Beetle

? Beetles
Colorado potato beetle, Leptinotarsa decemlineata
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 one of the most diverse groups of insects. Their order, Coleoptera (meaning "sheathed wing"), has more species in it than any other order in the entire animal kingdom. Nearly half of all described insect species are beetles, and new species are regularly discovered. Overall, there are about 400,000 known species of beetles, or about one-quarter of all names species in the plant and animal kingdoms. Estimates put the total number of beetle species—described and undescribed—at between 5 and 8 million.

The vast numbers of beetles led to the famous quip, perhaps apocryphal, from British geneticist J. B. S. Haldane, who, when asked what one could conclude as to the nature of God from a study of his creation, replied: "An inordinate fondness for beetles" (Gould 1996). Haldane himself was a noted atheist and for some time a Marxist and this quote reflects not only the vast numbers of beetled but also Haldane's skeptical view of natural theology.

Beetles can be found in almost all habitats, but are not known to occur in the sea or in the polar regions. They have a major impact on the ecosystem in three ways: feeding on plants and fungi, breaking down animal and plant debris, and eating other invertebrates. Certain species can be agricultural pests, for example the Colorado potato beetle (Leptinotarsa decemlineata), while other species are important controls of agricultural pests, for example the ladybirds (family Coccinellidae) consume aphids, fruit flies, thrips, and other plant-sucking insects that damage crops.

Anatomy

The anatomy of beetles is quite uniform. Beetles are generally characterised by a particularly hard exoskeleton, and the hard wing-cases (elytra) that tend to cover the hind part of the body and protect the second wings, the alae. The elytra are not used in flying, but generally must be raised in order to move the hindwings. In some cases, the ability to fly has been lost, characteristically in families such as Carabidae (ground beetles) and Curculionidae (snout beetles and true weevils). After landing, the hindwings are folded below the elytra.

In a few families, both the ability to fly and the wing-cases are absent, with the best known example being the "glowworms" of the family Phengodidae, in which the females are larviform throughout their lives.

The bodies of beetles are divided into three sections, the head, the thorax, and the abdomen, and these in themselves may be composed of several further segments.

The eyes are compound, and may display some remarkable adaptability, as in the case of the Whirligig beetles (family Gyrinidae), in which the eyes are split to allow a view both above and below the waterline. The dorsal appendage aids the beetle in stalking prey.

Like all insects, antennae and legs are both jointed.

Oxygen is taken in via a tracheal system: this takes air in through a series of tubes along the body, which 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

Larva of the cockchafer (Melolontha melolontha)

Beetles are endopterygotes—a superorder of insects of the subclass Pterygota that go through distinctive larval, pupal, and adult stages, or complete metamorphosis. The larva of a beetle is often called a grub and represents the principal feeding stage of the life-cycle.

The eggs of beetles are minute, but may be brightly colored. They are laid in clumps and there may be from several dozen to several thousand eggs laid by a single female.

Once the egg hatches, the larvae tend to feed voraciously, whether out in the open such as with Ladybird larvae, or within plants such as with leaf beetle larvae.

As with lepidoptera, beetle larvae pupate for a period, and from the pupa emerges a fully formed beetle or imago.

In some cases there are several transitory larvae stages and this is known as hypermetamorphosis; examples include the blister beetles (family Meloidae).

Physiology

There are few things that a beetle somewhere will not eat. Even inorganic matter may be consumed.

Some beetles are highly specialised in their diet; for example, the Colorado potato beetle (Leptinotarsa decemlineata) almost entirely colonizes plants of the potato family (Solanaceae). Others are generalists, eating both plants and animals. Ground beetles (family Carabidae) and rove beetles (family Staphylinidae) are entirely carnivorous and will catch and consume small prey such as earthworms and snails.

Decaying organic matter is a primary diet for many species, this can range from dung which is consumed by coprophagous species such as the scarab beetles (family Scarabaeidae), to dead animals which are eaten by necrophagous species such as the carrion beetles (family Silphidae).

Various techniques are employed by many species for retaining both air and water supplies. For example,Predaceous diving beetles (family Dytiscidae) employ a technique of retaining air, when diving, between the abdomen and the elytra.

Reproduction

The larval period of beetles varies between species but can be as long as several years. Adults have an extremely variable lifespan of weeks to years.

Beetles may display some 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 for example in species such as burying beetles (genus Nicrophorus) where localised conflicts between males and females rage until only one of each is left, thus ensuring reproduction by the strongest and fittest. Many beetles are territorial and will fiercly defend their small patch of territory from intruding males.

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.

Striped love beetle Eudicella gralli from the forests of Central Africa. The irridescent wing cases are used in marriage ceremonies.

Parental care

Parental care between species varies widely, ranging from the simple laying of eggs under a leaf to scarab beetles, which construct impressive underground structures complete with a supply of dung to house and feed their young.

There are other notable ways of caring for the eggs and young, such as those employed by leaf rollers, who bite sections of leaf causing it to curl inwards and then lay the eggs, thus protected, inside.

Predation

Beetles and larvae have a variety of strategies for avoiding being eaten.

Many employ simple camouflage to avoid being spotted by predators. These include the leaf beetles (family Chysomelidae) that have a green colouring very similar to their habitat on tree leaves.

A number of longhorn beetles (family Cerambycidae) bear a striking resemblance to wasps, thus benefitting from a measure of protection. Large ground beetles by contrast will tend to go on the attack, using their strong mandibles to forcibly persuade a predator to seek out easier prey.

Many species, including lady beetles and blister beetles, can secrete poisonous substances to make them unpalatable.

Evolutionary history and classification

Beetles entered the fossil record during the Lower Permian, about 265 million years ago.

The four extant suborders of beetle are these:

• Polyphaga is the largest suborder, containing more than 300,000 described species in more than 170 families, including rove beetles (Staphylinidae), scarab beetles (Scarabaeidae), blister beetles (Meloidae), stag beetles (Lucanidae), and true weevils (Curculionidae). These beetles can be identified by the cervical sclerites (hardened parts of the head used as points of attachment for muscles) absent in the other suborders.
• Adephaga contains about 10 families of predatory beetles, includes ground beetles (Carabidae), predacious diving beetles (Dytiscidae) and whirligig beetles (Gyrinidae). In these beetles the testes are tubular and the first abdominal sternum (a plate of the exoskeleton) is divided by the hind coxae (the basal joints of the beetle's legs).
• Archostemata contains four families of mainly wood-eating beetles, including reticulated beetles (Cupedidae) and telephone-pole beetles (Micromalthidae).
• Myxophaga contains about 100 described species in four families, mostly very small, including skiff beetles (Hydroscaphidae) and minute bog beetles (Sphaeriusidae).

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 an outgroup to those two, and Archostemata an outgroup to the other three.

The extraordinary number of beetle species poses special problems for classification, with some families consisting of thousands of species and needing further division into subfamilies and tribes.

See the article subgroups of the order Coleoptera for a complete list of families and [1] for a complete list of World families and subfamilies.

Impact on humans

Pests

Damage to beans by larvae of the common bean weevil, Acanthoscelides obtectus

There are several agricultural and household pests represented by the order, these include :

• The Colorado potato beetle (Leptinotarsa decemlineata) is a notorious pest of potato plants. Adults mate before overwintering deep in the soil, so that when they emerge the following spring, females can lay eggs immediately, once a suitable host plant has been found. As well as potatoes, this can be a number of plants from the potato family (Solanaceae) such as nightshade, tomato, aubergine and capsicum. Crops are destroyed and the beetle can only be treated by employing expensive pesticides, many of which it has begun to develop immunity to.
• The elm bark beetles, Hylurgopinus rufipes, elm leaf beetle Pyrrhalta luteola and Scolytus multistriatus (in the family Scolytidae) attack elm trees. They are important elm pests because they carry Dutch elm disease (the fungus Ophiostoma ulmi) as they move from infected breeding sites to feed on healthy elm trees. The spread of the fungus by the beetle has led to the devastastation of elm trees in many parts of the Northern Hemisphere, notably North America and Europe.
• The death watch beetle (Xestobium rufovillosum) is of some considerable importance as a pest of wooden structures in older buildings in Britain. It attacks hardwoods such as oak and chestnut, and always where some fungal decay has taken or is taking place. It is thought that the actual introduction of the pest into buildings takes place at the time of construction.
• Asian long-horned beetle
• Citrus long-horned beetle

Beneficial organisms

• The larvae of lady beetles (family Coccinellidae) are often found in aphid colonies. While both adult and larval lady beetles found on crops prefer aphids, they will, if aphids are scarce, use food from other sources, such as small caterpillars, young plant bugs, aphid honeydew, and plant nectar.
• Large ground beetles (family Carabidae) are predators of caterpillars and, on occasion, adult weevils, whereas smaller species attack eggs, small caterpillars, and other pest insects.

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

Scarab beetles in Egyptian culture

Ancient Egyptian scene depicting a scarab beetle

Several species of the dung beetles, most notably the Scarabaeus sacer (often referred to as "scarab"), enjoyed a sacred status among the ancient Egyptians, as the creature was likened to the god Khepri. Some scholars suggested that the people's practice of making mummy 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. During and following the New Kingdom, scarab amulets were often placed over the heart of the mummified deceased. The amulets were often inscribed with a spell from the Book of the Dead which entreated the heart to, "do not stand as a witness against me."

Collecting

Beetle collection at the Melbourne Museum, Australia

The study of beetles is called coleopterology, and its practitioners coleopterists. See the list of list of notable coleopterists for more information.

There is a thriving industry in the collection of wild caught species for amateur and professional collectors.

Taxonomy

This section classifies the subgroups of the order Coleoptera (beetles) down to the level of families, following the system in Arnett and Thomas' (2001, 2002) two volume set American Beetles. A few subfamilies, tribes and synonyms are shown here where a family has been reclassified.

Order Coleoptera (beetles)

• Suborder Adephaga Schellenberg 1806
  • Amphizoidae LeConte 1853 (trout-stream beetles)
  • Carabidae Latreille 1802 (ground beetles)
    • Cicindelinae, formerly Cicindelidae Latreille 1802 (tiger beetles)
    • Paussinae, formerly Paussidae Latreille 1807
  • Dytiscidae Leach 1815 (predacious diving beetles)
    

    Dytiscus latissimus, a predacious diving beetle
  • Gyrinidae Latreille 1802 (whirligig beetles)
  • Haliplidae Aube 1836 (crawling water beetles)
  • Hygrobiidae Régimbart 1878 (1837)
  • Noteridae C.G. Thomson 1860 (burrowing water beetles)
  • Rhysodidae Laporte 1840 (wrinkled bark beetles)
  • Trachypachidae C.G. Thomson 1857(false ground beetles)
• Suborder Archostemata Kolbe 1908
  • Crowsonellidae Iablokoff-Khnzorian 1983
  • Cupedidae Laporte 1836 (reticulated beetles)
    File:Archostemata.jpg

    Priacma serrata, a reticulated beetle
  • Micromalthidae Barber 1983 (telephone-pole beetles)
  • Ommatidae Sharp and Muir 1912
• Suborder Myxophaga Crowson 1955
  • Hydroscaphidae LeConte 1874 (skiff beetles)
  • Lepiceridae Hinton 1936
  • Sphaeriusidae Erichson 1845 (minute bog beetles) (= Microsporidae: Bulletin of Zoological Nomenclature 57(3): 182-184.)
  • Torridincolidae Steffan 1964
• Suborder Polyphaga Emery 1886
  • Infraorder Bostrichiformia Forbes 1926
    • Superfamily Bostrichoidea Latreille 1802
      • Anobiidae Fleming 1821 (death watch beetles)
        • Ptininae, formerly Ptinidae Latreille 1802 (spider beetles)
      • Bostrichidae Latreille 1802 (horned powder-post beetles)
        • Lyctinae, formerly Lyctidae Billberg 1820 (powder post beetles)
        • Endecatominae, formerly Endecatomidae LeConte 1861
      • Dermestidae Latreille 1804 (skin beetles)
        • Thorictinae, formerly Thorictidae Agassiz 1846
      • Jacobsoniidae Heller 1926 (Jacobson's beetles)
      • Nosodendridae Erichson 1846 (wounded-tree beetles)
    • Superfamily Derodontoidea LeConte 1861
      • Derodontidae LeConte 1861 (tooth-necked fungus beetles)
  • Infraorder Cucujiformia Lameere 1938
    • Superfamily Chrysomeloidea Latreille 1802
      • Bruchidae Latreille 1802 (pea and bean weevils)
      • Cerambycidae Latreille 1802 (long-horned beetles)
      • Chrysomelidae Latreille 1802 (leaf beetles)
        

        Colorado potato beetle, Leptinotarsa decemlineata, a leaf beetle
        • Cassidinae (tortoise beetles subfamily)
      • Megalopodidae Latreille 1802
      • Orsodacnidae C. G. Thomson 1869
    • Superfamily Cleroidea Latreille 1802
      • Acanthocnemidae Crowson 1964
      • Chaerosomatidae Crowson 1952
      • Cleridae Latreille 1802 (checkered beetles)
      • Melyridae Leach 1815 (soft-winged flower beetles)
      • Phloiophilidae Kiesenwetter 1863
      • Phycosecidae Crowson 1952
      • Prionoceridae Lacordaire 1857
      • Trogossitidae Latreille 1802 (bark-gnawing beetles)
    • Superfamily Cucujoidea Latreille 1802
      • Alexiidae Imhoff 1856
      • Biphyllidae LeConte 1861 (false skin beetles)
      • Boganiidae Sen Gupta and Crowson 1966
      • Bothrideridae Erichson 1845 (dry bark beetles)
      • Byturidae Jacquelin du Val 1858 (fruitworm beetles)
      • Cavognathidae Sen Gupta and Crowson 1966
      • Cerylonidae Billberg 1820 (minute bark beetles)
      • Coccinellidae Latreille 1807 (lady beetles)
      • Corylophidae LeConte 1852 (minute fungus beetles)
      • Cryptophagidae Kirby 1937 (silken fungus beetles)
      • Cucujidae Latreille 1802 (flat bark beetles)
      • Discolomatidae Horn 1878
      • Endomychidae Leach 1815 (handsome fungus beetles)
        • Merophysiinae, formerly Merophysiidae Seidlitz, 1872
      • Erotylidae Latreille 1802 (pleasing fungus beetles)
      • Helotidae Reitter 1876
      • Hobartiidae Sen Gupta and Crowson 1966
      • Kateretidae Erichson in Agassiz 1846 (short-winged flower beetles) (= Brachypteridae. ICZN Op. 1916, 1999).
      • Laemophloeidae Ganglbauer 1899 (lined flat bark beetles)
      • Lamingtoniidae Sen Gupta and Crowson 1966
      • Languriidae Crotch , 1873 (lizard beetles)
      • Latridiidae Erichson 1842 (minute brown scavenger beetles)
      • Monotomidae Laporte 1840 (root-eating beetles)
        • Rhizophaginae, formerly Rhizophagidae Redtenbacher 1845
      • Nitidulidae Latreille 1802 (sap-feeding beetles)
      • Passandridae Erichson 1845 (parasitic flat bark beetles)
      • Phalacridae Leach 1815 (shining flower beetles)
      • Phloeostichidae Reitter 1911
      • Propalticidae Crowson 1952
      • Protocucujidae Crowson 1954
      • Silvanidae Kirby 1937 (silvanid flat bark beetles)
      • Smicripidae Horn 1879 (palmetto beetles)
      • Sphindidae Jacquelin du Val 1860 (dry-fungus beetles)
    • Superfamily Curculionoidea Latreille 1802
      • Anthribidae Billberg 1820 (fungus weevils)
      • Attelabidae Billberg 1820 (tooth-nosed snout beetles)
      • Belidae Schönherr 1826 (primitive weevils)
        • Aglycyderinae, formerly Aglycyderidae Wollaston 1864
        • Oxycoryninae, formerly Oxycorynidae Schönherr 1840
      • Brentidae Billberg 1820 (straight-snouted weevils)
        • Apioninae, formerly Apionidae Schönherr, 1823
      • Caridae Thompson 1992
      • Curculionidae Latreille 1802 (snout beetles and true weevils)
        

        Citrus root weevil, Diaprepes abbreviatus
        • Scolytinae, formerly Scolytidae Latreille 1807 (typical bark beetles)
      • Ithyceridae Schönherr 1823 (New York weevils)
      • Nemonychidae Bedel 1882 (pine-flower snout beetles)
    • Superfamily Lymexyloidea Fleming 1821
      • Lymexylidae Fleming 1821 (ship-timber beetles)
    • Superfamily Tenebrionoidea Latreille 1802
      • Aderidae Winkler 1927 (antlike leaf beetles)
      • Anthicidae Latreille 1819 (antlike flower beetles)
      • Archeocrypticidae Kaszab 1964
      • Boridae C. G. Thomson 1859 (conifer bark beetles)
      • Chalcodryidae Watt 1974
      • Ciidae Leach 1819 (minute tree-fungus beetles) (= Cisidae)
      • Colydiidae Erichson 1842 (cylindrical bark beetles)
      • Melandryidae Leach 1815 (false darkling beetles)
      • Meloidae Gyllenhal 1810 (blister beetles)
        

        Lytta magister, a blister beetle
      • Monommatidae Blanchard 1845 (opossum beetles)
      • Mordellidae Latreille 1802 (tumbling flower beetles)
      • Mycetophagidae Leach 1815 (hairy fungus beetles)
      • Mycteridae Blanchard 1845 (palm and flower beetles)
        • Hemipeplinae, formerly Hemipeplidae Lacordaire, 1854
      • Oedemeridae Latreille 1810 (pollen-feeding beetles)
      • Perimylopidae St. George 1939
      • Prostomidae C. G. Thomson 1859 (jugular-horned beetles)
      • Pterogeniidae Crowson 1953
      • Pyrochroidae Latreille 1807 (fire-colored beetles)
        • Cononotini or Cononotidae
        • Pedilinae, formerly Pedilidae Lacordaire 1859
      • Pythidae Solier 1834 (dead log bark beetles)
      • Rhiphiphoridae Gemminger and Harold 1870 (wedge-shaped beetles)
      • Salpingidae Leach 1815 (narrow-waisted bark beetles)
        • Elacatini or Elacatidae
        • Inopeplinae, formerly Inopeplidae Grouvelle 1908
      • Scraptiidae Mulsant 1856 (false flower beetles)
      • Stenotrachelidae C. G. Thomson 1859 (false long-horned beetles)
        • Cephaloinae, formerly Cephaloidae LeConte 1852
      • Synchroidae Lacordaire 1859
      • Tenebrionidae Latreille 1802 (darkling beetles)
        

        Heliotautus ruficollis, a darkling beetle
        • Alleculinae, formerly Alleculidae Laporte 1840
        • Lagriinae, formerly Lagriidae Latreille 1825
        • Nilionini or Nilionidae Lacordaire 1859
        • Petriini or Petriidae
      • Tetratomidae Billberg 1820 (polypore fungus beetles)
      • Trachelostenidae Lacordaire 1859
      • Trictenotomidae Blanchard 1845
      • Ulodidae Pascoe 1869
      • Zopheridae Solier 1834 (ironclad beetles)
  • Infraorder Elateriformia Crowson 1960
    • Superfamily Buprestoidea Leach 1815
      • Buprestidae Leach 1815 (metallic wood-boring beetles)
        File:Emerald ash beetle.jpg

        Emerald ash borer, Agrilus planipennis, a metallic wood-boring beetle
      • Schizopodidae LeConte 1861
    • Superfamily Byrrhoidea Latreille 1804
      • Byrrhidae Latreille 1804 (pill beetles)
      • Callirhipidae Emden 1924 (cedar beetles)
      • Chelonariidae Blanchard 1845 (turtle beetles)
      • Cneoglossidae Champion 1897
      • Dryopidae Billberg 1820 (long-toed water beetles)
      • Elmidae Curtis 1830 (riffle beetles)
      • Eulichadidae Crowson 1973
      • Heteroceridae MacLeay 1825 (variegated marsh-loving beetles)
      • Limnichidae Erichson 1846 (minute marsh-loving beetles)
      • Lutrochidae Kasap and Crowson 1975 (robust marsh-loving beetles)
      • Psephenidae Lacordaire 1854 (water penny beetles)
      • Ptilodactylidae Laporte 1836 (toe-winged beetles)
    • Superfamily Dascilloidea Guerin-Meneville 1843
      • Dascillidae Guérin-Méneville 1843 (1834) (soft-bodied plant beetles)
      • Rhipiceridae Latreille 1834 (cicada parasite beetles)
    • Superfamily Elateroidea Leach 1815
      • Artematopodidae Lacordaire 1857 (soft-bodied plant beetles) = (Eurypogonidae)
      • Brachypsectridae Leconte and Horn 1883 (Texas beetles)
      • Cantharidae Imhoff 1856 (soldier beetles)
      • Cerophytidae Latreille 1834 (rare click beetles)
      • Drilidae Blanchard 1845
      • Elateridae Leach 1815 (click beetles)
        

        Alaus murina, a click beetle
        • Cebrioninae, formerly Cebrionidae Latreille 1802
      • Eucnemidae Eschscholtz 1829 (false click beetles)
      • Lampyridae Latreille 1817 (firefly beetles)
      • Lycidae Laporte 1836 (net-winged beetles)
      • Omalisidae Lacordaire 1857
      • Omethidae LeConte 1861 (false firefly beetles)
      • Phengodidae LeConte 1861 (glowworm beetles)
      • Plastoceridae Crowson 1972
      • Podabrocephalidae Pic 1930
      • Rhinorhipidae Lawrence 1988
      • Telegeusidae Leng 1920 (long-lipped beetles)
      • Throscidae Laporte 1840 (false metallic wood-boring beetles) = (Trixagidae)
    • Superfamily Scirtoidea Fleming 1821
      • Clambidae Fischer 1821 (minute beetles)
      • Decliniidae Nikitsky et al 1994
      • Eucinetidae Lacordaire 1857 (plate-thigh beetles)
      • Scirtidae Fleming 1821 (marsh beetles) (= Helodidae)
  • Infraorder Scarabaeiformia Crowson 1960
    • Superfamily Scarabaeoidea Latreille 1802
      • Belohinidae Paulian 1959
      • Bolboceratidae Laporte de Castelnau 1840
      • Ceratocanthidae White 1842 (= Acanthoceridae)
      • Diphyllostomatidae Holloway 1972 (false stag beetles)
      • Geotrupidae Latreille 1802 (earth-boring dung beetles)
      • Glaphyridae MacLeay 1819 (bumble bee scarab beetles)
      • Glaresidae Semenov-Tian-Shanskii and Medvedev 1932 (enigmatic scarab beetles)
      • Hybosoridae Erichson 1847 (scavenging scarab beetles)
      • Lucanidae Latreille 1804 (stag beetles)
        

        Lucanus cervus, a stag beetle
      • Ochodaeidae Mulsant and Rey 1871 (sand-loving scarab beetles)
      • Passalidae Leach 1815 (bess beetles)
      • Pleocomidae LeConte 1861 (rain beetles)
      • Scarabaeidae Latreille 1802 (scarab beetles)
        • Dynastinae, formerly Dynastidae MacLeay 1819 (rhinoceros beetles)
      • Trogidae MacLeay 1819 (hide beetles)
  • Infraorder Staphyliniformia Lameere 1900
    • Superfamily Histeroidea Gyllenhal 1808
      • Histeridae Gyllenhal 1808 (clown beetles)
      • Sphaeritidae Schuckard 1839 (false clown beetles)
      • Synteliidae Lewis 1882
    • Superfamily Hydrophiloidea Latreille 1802
      • Hydrophilidae Latreille 1802 (water scavenger beetles)
        • Georyssinae, formerly Georyssidae Laporte 1840
      • Epimetopidae
      • Helophoridae
    • Superfamily Staphylinoidea Latreille 1802
      • Agyrtidae C.G. Thomson 1859
      • Hydraenidae Mulsant 1844
      • Leiodidae Fleming 1821 (round fungus beetles) = (Anisotomidae)
        • Platypsyllinae Ritsema 1869 or Leptinidae
      • Ptiliidae Erichson 1845 (feather-winged beetles)
        File:Staphylinus olens2web.jpg

        Devil's coach horse beetle, Staphylinus olens, a rove beetle
        • Cephaloplectinae, formerly Limulodidae Sharp 1883 (horse-shoe crab beetles)
      • Scydmaenidae Leach 1815 (antlike stone beetles)
      • Silphidae Latreille 1807 (carrion beetles)
      • Staphylinidae Latreille 1802 (rove beetles)
        • Scaphidiinae, formerly Scaphidiidae Latreille 1807
        • Pselaphinae, formerly Pselaphidae Latreille 1802

Gallery

• 

  Anthrenus verbasci

• 

  Lady beetle

• 

  Tiger beetle

• 

  South African Dung Beetle

• Emerald ash beetle.jpg

  Emerald Ash Beetle

• 

  Flea beetle

• 

  Japanese beetle

• 

  Potato Beetle larvae

• 

  Spotted Flower Chafer

• Cerambycidae Coleoptera California Prionus.JPG

  California Prionus

• Beetle August 27-2005..png

  Pelidnota punctata

• 

  Bess Beetle

References

ISBN links support NWE through referral fees

• Poul Beckmann, Living Jewels: The Natural Design of Beetles ISBN 3791325280
• Arthur V. Evans, Charles Bellamy, and Lisa Charles Watson, An Inordinate Fondness for Beetles ISBN 0520223233
• Entomological Society of America, Beetle Larvae of the World ISBN 0643055061
• David Grimaldi, Michael S. Engel, Evolution of the Insects ISBN 0521821495
• Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2001-2)
• K. W. Harde, A Field Guide in Colour to Beetles ISBN 0706419375 Pages 7-24
• Gould, S. J. 1993. A special fondness for beetles. Natural history 1:4-12.

• Ross H. Arnett, Jr. and Michael C. Thomas, American Beetles (CRC Press, 2001–2002)

2001 Volume 1: Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia ISBN 0-8493-1925-0 2002 Volume 2: Polyphaga ISBN 0-8493-0954-9

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