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

Belongs within: Coleopterida.
Contains: Rhipidoceridae, 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 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. 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.

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.

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).

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
    |    |--Uralocoleus P02
    |    |--Tshekardocoleus magnus P02
    |    |--Moravocoleus perditus Kukalová 1969 KN13
    |    |--Votocoleus submissus K-P91
    |    `--Sylvacoleus P02
    |         |--S. richteri P02
    |         `--S. sharovi GE05
    `--+--+--Adephaga MW15
       |  `--+--Archostemata MW15
       |     `--Myxophaga MW15
       `--+--Ademosyne [Ademosynidae, Ademosynoidea] GE05
          |    |--A. speciosa K-P91
          |    `--A. wianamattensis Tillyard 1917 F71
          `--Polyphaga (see below for synonymy) MW15
               |  i. s.: Peltosyne triassica LB91
               |         Larvula cassa S02
               |         Heliotis hopei (n. d.) L09
               |         Tomocoleites Hong 2002 H02
               |           `--*T. longigastrulatus Hong 2002 H02
               |         Lobetus Kiesw. 1852 FS90
               |           |--L. abdominalis FS90
               |           `--L. guadeloupensis Fleutiaux & Sallé 1890 FS90
               |--Scirtoidea MW15
               `--+--Staphyliniformia [Haplogastra, Hydrophilii, Hydrophilomorpha, Staphylinomorpha] MW15
                  |    |--Staphylinoidea MW15
                  |    `--+--Hydrophiloidea MW15
                  |       `--Scarabaeoidea MW15
                  `--+--Elateriformia MW15
                     `--+--Bostrichoidea MW15
                        `--Cucujiformia (see below for synonymy) MW15
                             |  i. s.: Parandrexidae Z02c
                             |--Coccinelloidea MW15
                             `--+--Tenebrionoidea MW15
                                `--+--Cleroidea MW15
                                   `--+--Cucujoidea MW15
                                      `--Phytophaga MW15

Coleoptera incertae sedis:
  Homalaena dispersa F02
  Lagochile L01
  Tymbopiptus valeas Kuschel 1987 LR02
  Geodorcus capito E02
  Metabuprestium oyunchaiense Z02a
  Betulapion simile Z02a
  Rhyncholus kathrynae Z02a
  Megathopa Z02b
  Antliarrhinites Z02c
  Palaeosilpha fraasi Z02c
  Donaciella nagaokana S02
  Ryugadous ishikawai S86
  Kusumia takahashii S86
  Oxysoma A71
  Barypeithes araneiformis FFB88
  Sospita vigintiguttata D37
  Lychnuris I92
    |--L. atripennis I92
    |--L. miyako (Nakane 1981) I92
    `--L. rufa I92
  Apotomopterus maacki I92
    |--A. m. maacki I92
    `--A. m. aquatilis (Bates 1883) I92
  Thyestilla gebleri (Faldermann 1835) I92
  Chaetotrechiama procerus Uéno 1982 I92
  Rakantrechus I92
    |--R. elegans Uéno 1960 I92
    |--R. lallum Uéno 1970 I92
    `--R. mirabilis Uéno 1958 I92
  Awatrechus hygrobius Uéno 1958 I92
  Leptelmis gracilis Sharp 1888 I92
  Macroplea japana (Jacoby 1885) I92
  Ishikawatrechus intermedius Uéno 1957 I92
  Ceratoderus venustus Hisamatsu 1963 I92
  Eusclerus Sharp 1886 C92
  Prionomma White 1853 C92
  Limnozenus zealandicus TDC06
  Spelaeobates pharensis Müller 1901 UO05
  Crypta Stephens 1830 BR05
  Cylindra Illiger 1802 BR05
  ‘Limnorea’ Agassiz 1846 non Goldfuss 1826 BR05
  Pseudomesalia Ganglbauer 1900 BR05
  Selenites Hope 1840 BR05
  Permocupoides sojanensis GE05
  Notocupoides triassicus GE05
  Lissodes Berthold 1827 K02
  Dendrobias mandibularis R96
  Podischnus agenor R96
  Kubousia axillaris MG06
  Pityophthoridea Wickham 1916 W86
  Adipocephalus Wickham 1916 W86
  Pentaplatarthrus paussoides A61
  ‘Macropus’ Thunberg 1805 nec Shaw 1790 nec Latreille 1802 S61
  Coleomegilla maculata MB93
  Boldonia robiati V73
  Platyrrhinus Fabricius 1801 G69
  Crawfordia Pierce 1908 PH90
  Eusomus affinis Lucas 1846 E12
  Dischirius E12
    |--D. africanus Putz 1846 E12
    |--D. algiricus Putz 1846 E12
    |--D. numidicus Putz 1846 E12
    `--D. obsoletus Putz 1846 E12
  Myrmecobius Lucas 1846 E12
    `--M. agilis Lucas 1846 E12
  Nastus E12
    |--N. albomarginatus Lucas 1847 E12
    `--N. albopunctatus Lucas 1847 E12
  Parnus algiricus Lucas 1846 E12
  Hypophaeus angustus Lucas 1846 E12
  Brachyphylla barbara Lucas 1846 E12
  Anelastes barbarus Lucas 1846 E12
  Cataphronetis Lucas 1846 E12
    `--C. levaillanti Lucas 1846 E12
  Cerandria Lucas 1847 E12
  Philax E12
    |--P. costatipennis Lucas 1847 E12
    |--P. moreletii Lucas 1847 E12
    |--P. plicatus Lucas 1847 E12
    `--P. variolosus Lucas 1847 E12
  Ditomus dilaticollis Lucas 1846 E12
  Omazeus E12
    |--O. distinctus Lucas 1846 E12
    `--O. tingitanus Lucas 1846 E12
  Opilus dorsalis Lucas 1846 E12
  Acinopus E12
    |--A. elongatus Lucas 1846 E12
    |--A. lepeletieri Lucas 1846 E12
    `--A. mauritanicus Lucas 1846 E12
  Phytaecia E12
    |--P. erythrocnema Lucas 1847 E12
    |--P. malachitica Lucas 1847 E12
    |--P. rubricollis Lucas 1847 E12
    |--P. ventralis F89a
    |--P. virgula F89a
    `--P. warnieri Lucas 1847 E12
  Heterophaga Lucas 1847 E12
  Singilis mauritanica Lucas 1846 E12
  Pachypterus Lucas 1847 E12
    `--P. mauritanicus Lucas 1847 E12
  Colaspidea nitida Lucas 1846 E12
  Colaspidema E12
    |--C. pulchella Lucas 1846 E12
    `--C. signatipennis Lucas 1846 E12
  Thorectes E12
    |--T. puncticollis Lucas 1846 E12
    `--T. rotundatus Lucas 1846 E12
  Pachychila E12
    |--P. punctulata Lucas 1846 E12
    `--P. sabulosa Lucas 1846 E12
  Carterus rufipes Lucas 1846 E12
  Otophorus scolytoides Lucas 1846 E12
  Sybines sellatus Lucas 1846 E12
  Eutrapela suturalis Lucas 1847 E12
  Mazoreus testaceus Lucas 1846 E12
  Trypocladius Lucas 1847 E12
  Rhyzophagus unicolor Lucas 1846 E12
  Zarax Pascoe 1867 KA-Z11
  Tetracyphus Chevrolat 1881 KA-Z11
  Metapachylus Bates 1889 KA-Z11
  Labidomera clivicollis WP99
  Oulema melanopus G84
  Prionognathus Ferté-Sénèctere 1851 P61, H62
  Anisoceras Dejean 1833 P61
  Macronemus mimus B14
  Schizax senex B14
  Pelops Gistl 1834 CW92a
  Trionychus Burmeister 1847 CW92b
  Schizocoleidae GE05
  Rhombocoleidae GE05
  Permocupedidae GE05
    |--Permocupes sojanensis P02
    |--Kaltanicupes ponomarenkoi Pinto 1987 RJ93
    `--Afrocupes firmae EB99
  Asiocoleidae [Oborocoleidae] GE05
    |--Oborocoleus rohdendorfi Kukalová 1969 KN13
    `--Tetracoleus Ponomarenko 2009 KN13
  Tricoleidae GE05
  Taldycupedidae [Taldycupidae] GE05
    |--Taldycupes reticulatus P02
    `--Yuxianocoleus hebeiense Hong 1985 RJ93
  Rhipidoceridae M86
  Eros scutellaris Erichs. 1842 [incl. E. bremei Le Guillou 1844] M86
  Archicupes jacobsoni R70
  Permocrossos elongatus R70
  Permosyne R70
    |--P. affinis Tillyard 1924 F71
    |--P. belmontensis Tillyard 1924 F71
    |--P. mitchelli Tillyard 1924 F71
    `--P. pincombeae Tillyard 1924 F71
  Callopus marginatus Bosc 1802 B02
  Cylichnus Burmeister 1844 [=Cylichna (l. c.) non Lovén 1846] BR17
  Atopa Paykull 1799 BR17
  Faula Blanchard 1850 BR17
  Eriocampa ovata F92
  Jeannelia Raffray 1913 B61
  Labradorocoleidae RJ93
  Praelateridae [Praelateriidae] RJ93
  Siron Champion 1905 K18
  Praepodes vittatus B28
  Sitodrepa panicea R13
  Minotaurus typhoeus R13
  Phosphaenus hemipterus R13
  Anoxia R13
    |--A. australis B28
    |--A. matutinalis B28
    `--A. villosa B28
  Anthypna abdominalis R13
  Bolbites onitoides R13
  Rhytidosomus globulus H44
  Parnidae W00
    |--Potamophilus Germ. 1811 FS90
    |    `--P. caraibus Coq. 1851 FS90
    |--Pelonomus Er. 1847 FS90
    |    `--P. picipes Ol. 1791 FS90
    `--Parygrus W00
         |--P. hardwicki W00
         |--P. talpoides W00
         `--P. whiteheadi Waterhouse 1900 W00
  Lebidia octoguttata F89a
  Dolichus flavicornis F89a
  Librodor F89a
    |--L. forcipatus Fairmaire 1889 F89a
    `--L. japonicus F89a
  Pantheropterus davidis Fairmaire 1889 F89a
  Oxyomus striatocrenatus Fairmaire 1889 F89a
  Geotrypes F89a
    |--G. obscuratus Fairmaire 1889 F89a
    `--G. orientalis F89a
  Ectinoplia F89a
    |--E. davidis Fairmaire 1889 F89a
    |--E. pictipes Fairmaire 1889 F89a
    `--E. variolosa F89a
  Callistethus F89a
    |--C. iris F89a
    `--C. seminitidus Fairmaire 1889 F89a
  Stenonota Fairmaire 1889 F89a
    `--*S. semirugata Fairmaire 1889 F89a
  Macroma ochreipennis F89a
  Clinterocera F89a
    |--‘Callynomes’ davidis F89a
    `--C. discipennis Fairmaire 1889 F89a
  Campylus davidis Fairmaire 1889 F89a
  Dascyllus F89a
    |--D. longicornis F89a
    `--D. maculosus Fairmaire 1889 F89a
  Pyrocaelia moupinensis Fairmaire 1889 F89a
  Podabrus F89a
    |--P. aenescens Fairmaire 1889 F89a
    |--P. dimidiaticrus Fairmaire 1889 F89a
    `--P. semifumatus Fairmaire 1889 F89a
  Telephorops impressipennis F89a
  Tagonoides Fairm. 1886 [incl. Gnaptorina Reitter 1887] F89a
    `--‘Gnaptorina’ felicitana Reitter 1887 F89a
  Itagonia Reitter 1887 F89a
  Asidoblaps Fairm. 1886 F89a
  Plesiophthalmus pallidicrus Fairmaire 1889 F89a
  Xanthochroa F89a
    |--X. carniolica F89a
    |--X. fulvicrus Fairmaire 1889 F89a
    `--X. metallipennis Fairmaire 1889 F89a
  Anoncodes F89a
    |--A. fulvicollis F89a
    `--A. strangulata Fairmaire 1889 F89a
  Astycus chinensis Faust in Fairmaire 1889 F89a
  Stenura F89a
    |--S. aeneipennis Fairmaire 1889 F89a
    |--S. arcifera (Blanch. 1871) [=Strangalia arcifera] F89a
    |--S. basiplicata Fairmaire 1889 F89a
    |--S. lineigera Fairmaire 1889 F89a
    |--S. quadrifasciata F89a
    |--S. stricticollis Fairmaire 1889 F89a
    |--S. thibetana Blanch. 1871 F89a
    `--S. zonifera Blanch. 1871 F89a
  Pyresthes F89a
    |--P. hypomelas F89a
    `--P. quinquesignatus Fairmaire 1889 F89a
  Talmonus Fairmaire 1889 F89a
    `--*T. farinosus Fairmaire 1889 F89a
  Bromius vitis F89a
  Gonioctena quadriplagiata Fairmaire 1889 F89a
  Laphris emarginata F89a
  Eustetha F89a
    |--E. annulipennis Fairmaire 1889 F89a
    |--E. limbata F89a
    |--E. nigrofoveolata Fairmaire 1889 F89a
    `--E. nigropunctata Fairmaire 1889 F89a
  Tebalia Fairmaire 1889 F89a
    `--*T. caeruleata Fairmaire 1889 F89a
  Lobobrachus lacerdae L90
  Cyphus schoenherri L90
  Praimus pictus L90
  Metopocaelus maculicornis L90
  Ctenoscolis coecus L90
  Psammobius sabulosus [incl. Pleurophorus ovipennis] B89
  Riolus G89
    |--R. substriatus Grouvelle 1889 G89
    `--R. villosocostatus [=Limnius villosocostatus] G89
  Microdes G89
    |--M. coyei [=Helmis coyei] G89
    `--M. rioloides [=Macronychus rioloides] G89
  Dupophilus brevis [incl. Latelmis insignis] G89
  Latelmis G89
    |--L. damryi [=Limnius damryi] G89
    |--L. intermedia [=Limnius intermedius] G89
    `--L. sulcipennis [=Limnius sulcipennis] G89
  Cladodes Solier 1849 O89
  Issacaris Fairmaire 1889 F89b
    `--*I. petalophora Fairmaire 1889 F89b
  Dicrania aeneobrunnea F89b
  Danacaea DL90
    |--D. luctuosa Desbrochers des Loges 1890 DL90
    `--D. plumbea Desbrochers des Loges 1890 DL90
  Amauronia longula Desbrochers des Loges 1890 DL90
  Alexia F90
    |--A. algirica Fairmaire 1889 [=A. reitteri Desbrochers des Loges 1889 non Ormay 1888] F90
    `--A. reitteri Ormay 1888 F90
  Langelandia reitteri DL90
  Canidia Thomson 1857 KF22

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

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

Polyphaga [Bostrichiformia, Bostrichini, 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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