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Beetle
Fossil range: 318–0 Ma
Pennsylvanian - Recent
Phyllobius calcaratus, a species of weevil
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Subclass: Pterygota
Infraclass: Neoptera
Superorder: Endopterygota
Order: Coleoptera
Linnaeus, 1758
Suborders

Adephaga
Archostemata
Myxophaga
Polyphaga
See subgroups of the order Coleoptera

Beetles are the group of insects with the largest number of known species. They are classified in the order Coleoptera (pronounced /ˌkoʊliˈɒptərə/; 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.[1] About 40% of all described insect species are beetles (about 350,000 species[1]), and new species are frequently discovered. Estimates put the total number of species, described and undescribed, at between 5 and 8 million.[citation needed] The largest family also belongs to this order — the weevils, or snout beetles, Curculionidae.

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.

Contents

Description

The name "Coleoptera" was given by Aristotle for the hardened shield-like forewing (coleo = shield + ptera = wing).[1]

A cockchafer with its elytra raised, exposing the membranous flight wings, where the veins are visible
Trogodendron fasciculatum, a clerid beetle with bright yellow antennae
The extremely colourful Golden Stag Beetle

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.[1]

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

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 vertexes).

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).

Acilius sulcatus, a diving beetle showing hind legs adapted for life in water

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 driven by a tube-like heart attached to the top inside of the thorax.

Development

soil dwelling larva of a weevil, probably the vine weevil genus Otiorhynchus

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.

Beetles may be preyed upon by other insects such as robberflies

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

Punctate Flower Chafers mating
A flower beetle, Eudicella gralli, from the forests of Central Africa. The iridescent elytra are used in indigenous marriage ceremonies.

Beetles may display extremely intricate behaviour when mating. Pheromone communication 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 defense.

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

A Fiddler Beetle feeding from a flowering Cotoneaster glaucophyllus shrub

Besides being abundant and varied, the Coleoptera are able to exploit the wide diversity of food sources available in their many habitats. Some are omnivores, 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[2] while whirligig beetles simply carry an air bubble down with them whenever they dive.

Evolutionary history and classification

Sphaerius acaroides, a member of the small suborder Myxophaga

A 2007 study based on DNA of living beetles and maps of likely beetle evolution indicated that beetles may have originated during the Lower Permian, up to 299 million years ago.[3] In 2009, a fossil beetle was described from the Pennsylvanian of Mazon Creek, Illinois, pushing the origin of the beetles to an earlier date, between 318 to 299 million years ago.[4]

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 450,000 species of beetles — representing about 40% of all known insects. 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. This immense number of species allegedly led evolutionary biologist J. B. S. Haldane to quip when some theologians asked him what could be inferred about the mind of the Creator from the works of His Creation that God displayed "An inordinate fondness for beetles."[5]

Impact on humans

Pests

Colorado potato beetle (Leptinotarsa decemlineata) larvae

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

Beneficial organisms

Coccinella septempunctata, a beneficial beetle
  • Both the larvae and adults of some ladybugs (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.
  • Dung beetles (Coleoptera, Scarabidae) have been successfully used to reduce the populations of pestilent flies and parasitic worms that breed in cattle dung. The beetles make the dung unavailable to breeding pests by quickly rolling and burying it in the soil, with the added effect of improving soil fertility and nutrient cycling. The Australian Dung Beetle Project (1965–1985), led by Dr. George Bornemissza of the Commonwealth Scientific and Industrial Research Organisation introduced species of dung beetle to Australia from South Africa and Europe and effectively reduced the bush fly population by 90%.

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

Ancient Egyptian scene depicting a scarab beetle
Beetle collection at the Melbourne Museum, Australia

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.[citation needed] 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:

  • The Coleopterist (United Kingdom beetle fauna)
  • The Coleopterists Bulletin (published by The Coleopterists Society)
  • Elytron (published by the European Association of Coleopterology)

See also

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

  1. ^ a b c d James K. Liebherr and Joseph V. McHugh in Resh, V. H. & R. T. Cardé (Editors) 2003. Encyclopedia of Insects. Academic Press.
  2. ^ R. H. Arnett, Jr. & M. C. Thomas (2001). "Haliplidae". American Beetles, Volume 1. CRC Press, Boca Raton, Florida. pp. 138–143. ISBN 0-8493-1925-0. 
  3. ^ Modern Beetles Predate Dinosaurs, Dave Mosher, LiveScience.com, 26 December 2007.
  4. ^ Oliver Bethoux, The earliest beetle identified, Journal of Paleontology, vol. 83, no. 6 (Nov. 2009), p. 931.
  5. ^ Hutchinson, G. Evelyn (1959). "Homage to Santa Rosalia or Why Are There So Many Kinds of Animals?". The American Naturalist 93 (870): 145–159. doi:10.1086/282070. http://links.jstor.org/sici?sici=0003-0147(195905/06)93%3A870%3C145%3AHTSROW%3E2.0.CO;2-D. 
  6. ^ Inquirer.net, Beetles infest coconuts in Manila, 26 provinces
  7. ^ The Mountain Pine Beetle in British Columbia, Natural Resources Canada

External links


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Wiktionary

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Definition from Wiktionary, a free dictionary

Contents

Translingual

Etymology

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Wikipedia

From Ancient Greek κολεόπτερος (sheath-wing), from κολεόν (koleon), sheath) and πτερόν (pteron), wing).

Proper noun

Coleoptera

  1. a taxonomic order, within superorder Endopterygota - the beetles.

Hyponyms

Wikispecies-logo.svg
Wikispecies has information on:

Wikispecies

  • Adephaga (wrinkled bark beetles and ground beetles)
  • Archostemata
  • Myxophaga (minute beetles)
  • Polyphaga (most beetles)

Wikispecies

Up to date as of January 23, 2010

From Wikispecies

Coleoptera

Taxonavigation

Main Page
Cladus: Eukaryota
Supergroup: Unikonta
Cladus: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Cladus: Pterygota
Cladus: Metapterygota
Cladus: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordines: Adephaga - Archostemata - Myxophaga - Polyphaga

stem group

Familiae: †Permocupedidae - †Rhombocoleidae - †Triadocupedinae - †Tshekardocoleidae

incertae sedis

Familiae: †Labradorocoleidae - †Obrieniidae - †Taldycupedidae - ?†Oborocoleidae
Genera: †Adiphlebia - †Permocoleus

Name

Coleoptera

References

  • Arnett, R.H., jr.; Thomas, M.C.; Skelley, P.E.; Frank, J.H. (eds.) 2002: American beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press LLC, Boca Raton, Florida. limited preview
  • Béthoux, O. 2009: The earliest beetle identified. Journal of paleontology, 83: 931–937. doi: 10.1666/08-158.1
  • Beutel, R.G.; Ge, S.-Q.; Hörnschemeyer, T. 2008: On the head morphology of Tetraphalerus, the phylogeny of Archostemata and the basal branching events in Coleoptera. Cladistics, 24: 270-298.
  • Beutel, R.G.; Haas, F. 2000: Phylogenetic relationships of the suborders of Coleoptera (Insecta). Cladistics, 16: 103-141.
  • Beutel, R.G.; Haas, F. 2003: Basal splitting events in Coleoptera. Pp. 160-161 in: Klass, K.D. (ed.) Proceedings of the 1st Dresden meeting on insect phylogeny: “Phylogenetic relationships within the insect orders” (Dresden, September 19–21, 2003). Entomologische Abhandlungen, 61: 119-172.[1]
  • Beutel, R.G.; Leschen, R.A.B. (volume eds.) 2005: Coleoptera, Beetles. Volume 1: Morphology and systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim). In: Kristensen, N.P. & Beutel, R.G. (eds.) Handbook of zoology. A natural history of the phyla of the animal kingdom. Volume IV. Arthropoda: Insecta. Part 38. Berlin, New York: Walter de Gruyter. contents limited preview
  • Bishop, D.J.; Majka, C.G.; Bondrup-Nielsen, S.; Peck, S.B. 2009: Deadwood and saproxylic beetle diversity in naturally disturbed and managed spruce forests in Nova Scotia. In: Majka, C.G.; Klimaszewski, J. (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys, 22: 309–340. doi: 10.3897/zookeys.22.144
  • Caterino, M.S.; Shull, V.L.; Hammond, P.M.; Vogler, A.P. 2002: Basal relationships of Coleoptera inferred from 18S rDNA sequences. Zoologica scripta, 31: 41-49.
  • Crotch, G.R. 1871: List of all the Coleoptera described A.D. 1758-1821, referred to their modern genera. Cambridge. BUGZ
  • Cuccodoro, G.; Leschen, R.A.B. (eds.) 2003: Systematics of Coleoptera: papers celebrating the retirement of Ivan Löbl. Memoirs on entomology international, 17. contents
  • Dollin, P.E.; Majka, C.G.; Duinker, P.N. 2008: Saproxylic beetle (Coleoptera) communities and forest management practices in coniferous stands in southwest Nova Scotia, Canada. In: Majka, C.G.; Klimaszewski, J. (eds.) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys, 2: 291-336. Abstract PDF
  • Friedrich, F.; Beutel, R.G. 2006: The pterothoracic skeletomuscular system of Scirtoidea (Coleoptera: Polyphaga) and its implications for the high-level phylogeny of beetles. Journal of zoological systematics & evolutionary research, 44: 290-315.
  • Friedrich, F.; Farrell, B.D.; Beutel, R.G. 2008: The thoracic morphology of Archostemata and the relationships of the extant suborders of Coleoptera (Hexapoda). Cladistics, 24: 1-37.
  • Gerlach, J. (ed.) 2009: The Coleoptera of the Seychelles Islands. Pensoft Publishers, Sofia-Moscow. ISBN 9789546424983
  • Hunt, T.; Bergsten, J.; Levkanicova, Z.; Papadopoulou, A.; St. John, O.; Wild, R.; Hammond, P.M.; Ahrens, D.; Balke, M.; Caterino, M.S.; Gomez-Zurita, J.; Ribera, I.; Barraclough, T.G.; Bocakova, M.; Bocak, L.; Vogler, A.P. 2007: A comprehensive phylogeny of beetles reveals the evolutionary origins of a superradiation. Science, 318: 1913-1916. [2]
  • Lawrence, J.F. (coordinator) 1991: Order Coleoptera, in: Stehr, F.W. (ed.) Immature insects, 2. Kendall/Hunt Publishing Company, Iowa, USA.
  • Lawrence, J.F. 1982: Coleoptera. Pp. 482-553 in Parker, S.P. (ed.) Synopsis and classification of living organisms, 2. McGraw-Hill, USA.
  • Lawrence, J.F. 1999: The Australian Ommatidae (Coleoptera: Archostemata): new species, larva and discussion of relationships. Invertebrate taxonomy, 13: 369-390. [3]
  • Lawrence, J.F.; Newton, A.F., jr. 1982: Evolution and classification of beetles. Annual review of ecology and systematics, 13: 261-290. [4]
  • Lawrence, J.F.; Newton, A.F., jr. 1995: Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp. 779-1006 in Pakaluk, J.; Ślipiński, S.A. (eds.) Biology, phylogeny, and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Museum i Instytut Zoologii PAN, Warszawa.
  • Lawrence, J.F.; Ślipiński, S.A.; Pakaluk, J. 1995: From Latreille to Crowson: a history of the higher-level classification of beetles. Pp. 87-154 in Pakaluk, J.; Ślipiński, S.A. (eds.) Biology, phylogeny, and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Museum i Instytut Zoologii PAN, Warszawa.
  • Majka, C.G.; Klimaszewski, J. 2008: Editorial. How far have we come: 170 years of research on Canadian Coleoptera. In: Majka, C.G.; Klimaszewski, J. (eds.) Biodiversity, biosystematics, and ecology of Canadian Coleoptera. ZooKeys, 2: 1-10. Abstract PDF
  • Majka, C.G.; Sikes, D.S. 2009: Thomas L. Casey and Rhode Island's precinctive beetles: taxonomic lessons and the utility of distributional checklists. In: Majka, C.G.; Klimaszewski, J. (eds) Biodiversity, biosystematics, and ecology of Canadian Coleoptera II. ZooKeys, 22: 267–283. doi: 10.3897/zookeys.22.93
  • McKenna, D.D.; Farrell, B.D. 2009: Beetles (Coleoptera). Pp. 278-289 in Hedges, S.B. & Kumar, S. (eds.) The Timetree of Life. Oxford University Press. PDF
  • Ponomarenko, A.G. 1995: The geological history of beetles. Pp. 155-171 in Pakaluk, J.; Ślipiński, S.A. (eds.) Biology, phylogeny, and classification of Coleoptera: papers celebrating the 80th birthday of Roy A. Crowson. Museum i Instytut Zoologii PAN, Warszawa.
  • Ponomarenko, A.G. 2003: Ecological evolution of beetles (Insecta: Coleoptera). Acta zoologica cracoviensia, 46(suppl.-Fossil Insects): 319-328. [5]
  • Seago, A.E.; Brady, P.; Vigneron, J.-P.; Schultz, T.D. 2009: Gold bugs and beyond: a review of iridescence and structural colour mechanisms in beetles (Coleoptera). Journal of the Royal Society Interface, 6, S165–S184. doi:10.1098/rsif.2008.0354.focus
  • Vogler, A.P.; Caterino, M.S. 2003: The basal relationships of Coleoptera based on 18S rRNA sequences. Pp. 161-162 in: Klass, K.D. (ed.) Proceedings of the 1st Dresden meeting on insect phylogeny: “Phylogenetic relationships within the insect orders” (Dresden, September 19–21, 2003). Entomologische Abhandlungen, 61: 119-172.[6]
  • Watt, J.C. 1975: Notes on priority of family-group names in Coleoptera. Coleopterists bulletin, 29: 31-34.

links

Vernacular names

Български: Твърдокрили/Бръмбари
Česky: Brouci
Српски / Srpski: Тврдокрилци
Dansk: Biller
Deutsch: Käfer
Ελληνικά: Κολεόπτερα
English: beetles
Español: Escarabajo
Esperanto: Koleopteroj
Français: Coléoptères
한국어: 딱정벌레목
Italiano: Coleotteri
עברית: חיפושיות
Latina: Coleoptera
Magyar: Bogarak
Македонски: Тврдокрилци
Malagasy: Borera
Nederlands: Kevers
日本語: コウチュウ目 (鞘翅目)
Polski: Chrząszcze
Português: Coleópteros
Русский: Жесткокрылые/Жуки
Slovenščina: Hrošči
Suomi: Kovakuoriaiset
Svenska: Skalbaggar
Türkçe: Kın kanatlılar
Українська: Твердокрилі (жуки)
中文: 鞘翅目
Wikimedia Commons For more multimedia, look at Coleoptera on Wikimedia Commons.

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