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

Belongs within: Holometabola.
Contains: Rhipidoceridae, Adephaga, Archostemata, Myxophaga, Polyphaga.

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 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.

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
       `--+--Polyphaga MW15
          `--Ademosyne [Ademosynidae, Ademosynoidea] GE05
               |--A. speciosa K-P91
               `--A. wianamattensis Tillyard 1917 F71

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
  Ptomascopus aveyronensis Z02c
  Palaeosilpha fraasi Z02c
  Donaciella nagaokana S02
  Ryugadous ishikawai S86
  Kusumia takahashii S86
  Oxysoma A71
  Abax FFB88
    |--A. parallelepipedus FFB88
    `--A. parallelus FFB88
  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
  ‘Heteromorpha’ Kirby 1823 non Hübner 1822 NO08
  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
  Euryporus aeneiventris Lucas 1846 E12
  Eusomus affinis Lucas 1847 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 1847 E12
    `--M. agilis Lucas 1847 E12
  Nastus E12
    |--N. albomarginatus Lucas 1847 E12
    `--N. albopunctatus Lucas 1847 E12
  Parnus algiricus Lucas 1847 E12
  Hypophaeus angustus Lucas 1847 E12
  Hymenoplia E12
    |--H. aterrima Chevrolat 1847 E12
    `--H. cinctipennis Lucas 1847 E12
  Brachyphylla barbara Lucas 1847 E12
  Anelastes barbarus Lucas 1847 E12
  Cataphronetis Lucas 1847 E12
    `--C. levaillanti Lucas 1847 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 1847 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 1847 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 1847 E12
  Colaspidema E12
    |--C. pulchella Lucas 1847 E12
    `--C. signatipennis Lucas 1847 E12
  Thorectes E12
    |--T. puncticollis Lucas 1847 E12
    `--T. rotundatus Lucas 1847 E12
  Pachychila E12
    |--P. punctulata Lucas 1847 E12
    `--P. sabulosa Lucas 1847 E12
  Carterus rufipes Lucas 1846 E12
  Pachycerus rugosus Lucas 1847 E12
  Otophorus scolytoides Lucas 1847 E12
  Sybines sellatus Lucas 1847 E12
  Eutrapela suturalis Lucas 1847 E12
  Mazoreus testaceus Lucas 1846 E12
  Geobius tricornis Lucas 1847 E12
  Trypocladius Lucas 1847 E12
  Rhyzophagus unicolor Lucas 1847 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 M86
  Eros scutellaris Erichs. 1842 [incl. E. bremei Le Guillou 1844] M86
  Hapalochrys cinctus 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
  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

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

*Type species of generic name indicated


[A61] Andreae, H. 1961. Coleoptera: Cossyphodidae. In: Hanström, B., P. Brinck & G. Rudebeck (eds) South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 198–216. Almqvist & Wiksell: Uppsala.

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[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.

[B02] Bosc, C. 1802. Description d'un nouveau Callopus. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (Bulletin de la Société Philomathique, a ses correspondans): 12'.

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

[BR05] Bouchet, P., & J.-P. Rocroi. 2005. Classification and nomenclator of gastropod families. Malacologia 47 (1–2): 1–397.

[BR17] Bouchet, P., J.-P. Rocroi, B. Hausdorf, A. Kaim, Y. Kano, A. Nützel, P. Parkhaev, M. Schrödl & E. E. Strong. 2017. Revised classification, nomenclator and typification of gastropod and monoplacophoran families. Malacologia 61 (1–2): 1–526.

[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. Dynastinae. In: Houston, W. W. K. (ed.) Zoological Catalogue of Australia vol. 9. Coleoptera: Scarabaeoidea pp. 383–425. AGPS Press: Canberra.

[C92] Crawford, R. L. 1992. Catalogue of the genera and type species of the harvestman superfamily Phalangioidea (Arachnida). Burke Museum Contributions in Anthropology and Natural History 8: 1–60.

[D37] Dobzhansky, T. 1937. Genetics and the Origin of Species. Columbia University Press: New York.

[E02] Emberson, R. M. 2002. The beetle (Coleoptera) fauna of the Chatham Islands: Additions and corrections. New Zealand Entomologist 25: 69–77.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l’Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

[F02] Foord, M. 2002. Entomological Society conference field trip 2002. Weta 24: 37–38.

[FFB88] Franke, U., B. Friebe & L. Beck. 1988. Methodisches zur Ermittlung der Siedlungsdichte von Bodentieren aus Quadratproben und Barberfallen. Pedobiologia 32: 253–264.

[G60] Gatenby, J. B. 1960. The Australasian mycetophilid glowworms. Transactions of the Royal Society of New Zealand 88 (3): 577–593.

[G84] Gauld, I. D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum (Natural History).

[G69] Goodwin, G. G. 1969. Mammals from the State of Oaxaca, Mexico, in the American Museum of Natural History. Bulletin of the American Museum of Natural History 141 (1): 1–269, 40 pls.

[G76] Grasshoff, M. 1976. Zur Taxonomie und Nomenklatur mitteleuropäischer Radnetzspinnen der Familie Araneidae (Arachnida: Araneae). Senckenbergiana Biologica 57 (1–3): 143–154.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H62] Hass, W. H. 1962. Conodonts. In: Moore, R. C. (ed.) Treatise on Invertebrate Paleontology pt W. Miscellanea: Conodonts, Conoidal Shells of Uncertain Affinities, Worms, Trace Fossils and Problematica pp. W3–W69. Geological Society of America, and University of Kansas Press.

[Im92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[Iw92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[KN13] Kirejtshuk, A. G. & A. Nel. 2013. Skleroptera, a new order of holometabolous insects (Insecta) from the Carboniferous. Zoosystematica Rossica 22 (2): 247–257.

[K02] Krüger, M. 2002. Revision of the Afrotropical Ennominae of the Drepanogynis group IV: the genus Drepanogynis Guenée (Lepidoptera: Geometridae). Transvaal Museum Monograph 13: 1–220.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[KA-Z11] Kury, A. B., & M. A. Alonso-Zarazaga. 2011. Addenda and corrigenda to the “Annotated catalogue of the Laniatores of the New World (Arachnida, Opiliones)”. Zootaxa 3034: 47–68.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

Lawrence, J. F., B. P. Moore, J. E. Pyke & T. A. Weir. 1987. Zoological Catalogue of Australia vol. 4. Coleoptera: Archostemata, Myxophaga and Adephaga. Australian Government Publishing Service: Canberra.

[LR02] Leschen, R. A. B., & B. E. Rhode. 2002. A new genus and species of large extinct Ulodidae (Coleoptera) from New Zealand. New Zealand Entomologist 25: 57–64.

[L01] Lindquist, E. E. 2001. Poising for a new century: diversification in acarology. In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 17–34. CSIRO Publishing: Melbourne.

[MB93] Malecki, R. A., B. Blossey, S. D. Hight, D. Schroeder, L. T. Kok & J. R. Coulson. 1993. Biological control of purple loosestrife. BioScience 43 (10): 680-686.

[MG06] Manning, J. C., & P. Goldblatt. 2006. New species of Iridaceae from the Hantam-Roggeveld Centre of Endemism, and the Bokkeveld, Northern Cape, South Africa. Bothalia 36 (2): 139–145.

[M86] Masters, G. 1886. Catalogue of the described Coleoptera of Australia. Part IV. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 259–380.

[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.

[NO08] Nässig, W. A., & R. G. Oberprieler. 2008. An annotated catalogue of the genera of Eupterotidae (Insecta, Lepidoptera, Bombycoidea). Senckenbergiana Biologica 88 (1): 53–80.

[O81] O’Brien, C. O. 1981. A. A. Book of New Zealand Wildlife: A guide to the native and introduced animals of New Zealand. Lansdowne Press: Auckland.

[O49] Oliver, A. P. 1949. The fatty acid composition of Hemideina thoracica White. Transactions and Proceedings of the Royal Society of New Zealand 77 (3): 321–324.

[PH90] Petit, R. E., & M. G. Harasewych. 1990. Catalogue of the superfamily Cancellarioidea Forbes and Hanley, 1851 (Gastropoda: Prosobranchia). Nautilus 103 (Suppl. 1): 1–69.

[P61] Pettibone, M. H. 1961. New species of polychaete worms from the Atlantic Ocean, with a revision of the Dorvilleidae. Proceedings of the Biological Society of Washington 74: 167–186.

[P02] Ponomarenko, A. G. 2002. Superorder Scarabaeidea Laicharting, 1781. Order Coleoptera Linné, 1758. The beetles. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 164–176. Kluwer Academic Publishers: Dordrecht.

[R70] Riek, E. F. 1970. Fossil history. In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 168–186. Melbourne University Press.

[R96] Roff, D. A. 1996. The evolution of threshold traits in animals. Quarterly Review of Biology 71 (1): 3–35.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

[S61] Southcott, R. V. 1961. Studies on the systematics and biology of the Erythraeoidea (Acarina), with a critical revision of the genera and subfamilies. Australian Journal of Zoology 9: 367–610.

[S86] Spangler, P. J. 1986. Insecta: Coleoptera. In: Botosaneanu, L. (ed.) Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) pp. 622–631. E. J. Brill/Dr W. Backhuys: Leiden.

[TDC06] Timms, B. V., B. Datson & M. Coleman. 2006. The wetlands of the Lake Carey catchment, northeast Goldfields of Western Australia, with special reference to large branchiopods. Journal of the Royal Society of Western Australia 89 (4): 175–183.

[UO05] Ubick, D., & R. Ozimec. 2005. On the harvestman genus Lola Kratochvíl (Opiliones: Laniatores). Natura Croatica 14 (3): 161–174.

[V73] Vuilleumier, F. 1973. Insular biogeography in continental regions. II. Cave faunas from Tessin, southern Switzerland. Systematic Zoology 22 (1): 64–76.

[WP99] Walter, D. E. & H. C. Proctor. 1999. Mites: Ecology, Evolution and Behaviour. CABI Publishing: Wallingford (UK).

[WR-SG01] Wang, S. F., T. J. Ridsdill-Smith & E. L. Ghisalberti. 2001. Resistance in pasture legumes to redlegged earth mite Halotydeus destructor (Tucker) (Acari: Penthaleidae). In: Halliday, R. B., D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff (eds) Acarology: Proceedings of the 10th International Congress pp. 328–341. CSIRO Publishing: Melbourne.

[W86] Wood, S. L. 1986. A reclassification of the genera of Scolytidae (Coleoptera). Great Basin Naturalist Memoirs 10: 1–126.

[Z02a] Zherikhin, V. V. 2002a. Pattern of insect burial and conservation. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 17–63. Kluwer Academic Publishers: Dordrecht.

[Z02b] Zherikhin, V. V. 2002b. Insect trace fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 303–324. Kluwer Academic Publishers: Dordrecht.

[Z02c] Zherikhin, V. V. 2002c. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.

Last updated: 11 June 2017.

No comments:

Post a Comment

Markup Key:
- <b>bold</b> = bold
- <i>italic</i> = italic
- <a href="">FoS</a> = FoS