptera (beetles, weevils)
 Largest insect order; more than 350,000 species; size range from less than 1 mm to more than 12 cm (5 in.); modified front wings, called elytra, usually meet in a straight line down the middle of the back, covering membranous hind wings; hind wings usually longer than front wings, folded under front wings when at rest; mouthparts adapted for chewing; form of antennae variable; large compound eyes; hard outer skeleton; complete metamorphosis; found in almost all types of habitats; many plant feeders; many species of economic importance, either cause damage or benefit man; worldwide distribution.
      Suborder Archostemata
 Hind coxae rarely fused to metasternum; distinct notopleural suture between notum and pleural sclerites.

      Family Cupesidae (Cupedidae; reticulated beetles)
 Small and little-known; found under bark; about 30 species widely distributed.

      Family Micromalthidae
 Rare; 1 to 2 species; most complex life cycle among coleopterans.

      Suborder Adephaga
 Larval structure primitive; legs specialized for predatory life; hind coxae of legs immovably fixed to metasternum; distinct notopleural suture between notum and pleural sclerites; wing with base of Rs (radial sector) vein distinct.

      Family Cicindelidae (tiger beetles)
 Voracious and fierce, especially larvae; Cicindela; active; often brightly coloured; about 2,500 species; mostly tropical and subtropical.

      Family Rhysodidae (wrinkled bark beetles)
 Small, slender, brownish beetles; about 350 species, mostly tropical.

      Family Paussidae
 Sometimes a subfamily of Carabidae; about 400 species; tropical or subtropical; associated with ants.

      Family Carabidae (ground beetles)
 Usually dark, shiny, flattened; larvae and adults predatory; Calosoma feed on caterpillars; Brachinus, bombardier beetles, eject fluid from anus; about 30,000 species; worldwide distribution.

      Family Trachypachidae
 A few species in Europe and North America.

      Family Haliplidae (crawling water beetles)
 About 200 small aquatic species; wide geographical range.

      Family Amphizoidae (trout-stream beetles)
 About 5 species (Amphizoa) in Tibet, North America; feed on drowned insects.

      Family Hygrobiidae
 A few species (Hygrobia) widely distributed; aquatic; produce sound.

      Family Gyrinidae (whirligig beetles)
 About 700 species; surface swimmers; sometimes gregarious.

      Family Noteridae (burrowing water beetles)
 Similar to Dytiscidae; small; larvae burrow.

      Family Dytiscidae (true water beetles)
 Worldwide distribution; about 4,000 species; found in flowing and still water; Siettitia eyeless; Dytiscus best known.

      Suborder Myxophaga
 Wing with base of Rs vein absent; prothorax usually with distinct notopleural suture.

      Family Calyptomeridae
 Sometimes placed in Clambidae.

      Family Lepiceridae
 A few Central American species.

      Family Sphaeriidae (minute bog beetles)
 Less than 1 mm in length; 1 genus; a few widespread species; sometimes placed in Staphylinoidea.

      Family Hydroscaphidae (skiff beetles)
 Size about 1.5 mm; found in algae on rocks in streams; sometimes placed in Staphylinoidea; generic example: Hydroscapha; widely distributed.

      Suborder Polyphaga
 Includes the majority of beetles; wing with base of Rs vein absent; prothorax never with distinct notopleural suture.

      Superfamily Hydrophiloidea
 Head usually with Y-shaped line on front; antennae short, hairy and club-shaped at end; habits mostly aquatic; maxillary palp usually longer than antennae.

      Family Hydrophilidae (water scavenger beetles)
 About 2,000 species, numerous in tropics; aquatic and terrestrial; most live on decomposing vegetable matter; Hydrophilus is predatory; other examples: Cercyon, Helophorus.

      Superfamily Histeroidea
 Antennae geniculate (elbow-shaped) with last 3 segments club-shaped; wing with medio-cubital loop reduced; elytron truncate leaving 1 or 2 segments of abdomen exposed.

      Family Sphaeritidae
 One genus, about 4 species.

      Family Synteliidae
 One genus, a few species in Mexico and the Orient.

      Family Histeridae (hister beetles)
 Small, dark, shiny; found in decaying organic matter; predatory on small insects; about 3,900 species; wide distribution; examples: Hister, Niponius.

      Superfamily Staphylinoidea
 Very large group; antennae with last 3 segments rarely club-shaped; outer skeleton rarely very hard, shiny; wing veins M (media) and Cu (cubitus) not connected; elytron truncate, usually more than 2 abdominal segments exposed.

      Family Limulodidae (horseshoe crab beetles)
 One mm or less; in United States and Australia; ride on ants; 5 genera, about 37 species.

      Family Ptiliidae (feather-winged beetles)
 Among the smallest beetles; live in dung, rotting wood, fungi; about 400 species; temperate and neotropical regions.

      Family Leptinidae (mammal-nest beetles)
 Feed on eggs and young of small arthropods in small-mammal nests; wingless; widely distributed.

      Family Anisotomidae (Leiodidae; round fungus beetles)
 Small, shiny; habitats vary—caves, fungi, mammal nests; about 1,000 species.

      Family Dasyceridae
 About 10 species; Holarctic.

      Family Silphidae (carrion beetles, burying beetles)
 Relatively large, bright-coloured; usually feed on carrion; some predatory, some plant feeders; examples: Silpha, Nicrophorus; about 230 species; widely distributed.

      Family Scydmaenidae (antlike stone beetles)
 Under stones, logs; in ant nests; very small, hairy; widely distributed; about 1,200 species; example: Scydmaenus.

      Family Scaphidiidae (shining fungus beetles)
 In fungi, dead leaves, rotting wood; small, very shiny; about 250 species; example; Scaphidium.

      Family Staphylinidae (rove beetles)
 Short elytra; size variable; active; strong mandibles; in ant nests, predatory on other insects in decaying matter; over 20,000 species; widely distributed; examples: Stenus, Dinarda.

      Family Pselaphidae (short-winged mold beetles)
 Very small; diverse in form; live in ant nests; about 5,000 species; worldwide distribution but most abundant in tropics; example: Claviger.

      Superfamily Scarabaeoidea (Lamellicornia)
 Antennae 10-segmented with last 3 to 7 segments forming a lamellate (platelike) club; body stout; larvae without cerci (appendages at end of abdomen); males and females often differ in appearance; outgrowths on head and thorax produce bizarre forms; produce sound (stridulate).

      Family Lucanidae (stag beetles, pinching bugs)
 Large; variable in size; males with enormous mandibles (jaws); about 900 species; widely distributed; example: Lucanus.

      Family Passalidae (bess beetles)
 Large, dark, flattened; about 500 species, mostly in moist, warm forests; live in decaying wood.

      Family Trogidae (skin beetles)
 About 300 widely distributed species; example: Trox; dung or carrion feeders.

      Family Ceratocanthidae (Acanthoceridae)
 About 200 tropical species; associated with rotten wood.

      Family Geotrupidae (“dor” beetles)
 Large; over 600 species; widely distributed; habits variable; examples: Lethrus, Geotrupes.

      Family Scarabaeidae (scarab beetles, June beetles, tumblebugs, leaf-chafers)
 Variable in colour, size, habits; most feed on dung, carrion, other decaying matter; about 20,000 species; widely distributed; examples: Cetonia, Melolontha.

      Superfamily Dascilloidea
 Forecoxae projecting; abdomen with 5 visible segments; wing with radial cell short; anal cell of wing, if present, with 1 apical vein.

      Family Clambidae (fringed-wing beetles)
 Small, hairy; in decaying plant material; about 30 species; worldwide distribution; sometimes placed in Staphylinoidea.

      Family Eucinetidae
 About 25 widely distributed species; in rotten wood; example: Eucinetus.

      Family Scirtidae (Helodidae) (marsh beetles)
 Small, oval; on vegetation in swampy places; aquatic larvae; about 600 species; widely distributed; example: Scirtes.

      Family Dascillidae
 About 200 moderate-sized species; found on vegetation in moist places.

      Superfamily Byrrhoidea
 Forecoxae large; antennae more or less thickened at tip; body short, with legs and antennae retractable into grooves on under surface.

      Family Byrrhidae (pill beetles)
 Small, oval; found under debris, in sand, at grass roots; about 350 species; widely distributed; example: Byrrhus.

      Superfamily Dryopoidea
 Forecoxae sometimes slightly projecting; antennae usually filiform (threadlike) or short and broad; head with distinct labrum (upper lip); last tarsal segment of leg often as long as preceding 3 or 4 together.

      Family Psephenidae (water-penny beetles)
 Larvae flat, almost circular; a few species, mostly in India, North America.

      Family Eurypogonidae
 A few species in Northern Asia, North America.

      Family Ptilodactylidae
 About 200 tropical species; aquatic or in rotten wood.

      Family Chelonariidae
 About 50 species in tropics of Asia and America.

      Family Heteroceridae (variegated mud-loving beetles)
 About 500 widely distributed species; example: Heterocerus.

      Family Limnichidae (minute marsh-loving beetles)
 Similar to Dryopidae; a few widely distributed species.

      Family Dryopidae (long-toed water beetles)
 Small, downy; crawl on stream bottoms; few species; widely distributed.

      Family Elmidae (riffle beetles)
 Varied habitat; several hundred widely distributed species.

      Superfamily Rhipiceroidea
 Antennae flabellate (fanlike); noselike projection between mandibles; about 180 species; widely distributed; 2 families, Rhipiceridae (cedar beetles), Callirhipidae; example: Sandalus.

      Superfamily Buprestoidea
 Antenna short, serrate; abdomen weakly hardened. Family Buprestidae (metallic wood-boring beetles). Brightly coloured, metallic sheen; inhabit various hot, moist forests; about 15,000 species, mostly tropical; examples: Agrilus, Sphenoptera, Chrysobothris.

      Superfamily Elateroidea
 Forecoxae small; metasternum without transverse suture; larvae with no free labrum.

      Family Cerophytidae
 About 12 species in Europe and America; in hollow trees.

      Family Cebrionidae
 About 200 species; in mild regions; female often wingless.

      Family Elateridae (click beetles)
 About 7,000 species; widely distributed; can leap when lying on back; adults, plant feeders; larvae sometimes damage plants; examples: Pyrophorus, Agriotes, Athous

      Family Trixagidae (Throscidae; throscid beetles)
 Small, oblong; about 200 species; widely distributed.

      Family Perothopidae
 Medium-sized; on trunks or branches of old beech trees.

      Family Eucnemidae (false click beetles)
 Closely related to Elateridae; about 1,000 species, mostly in warm climates; example: Melasis.

      Superfamily Cantharoidea
 Larvae with grooved or channelled mandibles; front coxae large, projecting; 6 or 7 visible abdominal segments.

      Family Brachypsectridae
 A few species in Asia and California.

      Family Homalisidae
 A few species in Mediterranean region.

      Family Karumiidae
 A few species in Iran, South America.

      Family Drilidae
 About 80 species, mainly in Europe; larvae prey on snails.

      Family Phengodidae
 About 50 species in America; produce light.

      Family Lampyridae (lightning bugs, fireflies)
 Produce light in species-characteristic flashing rhythm; wingless females and most larvae called glowworms; about 2,000 species; widely distributed; examples: Lampyris, Photinus.

      Family Cantharidae (soldier beetles)
 Soft-bodied, predatory; about 3,500 species; widely distributed; examples: Cantharis, Rhagonycha.

      Family Lycidae (net-winged beetles)
 About 2,800 species, mostly tropical; often bright-coloured; distasteful to birds; example: Dulitocola.

      Superfamily Dermestoidea
 Last 3 antennal segments forming a club; tarsi of legs all 5 segmented, simple.

      Family Derodontidae
 About 12 widely distributed species.

      Family Nosodendridae
 About 30 widely distributed species; found under bark.

      Family Dermestidae (skin beetles, dermestid beetles)
 Many economically important species; mostly scavengers on plant and animal products; small to moderate-sized; hairy or with scales; examples: Dermestes, Anthrenus; widely distributed.

      Family Thorictidae
 About 80 species, commonly in Mediterranean region; associate with ants.

      Superfamily Bostrychoidea
 Larvae soft-bodied, lack specialized setae (hairs), maintain a C-shaped position; adult hard, head region hoodlike; members often associated with timber, destructive.

      Family Anobiidae (drugstore and death-watch beetles)
 Live in dry vegetable materials; some species destructive pests; examples: Xestobium, Stegobium, Lasioderma; about 1,100 widely distributed species.

      Family Ptinidae (spider beetles)
 Long legs; spiderlike appearance; sometimes infest stored products; about 500 widely distributed species.

      Family Bostrychidae (branch and twig borers)
 Attack living and dead wood; about 700 species; worldwide distribution; examples: Sinoxylon, Dinoderus.

      Family Lyctidae (powderpost beetles)
 Related to Bostrychidae; damage timber and furniture; about 90 widely distributed species.

      Family Psoidae (psoid branch and twig beetles)
 Medium-sized; sometimes cause extensive orchard damage; examples: Psoa, Polycaon.

      Superfamily Cleroidea
 Tarsi of legs always 5-segmented; forecoxae projecting or transverse; abdomen with 5 or 6 visible segments.

      Family Trogossitidae (bark-gnawing beetles)
 About 500 species, mostly tropical; vary in shape and habits; sometimes in stored products; example: Tenebroides.

      Family Chaetosomatidae
 Three genera in New Zealand.

      Family Cleridae (checkered beetles)
 Small; many brightly coloured; downy; most adults and larvae predatory on other insects; some adults pollen feeders; about 3,000 species, mainly tropical; examples: Corynetes, Necrobia.

      Family Melyridae (soft-winged flower beetles)
 About 4,000 species widely distributed; diverse; example: Malachius.

      Family Phloiophilidae
 Rare; 1 species in Britain.

      Superfamily Lymexyloidea
 Antennae short, more or less serrate; abdomen with 6 or 7 visible segments.

      Family Lymexylidae (ship-timber beetles)
 About 60 species; worldwide distribution; damage wood; examples: Lymexylon, Hylecoetus.

      Superfamily Cucujoidea
 Usually 5 visible abdominal segments; antennae filiform or clubbed, rarely serrate.

      Section Clavicornia
 All visible abdominal segments usually movable; sometimes less than 7 pairs of abdominal spiracles (respiratory holes)

      Family Passandridae
 Few species; mostly in warm climates.

      Family Smicripidae
 Sometimes placed in Nitidulidae; a few species in tropical America; example: Smicrips.

      Family Nitidulidae (sap beetles)
 Variable size, shape, habits; usually found around fermenting plant fluids or moldy plant materials; about 2,200 species; examples: Meligethes, Cybocephalus.

      Family Byturidae (fruitworm beetles)
 Small, hairy; few genera; damage raspberry blossoms and fruit; example: Byturus.

      Family Biphyllidae (false skin beetle)
 About 200 species; mostly tropical; example: Biphyllus.

      Family Rhizophagidae (root-eating beetles)
 Usually occur under bark; few species; sometimes placed in Nitidulidae.

      Family Cisidae (minute tree-fungus beetles)
 Occur under bark, in wood, or in dry woody fungi; about 360 species; widely distributed.

      Family Protocucujidae
 Two species; Chile and Australia; similar to Sphindidae.

      Family Sphindidae (dry-fungus beetles)
 Small, dark; occur in dry fungi; about 30 species; widely distributed.

      Family Cucujidae (flat bark beetles)
 Flat, medium-sized; predatory on mites and other insects; about 500 species; some species hypermetamorphic; example: Catogenus.

      Family Silvanidae (flat grain beetles)
 Closely related to Cucujidae; some feed on grain (Oryzaephilus); another genus, Silvanus.

      Family Hypocopridae
 Little known; few species.

      Family Helotidae
 About 80 species in warm parts of Asia.

      Family Phycosecidae
 Few species; examples: Phycosecis, Alfieriella; in Australia, Asia, Africa.

      Family Propalticidae
 About 20 species in warm regions of Old World.

      Family Cryptophagidae (silken fungus beetles)
 Mostly fungus feeders; sometimes in nests of bees and wasps; about 800 species; examples: Cryptophagus, Antherophagus.

      Family Phalacridae (shining flower beetles)
 Larvae develop in certain flower heads (e.g., goldenrod), about 500 species; widely distributed; example: Olibrus.

      Family Languriidae
 Feed on plant leaves and stems; example: Languria; about 400 species mostly in Asia and North America.

      Family Erotylidae (pleasing fungus beetles)
 Shiny; found with fungi; about 1,400 species; many in South America.

      Family Cerylonidae
 Often placed in Colydiidae; few species.

      Family Coccinellidae (ladybird beetles, ladybugs)
 Many predatory on aphids and coccids, a few serious plant pests (Epilachna); mostly beneficial; about 5,000 species, usually bright-coloured, spotted; widely distributed; another genus, Rodolia.

      Family Corylophidae
 About 300 species; widely distributed; minute in size.

      Family Endomychidae (handsome fungus beetles)
 Shiny, usually brightly coloured; feed on fungi (mold); about 600 species; mostly in tropical forests; examples: Endomychus, Mycetaea.

      Family Discolomidae
 About 30 tropical species; many wingless.

      Family Merophysiidae
 Few species, sometimes placed in Lathridiidae.

      Family Lathridiidae (minute brown scavenger beetles)
 Found in fungi, debris, flowers; about 600 species.

      Section Heteromera
 Abdomen with 7 pairs of spiracles; forecoxae usually projecting.

      Family Merycidae
 A few Australasian species; example: Meryx; affinities uncertain.

      Family Mycetophagidae (hairy fungus beetles)
 Mostly associated with fungi; often brightly marked; about 200 species.

      Family Colydiidae (cylindrical bark beetles)
 More than 1,400 species; especially in tropics, but many in New Zealand; diverse; example: Bothrideres.

      Family Pterogeniidae
 Two Indo-Malayan genera of uncertain affinities.

      Family Nilionidae
 About 40 species, mostly in tropics; associate with fungi on trees.

      Family Lagriidae
 Medium-sized with metallic sheen; found in foliage, on bark.

      Family Tenebrionidae (darkling beetles)
 Varied group; mostly plant scavengers; examples: Eleodes, Tenebrio; about 10,000 species; widely distributed.

      Family Alleculidae (Cistelidae; comb-clawed beetles)
 Found on flowers, foliage, fungi, under dead bark; 100 species; worldwide distribution; example: Omophlus.

      Family Monommidae
 Adults occur in leaf litter; larvae in rotten wood; about 100 species in warm regions; example: Monomma.

      Family Zopheridae
 Few species, mostly in America.

      Family Elacatidae (false tiger beetles)
 Widely distributed; umerous species; examples: Othnius, Eurystethus.

      Family Boridae
 Widely distributed small group; sometimes placed in Tenebrionidae.

      Family Inopeplidae
 One genus, Inopeplus; related to Salpingidae.

      Family Salpingidae (narrow-waisted bark beetles)
 Superficial resemblance to Carabidae (ground beetles); adults and larvae predatory; adults occur under rocks, or bark, in leaf litter, on vegetation; few species but widely distributed; examples: Salpingus, Lissodema.

      Family Cononotidae
 Similar to Salpingidae; several genera.

      Family Mycteridae
 Resemble Salpingidae.

      Family Hemipeplidae
 Few species in warm regions; live beneath palm leaves.

      Family Trictenotomidae
 About 12 species in forests of Oriental region.

      Family Pythidae
 Few species widely distributed in Eurasia and America; example: Pytho.

      Family Pyrochroidae (fire-coloured beetles)
 Adults large, found on foliage or flowers, under bark; about 100 species in north temperate region; example: Pyrochroa

      Family Tetratomidae
 Similar to Melandryidae.

      Family Melandryidae (false darkling beetles)
 Usually found under bark or logs; examples: Penthe, Osphya; about 400 species in woodlands of temperate regions.

      Family Scraptiidae
 About 200 species widely distributed; associated with rotten wood, fungi; example: Scraptia.

      Family Mordellidae (tumbling flower beetles)
 Wedgeshaped, hump-backed; common on flowers; active; about 1,500 species.

      Family Rhipiphoridae (wedge-shaped beetles)
 About 400 species, many with specialized parasitic habits on other insects; complicated life cycle; examples: Pelecotoma, Metoecus.

      Family Cephaloidae (false longhorn beetles)
 About 11 species in east Asia, North America.

      Family Meloidae (blister beetles, oil beetles)
 Body fluids contain cantharadin, sometimes used as a drug (Lytta); several important plant pests (Epicauta); many larvae beneficial, feed on grasshopper eggs; hypermetamorphic; complicated life cycle; about 2,500 species; widely distributed.

      Family Oedemeridae (false tiger beetles)
 Adults usually on flowers or foliage; larvae in moist decaying wood; about 600 species; widely distributed, but especially abundant in temperate regions; example: Nacerdes.

      Family Aderidae (antlike leaf beetles)
 About 350 species; usually found in deadwood or vegetable refuse; example: Aderus.

      Family Anthicidae (antlike flower beetles)
 Many occur in vegetable refuse; about 1,000 species; sometimes placed in Pedilidae; examples: Anthicus, Notoxus.

      Superfamily Chrysomeloidea
 Mostly wood or plant feeders; body shape very variable; antennae not clubbed.

      Family Cerambycidae (long-horned, wood-boring beetles)
 Some large in size; plant feeders; many brightly coloured; larval stage usually wood-boring, sometimes cause tree damage; about 25,000 species; worldwide distribution; examples: Macrotoma, Titanus, Clytus, Monochamus.

      Family Chrysomelidae (leaf beetles)
 Closely related to Cerambycidae; larvae usually plant feeders; many serious pest species; overwinter as adults; more than 26,000 species; widely distributed.

      Family Bruchidae (seed beetles, pea weevils)
 Most larvae live in leguminous seeds; examples: Acanthoscelides, Bruchus; damage stored seeds; about 900 species; widely distributed.

      Superfamily Curculionoidea (snout beetles)
 One of the largest and most highly evolved groups of coleopterans; head prolonged into beak or snout; mouthparts small; antennae usually clubbed and geniculate; larvae C-shaped; mostly plant feeders; of economic importance as pests.

      Family Anthribidae (fungus weevils)
 Found in deadwood and fungi; about 3,000 species, mostly in tropics; Brachytarsus predatory on scale insects.

      Family Nemonychidae (pine-flower snout beetles)
 Small group sometimes placed in Curculionidae or Attelabidae.

      Family Belidae
 Small group found in Australia, New Zealand, South America attached to a variety of plants.

      Family Oxycorynidae
 Small group; found in South America (Oxycorynus), East Indies (Metrioxena), tropical America (Allocorynus).

      Family Aglycyderidae (Proterrhinidae)
 About 200 species; Proterrhinus found in Pacific region; one Aglycyderes species in Canary Islands.

      Family Attelabidae (leaf-rolling weevils)
 Form leaf rolls on various trees; moderate number of species; widely distributed.

      Family Brenthidae
 About 2,000 species, mostly in wooded tropical countries; variable size range; males unlike females in structure.

      Family Apionidae
 Moderate number of species; Apion, Cylas; several species injure leguminous crops.

      Family Curculionidae (weevils)
 About 40,000 species, many with scales; many injurious species; worldwide distribution; Anthonomus (cotton boll weevil, apple blossom weevil); Calandra (granary weevil, rice weevil); Sitona species pests of leguminous crops.

      Family Scolytidae (bark beetles)
 Numerous species; damage trees; Scolytus (elm bark beetle); other genera: Ips; Dendroctonus; Hylastinus.

      Family Platypodidae (pin-hole borers or flat-footed ambrosia beetles)
 Seldom attack healthy trees; example: Platypus; fungus food called ambrosia.

Critical appraisal
      There are many different opinions among coleopterists concerning the relationships of the various groups of beetles; the groups that should be given family status; and the placement of families in superfamilies and suborders. Little information is available about many coleopteran groups, so that their taxonomic affinities are uncertain. Although Crowson includes Strepsiptera (twisted-wing parasites) as a superfamily of Coleoptera, opinion is divided concerning their status, and they have not been included here. Some classifications of Coleoptera recognize three suborders, Archostemmata, Adephaga, and Polyphaga, rather than the four used by Crowson.

Judson Linsley Gressitt

Additional Reading

In addition to information given in textbooks of entomology, the following works should be consulted: J.R. Dibb, Field Book of Beetles (1948); G. Taylor, Some British Beetles (1948); United States Department of Agriculture, “Insects,” 1952 Yearbook of Agriculture (1952), many chapters of this work are devoted to pestiferous beetles—their ravages and control; A.G. Bøving and F.C. Craighead, “An Illustrated Synopsis of the Principal Larval Forms of the Order Coleoptera,” Ent. Amer., vol. 11 (1930–31); J.H. Fabre, Souvenirs entomologiques, 10 vol. (1879–1907). Many of these detailed popular accounts of natural history are available in English. A standard anatomical and biological work, based upon a study of the water beetle Dytiscus, is E. Korschelt, Der Gelbrand Dytiscus marginalis, 2 vol. (1923–24). See also Ross H. Arnett, N.M. Downie and H.E. Jacques, How to Know the Beetles, 2nd ed. (1980); Richard E. White, A Field Guide to the Beetles of North America (1983).

R.A. Crowson, The Natural Classification of the Families of Coleoptera (1955, reprinted 1967), a descriptive synopsis of all families of beetles, with illustrated key, and many new arrangements; W.W. Fowler, Coleoptera: General Introduction and Cicindelidae and Paussidae (1912), a general treatment of morphology, including the internal anatomy of beetles, with a general discussion of organs, biology, and classification; J.T. Lacordaire, Histoire naturelle des insectes: genera des coléoptères, ou exposé méthodique et critique de tous les genres proposés jusqui'ici dans cet ordre d'insectes, 12 vol. (1854–76, reprinted 1954–72), the standard work on all the families and genera of beetles known at the time, still extremely useful, with keys to all categories (in French).Catalogs of the species of Coleoptera include the Catalogus Coleopterorum Europae, 2nd ed. (1906) by L. Heyden, E. Reitter, and J. Weise; C.W. Leng, Catalogue of the Coleoptera of America, North of Mexico (1920), with five supplements (1927–48); W. Junk, Coleopterorum Catalogus, 31 vol. (1910–40); R.E. Blackwelder (comp.), Checklist of the Coleopterous Insects of Mexico, Central America, the West Indies, and South America, 6 pt. (1944–57, reprinted 1982).Judson Linsley Gressitt

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Universalium. 2010.

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  • coleopteran — [kō΄lē äp′tər ən, käl΄ēäp′tər ən] n. [< ModL Coleoptera (< Gr koleopteros, sheath winged < koleos, sheath + pteron, wing: see FEATHER) + AN] BEETLE1 (sense 1) coleopterous adj …   English World dictionary

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