The Permian stem-beetle Sylvacoleus sharovi, from Grimaldi & Engel (2005).

Belongs within: Coleopterida.
Contains: Adephaga, Archostemata, Myxophaga, Scirtoidea, Staphylinoidea, Hydrophiloidea, Scarabaeoidea, Elateriformia, Bostrichoidea, Coccinelloidea, Tenebrionoidea, Cleroidea, Cucujoidea, Phytophaga.

The Coleoptera, beetles, are one of the most diverse groups of insects with over 300,000 species. The earliest beetles were probably inhabitants of subcortical and interstitial spaces in wood (Kirejtshuk & Nel 2013) but they have since diversified to a much wider range of habitats. Beetles are characterised by the transformation of the forewings into hardened elytra with the forewing venation entirely lost in modern Coleoptera. The lateral edge of the elytron often carries a distinct incurved section called the epipleuron, separated from the dorsal surface by a sharp ridge or carina. A median scutellum may remain visible between the elytral bases when closed. The elytra are often marked by longitudinal rows of punctures and/or impressed grooves called striae. In such cases, the spaces between puncture rows or striae are described as intervals. A short stria or puncture row may be present alongside the scutellum, dubbed the scutellary striole. The functional hind wings, if present, are usually folded beneath the elytra when not in use. To allow such folding, wing venation is highly modified in comparison to other insect orders with veins being partially de-sclerotised (Lawrence & Britton 1991).

Beetles also have a distinctly shaped pronotum, with the dorsal surface flattened and more or less separated from the sloping sides by lateral carinae (though these have been reduced or lost in many derived subgroups; Kirejtshuk & Nel 2013). A well developed articulation between prothorax and mesothorax allows the former to move independently of the rest of the thorax. The mesothorax and metathorax are fused to form the pterothorax, which broadly contacts the abdomen. The cavities for the mesocoxae may be surrounded by the meso- and metasternum (in which case they are described as ‘laterally closed’) or they may come into contact laterally with the pleural sclerites (‘laterally open’). Hind coxae are typically transverse and of limited or no mobility against the metasternum.

The head is primitively prognathous with some more derived subgroups becoming hypognathous. Ocelli are commonly absent, one or two being present in select subgroups. Antennae usually have eleven segments or less with a distinct scape and pedicel. Antennal segments may vary significantly in form between taxa, such as individual segments being serrate or pectinate (more or less extended to one side so the antenna has a saw- or comb-like appearance), or terminal segments being enlarged to form a club.

The abdomen comprises nine segments in females and ten in males, with the ninth segment being modified into the genital segment and the male tenth often being reduced or fused to the ninth (Lawrence & Britton 1991). Those tergites covered by the elytra at rest are weakly sclerotised. Male genitalia comprise an aedeagus divided between a basal tegmen and distal penis. The gonopore emerges through an internal sac that is usually membranous but may bear sclerotised spicules. The ovipositor has a pair of appendages attached ventrally to the paraprocts, with each appendage divided between a basal gonocoxite and distal gonostylus (Lawrence & Britton 1991).

In the Tshekardocoleidae and other stem-Coleoptera families, the forewings retained at least some of their original venation (the forewings of Tshekardocoleidae were also probably leathery rather than entirely sclerotised) and some authors have recognised the Tshekardocoleidae as a separate order Protocoleoptera. The spaces between the veins and crossveins were occupied by a system of dense, square cells that at least superficially resemble the series of square punctures found in some basal crown-beetle families (Grimaldi & Engel 2005). The Early Permian Tshekardocoleidae retained the full complement of veins and possessed very broad costal and subcostal areas on the elytra (Kirejtshuk & Nel 2013). They also differ from later beetles in having antennae with 13 segments (versus 11 or less) and small procoxae with widely separated bases (versus closely placed procoxae) (Grimaldi & Engel 2005). A short ovipositor is known to have been present in at least on genus, Moravocoleus; larvae may have been woodborers (Grimaldi & Engel 2005). Members of the genera Tshekardocoleus and Sylvacoleus lack a common stem, or have at most a very short stem, to veins M and CuA; this stem is present and elongate in Moravocoleus. Vein Rs has two posterior branches in Tshekardocoleus but is unbranched in Sylvacoleus.

More derived families such as Permocupedidae and Taldycupedidae with reduced venation, and usually with reduced costal areas, appeared in the Late Permian. The Permocupedidae had most veins distinguishable but lacked vein Rs; veins M and CuA arose independently. Members of the Asiocoleidae have vein M reduced but other veins remain elongate. Taldycupes reticulatus was a small, elongate and flattened beetle with a prognathous head and small paranotals on the prothorax. In the Schizocoleidae, the veins are either distinct or replaced by punctate grooves, often not distinct from interstitial veins, and there are four to six rows of cells in the anterior of the elytra. The Tricoleidae are known from the Triassic and Jurassic, and possessed three main veins on the elytra. Unfortunately, interpretation of the later beetle fossil record is often hindered by the loss of veins resulting in elytra preserving few diagnostic features (Grimaldi & Engel 2005).

Living species of Coleoptera are divided between the large subgroups Adephaga and Polyphaga, and the two smaller taxa Archostemata and Myxophaga. Relationships between these subgroups have been subject to some contention; the Archostemata (which retain the potentially primitive wood-boring habit) are commonly regarded as basal to the other lineages but a recent molecular analysis by McKenna et al. (2015) supported a sister relationship between the Polyphaga and all other beetles. Potential synapomorphies of an Adephaga-Archostemata-Myxophaga clade include wing-folding type, lack of cervical sclerites, and the absence of a basal piece on the aedeagus (Lawrence et al. 1987). The Polyphaga are by far the largest of the major coleopteran subgroups and are characterised by the absence of the notopleural sutures at the sides of the prothorax. The Ademosynidae, known from the Triassic of Australia and South America, have been suggested to be stem-group Polyphaga (Grimaldi & Engel 2005). Ademosynids were small oval or elongate-oval beetles with costate elytra lacking remnant veins.

Polyphagans are first represented in the fossil record by the Triassic Peltosyne triassica from Central Asia (Lawrence & Britton 1991). The Jurassic Larvula cassa may represent an early aquatic polyphagan larva, but its affinities are uncertain (Sinitshenkova 2002). Tomocoleites longigastrulatus was described from Chinese Eocene amber by Hong (2002) as a likely polyphagan, albeit one that cannot be easily linked to any living subgroup. The Polyphaga can mostly be divided between the major clades Staphyliniformia, Elateriformia, Bostrichoidea and Cucujiformia, with a smaller clade Scirtoidea recently supported as the sister clade to other living Polyphaga by McKenna et al. (2015). The Staphyliniformia, usually defined as including Staphylinoidea and Hydrophiloidea, should probably also include the Scarabaeoidea. Members of this expanded clade have six or seven visible abdominal ventrites with the anteriormost ventrite divided by the metathorax so it is present as two lateral triangular pieces. McKenna et al.‘s (2015) molecular analysis also supported a sister-group relationship between the Bostrichoidea and Cucujiformia. Such a relationship had previously been suggested on the basis of the presence in members of these groups of cryptonephridial Malpighian tubules, in which the ends of the tubules are attached to the rectum (Lawrence & Britton 1991). This arrangement may function in water retention, allowing the tubules to resorb water from the beetle’s waste. Synapomorphies of the Cucujiformia include the absence of plates on the hind coxae, absence of functional spiracles on abdominal segment 8, reduction of the pregenital segments 9 and 10 (Lawrence & Britton 1991). The Jurassic Parandrexidae are an extinct endophytic family of cucujiforms (Zherikhin 2002c).

Characters (from Grimaldi & Engel 2005): Ocelli often absent. Forewings heavily sclerotised, transformed into elytra, with venation reduced (absent in all Recent taxa); metathorax reduced; hind wings folded lengthwise and crosswise to be tucked under elytra when at rest, with reduced venation. Prothorax freely articulating with pterothorax, always large and shieldlike. Abdomen with sternites heavily sclerotised, tergites usually less sclerotised; cerci absent.

Coleoptera (see below for synonymy)
|--Tshekardocoleidae [Protocoleoptera]MW15
| |--UralocoleusP02
| |--Tshekardocoleus magnusP02
| |--Moravocoleus perditus Kukalová 1969KN13
| |--Votocoleus submissusK-P91
| `--SylvacoleusP02
| |--S. richteriP02
| `--S. sharoviGE05
| `--+--ArchostemataMW15
| `--MyxophagaMW15
`--+--Ademosyne [Ademosynidae, Ademosynoidea]GE05
| |--A. speciosaK-P91
| `--A. wianamattensis Tillyard 1917F71
`--Polyphaga (see below for synonymy)MW15
| i. s.: Peltosyne triassicaLB91
| Larvula cassaS02
| Heliotis hopei (n. d.)L09
| Tomocoleites Hong 2002H02
| `--*T. longigastrulatus Hong 2002H02
| Lobetus Kiesw. 1852FS90
| |--L. abdominalisFS90
| `--L. guadeloupensis Fleutiaux & Sallé 1890FS90
`--+--Staphyliniformia [Haplogastra, Hydrophilii, Hydrophilomorpha, Staphylinomorpha]MW15
| |--StaphylinoideaMW15
| `--+--HydrophiloideaMW15
| `--ScarabaeoideaMW15
`--Cucujiformia (see below for synonymy)MW15
| i. s.: ParandrexidaeZ02c
| `--CleroideaRS15

Coleoptera incertae sedis:
Homalaena dispersaF02
Tymbopiptus valeas Kuschel 1987LR02
Geodorcus capitoE02
Metabuprestium oyunchaienseZ02a
Palaeosilpha fraasiZ02c
Ryugadous ishikawaiS86
Kusumia takahashiiS86
Sospita vigintiguttataD37
|--L. atripennisI92
|--L. miyako (Nakane 1981)I92
`--L. rufaI92
Apotomopterus maackiI92
|--A. m. maackiI92
`--A. m. aquatilis (Bates 1883)I92
Thyestilla gebleri (Faldermann 1835)I92
Chaetotrechiama procerus Uéno 1982I92
|--R. elegans Uéno 1960I92
|--R. lallum Uéno 1970I92
`--R. mirabilis Uéno 1958I92
Awatrechus hygrobius Uéno 1958I92
Leptelmis gracilis Sharp 1888I92
Ishikawatrechus intermedius Uéno 1957I92
Eusclerus Sharp 1886C92
Prionomma White 1853C92
Spelaeobates pharensis Müller 1901UO05
Crypta Stephens 1830BR05
Cylindra Illiger 1802BR05
‘Limnorea’ Agassiz 1846 non Goldfuss 1826BR05
Pseudomesalia Ganglbauer 1900BR05
Selenites Hope 1840BR05
Permocupoides sojanensisGE05
Notocupoides triassicusGE05
Lissodes Berthold 1827K02
Podischnus agenorR96
Kubousia axillarisMG06
Pityophthoridea Wickham 1916W86
Adipocephalus Wickham 1916W86
‘Macropus’ Thunberg 1805 nec Shaw 1790 nec Latreille 1802S61
Boldonia robiatiV73
Platyrrhinus Fabricius 1801G69
Crawfordia Pierce 1908PH90
Eusomus affinis Lucas 1846E12
|--D. africanus Putz 1846E12
|--D. algiricus Putz 1846E12
|--D. numidicus Putz 1846E12
`--D. obsoletus Putz 1846E12
Myrmecobius Lucas 1846E12
`--M. agilis Lucas 1846E12
|--N. albomarginatus Lucas 1847E12
`--N. albopunctatus Lucas 1847E12
Hypophaeus angustus Lucas 1846E12
Brachyphylla barbara Lucas 1846E12
Cataphronetis Lucas 1846E12
`--C. levaillanti Lucas 1846E12
|--P. costatipennis Lucas 1847E12
|--P. moreletii Lucas 1847E12
|--P. plicatus Lucas 1847E12
`--P. variolosus Lucas 1847E12
|--O. distinctus Lucas 1846E12
`--O. tingitanus Lucas 1846E12
Opilus dorsalis Lucas 1846E12
|--P. erythrocnema Lucas 1847E12
|--P. malachitica Lucas 1847E12
|--P. rubricollis Lucas 1847E12
|--P. ventralisF89a
|--P. virgulaF89a
`--P. warnieri Lucas 1847E12
Heterophaga Lucas 1847E12
Singilis mauritanica Lucas 1846E12
Pachypterus Lucas 1847E12
`--P. mauritanicus Lucas 1847E12
|--T. puncticollis Lucas 1846E12
`--T. rotundatus Lucas 1846E12
Carterus rufipes Lucas 1846E12
Otophorus scolytoides Lucas 1846E12
Sybines sellatus Lucas 1846E12
Eutrapela suturalis Lucas 1847E12
Mazoreus testaceus Lucas 1846E12
Trypocladius Lucas 1847E12
Zarax Pascoe 1867KA-Z11
Tetracyphus Chevrolat 1881KA-Z11
Metapachylus Bates 1889KA-Z11
Prionognathus Ferté-Sénèctere 1851P61, H62
Anisoceras Dejean 1833P61
Macronemus mimusB14
Pelops Gistl 1834CW92a
Trionychus Burmeister 1847CW92b
|--Permocupes sojanensisP02
|--Kaltanicupes ponomarenkoi Pinto 1987RJ93
`--Afrocupes firmaeEB99
Asiocoleidae [Oborocoleidae]GE05
|--Oborocoleus rohdendorfi Kukalová 1969KN13
`--Tetracoleus Ponomarenko 2009KN13
Taldycupedidae [Taldycupidae]GE05
|--Taldycupes reticulatusP02
`--Yuxianocoleus hebeiense Hong 1985RJ93
|--Permocrossos elongatusR70
|--P. affinis Tillyard 1924F71
|--P. belmontensis Tillyard 1924F71
|--P. mitchelli Tillyard 1924F71
`--P. pincombeae Tillyard 1924F71
Callopus marginatus Bosc 1802B02
Cylichnus Burmeister 1844 [=Cylichna (l. c.) non Lovén 1846]BR17
Eriocampa ovataF92
Jeannelia Raffray 1913B61
Praelateridae [Praelateriidae]RJ93
Siron Champion 1905K18
Praepodes vittatusB28
Minotaurus typhoeusR13
Bolbites onitoidesR13
Lebidia octoguttataF89a
Dolichus flavicornisF89a
Pantheropterus davidis Fairmaire 1889F89a
|--G. obscuratus Fairmaire 1889F89a
`--G. orientalisF89a
|--E. davidis Fairmaire 1889F89a
|--E. pictipes Fairmaire 1889F89a
`--E. variolosaF89a
Stenonota Fairmaire 1889F89a
`--*S. semirugata Fairmaire 1889F89a
|--‘Callynomes’ davidisF89a
`--C. discipennis Fairmaire 1889F89a
|--D. longicornisF89a
`--D. maculosus Fairmaire 1889F89a
Pyrocaelia moupinensis Fairmaire 1889F89a
Telephorops impressipennisF89a
Tagonoides Fairm. 1886 [incl. Gnaptorina Reitter 1887]F89a
`--‘Gnaptorina’ felicitana Reitter 1887F89a
Itagonia Reitter 1887F89a
Asidoblaps Fairm. 1886F89a
Plesiophthalmus pallidicrus Fairmaire 1889F89a
|--A. fulvicollisF89a
`--A. strangulata Fairmaire 1889F89a
Astycus chinensis Faust in Fairmaire 1889F89a
|--P. hypomelasF89a
`--P. quinquesignatus Fairmaire 1889F89a
Talmonus Fairmaire 1889F89a
`--*T. farinosus Fairmaire 1889F89a
Laphris emarginataF89a
|--E. annulipennis Fairmaire 1889F89a
|--E. limbataF89a
|--E. nigrofoveolata Fairmaire 1889F89a
`--E. nigropunctata Fairmaire 1889F89a
Tebalia Fairmaire 1889F89a
`--*T. caeruleata Fairmaire 1889F89a
Lobobrachus lacerdaeL90
Praimus pictusL90
Metopocaelus maculicornisL90
Ctenoscolis coecusL90
|--R. substriatus Grouvelle 1889G89
`--R. villosocostatus [=Limnius villosocostatus]G89
|--M. coyei [=Helmis coyei]G89
`--M. rioloides [=Macronychus rioloides]G89
Dupophilus brevis [incl. Latelmis insignis]G89
|--L. damryi [=Limnius damryi]G89
|--L. intermedia [=Limnius intermedius]G89
`--L. sulcipennis [=Limnius sulcipennis]G89
Cladodes Solier 1849O89
Issacaris Fairmaire 1889F89b
`--*I. petalophora Fairmaire 1889F89b
|--D. luctuosa Desbrochers des Loges 1890DL90
`--D. plumbea Desbrochers des Loges 1890DL90
Amauronia longula Desbrochers des Loges 1890DL90
|--A. algirica Fairmaire 1889 [=A. reitteri Desbrochers des Loges 1889 non Ormay 1888]F90
`--A. reitteri Ormay 1888F90
Langelandia reitteriDL90
Canidia Thomson 1857KF22
Nyctaea Berthold 1827V99
Phyllophorus gigas [=Elater gigas]H42
Piezophyllus Hope 1842H42
|--P. shuckhardi Hope 1842 [incl. Tetralobus dumolinii ms]H42
`--P. spencei Hope 1842H42
Leptophyllus Hope 1842H42
`--*L. strachani Hope 1842H42
Tetrophthalmus Kirby 1827PB96

Coleoptera [Archecoleoptera, Asiocoleoidea, Cupedina, Cupedomorpha, Eleutherata, Malacodermi, Pantophaga, Permocupedoidea, Scarabaeida, Scarabaeidea, Schizophoromorpha]

Cucujiformia [Cucujomorpha, Erotylenae, Nitidulariae, Pseudotetramera, Trogossitarii, Xylophagi]MW15

Polyphaga [Bostrichiformia, Clavicornes, Clavicornia, Dermestoidea, Diaperialae, Heterolytra, Heterophaga, Malacodermata, Malacodermidae, Microsomata, Necrophagi, Saprophaga, Scarabaeina, Scarabaeomorpha, Serricornia, Symphiogastra, Xylophaga]MW15

*Type species of generic name indicated


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