Definitions

beetle

beetle

[beet-l]
beetle, common name for insects of the order Coleoptera, which, with more than 300,000 described species, is the largest of the insect orders. Beetles have chewing mouthparts and well-developed antennae. They are characterized by a front pair of hard, opaque, waterproof wings called elytra, which usually meet in a straight line down the middle of the back. The elytra cover the rear pair of membranous flight wings, protecting them and the body from mechanical damage and desiccation. Beetles are poor flyers compared with many other insects, but they are well adapted for surviving rigorous conditions. They are found everywhere except in oceans and near the poles, and they occupy nearly every kind of habitat. Most are terrestrial, but some are underground tunnelers and some live in water. These water beetles are often confused with water bugs, but the latter all have sucking mouthparts. Beetles range in size from under 1/32 in. (1 mm) to over 6 in. (15 cm) long; tropical species are the largest. Most are dull, but members of several beetle families are brilliantly colored, some with a metallic or iridescent sheen. The majority of beetles are plant eaters, but there are also many predators and scavengers and a few parasites. Many beetles are highly destructive pests of crops and gardens (e.g., Japanese beetle, potato beetle, boll weevil), but others are beneficial predators of harmful insects (e.g., ladybird beetles). The largest of the many beetle families is the scarab beetle family, with over 20,000 species; among these are the dung beetles, which are invaluable scavengers. Weevils are plant-eating beetles with mouthparts elongated into snouts bearing jaws at their ends. The fireflies are luminescent beetles. Blister beetles, including the so-called Spanish fly, produce irritating secretions. Beetles are classified in the phylum Arthropoda, class Insecta, order Coleoptera.

Tiger beetle (Cicindela).

Any of some 2,000 species (family Cicindelidae) of voracious beetles, found worldwide but mostly in the tropics and subtropics. The larva waits at the top of its burrow (up to 2 ft, or 0.7 m, deep) and grasps approaching insect prey with sicklelike jaws. Hooks on the abdomen anchor it so that the struggling victim cannot pull away, and the prey is dragged into the burrow and eaten. The slender, long-legged adults, less than an inch (25 mm) long, have long jaws that can inflict a painful bite. Many are iridescent blue, green, orange, or scarlet.

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or snout beetle

Any of about 40,000 beetle species in the largest family of beetles, Curculionidae, which is also the largest family in the animal kingdom. Most weevils have long, elbowed antennae that may fold into special grooves on the prominent snout. Many species are wingless. Most species are less than 0.25 in. (6 mm) long, are plainly coloured and marked, and feed exclusively on plants. Some species are more than 3 in. (80 mm) long. The larvae may feed on only a certain part of a plant or a single plant species; adults are less specialized. The family includes many destructive pests, including the boll weevil.

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Boll weevil (Anthonomus grandis)

Small beetle (Anthonomus grandis) found almost everywhere cotton is cultivated. It is the most serious cotton pest in North America. Adults vary in size according to how much food they received as larvae, but they average about 0.25 in. (6 mm) long, including the long, curved snout. In the spring adults deposit eggs in cotton buds or fruit. After hatching, the larvae live within the cotton boll, destroying the seeds and surrounding fibres. Because the larvae and pupae remain inside the cotton bolls, they cannot be killed with insecticides. The boll weevil destroys an estimated three to five million bales of cotton annually.

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Any of about 30,000 beetle species (family Scarabaeidae), found worldwide, that are compact, heavy-bodied, and oval. Each antenna terminates in three flattened plates that fit together to form a club. The outer edges of the front legs may be toothed or scalloped. Species range from about 0.2 to 4.8 in. (5 to 120 mm) long and include one of the heaviest known insects. One species of dung beetle, Scarabaeus sacer, was sacred to the ancient Egyptians. Many species are agricultural pests (e.g., chafer, Japanese beetle, June beetle); many are popular with insect collectors because they are large and have beautifully coloured, hard, highly polished forewings.

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Destructive species (Lema trilineata) of leaf beetle (family Chrysomelidae). Less than 0.25 in. (6 mm) long, it is yellow and has three black stripes on its wing covers. Eggs are laid on the underside of a potato leaf, on which both larvae and adults feed. The larvae are camouflaged by excrement the beetles pile on their back. Two generations are produced each year; the second overwinters in the ground in the pupal stage. Seealso Colorado potato beetle.

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or ladybird beetle

Any of the approximately 5,000 widely distributed beetles of the family Coccinellidae. The name originated in the Middle Ages, when the beetle was dedicated to the Virgin Mary and called “beetle of Our Lady.” Ladybugs are hemispheric and are usually 0.3–0.4 in. (8–10 mm) long. They have short legs and are usually brightly coloured with black, yellow, or reddish markings. Several generations are produced each summer. Ladybugs are often used to control such insect pests as aphids, scales, and mites, which they eat. Several species of ladybugs feed on plants.

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Any member of about 700 species (family Dermestidae) of widely distributed beetles that are household pests. Usually brown or black, some are brightly coloured or patterned, and they vary in shape from elongated to oval. Dermestids range from 0.05 to 0.5 in. (1 to 12 mm) long and are covered with hairs or scales that easily flake off. The wormlike larvae feed on furs, skins, feathers, horn, and hair; some feed on cheese and dried meats or on carpets, rugs, furniture, and clothing. Two are museum pests that have destroyed collections of stuffed animals; museums and collectors must either have pestproof display shelves or continuously apply pesticides. The larvae of carrion-feeding species are sometimes used to clean the soft tissue attached to animal skeletons.

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Any member of the beetle subfamily Alticinae (family Chrysomelidae), found worldwide. It is tiny (less than 0.25 in. [6 mm] long) and dark or metallic in colour. The enlarged hind legs are adapted for jumping. Flea beetles are important pests of cultivated plants (e.g., grapes, cucumbers, melons, tobacco, potatoes, and tomatoes). The adults feed on the leaves, the larvae on the roots. Some flea beetles carry plant diseases (e.g., early potato blight).

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Any member of one subfamily (Scarabaeinae) of scarab beetles, which shapes manure into a ball (sometimes as large as an apple) with its scooperlike head and paddle-shaped antennae. They vary from 0.2 to more than 1 in. (5–30 mm) long. In early summer it buries itself and the ball and feeds on it. Later in the season the female deposits eggs in dung balls, on which the larvae will later feed. They are usually round with short wing covers (elytra) that expose the end of the abdomen. They can eat more than their own weight in 24 hours and are considered helpful because they hasten the conversion of manure to substances usable by other organisms.

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Deathwatch beetle (Xestobium refuvillosum)

Borer insect (beetle species Xestobium refuvillosum) that tends to be small (less than 0.5 in., or 1–9 mm) and cylindrical. When disturbed, it usually pulls in its legs and plays dead. It makes a ticking or clicking sound by bumping its head or jaws against the sides of the tunnels it creates as it bores into old furniture and wood, a sound that, according to superstition, forecasts a death.

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Any of several leaf beetles (genus Diabrotica) that are important pests. They are greenish yellow, marked with black spots or stripes, and 0.1–0.5 in. (2.5–11 mm) long. The striped cucumber beetle and spotted cucumber beetle both feed on garden plants, and their larvae feed on the roots.

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Blister beetle (Lytta magister).

Any of approximately 2,000 species of beetles (family Meloidae) that secrete an irritating substance, cantharidin, which is used medically as a topical skin irritant to remove warts. In the past, cantharidin was often used to induce blisters, a common remedy for many ailments, and the dried remains of Spanish fly (Lytta vesicatoria) were a major ingredient in so-called love potions. Adult blister beetles, which are often brightly coloured, range between 0.1 and 0.8 in. (3–20 mm) in length. Blister beetles are both helpful and harmful to humans; the larvae eat grasshopper eggs, but the adults destroy crops.

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Bark beetle (Dendroctonus valens)

Any member of the beetle family Scolytidae, many of which severely damage trees. Bark beetles are cylindrical, brown or black, and usually less than 0.25 in. (6 mm) long. A male and females (as many as 60 females with each male) bore into a tree and form a chamber where each female deposits her eggs. The emerging larvae bore away from the chamber, forming a characteristic series of tunnels. Different species attack particular trees, damaging roots, stems, seeds, or fruits. Some species transmit disease (e.g., elm bark beetles carry spores of the fungal Dutch elm disease).

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Large European beetle (Melolontha melolontha) that damages foliage, flowers, and fruit as an adult and plant roots as a larva. In Britain, the name refers more broadly to any of the beetles in this subfamily (Melolonthinae), which are known in North America as June beetles. Seealso chafer, scarab beetle.

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or May beetle or June bug

June beetle (Phyllophaga rugosa).

Any insect of the genus Phyllophaga, belonging to a widely distributed, plant-feeding scarab beetle subfamily (Melolonthinae). These red-brown beetles commonly appear in the Northern Hemisphere on warm spring evenings and are attracted to lights. Heavy-bodied, they are 0.5–1 in. (1.2–2.5 cm) long and have shiny wing covers. They feed on foliage and flowers at night, sometimes causing considerable damage. The larvae live in the soil, and can destroy crops and kill lawns and pastures by severing the grasses from their roots; they are considered excellent fish bait.

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Scarab beetle (Popillia japonica) that is a major pest of plants. Introduced accidentally from Japan into the U.S. in 1916, Japanese beetles are known to feed on more than 200 species of plant. Their larvae feed underground on roots; adults feed on flowers, fruit, and foliage. They range from Maine to South Carolina, and infestations have occurred in other parts of North America. The adult, about 0.4 in. (10 mm) long, is bright metallic green with coppery-brown wing covers. Control efforts include the use of poisonous sprays and a disease-inducing bacterium and introduction of the beetle's natural enemies (certain parasitic wasp and fly species).

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or potato bug

Leaf beetle (Leptinotarsa decemlineata, family Chrysomelidae) native to western North America. It began feeding on the leaves of cultivated potatoes when the plants were introduced into western North America, and by 1874 it had become an important and widespread pest. It has a hemispherical body, about 0.4 in. (10 mm) long, and is orange-red or yellow, with black stripes on the wing covers. Depending on climate, potato beetles may produce one to three generations each year.

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Beetles are the group of insects with the largest number of known species. They are placed in the order Coleoptera (from Greek κολεός, koleos, "sheath"; and πτερόν, pteron, "wing", thus "sheathed wing"), which contains more described species than in any other order in the animal kingdom, constituting about 25% of all known life-forms. 40% of all described insect species are beetles (about 350,000 species), and new species are frequently discovered. Estimates put the total number of species, described and undescribed, at between 5 and 8 million.

Beetles can be found in almost all habitats, but are not known to occur in the sea or in the polar regions. They interact with their ecosystems in several ways. They often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Some species are prey of various animals including birds and mammals. Certain species are agricultural pests, such as the Colorado potato beetle Leptinotarsa decemlineata, the boll weevil Anthonomus grandis, the red flour beetle Tribolium castaneum, and the mungbean or cowpea beetle Callosobruchus maculatus, while other species of beetles are important controls of agricultural pests. For example, beetles in the family Coccinellidae ("ladybirds" or "ladybugs") consume aphids, scale insects, thrips, and other plant-sucking insects that damage crops.

Description

The name "Coleoptera" was given by Aristotle for the hardened shield-like forewings (coleo = shield + ptera = wing).

Other characters of this group which are believed to be monophyletic include a holometabolous life cycle; having a prothorax that is distinct from and freely articulating with the mesothorax; the meso- and meta-thoracic segments fusing to form a pterothorax; a depressed body shape with the legs on the ventral surface; the coxae of legs recessed into cavities formed by heavily sclerotized thoracic sclerites; the abdominal sternites more sclerotized than the tergites; antennae with 11 or fewer segments; and terminal genitalic appendages retracted into the abdomen and invisible at rest.

The general anatomy of beetles is quite uniform, although specific organs and appendages may vary greatly in appearance and function between the many families in the order. Like all insects, beetles' bodies are divided into three sections: the head, the thorax, and the abdomen. When viewed from below, the thorax is that part from which all three pairs of legs and both pairs of wings arise. The abdomen is everything posterior to the thorax. When viewed from above, most beetles appear to have three clear sections, but this is deceptive: on the beetle's upper surface, the middle "section" is a hard plate called the pronotum, which is only the front part of the thorax; the back part of the thorax is concealed by the beetle's wings. Like all arthropods, beetles are segmented organisms, and all three of the major sections of the body are themselves composed of several further segments, although these are not always readily discernible. This further segmentation is usually best seen on the abdomen.

Beetles are generally characterised by a particularly hard exoskeleton and hard forewings (elytra). The beetle's exoskeleton is made up of numerous plates called sclerites, separated by thin sutures. This design creates the armoured defences of the beetle while maintaining flexibility. The elytra are not used for flight, but tend to cover the hind part of the body and protect the second pair of wings (alae). The elytra must be raised in order to move the hind flight wings. A beetle's flight wings are crossed with veins and are folded after landing, often along these veins, and are stored below the elytra.

In some beetles, the ability to fly has been lost. These include the ground beetles (family Carabidae) and some "true weevils" (family Curculionidae), but also some desert and cave-dwelling species of other families. Many of these species have the two elytra fused together, forming a solid shield over the abdomen. In a few families, both the ability to fly and the elytra have been lost, with the best known example being the glow-worms of the family Phengodidae, in which the females are larviform throughout their lives.

Beetles have mouthparts similar to those of grasshoppers. Of these parts, the most commonly known are probably the mandibles, which appear as large pincers on the front of some beetles. The mandibles are a pair of hard, often tooth-like structures that move horizontally to grasp, crush, or cut food or enemies (see defence, below). Two pairs of finger-like appendages are found around the mouth in most beetles, serving to move food into the mouth. These are the maxillary and labial palpi.

The eyes are compound and may display remarkable adaptability, as in the case of whirligig beetles (family Gyrinidae), in which the eyes are split to allow a view both above and below the waterline. Other species also have divided eyes — some longhorn beetles (family Cerambycidae) and weevils — while many beetles have eyes that are notched to some degree. A few beetle genera also possess ocelli, which are small, simple eyes usually situated farther back on the head (on the vertex).

Beetles' antennae are primarily organs of smell, but may also be used to feel out a beetle's environment physically. They may also be used in some families during mating, or among a few beetles for defence. Antennae vary greatly in form within the Coleoptera, but are often similar within any given family. In some cases, males and females of the same species will have different antennal forms. Antennae may be clavate (flabellate and lamellate are sub-forms of clavate, or clubbed antennae), filiform, geniculate, moniliform, pectinate, or serrate. For images of these antennal forms see antenna (biology).

The legs, which are multi-segmented, end in two to five small segments called tarsi. Like many other insect orders beetles bear claws, usually one pair, on the end of the last tarsal segment of each leg. While most beetles use their legs for walking, legs may be variously modified and adapted for other uses. Among aquatic families — Dytiscidae, Haliplidae, many species of Hydrophilidae and others — the legs, most notably the last pair, are modified for swimming and often bear rows of long hairs to aid this purpose. Other beetles have fossorial legs that are widened and often spined for digging. Species with such adaptations are found among the scarabs, ground beetles, and clown beetles (family Histeridae). The hind legs of some beetles, such as flea beetles (within Chrysomelidae) and flea weevils (within Curculionidae), are enlarged and designed for jumping.

Oxygen is obtained via a tracheal system. Air enters a series of tubes along the body through openings called spiracles, and is then taken into increasingly finer fibres. Pumping movements of the body force the air through the system.

Beetles have hemolymph instead of blood, and the open circulatory system of the beetle is powered by a tube-like heart attached to the top inside of the thorax.

Development

Beetles are endopterygotes with complete metamorphosis.

A single female may lay from several dozen to several thousand eggs during her lifetime. Eggs are usually laid according to the substrate the larva will feed on upon hatching. Among others, they can be laid loose in the substrate (e.g. flour beetle), laid in clumps on leaves (e.g. Colorado potato beetle), or individually attached (e.g. mungbean beetle and other seed borers) or buried in the medium (e.g. carrot weevil).

The larva is usually the principal feeding stage of the beetle life cycle. Larvae tend to feed voraciously once they emerge from their eggs. Some feed externally on plants, such as those of certain leaf beetles, while others feed within their food sources. Examples of internal feeders are most Buprestidae and longhorn beetles. The larvae of many beetle families are predatory like the adults (ground beetles, ladybirds, rove beetles). The larval period varies between species but can be as long as several years.

Beetle larvae can be differentiated from other insect larvae by their hardened, often darkened head, the presence of chewing mouthparts, and spiracles along the sides of the body. Like adult beetles, the larvae are varied in appearance, particularly between beetle families. Beetles whose larvae are somewhat flattened and are highly mobile are the ground beetles, some rove beetles, and others; their larvae are described as campodeiform. Some beetle larvae resemble hardened worms with dark head capsules and minute legs. These are elateriform larvae, and are found in the click beetle (Elateridae) and darkling beetle (Tenebrionidae) families. Some elateriform larvae of click beetles are known as wireworms. Beetles in the families of the Scarabaeoidea have short, thick larvae described as scarabaeiform, but more commonly known as grubs.

All beetle larvae go through several instars, which are the developmental stages between each moult. In many species the larvae simply increase in size with each successive instar as more food is consumed. In some cases, however, more dramatic changes occur. Among certain beetle families or genera, particularly those that exhibit parasitic lifestyles, the first instar (the planidium) is highly mobile in order to search out a host, while the following instars are more sedentary and remain on or within their host. This is known as hypermetamorphosis; examples include the blister beetles (family Meloidae) and some rove beetles, particularly those of the genus Aleochara.

As with all endopterygotes, beetle larvae pupate, and from this pupa emerges a fully formed, sexually mature adult beetle, or imago. Adults have an extremely variable lifespan, from weeks to years, depending on the species.

Physiology and behaviour

Reproduction

Beetles may display extremely intricate behaviour when mating. Smell is thought to be important in the location of a mate.

Conflict can play a part in the mating rituals of species such as burying beetles (genus Nicrophorus) where conflicts between males and females rage until only one of each is left, thus ensuring reproduction by the strongest and fittest. Many male beetles are territorial and will fiercely defend their small patch of territory from intruding males. In such species, the males may often have horns on the head and/or thorax, making their overall body lengths greater than those of the females, unlike most insects.

Pairing is generally short but in some cases will last for several hours. During pairing sperm cells are transferred to the female to fertilise the egg.

Parental care varies between species, ranging from the simple laying of eggs under a leaf to certain scarab beetles, which construct underground structures complete with a supply of dung to house and feed their young. Other beetles are leaf rollers, biting sections of leaves to cause them to curl inwards, then laying their eggs, thus protected, inside.

Defense

Beetles and their larvae have a variety of strategies to avoid being attacked by predators or parasitoids. These include camouflage, mimicry, toxicity, and active defence.

Camouflage involves the use of colouration or shape to blend into the surrounding environment. This sort of protective coloration is common and widespread among beetle families, especially those that feed on wood or vegetation, such as many of the leaf beetles (family Chrysomelidae) or weevils. In some of these species, sculpturing or various coloured scales or hairs cause the beetle to resemble bird dung or other inedible objects. Many of those that live in sandy environments blend in with the coloration of the substrate.

Another defence that often uses colour or shape to deceive potential enemies is mimicry. A number of longhorn beetles (family Cerambycidae) bear a striking resemblance to wasps, which helps them avoid predation even though the beetles are in fact harmless. This defence can be found to a lesser extent in other beetle families, such as the scarab beetles. Beetles may combine their colour mimicry with behavioural mimicry, acting like the wasps they already closely resemble. Many beetle species, including ladybirds, blister beetles, and lycid beetles can secrete distasteful or toxic substances to make them unpalatable or even poisonous. These same species often exhibit aposematism, where bright or contrasting colour patterns warn away potential predators, and there are, not surprisingly, a great many beetles and other insects that mimic these chemically-protected species.

Large ground beetles and longhorn beetles may defend themselves using strong mandibles and/or spines or horns to forcibly persuade a predator to seek out easier prey. Others, such as bombardier beetles (within Carabidae), may spray chemicals from their abdomen to repel predators.

Feeding

Besides being abundant and varied, the Coleoptera are able to exploit the wide diversity of food sources available in their many habitats. Some are generalists, eating both plants and animals. Other beetles are highly specialised in their diet. Many species of leaf beetles, longhorn beetles, and weevils are very host specific, feeding on only a single species of plant. Ground beetles and rove beetles (family Staphylinidae), among others, are primarily carnivorous and will catch and consume many other arthropods and small prey such as earthworms and snails. While most predatory beetles are generalists, a few species have more specific prey requirements or preferences.

Decaying organic matter is a primary diet for many species. This can range from dung, which is consumed by coprophagous species such as certain scarab beetles (family Scarabaeidae), to dead animals, which are eaten by necrophagous species such as the carrion beetles (family Silphidae). Some of the beetles found within dung and carrion are in fact predatory, such as the clown beetles, preying on the larvae of coprophagous and necrophagous insects.

Adaptations to the environment

Aquatic beetles use several techniques for retaining air beneath the water's surface. Beetles of the family Dytiscidae hold air between the abdomen and the elytra when diving. Hydrophilidae have hairs on their under surface that retain a layer of air against their bodies. Adult crawling water beetles use both their elytra and their hind coxae (the basal segment of the back legs) in air retention  while whirligig beetles simply carry an air bubble down with them whenever they dive.

Evolutionary history and classification

While some authorities believe modern beetles began about 140 million years ago, research announced in 2007 showed that beetles may have entered the fossil record during the Lower Permian, about 265 to 300 million years ago.

The four extant suborders of beetle are these:

These suborders diverged in the Permian and Triassic. Their phylogenetic relationship is uncertain, with the most popular hypothesis being that Polyphaga and Myxophaga are most closely related, with Adephaga as the sister group to those two, and Archostemata as sister to the other three collectively.

There are about 350,000 species of beetles. Such a large number of species poses special problems for classification, with some families consisting of thousands of species and needing further division into subfamilies and tribes.

Impact on humans

Pests

Many agricultural, forestry, and household insect pests are beetles. These include the following:

Beneficial organisms

  • Both the larvae and adults of some ladybirds (family Coccinellidae) are found in aphid colonies. Other lady beetles feed on scale insects and mealybugs. If normal food sources are scarce they may feed on other things, such as small caterpillars, young plant bugs, honeydew and nectar.
  • Ground beetles (family Carabidae) are common predators of many different insects and other arthropods, including fly eggs, caterpillars, wireworms and others.
  • Plant-feeding beetles are often important beneficial insects, controlling problem weeds. Some flea beetles of the genus Aphthona feed on leafy spurge, a considerable weed of rangeland in western North America.

Some farmers develop beetle banks to foster and provide cover for beneficial beetles.

Beetles of the Dermestidae family are often used in taxidermy to clean bones of remaining flesh.

Beetles in ancient Egypt and other cultures

Several species of dung beetle, most notably Scarabaeus sacer (often referred to as "scarab"), enjoyed a sacred status among the ancient Egyptians, as the creatures were likened to the major god Khepri. Some scholars suggest that the Egyptians' practice of making mummies was inspired by the brooding process of the beetle. Many thousands of amulets and stamp seals have been excavated that depict the scarab. In many artifacts, the scarab is depicted pushing the sun along its course in the sky, much as scarabs push or roll balls of dung to their brood sites. During and following the New Kingdom, scarab amulets were often placed over the heart of the mummified deceased.

Some tribal groups, particularly in tropical parts of the world, use the colourful, iridescent elytra of certain beetles, especially certain Scarabaeidae, in ceremonies and as adornment.

Study and collection

The study of beetles is called coleopterology (from Coleoptera, see above, and Greek -λογία, -logia), and its practitioners are coleopterists (see this list). Coleopterists have formed organisations to facilitate the study of beetles. Among these is The Coleopterists Society, an international organisation based in the United States. Such organisations may have both professionals and amateurs interested in beetles as members.

Research in this field is often published in peer-reviewed journals specific to the field of coleopterology, though journals dealing with general entomology also publish many papers on various aspects of beetle biology. Some of the journals specific to beetle research are:

There is a thriving industry in the collection of beetle specimens for amateur and professional collectors. Many coleopterists prefer to collect beetle specimens for themselves, recording detailed information about each specimen and its habitat. Such collections add to the body of knowledge about the Coleoptera. Some countries have established laws governing or prohibiting the collection of certain rare (and often much sought after) species. One such beetle whose collection is illegal or restricted is the American burying beetle, Nicrophorus americanus.

See also

  • Heteroptera - insect suborder that is superficially similar to beetles

References

General references

  • Poul Beckmann, Living Jewels: The Natural Design of Beetles ISBN 3-7913-2528-0
  • Arthur V. Evans, Charles Bellamy, and Lisa Charles Watson, An Inordinate Fondness for Beetles ISBN 0-520-22323-3
  • Entomological Society of America, Beetle Larvae of the World ISBN 0-643-05506-1
  • David Grimaldi, Michael S. Engel, Evolution of the Insects ISBN 0-521-82149-5
  • Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2001-2). ISBN 0-8493-1925-0
  • K. W. Harde, A Field Guide in Colour to Beetles ISBN 0-7064-1937-5 Pages 7-24
  • White, R.E. 1983. Beetles. Houghton Mifflin Company, New York, NY. ISBN 0-395-91089-7

Cited references

External links

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