Antiochrus brunneus, from here.

Belongs within: Coleoptera.
Contains: Trogidae, Geotrupidae, Lampriminae, Lucaninae, Ceratognathus, Aclopinae, Melolonthinae, Bolboceratinae, Passalinae, Aulacocyclus, Scarabaeinae, Aphodiinae, Liparochrus.

The Scarabaeoidea include the scarabs and related beetles. Scarabaeoids have distinctive antennae with an asymmetrical lamellate club, and most species have broad flattened tibiae for digging. The majority of species in the group are currently included in the family Scarabaeidae, species of which usually have the labrum and/or the mandibles concealed by the clypeus and tergite VIII of the abdomen exposed to form a pygidium. Phylogenetic analyses, however, suggest the Scarabaeidae may be paraphyletic to at least some of the other scarabaeoid families. The Lucanidae, stag beetles, differ from other families in having the segments of the antennal club thick and not capable of being closely apposed to each other. The Passalidae, bessbugs, live in groups in rotting logs; they exhibit some social behaviour, with adults preparing food for the larvae. Members of Lucanidae and Passalidae have antennae with relatively thick segments in the club that are not capable of close apposition. The Glaphyridae, bumble bee scarab beetles, are actively flying, flower-visiting scarab beetles that are often brightly coloured bee mimics. The Hybosoridae are carrion-feeding beetles with sharply curved mandibles visible beyond the labrum, and a compact antennal club with closely appressed lamellae. The Scarabaeinae and Aphodiinae form a clade of fungus- and dung-feeding species with abdominal spiracles all within the pleural membrane, oblique or longitudinal mid coxae, and often weakly sclerotised mandibles.

Synapomorphies (from Grimaldi & Engel 2005): Burrowing adaptations, including large forecoxae and tibiae with series of teeth; hind wings with reduced venation and spring mechanism for folding them; adult antennae with club or lamellate apex bearing series of flattened extensions; larvae scarabaeiform, thick and C-shaped with well-developed legs and no urogomphi.

<==Scarabaeoidea [Lamellicornia, Scarabaeides, Scarabaeiformia]
    |--+--+--Glaresis [Glaresidae] MW15
    |  |  `--+--Trogidae MW15
    |  |     `--Geotrupidae MW15
    |  `--+--Taurocerastinae CW92c
    |     |    |--Taurocerastes patagonicus Philippi 1866 MW15, B14
    |     |    `--Frickius B14
    |     `--+--Diphyllostoma [Diphyllostomatidae] MW15
    |        `--Lucanidae [Lucanides, Pectinicornia] MW15
    |             |  i. s.: Neolucanus B66
    |             |           |--N. castanopterus B66
    |             |           |--N. championi B66
    |             |           |--N. laticollis B43
    |             |           |--N. sinicus B66
    |             |           `--N. swinhoei Parry in Bates 1866 B66
    |             |         Casignetis AY04
    |             |         Chiasognathus [Chiasognathinae] RD77
    |             |           `--C. grantii RD77
    |             |         Sinodendron cylindricum RD77, L02
    |             |         Calcodes maculosus (Didier 1930) [=Neolucanus maculosus] B43
    |             |         Prismognathus B43
    |             |           |--P. branczicki Nonfried 1905 B43
    |             |           |--P. dauricus (Motschulsky 1880) I92
    |             |           |--P. subaeneus B43
    |             |           `--P. tokui Kurosawa 1975 I92
    |             |         Aesalus scarabaeoides G20 [=Lucanus scarabaeoides L02]
    |             |         Dorcasoides P92
    |             |         Paleognathus P92
    |             |--Lampriminae MC92
    |             |--Lucaninae MC92
    |             |--Aesalinae MC92
    |             |--Nicaginae MC92
    |             |    |--Ceratognathus MC92
    |             |    `--Nicagus MW15
    |             |--Dorcus E30 [Dorcinae B27]
    |             |    |--D. helmsi O81
    |             |    `--D. parallelopipedus E30
    |             |--Platycerus Geoffroy 1762 P92, BR17 [Platycerinae]
    |             |    |--P. caraboides G20
    |             |    `--P. foveicollis Dupont in Boisduval 1835 B35
    |             `--Syndesinae MC92
    |                  |--Psilodon M78
    |                  `--Syndesus Macleay 1819 MC92
    |                       |--*S. cornutus (Fabricius 1801) MC92 (see below for synonymy)
    |                       `--S. macleayi Boileau 1905 MC92
    `--Scarabaeidae [Copridae, Coprophagi, Melonthidae, Rutelidae] MW15
         |  i. s.: Odontria striata WH02
         |         Holcorobeus GE05
         |           |--H. nigrimontanus GE05
         |           `--H. vittatus GE05
         |         Costelytra zealandica (White 1846) H82
         |         Trichillium brachyporum A71
         |         ‘Benedictia’ Sanderson 1939 non Dybowski 1875 N02
         |         Proteroscarabaeus Grabau 1923 N02
         |         Plusiotes resplendens GE05
         |         Schizonycha G89
         |           |--S. carpentariae B35
         |           `--S. ruficollis G89
         |         Temnorhynchus baal Reiche 1856 G89
         |         Stalagmosoma cynanchi [=Cetonia cynanchi] G89
         |         Canthochilum L88
         |         Ipsellissus L88
         |         Pycnopanelus L88
         |         Canthonella L88
         |         Pleurophorus caesus C01
         |         Epicometis hirta C01
         |         Oncerinae N02
         |           |--Oncerus floralis Leconte 1856 S38
         |           `--Nefoncerus Saylor 1938 S38
         |                `--*N. convergens (Horn 1894) [=Oncerus convergens] S38
         |         Aclopinae CW92f
         |         Stephanucha thoracica AT01
         |         Pseudosinghala A01
         |           |--P. conjuga Arrow 1901 A01
         |           `--P. regalis Arrow 1901 A01
         |         Hylamorpha elegans [incl. H. rufimana, Sulcipalpus subviolaceus] A01
         |         Trichorhyssemus riparius W87
         |         Aulocnocnemis A06
         |         Hybalus [Orphninae] A06
         |           `--H. graecus A06
         |         Chaunanthus A06
         |         Panotrogus nepalensis A06
         |--+--Melolonthinae MW15
         |  `--+--Bolboceratinae MW15
         |     `--+--Passalopalpus Boucher & Bai in Boucher, Bai et al. 2016 [Passalopalpidae] BB16
         |        |    `--*P. cheni Boucher, Bai & Zhang in Boucher, Bai et al. 2016 BB16
         |        `--Passalidae [Passalides] MW15
         |             |  i. s.: Odontotaenius disjunctus S01
         |             |         Popilius disjunctus A-B91
         |             |         Vellejus compergus Mac86
         |             |         Laches comptonii Kaup 1868 Mac86
         |             |--Passalinae CW92a
         |             `--Aulacocyclinae BB16
         |                  |--Aulacocyclus BB16
         |                  `--Ceracupini BB16
         `--+--+--Scarabaeinae MW15
            |  `--Aphodiinae MW15
            `--+--+--Ochodaeus [Ochodaeidae] MW15
               |  `--Glaphyridae [Glaphyrides, Glaphyrinae] MW15
               |       |--Lichnanthe MW15
               |       `--Glaphyrus E12
               |            |--G. (Anisonix) crinitus G20
               |            |--G. (Glaphyrus) serratulae G20 [=Melolontha serratulae L02]
               |            |--G. viridicollis Lucas 1846 E12
               |            `--G. (Amphicoma) vulpes G20
               `--Hybosoridae [Hybosorides, Hybosorinae] MW15
                    |--Hybosorus illigeri Reiche 1853 G89
                    |--Liparochrus CW92e
                    |--Phaeochrous Castelnau 1840 CW92e
                    |    |--*P. imaginatus Castelnau 1840 CW92e
                    |    |--P. australicus Kuijten 1978 CW92e
                    |    `--P. emarginatus Castelnau 1840 CW92e (see below for synonymy)
                    `--Antiochrus Sharp 1873 CW92e
                         |--*A. brunneus Sharp 1873 [incl. Liparochrus oblongus Harold 1873] CW92e
                         |--A. aberrans (Fairmaire 1877) [=Liparochrus aberrans] CW92e
                         |--A. freyi Petrovitz 1963 CW92e
                         |--A. politulus (Macleay 1888) [=Liparochrus politulus] CW92e
                         |--A. septendrionalis Petrovitz 1968 CW92e
                         |--A. setosus Petrovitz 1963 CW92e
                         `--A. similis Petrovitz 1971 CW92e

Scarabaeoidea incertae sedis:
  Lithoscarabaeus N02
  Cretogeotrupes N02
  Cretochodaeus N02
  Cretobolbus N02
  Cretaegialia N02
  Cretoscarabaeus N02
  Cretomelolontha N02
  Cretohybosorus N02
  Avitortor N02
  Prototrox N02
  Cretaesalus N02
  Cretorabaeus N02
  Cretanaides N02
  Euchlora B66
    |--E. albopilosa B66
    |--E. castaneoventris Bates 1866 B66
    |--E. expansa Bates 1866 B66
    |--E. lucidula B35
    `--E. trachypyga Bates 1866 B66
  Afroglaresis CW92b
  Cryptogenius CW92b
  Antemnacrassa MW15
  Belohinidae B14
  Ceratocanthidae [Acanthoceridae, Acanthocerinae] CW92d
    |--Acanthocerus spinicornis Mas86
    |--Cyphopisthes Gestro 1899 CW92d
    |    |--*C. amphicyllis (Sharp 1875) [=Synarmostes amphicyllis] CW92d
    |    `--C. descarpentriesi Paulian 1977 CW92d
    `--Pterorthochaetes Gestro 1899 CW92d
         |--*P. gestroi (Harold 1874) [=Synarmostes gestroi] CW92d
         |--P. cribricollis Gestro 1899 CW92d
         `--P. simplex Gestro 1899 CW92d
  Pachylistes chinensis WK86
  Epichloris B35
  Areoda B35
  Caelidia marginata B35
  Micronyx chlorophyllus B35
  Popilia nigrita Boisduval 1835 B35
  Apogonia conspersa Boisduval 1835 B35
  Dicheros plagiatus B35

Phaeochrous emarginatus Castelnau 1840 CW92e [incl. Silphoides hirtipes Macleay 1864 CW92e, P. hirtipes Mas86]

*Syndesus cornutus (Fabricius 1801) MC92 [=Sinodendron cornutus MC92; incl. Lucanus parvus Donovan 1805 MC92, Lamprima parva B35]

*Type species of generic name indicated


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