Protocoleoptera

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Protocoleoptera
Temporal range: Asselian–Anisian
Permian beetle diversity.jpg
(A, B) Moravocoleus permianus (Tshekardocoleidae) (C,D) Permocupes sojanensis (Permocupedidae) (E) Tshekardocoleidae, Sylvacoleus richteri (F) Taldycupedidae, Taldycupes reticulatus
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Coleoptera
Suborder: Protocoleoptera
Tillyard, 1924 [1]
Families

see text

The Protocoleoptera are a paraphyletic group of extinct beetles, containing the earliest and most primitive lineages of beetles. They represented the dominant group of beetles during the Permian, but were largely replaced by modern beetle groups during the following Triassic. Protocoleopterans typically possess prognathous (horizontal) heads, distinctive elytra with regular window punctures, culticles with tubercles or scales, as well as a primitive pattern of ventral sclerites, similar to the modern archostematan families Ommatidae and Cupedidae. They are thought to have been xylophagous and wood boring. [2]

Contents

Nomenclature

Protocoleoptera was originally proposed by Robert John Tillyard in 1924 for the extinct genus Protocoleus , assigned to the family Protocoleidae. Protocoleus is now considered a member of the extinct order Protelytroptera (a stem-group of the modern Dermaptera, the earwigs), which would make Protocoleoptera in this sense a synonym of the order. [3] Roy Crowson later reused the name "Protocoleoptera" to refer to Early Permian beetles such as the Tshekardocoleidae, while establishing the Archecoleoptera for Late Permian beetles. [4] Because Protocoleoptera in its original sense is a synonym of Protelytroptera, Cai et al. (2022) proposed a replacement name, Alphacoleoptera Engel, Cai & Tihelka, 2022. [5] Some authors reject the use of Protocoleoptera and Archecoleoptera and include their members within a broad concept of the suborder Archostemata instead. [3] [6]

Taxonomy

The taxonomic naming scheme of early beetles currently has no consensus, with several separate classification schemes proposed for higher level-clades within the stem-group. [5] [7] It is generally agreed that Tshekardocoleidae is the earliest diverging group among the major families. [2] [7]

Related Research Articles

<span class="mw-page-title-main">Cupedidae</span> Family of beetles

The Cupedidae are a small family of beetles, notable for the square pattern of "windows" on their elytra, which give the family their common name of reticulated beetles.

<span class="mw-page-title-main">Telephone-pole beetle</span> Species of beetle

The telephone-pole beetle is a beetle native to the eastern United States, and the only living representative of the otherwise extinct family Micromalthidae. They have an unusual life cycle involving asexually reproducing (parthenogenetic) female larvae, as well as non-functional "ghost adults".

<span class="mw-page-title-main">Archostemata</span> Suborder of beetles

The Archostemata are the smallest suborder of beetles, consisting 45 living species in five families. They are an ancient lineage with a number of primitive characteristics. Antennae may be thread-shaped (filiform) or like a string of beads (moniliform). This suborder also contains the only beetles where both sexes are paedogenic, Micromalthus debilis. Modern archostematan beetles are considered rare, but were more diverse during the Mesozoic. The term "Archostemata" is used more broadly by some authors to include both modern archostematans as well as stem-group beetles like "protocoleopterans", which some modern archostematans closely resemble to due to their plesiomorphic morphology. Genetic research suggests that modern archostematans are a monophyletic group. Some genetic studies have recovered archostematans as the sister group of Myxophaga.

<span class="mw-page-title-main">Trachypachidae</span> Family of beetles

The Trachypachidae are a family of beetles that generally resemble small ground beetles, but that are distinguished by the large coxae of their rearmost legs. There are only six known extant species in the family, with four species of Trachypachus found in northern Eurasia and northern North America, and two species of Systolosoma in Chile and Argentina. They were much more diverse in the past, with dozens of described species from the Mesozoic.

<span class="mw-page-title-main">Myxophaga</span> Suborder of beetles

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.

<span class="mw-page-title-main">Jurodidae</span> Family of beetles

Jurodidae is a family of beetles that was originally described for the extinct genus Jurodes, known from the Middle-Late Jurassic of Asia. In 1996, a living species, Sikhotealinia zhiltzovae was discovered in the Sikhote-Alin mountains in southeastern Siberia, and assigned to this family. Their placement is uncertain, but are usually considered archostematans. In one study, Sikhotealinia and Jurodes were considered a sister group to all other archostematan beetles. However, other authors have considered their placement within beetles as a whole uncertain, due to their mix characteristics of typical Archostemata, as well as Polyphaga and Adephaga.

<span class="mw-page-title-main">Scirtoidea</span> Superfamily of beetles

Scirtoidea is a superfamily of beetles. It is traditionally considered to consist of four families: Clambidae, Decliniidae, Eucinetidae and Scirtidae. However, genetic studies have suggested that Clambidae and Eucinetidae belong to a separate superfamily Clamboidea, which also includes Derodontidae. Scirtoidea and Clamboidea are the two earliest diverging lineages of living polyphagans.

<span class="mw-page-title-main">Ommatidae</span> Family of beetles

The Ommatidae are a family of beetles in the suborder Archostemata. The Ommatidae are considered the extant beetle family that has most ancestral characteristics. There are only seven extant species, confined to Australia and South America. However, the geographical distribution was much wider during the Mesozoic spanning across Eurasia and Australia, suggesting that they were widespread on Pangea. So far, over 26 extinct genera containing over 170 species of these beetles have been described. Three extant genera have been assigned to this family: Omma,Tetraphalerus and Beutelius. The family is considered to be a subfamily of Cupedidae by some authors, but have been found to be more closely related to Micromalthidae in molecular phylogenies. A close relationship with Micromalthidae is supported by several morphological characters, including those of the mandibles and male genitalia. Due to their rarity, their ecology is obscure, it is likely that their larvae feed on deadwood.

<span class="mw-page-title-main">Sphindidae</span> Family of beetles

Sphindidae is a family of beetles, in the suborder Polyphaga. They are called slime mold beetles due to their exclusive feeding on slime molds during adult and larval stages, other aspects of their life history are obscure. Palaeontological discoveries since 2015 have added to the geologic history of Sphindidae, including the discovery of Libanopsis, placed in the extinct subfamily Libanopsinae.

Rhabdocupes is an extinct genus of beetles in the family Ommatidae, known from the Late Triassic Madygen Formation of Kyrgyzstan, containing the following species:

Zygadenia is an extinct genus of archostematan beetles from the Jurassic to Cretaceous. It is considered to be a senior synonym of Notocupes by Kirejtshuk (2020), but other researchers suggest to reserve the genus Zygadenia as a form taxon for isolated elytra that probably belong to the genus Notocupes, while retaining Notocupes as a valid genus for complete body fossils.

<i>Omma</i> Genus of beetles

Omma is a genus of beetles in the family Ommatidae. Omma is an example of a living fossil. The oldest species known, O. liassicum, lived during the final stage of the Triassic (Rhaetian), over 200 million years ago, though the placement of this species in Omma has been questioned. Numerous other fossil species are known from the Jurassic and Cretaceous of Europe and Asia. The only living species is Omma stanleyi, which is endemic to Australia. Three other extant species endemic to Australia that were formerly part of this genus were moved to the separate genus Beutelius in 2020.Omma stanleyi is strongly associated with wood, being found under Eucalyptus bark and exhibiting thanatosis when disturbed. Its larval stage and many other life details are unknown due to its rarity. Males are typically 14–20 mm in length, while females are 14.4-27.5 mm. Omma stanleyi occurs throughout eastern Australia from Victoria to Central Queensland.

<span class="mw-page-title-main">Permocupedidae</span> Extinct family of beetles

Permocupedidae is a family of Protocoleopteran stem group beetles. They first appeared during the Early Permian, and were one of the dominant groups of beetles during the Middle Permian. They became rare in the Late Permian, with only one species known from the Triassic, Frankencupes ultimus from the Middle Triassic (Anisian) Röt Formation of Germany. They are thought to have been xylophagous, which is presumed to be the ancestral ecology of beetles.

Ademosynidae is an extinct family of beetles, known from the Late Triassic to Early Cretaceous. There are at least six genera and 11 species in Ademosynidae. Members of the family were small oval beetles, with a length generally smaller than 1 cm. Characteristics of the family include a pronotum without anterior angles and a rounded anterior margin, and elytra with 9–12 punctate striae. The systematic position of the family within the order Coleoptera is currently uncertain: they can be considered early examples of the suborder Polyphaga, or unusual examples of Archostemata.

<span class="mw-page-title-main">Permopsocida</span> Extinct genus of insects

Permopsocida is an extinct order of insects known from the Early Permian to the Mid-Cretaceous. It is part of Paraneoptera, alongside bark lice, bugs and thrips. Within Paraneoptera it is considered to be closer to the clade containing bugs and thrips rather than bark lice, with an estimated divergence during the Late Carboniferous. The group was first named as a suborder by Robert John Tillyard in 1926, and was raised to a full order by Huang et al. in 2016. It is currently divided up into three families, Psocidiidae which is known from the Permian to Liassic. Permopsocidae which is only known from the Permian, and Archipsyllidae, which is known from the Late Triassic to mid-Cretaceous (Cenomanian). While most members of the group are known from compression fossils, several members of Archipsyllidae are 3 dimensionally preserved in Burmese amber, which has helped clarify the morphology and phylogenetic position of the group. The morphology of the mouthparts suggests that they were capable of suction feeding and chewing, with preserved angiosperm pollen grains in the gut of Psocorrhyncha suggesting that at least some members of the group were pollenivorous.

<span class="mw-page-title-main">Tshekardocoleidae</span> Extinct family of beetles

Tshekardocoleidae is an extinct family of beetles, known from the Permian. They represent amongst the oldest beetles. Like other primitive beetles, they are thought to have been xylophagous. They first appeared during the Cisuralian, before becoming extinct at the beginning of the Guadalupian. A claimed Jurassic record is doubtful. The oldest known beetle, Coleopsis, was originally assigned to this family, but is now assigned to its own family Coleopsidae.

<span class="mw-page-title-main">Taldycupedidae</span> Extinct family of beetles

Taldycupedidae is an extinct family of beetles primarily known from the Permian period.

Lepidomma is an extinct genus of ommatid beetle. The genus was first described in 2019 for the species L. tianae. Lepidomma was synonymised with Clessidromma by Kirejtshuk, 2020. This synonymy was disputed by Li et al. (2021), who maintained Lepidomma as a separate genus from Clessidromma. Three additional species of Lepidomma were described in 2020 and 2022. All four species are known from the Cenomanian aged Burmese amber of Myanmar.

<i>Notocupes</i> Extinct genus of beetles

Notocupes is an extinct genus of medium-sized archostematan beetles from the Mesozoic Era of Eurasia, including over 50 described species. Historically, the genus was classified as a member of the family Ommatidae, but the presence of characters such as the horizontal mandibular cutting edge, separated procoxae and overlapping abdominal sternites indicate that the genus may have a closer affinity with the family Cupedidae. Notocupes is considered to be a junior synonym of Zygadenia by Kirejtshuk (2020), but other researchers suggest to reserve the genus Zygadenia as a form taxon for isolated elytra that probably belong to the genus Notocupes, while retaining Notocupes as a valid genus for complete body fossils. Most species of Notocupes were described from compression fossils. An additional three species were described from Cenomanian-aged Burmese amber, which were treated as a separate genus, Echinocups, by Kirejtshuk (2020), but Li et al. (2023) consider Echinocups to be a junior synonym of Notocupes. Notocupes has a flattened body, which may suggest that it occupied narrow habitats, such as living under bark. Some species had serrated/spined margins of the carapace, which may have served as a defense against predators, or served as camouflage to resemble bark.

References

  1. Tillyard, R.J. (1924). "Upper Permian Coleoptera and a new order from the Belmont Beds, New South Wales". Proceedings of the Linnean Society of New South Wales . 49: 429–435.
  2. 1 2 Zhao, Xianye; Yu, Yilun; Clapham, Matthew E; Yan, Evgeny; Chen, Jun; Jarzembowski, Edmund A; Zhao, Xiangdong; Wang, Bo (2021-11-08). Perry, George H; Fikacek, Martin (eds.). "Early evolution of beetles regulated by the end-Permian deforestation". eLife. 10: e72692. doi: 10.7554/eLife.72692 . ISSN   2050-084X. PMC   8585485 . PMID   34747694.
  3. 1 2 Alexander G. Kirejtshuk; Markus Poschmann; Jakub Prokop; Romain Garrouste; André Nel (2014). "Evolution of the elytral venation and structural adaptations in the oldest Palaeozoic beetles (Insecta: Coleoptera: Tshekardocoleidae)". Journal of Systematic Palaeontology. 12 (5): 575–600. doi:10.1080/14772019.2013.821530. S2CID   85163674.
  4. Crowson, R.A. (1981). "Chapter 20: Evolutionary History of Beetles". The Biology of the Coleoptera (PDF). London: Academic Press. pp. 658–688.
  5. 1 2 Cai, C.; Tihelka, E.; Giacomelli, M.; Lawrence, J.F.; Ślipiński, A.; Kundrata, R.; Yamamoto, S.; Thayer, M.K.; Newton, A.F.; Leschen, R.A.B.; Gimmel, M.L.; Lü, L.; Engel, M.S.; Bouchard, P.; Huang, D.; Pisani, D.; Donoghue, P.C.J. (2022). "Integrated phylogenomics and fossil data illuminate the evolution of beetles". Royal Society Open Science . 9 (3): 211771. doi: 10.1098/rsos.211771 . PMC   8941382 . PMID   35345430.
  6. Kirejtshuk, Alexander G. (2020-02-17). "Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea". Geosciences. 10 (2): 73. Bibcode:2020Geosc..10...73K. doi: 10.3390/geosciences10020073 . ISSN   2076-3263.
  7. 1 2 Boudinot, Brendon Elias; Yan, Evgeny Viktorovich; Prokop, Jakub; Luo, Xiao‐Zhu; Beutel, Rolf Georg (2022-07-28). "Permian parallelisms: Reanalysis of †Tshekardocoleidae sheds light on the earliest evolution of the Coleoptera". Systematic Entomology. 48: 69–96. doi: 10.1111/syen.12562 . ISSN   0307-6970. S2CID   251171914.