Coleoptera

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


Belongs within: Holometabola.
Contains: Rhipidoceridae, Adephaga, Archostemata, Myxophaga, Scirtoidea, Staphylinoidea, Scarabaeoidea, Hydrophiloidea, Elateriformia, Bostrichoidea, Coccinelloidea, Tenebrionoidea, Cleroidea, Cucujoidea, Nodostoma, Chrysomeloidea, Belidae, Anthribidae, Nemonychidae, Attelabidae, Brentidae, Brachyceridae, Dryophthoridae, Curculionidae.

The Coleoptera are the beetles, one of the most diverse groups of insects. 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. They 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).

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). More derived families such as Permocupedidae and Taldycupedidae with reduced venation, and usually with reduced costal areas, appeared in the Late Permian. The Tshekardocoleidae 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).

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 had costate elytra without remnant veins. 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 Phytophaga is a major clade of beetles in the Cucujiformia, including the Chrysomelidae (leaf beetles), Cerambycidae (longicorn beetles) and Curculionoidea (weevils). Phytophagans are united by the structure of the pseudotetramerous tarsus, with the fourth tarsal segment minute and concealed between lobes of the third, and of the internal male reproductive organs, with a reduced aedeagus (Grimaldi & Engel 2005). The clade's name reflects the plant-feeding habits of the vast majority of its species; more than 99% of the more than 110,000 described species are plant feeders.

Members of the Curculionoidea have the front of the head extended into an elongate rostrum (secondarily reduced in some subgroups) with a concordant reduction and fusion of many of the mouthparts (such as reduction or loss of the labrum); they also usually have distinctly clubbed antennae with a long scape lying in a scrobe (groove on the side of the rostrum) (Grimaldi & Engel 2005). The classification of the Curculionoidea has been subject to much debate between authors. For instance, members of the Caridae, a small group of weevils found in Australia and South America, have been variously classified within the Attelabidae, Belidae or Brentidae, and are now generally treated as their own family (Bouchard 2014). Two smaller families, the Brachyceridae and Dryophthoridae, are similar in external appearance to the much more diverse Curculionidae and have often been included in the latter family, but have been separated in recent years largely on the basis of differences in genital morphology.

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] GE05
    |    |--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
               |         Eledona Latreille 1802 L02
               |           `--*E. agricola [=Bolitophagus agricola] L02
               |         Heliotis hopei (n. d.) L09
               |         Tomocoleites Hong 2002 H02
               |           `--*T. longigastrulatus Hong 2002 H02
               |--Scirtoidea MW15
               `--+--Staphyliniformia [Haplogastra, Hydrophilomorpha, Staphylinomorpha] MW15
                  |    |--Staphylinoidea MW15
                  |    `--+--Scarabaeoidea MW15
                  |       `--Hydrophiloidea MW15
                  `--+--Elateriformia MW15
                     `--+--Bostrichoidea MW15
                        `--Cucujiformia [Cucujomorpha, Nitidulariae, Trogossitarii, Xylophagi] MW15
                             |  i. s.: Parandrexidae Z02c
                             |         Cercus Latreille 1802 [incl. Cateretes] L02
                             |           |--*C. pedicularius (Linnaeus 1758) L02, L58 [=Dermestes pedicularius L02]
                             |           |--C. barbarus Lucas 1846 E12
                             |           `--C. bicolor Lucas 1846 E12
                             |--Coccinelloidea MW15
                             `--+--Tenebrionoidea MW15
                                `--+--Cleroidea MW15
                                   `--+--Cucujoidea MW15
                                      `--Phytophaga [Bruchelae, Chrysomeliformia, Chrysomelomorpha] MW15
                                           |  i. s.: Diapromorpha J89a
                                           |           |--D. melanophthalma [=Aspidolopha melanophthalma] J89a
                                           |           `--D. melanopus J89a
                                           |         Gynandrophthalma J89a
                                           |           |--G. divisa Jacoby 1889 J89a
                                           |           `--G. sexmaculata Jacoby 1889 J89a
                                           |         Melitonoma J89a
                                           |           |--M. affine Jacoby 1889 J89a
                                           |           `--M. indica Jacoby 1889 J89a
                                           |         Chrysolampra J89a
                                           |           |--C. curvipes Jacoby 1889 J89a
                                           |           `--C. splendens J89a
                                           |         Nodostoma J89b
                                           |         Aulacolepis decorata J89a
                                           |         Demotina J89a
                                           |           |--D. balyi Jacoby 1889 J89a
                                           |           `--D. serraticollis J89a
                                           |         Heteraspis J89b
                                           |           |--H. clypeata Jacoby 1889 J89a
                                           |           |--H. hebe J89a
                                           |           |--H. hirta J89b
                                           |           |--H. quadrifasciata Jacoby 1889 J89a
                                           |           |--H. variegata Jacoby 1889 J89a
                                           |           `--H. vestita J89a
                                           |         Acrothinium violaceum Jacoby 1889 J89a
                                           |         Corynodes J89a
                                           |           |--C. approximans J89a
                                           |           |--C. chalybeus J89a
                                           |           |--C. fulvicollis Jacoby 1889 J89a
                                           |           |--C. gratiosus J89a
                                           |           |--C. igneicollis Jacoby 1889 J89a
                                           |           |--C. mouhoti J89a
                                           |           |--C. peregrinus J89a
                                           |           |--C. pretiosus J89a
                                           |           `--C. pyrospilotus J89a
                                           |         Agasta formosa J89a
                                           |         Eumela cyanicollis J89a
                                           |         Lygaria westermanni J89a
                                           |         Plagiodera marginipennis Jacoby 1889 J89a
                                           |         Sphaerophysa piceicollis Jacoby 1889 J89a
                                           |         Enneamera J89b
                                           |           |--E. apicalis Jacoby 1889 J89a
                                           |           `--E. sumatrensis J89b
                                           |         Pseudodera J89a
                                           |           |--P. bifasciata Jacoby 1889 J89a
                                           |           `--P. xanthospila J89a
                                           |         Clitea picta J89a
                                           |         Podontia J89a
                                           |           |--P. affinis J89a
                                           |           `--P. quattuordecimpunctata J89a
                                           |         Bonesia J89a
                                           |           |--B. balyi Jacoby 1889 J89a
                                           |           `--B. clarki J89a
                                           |         Morphosphaera sumatrana J89a
                                           |         Clitena cyanea J89a
                                           |         Hoplasoma J89b
                                           |           |--H. ceylonensis J89b
                                           |           `--H. ventralis J89a
                                           |         Paridea thoracica J89a
                                           |         Pseudocophora J89a
                                           |           |--P. birmanica Jacoby 1889 J89a
                                           |           `--P. buqueti J89a
                                           |         Cneorane fulvicornis Jacoby 1889 J89a
                                           |         Mimastra J89a
                                           |           |--M. chenelli J89a
                                           |           |--M. gracilicornis Jacoby 1889 J89a
                                           |           `--M. polita Jacoby 1889 J89a
                                           |         Malaxia viridis [=Glyptolus viridis] J89a
                                           |         Sastroides birmanica Jacoby 1889 J89a
                                           |         Menippus J89a
                                           |           |--M. cervinus J89a
                                           |           |--M. cynicus J89a
                                           |           `--M. dimidiaticornis Jacoby 1889 J89a
                                           |         Haplosonyx J89b
                                           |           |--H. nigricollis J89b
                                           |           |--H. parvulus J89b
                                           |           |--H. scutellata J89a
                                           |           `--H. sublaevicollis Jacoby 1889 J89a
                                           |         Sphenoraia J89a
                                           |           |--S. bicolor J89a
                                           |           |--S. duodecimmaculata Jacoby 1889 J89a
                                           |           `--S. fulgida J89a
                                           |         Leptarthra J89a
                                           |           |--L. dohrni J89a
                                           |           `--L. fraternalis J89a
                                           |         Cynorta melanocephala Jacoby 1889 J89a
                                           |         Antipha J89a
                                           |           |--A. birmanica Jacoby 1889 J89a
                                           |           |--A. bretinghami J89a
                                           |           |--A. concolor Jacoby 1889 J89a
                                           |           |--A. discoidalis J89a
                                           |           |--A. postica J89a
                                           |           `--A. puncticollis Jacoby 1889 J89a
                                           |         Ochralea J89a
                                           |           |--O. divisa Jacoby 1889 J89a
                                           |           `--O. nigripes J89a
                                           |         Aenidea J89b
                                           |           |--A. apicalis Jacoby 1889 J89b
                                           |           `--A. sumatrensis J89a
                                           |         Syoplia javanensis J89b
                                           |         Dorydea J89a
                                           |           |--D. basalis Jacoby 1889 J89a
                                           |           |--D. indica J89a
                                           |           `--D. insignis J89a
                                           |         Candezea feae Jacoby 1889 J89a
                                           |         Platyxantha gracilis Jacoby 1889 J89a
                                           |         Solenia J89a
                                           |           |--S. collaris J89a
                                           |           `--S. indica Jacoby 1889 [=Euphyma indica] J89a
                                           |         Pyropida sumptuosa J89b
                                           |         Imolia nigrofasciata J89b
                                           |         Lina B66
                                           |           |--L. aenea B66
                                           |           `--L. formosana Bates 1866 B66
                                           |         Rhinosimus planirostris [=Anthribus planirostris] L02
                                           |         Aesernia Mac86b
                                           |--Chrysomeloidea GE05
                                           `--Curculionoidea (see below for synonymy) GE05
                                                |  i. s.: Episomus pauperatus P66
                                                |         Etheophanus C88
                                                |           |--E. striatus C88
                                                |           `--+--E. optandus C88
                                                |              `--+--E. nitidellus C88
                                                |                 `--E. pinguis C88
                                                |         Dysticheus rotundicollis BM76
                                                |         Minyomerus languidus BM76
                                                |         Dyslobus lecontei BM76
                                                |         Mahakamia kampmeinerti B14
                                                |         Sciopithes obscurus WZ10
                                                |         Metamasius quadrilineatus WZ10
                                                |         Proterhinus O94 [Aglycyderidae W86, Proterhinidae]
                                                |         Eobelidae W86
                                                |         Allocorynidae LB91
                                                |         Cretonanophyes P02 [Eccoptarthridae GE05]
                                                |           `--C. longirostris P02
                                                |         Rhina Latreille 1802 L02
                                                |           |--R. barbicornis [=Curculio barbicornis] L02
                                                |           `--R. cerasi (Linnaeus 1758) L02, L58 [=Curculio cerasi L02]
                                                |         Campyloscelinae Z93
                                                |           |--Phaenomerus [Phaenomerini] Z93
                                                |           `--Tomicoproctus Z93
                                                |         Aades cultratus (Fabricius 1775) Z93 (see below for synonymy)
                                                |         Alatidotasia exclamationis (Fabricius 1775) (see below for synonymy) Z93
                                                |         Catoptes Z93
                                                |           |--C. brevicornis (Broun 1904) (see below for synonymy) E02
                                                |           `--C. interruptus (Fabricius 1781) (see below for synonymy) Z93
                                                |         Phytoscaphus triangularis (see below for synonymy) Z93
                                                |         Aoplocnemis clavipes (Fabricius 1775) [=Lixus clavipes] Z93
                                                |         Cecyropa modesta M94
                                                |         Mecysmoderes Schoenherr 1837 M94
                                                |           `--M. stylicornis M94
                                                |         Peribleptus Schoenherr 1843 M94
                                                |           |--P. dealbatus (Boisduval 1835) (see below for synonymy) M94
                                                |           `--P. scalptus M94
                                                |         Rhynchitoma MW15
                                                |--Belidae MW15
                                                `--+--Anthribidae GC-P14
                                                   `--+--Nemonychidae GC-P14
                                                      `--+--Attelabidae GC-P14
                                                         `--+--Caridae [Carinae] MW15
                                                            |    |--Cretocar luzzii GE05
                                                            |    |--Caenominurus topali ML05
                                                            |    `--Car Blackburn 1897 Z94
                                                            |         |--*C. condensatus Blackburn 1897 M94
                                                            |         |--C. intermedius B14
                                                            |         `--C. pini B14
                                                            `--+--Brentidae MW15
                                                               `--+--Brachyceridae MW15
                                                                  `--+--+--Dryophthoridae GC-P14
                                                                     |  `--Curculionidae MW15
                                                                     `--Ocladius [Ocladiinae] GC-P14
                                                                          |--O. bifasciatus Tourn. 1875 (see below for synonymy) G89
                                                                          |--O. holomelas G89
                                                                          |--O. salicorniae G89
                                                                          |--O. sapeti Gestro 1889 G89
                                                                          `--O. sharpi G89

Coleoptera incertae sedis:
  Homalaena dispersa F02
  Lagochile L01
  Cerotoma trifurcata WR-SG01
  Tymbopiptus valeas Kuschel 1987 LR02
  Geodorcus capito E02
  Metabuprestium oyunchaiense Z02a
  Betulapion simile Z02a
  Electrotribus 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
  Osmoderma opicum Lewis 1887 Iw92
  Prismognathus Iw92
    |--P. dauricus (Motschulsky 1880) Iw92
    `--P. tokui Kurosawa 1975 Iw92
  Lychnuris Iw92
    |--L. atripennis Iw92
    |--L. miyako (Nakane 1981) Iw92
    `--L. rufa Iw92
  Acalolepta degener (Bates 1873) Iw92
  Apotomopterus maacki Iw92
    |--A. m. maacki Iw92
    `--A. m. aquatilis (Bates 1883) Iw92
  Thyestilla gebleri (Faldermann 1835) Iw92
  Chaetotrechiama procerus Uéno 1982 Iw92
  Rakantrechus Iw92
    |--R. elegans Uéno 1960 Iw92
    |--R. lallum Uéno 1970 Iw92
    `--R. mirabilis Uéno 1958 Iw92
  Awatrechus hygrobius Uéno 1958 Iw92
  Leptelmis gracilis Sharp 1888 Iw92
  Macroplea japana (Jacoby 1885) Iw92
  Ishikawatrechus intermedius Uéno 1957 Iw92
  Ceratoderus venustus Hisamatsu 1963 Iw92
  Eusclerus Sharp 1886 C92
  Prionomma White 1853 C92
  Limnozenus zealandicus TDC06
  Sternolopus immarginatus TDC06
  Spelaeobates pharensis Müller 1901 UO05
  Lasiorrhynchus barbicornis O81
  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
  Zygia Fabricius 1775 G76
    `--Z. oblonga L02
  Eulasia Truqui 1847 K02
  Lissodes Berthold 1827 K02
  Dendrobias mandibularis R96
  Podischnus agenor R96
  Kubousia axillaris MG06
  Pachymerus dactris O49
  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
  Photinus pyralis G60
  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
  Hymenoplia E12
    |--H. aterrima Chevrolat 1847 E12
    `--H. cinctipennis 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. warnieri Lucas 1847 E12
  Omophlus E12
    |--O. erythrogaster Lucas 1846 E12
    `--O. maroccanus Lucas 1847 E12
  Heterophaga Lucas 1847 E12
  Singilis mauritanica Lucas 1846 E12
  Pachypterus Lucas 1847 E12
    `--P. mauritanicus Lucas 1847 E12
  Stenopterus mauritanius 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
  Pachycerus rugosus 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
  Tetraonyx frontalis Im92
  Oulema melanopus G84
  Prionognathus Ferté-Sénèctere 1851 P61, H62
  Anisoceras Dejean 1833 P61
  Macronemus mimus B14
  Neoclytus conjunctus B14
  Schizax senex B14
  Pelops Gistl 1834 CW92a
  Trionychus Burmeister 1847 CW92b
  Schizocoleidae GE05
  Rhombocoleidae GE05
  Permocupes P02 [Permocupedidae GE05]
    `--P. sojanensis P02
  Asiocoleidae GE05
    |--Oborocoleus rohdendorfi Kukalová 1969 KN13
    `--Tetracoleus Ponomarenko 2009 KN13
  Tricoleidae GE05
  Taldycupes P02 [Taldycupedidae GE05]
    `--T. reticulatus P02
  Mastigus L02
    |--M. palpalis L02
    `--M. spinicornis [=Ptinus spinicornis] L02
  Rhipidoceridae Mas86
  Eros scutellaris Erichs. 1842 [incl. E. bremei Le Guillou 1844] Mas86
  Hapalochrus cinctus Mas86
  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
  Apate B35
    |--A. jesuita B35
    |--A. minuta B35
    `--A. religiosa B35
  Cylichnus Burmeister 1844 [=Cylichna (l. c.) non Lovén 1846] BR17
  Atopa Paykull 1799 BR17
  Faula Blanchard 1850 BR17
  Pachnoda Burmeister 1842 [Pachnodina] BR17
  Eriocampa ovata F92

Aades cultratus (Fabricius 1775) Z93 [=Curculio cultratus Z93, Aterpus cultratus B35; incl. C. bicristatus Fabricius 1801 Z93]

Alatidotasia exclamationis (Fabricius 1775) [=Curculio exclamationis] Z93

Catoptes brevicornis (Broun 1904) [=Platyomidia brevicornis non Catoptes brevicornis Sharp 1886 (not preoc. if in dif. gen.)] E02

Catoptes interruptus (Fabricius 1781) [=Curculio interruptus] Z93

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

Curculionoidea [Adelognatha, Cryptoplides, Curculionites, Heteromorphi, Rhynchophora] GE05

Ocladius bifasciatus Tourn. 1875 [=Oecladius bifasciatus; incl. Oc. setipes Ancey 1881] G89

Peribleptus dealbatus (Boisduval 1835) [=Alcides dealbatus, Paepalosomus dealbatus] M94

Phytoscaphus triangularis [incl. Curculio lanatus Fabricius 1801] Z93

Polyphaga [Bostrichiformia, Bostrichini, Diaperialae, Heterophaga, Malacodermata, Malacodermidae, Necrophagi, Scarabaeina, Scarabaeomorpha]

*Type species of generic name indicated

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