Adephaga

The Adephaga (from Greek ἀδηφάγος, adephagos, "gluttonous"), with more than 40,000 recorded species in 10 families, are a suborder of highly specialized beetles and the second-largest suborder of the order Coleoptera. Members of this suborder are adephagans, a term which notably include ground beetles, tiger beetles, predacious diving beetles, and whirligig beetles. The majority of the species belongs to the family of carabids, or ground beetles (Carabidae).

Adephaga
Temporal range: Changhsingian/InduanHolocene, 251.2–0 Ma[1]
Dytiscus latissimus
Dytiscus latissimus, a predaceous diving beetle
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Adephaga
Schellenberg, 1806
Families

Extant families:

Amphizoidae
Aspidytidae
Carabidae
Dytiscidae
Gyrinidae
Haliplidae
Hygrobiidae
Meruidae Spangler & Steiner 2005
Noteridae
Rhysodidae
Trachypachidae

Extinct families:

Colymbothetidae
Coptoclavidae
Liadytidae
Parahygrobiidae
Triaplidae
76-Indian-Insect-Life - Harold Maxwell-Lefroy - Adephaga
Image by Harold Maxwell-Lefroy - Adephaga

Anatomy

Adephagans have simple antennae with no pectination or clubs. The galeae of the maxillae usually consist of two segments. Adult adephagans have visible notopleural sutures. The first visible abdominal sternum is completely separated by the hind coxae, which is one of the most easily recognizable traits of adephagans. Five segments are on each foot.

Wings

The transverse fold of the hind wing is near the wing tip. The median nervure ends at this fold, where it is joined by a cross nervure.

Internal organs

Adephagans have four Malpighian tubules. Unlike in other beetles, yolk chambers alternate with egg chambers in the ovarian tubes of adephagans. The coiled, tubular testes consist of a single follicle, and the ovaries are polytrophic.

Chemical glands

All families of adephagan have paired pygidial glands located posterodorsally in the abdomen, which are used for secreting chemicals. The glands consist of complex invaginations of the cuticle lined with epidermal cells contiguous with the integument. The glands have no connection with the rectum and open on the eighth abdominal tergum.

Secretions pass from the secretory lobes, which are aggregations of secretory cells, through a tube to a reservoir lined with muscles. This reservoir then narrows to a tube leading to an opening valve. The secretory lobes differ structurally from one taxon to another; it may be elongated or oval, branched basally or apically, or unbranched.

Delivery of glandular compounds

The secretion is realized in one of these manners:

  • Oozing: if the gland is not muscle-lined, the discharge is limited in amount.
  • Spraying: if the gland is muscle-lined, which is typically the case of carabids, the substances are ejected more or less forcefully.
  • Crepitation is only associated with the Brachininae carabids and several related species. See bombardier beetle for a detailed description of the mechanism.

The secretions differ in the chemical constituents, according to the taxa. Gyrinids, for instance, secrete norsesquiterpenes such as gyrinidal, isogyrinidal, gyrinidione, or gyrinidone. Dytiscids discharge aromatic aldehydes, esters, and acids, especially benzoic acid. Carabids typically produce carboxylic acids, particularly formic acid, methacrylic acid, and tiglic acid, but also aliphatic ketones, saturated esters, phenols, aromatic aldehydes, and quinones. Accessory glands or modified structures are present in some taxa: the Dytiscidae and Hygrobiidae also possess paired prothoracic glands secreting steroids; and the Gyrinidae are unique in the extended shape of the external opening of the pygidial gland.

The function of many compounds remain unknown, yet several hypotheses have been advanced:

  • As toxins or deterrent against predators, some compounds indirectly play this role by easing the penetration of the deterrent into the predator's integument.
  • Antimicrobial and antifungal agents (especially in Hydradephaga)
  • A means to increase wetability of the integument (especially in Hydradephaga)
  • Alarm pheromones (especially in Gyrinidae)
  • Propellant on water surfaces (especially in Gyrinidae)
  • Conditioning plant tissues associated with oviposition

Distribution and habitat

Habitats range from caves to rainforest canopy and alpine habitats. The body forms of some are structurally modified for adaptation to habitats: members of the family Gyrinidae live at the air-water interface, rhysodines live in heartwood, and paussine carabids inhabit ant nests.

Feeding

Most species are predators. Other less-typical forms of feeding include: algophagy (family Haliplidae), seed-feeding (arpaline carabids), mycophagy (rhysodine carabids), and snail-feeding (licinine and cychrine carabids). Some species are ectoparasitoids of insects (brachinine and lebiine carabids) or of millipedes (peleciine carabids).

Reproduction and larval stage

Some species are ovoviviparous, such as pseudomorphine carabids.

The larvae are active, with well-chitinized cuticle, often with elongated cerci and five-segmented legs, the foot-segment carrying two claws. Larvae have a fused labrum and no mandibular molae.

Phylogeny

Adephagans diverged from their sister group in the Late Permian, the most recent common ancestor of living adephagans probably existing in the early Triassic, around 240 million years ago. Both aquatic and terrestrial representatives of the suborder appear in fossil records of the late Triassic. The Jurassic fauna consisted of trachypachids, carabids, gyrinids, and haliplid-like forms. The familial and tribal diversification of the group spans the Mesozoic, with a few tribes radiating explosively during the Tertiary.

The phylogeny of adephagans is disputed. The group is usually divided into two main groups:

  • The Geadephaga comprise the two terrestrial families Carabidae and Trachypachidae (the Trachypachidae are sometimes considered a subfamily of the Carabidae).
  • The Hydradephaga, gathering all other families, are aquatic.

This division is often criticized, as mounting evidence is pointing out that the two groups are not monophyletic.

See also

References

  • ZoolScripta37:647
  • "Adephaga". Integrated Taxonomic Information System.
  • Adephaga Tree of Life
  1. ^ Wang, Yan-hui; Engel, Michael S.; Rafael, José A.; Wu, Hao-yang; Rédei, Dávid; Xie, Qiang; Wang, Gang; Liu, Xiao-guang; Bu, Wen-jun (2016). "Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda)". Scientific Reports. 6: 38939. doi:10.1038/srep38939. PMC 5154178. PMID 27958352.
Amphizoa

Amphizoa is a genus of aquatic beetles in the suborder Adephaga, placed in its own monogeneric family, Amphizoidae. There are five known species of Amphizoa, three in western North America and two in eastern palearctic. They are sometimes referred to by the common name troutstream beetles.

Aspidytidae

Aspidytidae is a family of beetles of the suborder Adephaga, first recorded in 2002 from specimens in South Africa and China.

Beetle

Beetles are a group of insects that form the order Coleoptera, in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 species, is the largest of all orders, constituting almost 40% of described insects and 25% of all known animal life-forms; new species are discovered frequently. The largest of all families, the Curculionidae (weevils) with some 83,000 member species,

belongs to this order. Found in almost every habitat except the sea and the polar regions, they interact with their ecosystems in several ways: beetles often feed on plants and fungi, break down animal and plant debris, and eat other invertebrates. Some species are serious agricultural pests, such as the Colorado potato beetle, while others such as Coccinellidae (ladybirds or ladybugs) eat aphids, scale insects, thrips, and other plant-sucking insects that damage crops.

Beetles typically have a particularly hard exoskeleton including the elytra, though some such as the rove beetles have very short elytra while blister beetles have softer elytra. The general anatomy of a beetle is quite uniform and typical of insects, although there are several examples of novelty, such as adaptations in water beetles which trap air bubbles under the elytra for use while diving. Beetles are endopterygotes, which means that they undergo complete metamorphosis, with a series of conspicuous and relatively abrupt changes in body structure between hatching and becoming adult after a relatively immobile pupal stage. Some, such as stag beetles, have a marked sexual dimorphism, the males possessing enormously enlarged mandibles which they use to fight other males. Many beetles are aposematic, with bright colours and patterns warning of their toxicity, while others are harmless Batesian mimics of such insects. Many beetles, including those that live in sandy places, have effective camouflage.

Beetles are prominent in human culture, from the sacred scarabs of ancient Egypt to beetlewing art and use as pets or fighting insects for entertainment and gambling. Many beetle groups are brightly and attractively coloured making them objects of collection and decorative displays. Over 300 species are used as food, mostly as larvae; species widely consumed include mealworms and rhinoceros beetle larvae. However, the major impact of beetles on human life is as agricultural, forestry, and horticultural pests. Serious pests include the boll weevil of cotton, the Colorado potato beetle, the coconut hispine beetle, and the mountain pine beetle. Most beetles, however, do not cause economic damage and many, such as the lady beetles and dung beetles are beneficial by helping to control insect pests.

Copelatinae

Copelatinae is a subfamily of diving beetles, in the family Dytiscidae. The subfamily contains seven genera:

Agaporomorphus, Aglymbus, Copelatus, Exocelina, Lacconectus, Liopterus, and Madaglymbus. Of these, the largest is Copelatus, which has about 470 described species found worldwide, but most diverse in tropical South America, Africa and South-East Asia.

Coptoclavella

Coptoclavella is an extinct genus of beetles in the family Coptoclavidae. There are about seven described species in Coptoclavella.

Coptoclavidae

Coptoclavidae is an extinct family of beetles in the suborder Adephaga. The Coptoclavidae lived from the Jurassic

to the Early Cretaceous.

Ground beetle

Ground beetles are a large, cosmopolitan family of beetles, Carabidae, with more than 40,000 species worldwide, around 2,000 of which are found in North America and 2,700 in Europe. It is one of the ten most speciose animal families, as of 2015.

Haliplidae

The Haliplidae are a family of water beetles who swim using an alternating motion of the legs. They are therefore clumsy in water (compared e.g. with the Dytiscidae or Hydrophilidae), and prefer to get around by crawling. The family consists of about 200 species in 5 genera, distributed wherever there is freshwater habitat; it is the only extant member of superfamily Haliploidea. They are also known as crawling water beetles or haliplids.

Hygrobia

Hygrobia is a genus of aquatic beetles native to Europe, North Africa, China and Australia. It is the only genus in the family Hygrobiidae, also known as the Paelobiidae. These are known commonly as squeak beetles or screech-beetles.There are six known living species, with a highly disjunct distribution, and one extinct species, Hygrobia cretzschmari.

Meru (beetle)

Meruidae is a recently described family of aquatic beetles in the suborder Adephaga, with only one genus and species, Meru phyllisae. This beetle species was first found in the early 1980s. At 0.8 mm, it is one of the smallest adephagan beetles in the world. A recent survey of aquatic beetles of Venezuela prove that Meru is most common during the wet season, when larger areas of granitic rock surface are covered with water film, which the adult beetles as well as the larvae inhabit.

Myxophaga

Myxophaga is the second smallest suborder of the Coleoptera after Archostemata, consisting of roughly 65 species of small to minute beetles in four families. The members of this suborder are aquatic and semiaquatic, and feed on algae.

Noteridae

Noteridae is a family of water beetles closely related to the Dytiscidae, and formerly classified with them. They are mainly distinguished by the presence of a distinctive "noterid platform" underneath, in the form of a plate between the second and third pair of legs. The family consists of about 230 species in 14 genera, and is found worldwide, more commonly in the tropics. They are sometimes referred to as burrowing water beetles.

These beetles are relatively small, ranging from 1 to 5 mm, with smooth oval bodies ranging from light brown to a darker reddish brown. The head is short and somewhat covered by the prothorax.

Both adults and larvae are aquatic, and are commonly found around plants. They have a habit of burrowing through pond and marsh substrate, thus the common name, and are primarily carnivorous, with some scavenging observed.

Noterus clavicornis

Noterus clavicornis is a species of beetle belonging to the family Noteridae.

Noterus crassicornis

Noterus crassicornis is a genus of beetle native to the Palearctic (including Europe) and the Near East. In Europe, it is only found in Austria, Belarus, Belgium, Bosnia and Herzegovina, Bulgaria, Croatia, Cyprus, the Czech Republic, mainland Denmark, Estonia, European Turkey, Finland, mainland France, Germany, Great Britain including the Isle of Man, Hungary, the Republic of Ireland, mainland Italy, Latvia, Lithuania, Luxembourg, Moldova, Northern Ireland, North Macedonia, mainland Norway, Poland, Russia, Sicily, Slovakia, Slovenia, Sweden, the Netherlands, Ukraine and Yugoslavia.

Rhysodidae

Rhysodidae (sometimes called wrinkled bark beetles) is a family of beetles, consisting of more than 350 species in about 20 genera.

These beetles are elongate, in size ranging from 5–8 mm, and color ranging from a reddish brown to black. Both the thorax and the elytra are deeply grooved lengthwise, thus giving these beetles their common name. The head is also grooved, and posteriorly constricted into a short but visible "neck". The 11-segment antennae are short, resembling a string of beads, while the mandibles lack cutting edges and are thus nonfunctional. The front legs are short and strongly built,

Adults and larvae live in moist rotten wood that is infested with slime moulds, which are believed to be their diet. Instead of using their mandibles to bite, they use the anterior edge of the mentum and swivel their heads to cut off pieces of food. Adults do not make burrows, instead just squeezing between the cell layers of the decomposed wood, generally leaving no visible trace of their passage, while larvae live in short tunnels.

They occur on all continents with forested areas, the richest fauna being found in New Guinea, Indonesia, the Philippines, and northern South America.

Classification remains controversial, with specialist Ross T. Bell arguing for placement as a tribe Rhysodini of Carabidae, while R. G. Beutel and others argue that larval characteristics indicate that rhysodids are their own family. Recent DNA analysis supports the placement of Rhysodidae within the family Carabidae.The following genera have generally been treated as members of the family Rhysodidae, but are now sometimes considered members of Carabidae instead.

Arrowina Bell & Bell, 1978 (Palearctic, Indomalaya)

Clinidium Kirby 1835

Dhysores Grouvelle 1903 (Africa)

Grouvellina Bell & Bell 1978 (Madagascar, Comoros)

Kaveinga Bell & Bell 1978 (Australasia)

Kupeus Bell & Bell 1982 (New Zealand)

Leoglymmius Bell & Bell, 1978 (Australia)

Medisores Bell & Bell, 1987 (South Africa)

Neodhysores Bell & Bell 1978 (South America)

Omoglymmius Ganglbauer, 1891

Plesioglymmius Bell & Bell, 1978

Rhysodes Germar 1822 (Paleac)

Rhyzodiastes Fairmaire 1895

Shyrodes Grouvelle, 1903 (Indomalaya)

Sloanoglymmius Bell & Bell, 1991 (Australia)

Srimara Bell & Bell, 1978 (Vietnam)

Tangarona Bell & Bell 1982 (New Zealand)

Xhosores Bell & Bell, 1978 (South Africa)

Yamatosa Bell & Bell, 1979

Suphisellus

Suphisellus is a genus of beetles in the family Noteridae, containing the following species:

Suphisellus anticicollis Guignot, 1950:153

Suphisellus balzani (Régimbart, 1889a:259) (Canthydrus balzani)

Suphisellus bicolor (Say, 1830:33) (Noterus bicolor)

Suphisellus binotatus (Fleutiaux & Sallé, 1890:370) (Canthydrus binotatus)

Suphisellus brevicornis (Sharp, 1882a:273) (Canthydrus brevicornis)

Suphisellus bruchi (Zimmermann, 1919:115) (Canthydrus bruchi)

Suphisellus brunneus Guignot, 1950:152

Suphisellus canthydroides Guignot, 1940:9

Suphisellus cribrosus (Régimbart, 1903:59) (Canthydrus cribrosus)

Suphisellus curtus (Sharp, 1882a:272) (Canthydrus curtus)

Suphisellus dilutus (Sharp, 1882a:272) (Canthydrus dilutus)

Suphisellus epleri Arce-Pérez & Baca, 2017:278

Suphisellus flavolineatus (Régimbart, 1889a:262) (Canthydrus flavolineatus)

Suphisellus flavopictus (Régimbart, 1889a:260) (Canthydrus flavopictus)

Suphisellus gibbulus (Aubé, 1838:414) (Suphis gibbulus)

Suphisellus globosus (Régimbart, 1903:62) (Canthydrus globosus)

Suphisellus grammicus (Sharp, 1882a:274) (Canthydrus grammicus)

Suphisellus grammopterus (Régimbart, 1889c:390) (Canthydrus grammopterus)

Suphisellus grossus (Sharp, 1882a:270) (Canthydrus grossus)

Suphisellus hieroglyphicus Zimmermann, 1921:187

Suphisellus insularis (Sharp, 1882a:270) (Canthydrus insularis)

Suphisellus levis (Fall, 1909:99) (Canthydrus levis)

Suphisellus lineatus (Horn, 1871:329) (Suphis lineatus)

Suphisellus majusculus (Sharp, 1882b:6) (Canthydrus majusculus)

Suphisellus melzeri Zimmermann, 1925:254

Suphisellus minimus Gschwendtner, 1922:135

Suphisellus neglectus Young, 1979:419

Suphisellus nigrinus (Aubé, 1838:411) (Hydrocanthus nigrinus)

Suphisellus obesus (Régimbart, 1903:59) (Canthydrus obesus)

Suphisellus obscuripennis (Régimbart, 1889a:257) (Canthydrus obscuripennis)

Suphisellus ovatus (Sharp, 1882a:270) (Canthydrus ovatus)

Suphisellus parsonsi Young, 1952:157

Suphisellus penthimus Guignot, 1957:41

Suphisellus pereirai Guignot, 1958:37

Suphisellus phenax Guignot, 1954:198

Suphisellus pinguiculus (Régimbart, 1903:62) (Canthydrus pinguiculus)

Suphisellus puncticollis (Crotch, 1873:397) (Suphis puncticollis)

Suphisellus remator (Sharp, 1882a: 272) (Canthydrus remator)

Suphisellus rotundatus (Sharp, 1882a:270) (Canthydrus rotundatus)

Suphisellus rubripes (Boheman, 1858:19) (Hydrocanthus rubripes)

Suphisellus rufulus (Zimmermann, 1921:188)

Suphisellus sculpturatus (Sharp, 1882a:269) (Canthydrus sculpturatus)

Suphisellus sexnotatus (Régimbart, 1889a:259) (Canthydrus sexnotatus)

Suphisellus similis Zimmermann, 1921:188

Suphisellus simoni (Régimbart, 1889b:383) (Canthydrus simoni)

Suphisellus subsignatus (Sharp, 1882a:271) (Canthydrus subsignatus)

Suphisellus tenuicornis (Chevrolat, 1863:199) (Hydrocanthus tenuicornis)

Suphisellus transversus (Régimbart, 1903:61) (Canthydrus transversus)

Suphisellus vacuifer Guignot, 1958:37

Suphisellus varians (Sharp, 1882b:5) (Canthydrus varians)

Suphisellus variicollis Zimmermann, 1921:187

Suphisellus vicinus (Sharp, 1882a:2699) (Canthydrus vicinus)

Trachypachidae

The Trachypachidae (or sometimes false ground beetles) are a family of beetles that generally resemble small ground beetles, but that are distinguished by the large coxae of their rearmost legs. There are only six known extant species in the family, with four species of Trachypachus found in northern Eurasia and northern North America, and two species of Systolosoma in Chile. They were much more diverse in the past, with many members belonging to the extinct subfamily Eodromeinae, the first fossils known of this family are of the genera Petrodromeus and Permunda from the Permian-Triassic boundary of Russia.Their habits are similar to those of the ground beetles; they are usually found in the leaf litter of conifer forests.

To quote G. E. Ball, "the most interesting thing about this small family is its uncertain phylogenetic relationships"; it has been classified as a subfamily of Carabidae, and as a family been placed in several different positions relative to other beetle families.

Walther Horn

Walther Hermann Richard Horn (19 October 1871 – 10 July 1939) was a German entomologist who specialised in beetles (Coleoptera). He was born in Berlin, where he also died. He is not to be confused with the American entomologist George Henry Horn, another entomologist that studied Coleoptera.

Walther Horn was first a physician then Director of the German Entomological Institute.

Whirligig beetle

The whirligig beetles are a family (Gyrinidae) of water beetles that usually swim on the surface of the water if undisturbed, though they swim underwater when threatened. They get their common name from their habit of swimming rapidly in circles when alarmed, and are also notable for their divided eyes which are believed to enable them to see both above and below water. The family includes some 700 extant species worldwide, in 15 genera, plus a few fossil species. Most species are very similar in general appearance, though they vary in size from perhaps 3 mm to 18 mm in length. They tend to be flattened and rounded in cross section, in plain view as seen from above, and in longitudinal section. In fact their shape is a good first approximation to an ellipsoid, with legs and other appendages fitting closely into a streamlined surface.

Extant Coleoptera families

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