Scarabaeoidea

Antiochrus brunneus, from here.


Belongs within: Staphyliniformia.
Contains: Trogidae, Geotrupidae, Lampriminae, Lucaninae, Ceratognathus, Rutelinae, Aclopinae, Cetoniinae, Melolonthinae, Dynastinae, Bolboceratinae, Aulacocyclus, Passalinae, 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.

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 B14
    |     |    |--Taurocerastes patagonicus Philippi 1866 MW15, B14
    |     |    `--Frickius B14
    |     `--+--Diphyllostoma [Diphyllostomatidae] MW15
    |        `--Lucanidae [Pectinicornia] MW15
    |             |  i. s.: Neolucanus B66
    |             |           |--N. castanopterus B66
    |             |           |--N. championi B66
    |             |           |--N. sinicus B66
    |             |           `--N. swinhoei Parry in Bates 1866 B66
    |             |         Casignetis AY04
    |             |--Lampriminae B14
    |             |--Lucaninae B14
    |             |--Nicaginae MC92
    |             |    |--Ceratognathus MC92
    |             |    `--Nicagus MW15
    |             |--Syndesus Macleay 1819 [Syndesinae] MC92
    |             |    |--*S. cornutus (Fabricius 1801) [=Sinodendron cornutus; incl. Lucanus parvus Donovan 1805] MC92
    |             |    `--S. macleayi Boileau 1905 MC92
    |             `--Dorcus E30 [Dorcinae B27]
    |                  |--D. helmsi O81
    |                  `--D. parallelopipedus E30
    `--Scarabaeidae [Copridae, Coprophagi] MW15
         |  i. s.: Odontria striata WH02
         |         Holcorobeus GE05
         |           |--H. nigrimontanus GE05
         |           `--H. vittatus GE05
         |         Costelytra zealandica (White 1846) H82
         |         Trichillium brachyporum A71
         |         Acoma Casey 1881 N02
         |         ‘Benedictia’ Sanderson 1939 non Dybowski 1875 N02
         |         Proteroscarabaeus Grabau 1923 N02
         |         Plusiotes resplendens GE05
         |         Schizonycha ruficollis G89
         |         Temnorhynchus baal Reiche 1856 G89
         |         Stalagmosoma cynanchi [=Cetonia cynanchi] G89
         |         Tropinota squalida [=Scarabaeus squalidus] G89
         |         Canthochilum L88
         |         Ipsellissus L88
         |         Pycnopanelus L88
         |         Canthonella L88
         |         Serica C01
         |           |--S. brunnea V09
         |           `--S. holosericea C01
         |         Pleurophorus caesus C01
         |         Phyllopertha horticola C01
         |         Anisoplia villosa C01
         |         Epicometis hirta C01
         |         Potosia C01
         |           |--P. cuprea [incl. P. floricola] C01
         |           `--P. hungarica [incl. P. hungarica var. armeniaca] C01
         |         Oncerus [Oncerinae] N02
         |         Pachypodinae N02
         |         Rutelinae F55
         |         Aclopinae CW92d
         |--+--+--+--Cetoniinae MW15
         |  |  |  `--Melolonthinae MW15
         |  |  `--+--Dynastinae MW15
         |  |     `--Pleocominae [Pleocomidae] MW15
         |  |          |--Crecotoma Nikolajev 2002 [Cretocomini] N02
         |  |          |    `--*C. tologoica Nikolajev 2002 N02
         |  |          `--Pleocoma Le Conte 1856 [Pleocomini] N02
         |  |               `--P. australis Fall 1911 B14
         |  `--+--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
         |             |--Aulacocyclinae BB16
         |             |    |--Aulacocyclus BB16
         |             |    `--Ceracupini BB16
         |             `--Passalinae CW92a
         `--+--+--Scarabaeinae MW15
            |  `--Aphodiinae MW15
            `--+--+--Ochodaeus [Ochodaeidae] MW15
               |  `--Glaphyridae [Glaphyrides, Glaphyrinae] MW15
               |       |--Lichnanthe MW15
               |       `--Glaphyrus E12
               |            |--G. serratulae [=Melolontha serratulae] L02
               |            `--G. viridicollis Lucas 1847 E12
               `--Hybosoridae [Hybosorides, Hybosorinae] MW15
                    |--Hybosorus illigeri Reiche 1853 G89
                    |--Liparochrus CW92c
                    |--Phaeochrous Castelnau 1840 CW92c
                    |    |--*P. imaginatus Castelnau 1840 CW92c
                    |    |--P. australicus Kuijten 1978 CW92c
                    |    `--P. emarginatus Castelnau 1840 CW92c (see below for synonymy)
                    `--Antiochrus Sharp 1873 CW92c
                         |--*A. brunneus Sharp 1873 [incl. Liparochrus oblongus Harold 1873] CW92c
                         |--A. aberrans (Fairmaire 1877) [=Liparochrus aberrans] CW92c
                         |--A. freyi Petrovitz 1963 CW92c
                         |--A. politulus (Macleay 1888) [=Liparochrus politulus] CW92c
                         |--A. septendrionalis Petrovitz 1968 CW92c
                         |--A. setosus Petrovitz 1963 CW92c
                         `--A. similis Petrovitz 1971 CW92c

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
  Lithanomala N02
  Cretoserica N02
  Cretanaides N02
  Euchlora B66
    |--E. albopilosa B66
    |--E. castaneoventris Bates 1866 B66
    |--E. expansa Bates 1866 B66
    `--E. trachypyga Bates 1866 B66
  Afroglaresis CH92
  Cryptogenius CH92
  Antemnacrassa MW15
  Belohinidae GE05
  Ceratocanthidae [Acanthoceridae, Acanthocerinae] GE05
    |--Acanthocerus spinicornis Mas86
    |--Cyphopisthes Gestro 1899 CW92b
    |    |--*C. amphicyllis (Sharp 1875) [=Synarmostes amphicyllis] CW92b
    |    `--C. descarpentriesi Paulian 1977 CW92b
    `--Pterorthochaetes Gestro 1899 CW92b
         |--*P. gestroi (Harold 1874) [=Synarmostes gestroi] CW92b
         |--P. cribricollis Gestro 1899 CW92b
         `--P. simplex Gestro 1899 CW92b
  Sinodendron cylindricum L02
  Cremastocheilus castaneus L02

Phaeochrous emarginatus Castelnau 1840 CW92c [incl. Silphoides hirtipes Macleay 1864 CW92c, P. hirtipes Mas86]

*Type species of generic name indicated

REFERENCES

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[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales & G. S. Taylor. 2004. Insects ‘Down Under’—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216–234.

[B66] Bates, H. W. 1866. On a collection of Coleoptera from Formosa sent home by R. Swinhoe, Esq., H. B. M. Consul, Formosa. Proceedings of the Zoological Society of London 1866: 339–355.

[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature's gems. Ivy Press: Lewes (United Kingdom).

[BB16] Boucher, S., M. Bai, B. Wang, W. Zhang & X. Yang. 2016. Passalopalpidae, a new family from the Cretaceous Burmese amber, as the possible sister group of Passalidae Leach (Coleoptera: Scarabaeoidea). Cretaceous Research 64: 67–78.

[B27] Brookes, A. E. 1927. A new genus and three new species of Coleoptera. Transactions and Proceedings of the New Zealand Institute 57: 563–566.

[CH92] Cassis, G., W. W. K. Houston, T. A. Weir & B. P. Moore. 1992. Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea. AGPS Press: Canberra.

[CW92a] Cassis, G., & T. A. Weir. 1992a. Passalidae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 20–29. AGPS Press: Canberra.

[CW92b] Cassis, G., & T. A. Weir. 1992b. Ceratocanthidae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 65–66. AGPS Press: Canberra.

[CW92c] Cassis, G., & T. A. Weir. 1992c. Hybosoridae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 67–75. AGPS Press: Canberra.

[CW92d] Cassis, G., & T. A. Weir. 1992d. Aclopinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 78–80. AGPS Press: Canberra.

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