Belongs within: Coleoptera.
Contains: Ceratocanthidae, Geotrupidae, Hybosoridae, Ochodaeidae, Glaphyridae, Scarabaeidae, Passalidae, Lucanidae, Aesalinae, Trogidae.
The Scarabaeoidea, scarabs and related beetles, are a group of beetles characterised by distinctive antennae with an asymmetrical lamellate club. In many families, except Lucanidae, Passalidae and Diphyllostoma, the segments of the antennal club are flattened and capable of close apposition. Most species also have broad flattened fore tibiae for digging. The terminal abdominal tergite (T8) forms a broad pygidium that is often exposed by the elytra. Larvae are subterranean or concealed feeders and typically adopt a characteristic C-shaped pose.
Smaller taxa in the Scarabaeoidea recognised at family level include the Californian genus Diphyllostoma, distinguished by unopposable antennal club segments, lack of a canthus across the eye, and seven visible abdominal ventrites. Females of this genus have greatly reduced eyes compared to males and vestigial wings (Jameson & Ratcliffe 2002). Glaresis is a cosmopolitan genus of small beetles found in dry, sandy areas and attracted to lights, though their biology is otherwise little known. They have five ventrites on the abdomen and eyes divided by a canthus. Adults stridulate weakly when disturbed. About fifty species have been described from all continents except Australia (Jameson 2002). The Belohinidae are represented by a single species from southern Madagascar.
Passalopalpus cheni is a fossil species described from Cretaceous Burmese amber, closely resembling modern Passalidae but differing in having seven rather than five visible abdominal ventrites and nine- rather than ten-segmented antennae (Boucher et al. 2016).
Synapomorphies (Grimaldi & Engel 2005): Burrowing adaptations, including large fore coxae 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]
|--+--GeotrupidaeMF15
| `--+--+--HybosoridaeMF15
| | `--+--OchodaeidaeMW15
| | `--GlaphyridaeMW15
| `--+--ScarabaeidaeMF15
| `--+--PassalidaeMF15
| `--Passalopalpus Boucher & Bai in Boucher, Bai et al. 2016 [Passalopalpidae]BB16
| `--*P. cheni Boucher, Bai & Zhang in Boucher, Bai et al. 2016BB16
`--+--LucanidaeMF15
`--+--+--AesalinaeMF15
| `--Diphyllostoma Fall 1901MF15, JR02 [incl. Phyllostoma Fall 1901 non Cuvier 1800JR02; Diphyllostomatidae]
| |--D. fimbriataLC20
| `--D. nigricollis Fall 1912F12
`--+--TrogidaeMF15
`--Glaresis Erichson 1848MF15, J02 (see below for synonymy)
|--G. beckeriLC20
|--G. dakotensisJ02
|--G. ecostataLC20
`--G. orthochilusLC20
Scarabaeoidea incertae sedis:
LithoscarabaeusN02
CretogeotrupesN02
CretochodaeusN02
CretobolbusN02
CretaegialiaN02
CretoscarabaeusN02
CretomelolonthaN02
CretohybosorusN02
AvitortorN02
PrototroxN02
CretaesalusN02
CretorabaeusN02
CretanaidesN02
CryptogeniusCW92a
Antemnacrassa albosulcataLC20
BelohinidaeMW15
CeratocanthidaeCW92b
Pachylistes chinensisWK86
EpichlorisB35
Caelidia marginataB35
Micronyx chlorophyllusB35
PopiliaB35
|--P. cinnabarinaF89
|--P. discipennis [incl. P. discipennis var. unicolor Fairmaire 1889]F89
|--P. diversaF89
|--P. fallaciosa Fairmaire 1889F89
|--P. inconstans Fairmaire 1889F89
|--P. miniatipennis Fairmaire 1889F89
|--P. nigrita Boisduval 1835B35
`--P. pustulataF89
LepitrixLC40
|--L. abbreviatusLC40
|--L. capicolaLC40
|--L. impuncta [=Melolontha impuncta]LC40
|--L. lineatusLC40
`--L. nigripesLC40
Chasme decoraLC40
Pachycnema crassipesLC40
AnisonyxLC40
|--A. crinitusLC40
|--A. ignitus Klug in Laporte de Castelnau 1840LC40
|--A. lepidota [=Melolontha lepidota]LC40
|--A. lynxLC40
|--A. nasua [=Melolontha nasua]LC40
`--A. ursusLC40
Glaresis Erichson 1848MF15, J02 [incl. Afroglaresis Petrovitz 1968J02, Eoglaresis Semenov-Tian-Shanskii & Medvedev 1932J02; Glaresidae, Glaresini]
*Type species of generic name indicated
References
[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d’Urville. Faune entomologique de l’océan Pacifique, avec l’illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.
[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.
[CW92a] Cassis, G., & T. A. Weir. 1992a. Trogidae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 30–40. 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.
[F89] Fairmaire, L. 1889. Coléoptères de l’intérieur de la Chine. 5e partie. Annales de la Société Entomologique de France, 6e série 9: 5–84.
[F12] Fall, H. C. 1912. New Coleoptera chiefly from the southwest.—V. Canadian Entomologist 44 (2): 40–48.
Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.
[J02] Jameson, M. L. 2002. Glaresidae Semenov-Tian-Shanskii and Medvedev 1932. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 15–16. CRC Press.
[JR02] Jameson, M. L., & B. C. Ratcliffe. 2002. Diphyllostomatidae Holloway 1972. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 10–11. CRC Press.
[LC40] Laporte de Castelnau, F. 1840. Histoire Naturelle des Insectes Coléoptères vol. 2. P. Duménil: Paris.
[LC20] Lü, L., C.-Y. Cai, X. Zhang, A. F. Newton, M. K. Thayer & H.-Z. Zhou. 2020. Linking evolutionary mode to palaeoclimate change reveals rapid radiations of staphylinoid beetles in low-energy conditions. Current Zoology 66 (4): 435–444.
[MF15] McKenna, D. D., B. D. Farrell, M. S. Caterino, C. W. Farnum, D. C. Hawks, D. R. Maddison, A. E. Seago, A. E. Z. Short, A. F. Newton & M. K. Thayer. 2015. Phylogeny and evolution of Staphyliniformia and Scarabaeiformia: forest litter as a stepping stone for diversification of nonphytophagous beetles. Systematic Entomology 40: 35–60.
[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.
[N02] Nikolajev, G. V. 2002. To possible assignment of a new genus of lamellicorn beetles from the Lower Cretaceous of Mongolia to the subfamily Pleocominae (Coleoptera, Scarabaeidae). Paleontologicheskii Zhurnal 2002 (3): 51–55 (transl. Paleontological Journal 36 (3): 279–282).
[WK86] Walter, D. E., & G. W. Krantz. 1986. Description of the Macrocheles kraepelini species complex (Acari: Macrochelidae) with two new species. Canadian Journal of Zoology 64: 212–217.