Myxophaga

Suborder of beetles

Myxophaga

Temporal range: Carnian–Recent

PreꞒ

Sphaerius acaroides
Scientific classification
Kingdom:  | Animalia
Phylum:  | Arthropoda
Class:  | Insecta
Order:  | Coleoptera
Suborder:  | Myxophaga
Crowson, 1955
Families

Hydroscaphidae
Lepiceridae
Sphaeriusidae
Torridincolidae
†Triamyxidae

Myxophaga is the second-smallest suborder of the Coleoptera after Archostemata, consisting of roughly 65 species of small to minute beetles in four families. The members of this suborder are aquatic and semiaquatic, and feed on algae.

## Contents

* 1 Description
* 2 Taxonomy
* 3 Distribution
* 4 Fossil record
* 5 See also
* 6 Notes
* 7 References
* 8 External links

## Description[edit]

Myxophaga have several diagnostic features: the antennae are more or less distinctly clubbed with usually fewer than nine segments, mesocoxal cavities are open laterally and bordered by a mesepimeron and metanepisternum, the hind wings are rolled apically in the resting positions. Internally, they are characterised by the presence of six malpighian tubules and the testes are tube-like and coiled.[1]

Beetles of this suborder are adapted to feed on algae. Their mouthparts are characteristic in lacking galeae and having a mobile tooth on their left mandible.[2]

## Taxonomy[edit]

There are four extant families in the suborder Myxophaga divided between two superfamilies,[3] containing about 65 described species,[4] and at least one extinct family.[a]

Superfamily Lepiceroidea Hinton, 1936

* Family Lepiceridae Hinton, 1936
* Lepicerus Hinton, 1936
* †Lepiceratus Jałoszyński et al. 2020

Superfamily Sphaeriusoidea Erichson, 1845

* Family Hydroscaphidae LeConte, 1874
* Hydroscapha LeConte, 1874
* Scaphydra Reichardt, 1973
* Yara Reichardt & Hinton, 1976
* Confossa Short, Joly, García & Maddison, 2015
* Family Sphaeriusidae Erichson, 1845
* Sphaerius Waltl, 1838
* †Burmasporum Kirejtshuk 2009
* Family Torridincolidae Steffan, 1964
* Claudiella Reichardt & Vanin, 1976
* Delevea Reichardt, 1976
* Iapir Py-Daniel, da Fonseca & Barbosa, 1993
* Incoltorrida Steffan, 1973
* Satonius Endrödy-Younga, 1997
* Torridincola Steffan, 1964
* Ytu Reichardt, 1973

Family †Triamyxidae Qvarnström et al. 2021

* †Triamyxa Qvarnström et al. 2021

Unplaced in family

* †Leehermania Chatzimanolis et al. 2012

## Distribution[edit]

Hydroscapha natans

Living members of Lepiceridae are confined to northern South America and Central America. Members of Sphaeriusidae occur on all continents except Antarctica, while Hydroscaphidae occurs on all continents except Australia and Antarctica. Torridincolidae occurs in Africa, Asia, and South America.[5]

## Fossil record[edit]

Little about the fossil record of Myxophaga is known. Fossils from the early Late Cretaceous (Cenomanian) aged Burmese amber from Myanmar, have been assigned to the extant genus Lepicerus and extinct genus Lepiceratus within Lepiceridae,[6][7][8] as well as Burmasporum, which belongs to Sphaerusidae.[9] No impression fossils of myxophagan beetle were described until 2012, probably because of their small body size and specialized habitat.[10] A fossil specimen, assigned to the living genus Hydroscapha in Hydroscaphidae, was described in 2012 from the Yixian Formation in the Jehol Biota, dating from the Early Cretaceous (Aptian).[10] In 2019, Leehermania from the Late Triassic (Norian) Cow Branch Formation of North Carolina, which had previously been interpreted as the oldest known rove beetle was reinterpreted as an early diverging relative of the family Hydroscaphidae, making it the then oldest known Myxophagan.[11] In 2021, numerous specimens of a new myxophagan were described from a coprolite found in Late Triassic (Carnian) aged sediments in Poland. The new taxon Triamyxa, was placed in its own monotypic family Triamyxidae, and was resolved as either the most basal myxophagan or sister to Hydroscaphidae.[12]

## See also[edit]

* List of subgroups of the order Coleoptera

## Notes[edit]

1. ^ † Denotes that the group is extinct

## References[edit]

1. ^ Beutel, R.; Leschen, R. (2005). "Morphology and Systematics (Archostemata, Adephaga, Myxophaga, Polyphaga partim)". Band 4 Part 38: Arthropoda, Hälfte: Insecta, Coleoptera, Beetles. Handbuch der Zoologie/Handbook of Zoology. Vol. 1\. Walter de Gruyter. ISBN 3-11-017130-9.
2. ^ Beutel & Leschen 2005, p. 43
3. ^ "Myxophaga". Tree of Life Web Project. 2007. Retrieved 2017-05-26.
4. ^ Mesaros, Gabor (2013). "Sphaeriusidae (Coleoptera, Myxophaga): A new beetle family to the fauna of Serbia". Bulletin of the Natural History Museum (6): 71–74. doi:10.5937/bnhmb1306071m.
5. ^ Sampaio, Brunno H.L.; Short, Andrew E.Z. (2018), "Families Hydroscaphidae and Torridincolidae", Thorp and Covich's Freshwater Invertebrates, Elsevier, pp. 519–525, doi:10.1016/b978-0-12-804223-6.00022-6, ISBN 978-0-12-804223-6, retrieved 2021-07-26
6. ^ Kirejtshuk, A. G.; Poinar, G. (2006). "Haplochelidae, a new family of cretaceous beetles (Coleoptera: Myxophaga) from Burmese amber". Proceedings of the Entomological Society of Washington. 108: 155–164.
7. ^ Ge, S. Q.; Friedrich, F.; Beutel, R. G. (2010). "On the systematic position and taxonomic rank of the extinct myxophagan dagger Haplochelus (Coleoptera)". Insect Systematics and Evolution. 41 (4): 329–338. doi:10.1163/187631210X537385.
8. ^ Jałoszyński, Paweł; Luo, Xiao-Zhu; Hammel, Jörg U.; Yamamoto, Shûhei & Beutel, Rolf G. (2020). "The mid-Cretaceous †Lepiceratus gen. nov. and the evolution of the relict beetle family Lepiceridae (Insecta: Coleoptera: Myxophaga)". Journal of Systematic Palaeontology. 18 (13). doi:10.1080/14772019.2020.1747561
9. ^ Kirejtshuk, A. G. (2009). "A new genus and species of Sphaeriusidae (Coleoptera, Myxophaga) from Lower Cretaceous Burmese amber" (PDF). Denisia. 26: 99–102.
10. ^ a b Cai, C.; Short, A. E. Z.; Huang, D. (2012). "The First Skiff Beetle (Coleoptera: Myxophaga: Hydroscaphidae) from Early Cretaceous Jehol Biota". Journal of Paleontology. 86 (1): 116–9. doi:10.1666/11-050.1. S2CID 140170420.
11. ^ Fikáček, Martin; Beutel, Rolf G.; Cai, Chenyang; Lawrence, John F.; Newton, Alfred F.; Solodovnikov, Alexey; Ślipiński, Adam; Thayer, Margaret K.; Yamamoto, Shûhei (January 2020). "Reliable placement of beetle fossils via phylogenetic analyses – Triassic Leehermania as a case study (Staphylinidae or Myxophaga?)". Systematic Entomology. 45 (1): 175–187. doi:10.1111/syen.12386. ISSN 0307-6970.
12. ^ Qvarnström, Martin; Fikáček, Martin; Vikberg Wernström, Joel; Huld, Sigrid; Beutel, Rolf G.; Arriaga-Varela, Emmanuel; Ahlberg, Per E.; Niedźwiedzki, Grzegorz (June 2021). "Exceptionally preserved beetles in a Triassic coprolite of putative dinosauriform origin". Current Biology: S0960982221006746. doi:10.1016/j.cub.2021.05.015.

## External links[edit]

* "Myxophaga". Atlas of Living Australia.
* Data related to Myxophaga at Wikispecies

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Extant Coleoptera families

* Kingdom: Animalia
* Phylum: Arthropoda
* Class: Insecta
* Subclass: Pterygota
* Infraclass: Neoptera
* Superorder: Endopterygota

Suborder Archostemata

* Crowsoniellidae (Crowsoniella relicta)
* Cupedidae (reticulated beetles)
* Jurodidae (Sikhotealinia zhiltzovae)
* Micromalthidae (telephone-pole beetle)
* Ommatidae

Suborder Adephaga

| Extant families|

* Amphizoidae (trout-stream beetles)
* Aspidytidae
* Carabidae (ground beetles)
* Dytiscidae (predaceous diving beetles)
* Gyrinidae (whirligig beetles)
* Haliplidae (crawling water beetles)
* Hygrobiidae
* Meruidae (Meru phyllisae)
* Noteridae (burrowing water beetles)
* Rhysodidae (wrinkled bark beetles)
* Trachypachidae (false ground beetles)

|

Suborder Myxophaga

* Hydroscaphidae (skiff beetles)
* Lepiceridae
* Sphaeriusidae
* Torridincolidae

Suborder Polyphaga

| Bostrichiformia|

| Bostrichoidea|

* Bostrichidae (auger beetles)
* Dermestidae (skin beetles)
* Jacobsoniidae (Jacobson's beetles)
* Nosodendridae (wounded-tree beetles)
* Ptiniidae (furniture beetles, death watch beetles, spider beetles)

|

Derodontoidea|

* Derodontidae (tooth-necked fungus beetles)

Cucujiformia|

| Chrysomeloidea|

* Cerambycidae (longhorn beetles)
* Chrysomelidae (leaf beetles)
* Disteniidae
* Megalopodidae
* Orsodacnidae
* Oxypeltidae
* Vesperidae

|

Cleroidea|

* Acanthocnemidae (Acanthocnemus nigricans)
* Chaetosomatidae
* Cleridae (checkered beetles)
* Mauroniscidae
* Melyridae (soft-wing flower beetles)
* Metaxinidae
* Phloiophilidae (Phloiophilus edwardsi)
* Phycosecidae
* Prionoceridae
* Thanerocleridae
* Trogossitidae (bark-gnawing beetles)

Cucujoidea|

* Alexiidae
* Biphyllidae (false skin beetles)
* Boganiidae
* Bothrideridae (dry bark beetles)
* Byturidae (fruitworm beetles)
* Cavognathidae
* Cerylonidae (minute bark beetles)
* Coccinellidae (lady beetles, or God's cows)
* Corylophidae (minute fungus beetles)
* Cryptophagidae (silken fungus beetles)
* Cucujidae (flat bark beetles)
* Cyclaxyridae
* Discolomatidae
* Endomychidae (handsome fungus beetles)
* Erotylidae (pleasing fungus beetles)
* Helotidae
* Hobartiidae
* Kateretidae (short-winged flower beetles)
* Laemophloeidae (lined flat bark beetles)
* Lamingtoniidae (Lamingtonium binnaberrense)
* Latridiidae (minute brown scavenger beetles)
* Monotomidae (root-eating beetles)
* Myraboliidae
* Nitidulidae (sap beetles)
* Passandridae (parasitic flat bark beetles)
* Phalacridae (shining flower beetles)
* Phloeostichidae
* Propalticidae
* Protocucujidae
* Silvanidae (silvanid flat bark beetles)
* Smicripidae (palmetto beetles)
* Sphindidae (dry-fungus beetles)

Curculionoidea
(weevils)|

* Anthribidae (fungus weevils)
* Attelabidae (leaf-rolling weevils)
* Belidae (primitive weevils)
* Brentidae (straight snout weevils, New York weevil)
* Caridae
* Curculionidae (true weevils, bark beetles, ambrosia beetles)
* Nemonychidae (pine flower weevils)

Lymexyloidea|

* Lymexylidae (ship-timber beetles)

Tenebrionoidea|

* Aderidae (ant-like leaf beetles)
* Anthicidae (ant-like flower beetles)
* Archeocrypticidae (cryptic fungus beetles)
* Boridae (conifer bark beetles)
* Chalcodryidae
* Ciidae (minute tree-fungus beetles)
* Melandryidae (false darkling beetles)
* Meloidae (blister beetles)
* Mordellidae (tumbling flower beetles)
* Mycetophagidae (hairy fungus beetles)
* Mycteridae (palm and flower beetles)
* Oedemeridae (false blister beetle)
* Perimylopidae, or Promecheilidae
* Prostomidae (jugular-horned beetles)
* Pterogeniidae
* Pyrochroidae (fire-coloured beetles)
* Pythidae (dead log bark beetles)
* Ripiphoridae (wedge-shaped beetles)
* Salpingidae (narrow-waisted bark beetles)
* Scraptiidae (false flower beetles)
* Stenotrachelidae (false longhorn beetles)
* Synchroidae (synchroa bark beetles)
* Tenebrionidae (darkling beetles)
* Tetratomidae (polypore fungus beetles)
* Trachelostenidae
* Trictenotomidae
* Ulodidae
* Zopheridae (ironclad beetles, cylindrical bark beetles)

Elateriformia|

| Buprestoidea|

* Buprestidae (jewel beetles, or metallic wood-boring beetles)
* Schizopodidae

|

Byrrhoidea|

* Byrrhidae (pill beetles)
* Callirhipidae (cedar beetles)
* Chelonariidae (turtle beetles)
* Cneoglossidae
* Dryopidae (long-toed water beetles)
* Elmidae (riffle beetles)
* Eulichadidae (forest stream beetles)
* Heteroceridae (variegated mud-loving beetles)
* Limnichidae (minute mud beetles)
* Lutrochidae (travertine beetles)
* Psephenidae (water-penny beetles)
* Ptilodactylidae

Dascilloidea|

* Dascillidae (soft bodied plant beetles)
* Rhipiceridae (cicada beetle, cicada parasite beetles)

Elateroidea|

* Artematopodidae (soft-bodied plant beetles)
* Brachypsectridae (Texas beetles)
* Cantharidae (soldier beetles)
* Cerophytidae (rare click beetles)
* Elateridae (click beetles)
* Eucnemidae (false click beetles)
* Jurasaidae
* Lampyridae (fireflies)
* Lycidae (net-winged beetles)
* Omethidae (false fireflies, long-lipped beetles)
* Phengodidae (glowworm beetles)
* Podabrocephalidae
* Rhagophthalmidae
* Sinopyrophoridae
* Throscidae (false metallic wood-boring beetles)

Rhinorhipoidea|

* Rhinorhipidae (Rhinorhipus tamborinensis)

Scirtoidea|

* Clambidae
* Decliniidae (Declinia relicta)
* Eucinetidae (plate-thigh beetles)
* Scirtidae

Scarabaeiformia|

| Scarabaeoidea|

* Belohinidae (Belohina inexpectata)
* Diphyllostomatidae (false stag beetles)
* Geotrupidae (dor beetles)
* Glaphyridae (bumble bee scarab beetles)
* Glaresidae (enigmatic scarab beetles)
* Hybosoridae (scavenger scarab beetles)
* Lucanidae (stag beetles)
* Ochodaeidae (sand-loving scarab beetles)
* Passalidae (betsy beetles)
* Pleocomidae (rain beetles)
* Scarabaeidae (scarabs)
* Trogidae (hide beetles)

|

Staphyliniformia|

| Histeroidea|

* Histeridae (clown beetles)
* Sphaeritidae (false clown beetles)
* Synteliidae

|

Hydrophiloidea|

* Epimetopidae
* Georissidae (minute mud-loving beetles)
* Helophoridae
* Hydrochidae
* Hydrophilidae (water scavenger beetles)
* Spercheidae

Staphylinoidea|

* Agyrtidae (primitive carrion beetles)
* Hydraenidae
* Leiodidae (round fungus beetles)
* Ptiliidae (feather-winged beetles)
* Silphidae (carrion beetles)
* Staphylinidae (rove beetles)

List of subgroups of the order Coleoptera

Taxon identifiers|

* Wikidata: Q133005
* Wikispecies: Myxophaga
* AFD: Myxophaga
* BugGuide: 40604
* EoL: 2650686
* Fauna Europaea: 10998
* Fauna Europaea (new): 42f89ae7-71c5-45ac-8492-825030358e4e
* Fossilworks: 69191
* GBIF: 4577555
* iNaturalist: 340472
* ITIS: 112736
* NBN: NHMSYS0001719091
* NCBI: 63907

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