Biology:Coleopsis

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Coleopsis is an extinct genus of stem-group beetles. It contains a single species, Coleopsis archaica, and is the only member of the family Coleopsidae and superfamily Coleopsoidea. It is known from a single specimen from the Early Permian of southwestern Germany, estimated to be about 297 million years old. It is currently the oldest known beetle.[1]

Etymology

The generic name, Coleopsis, is derived from the Greek words κολεός (koleós, meaning "sheath, scabbard") and ὄψῐς (ópsĭs, meaning "aspect, appearance"). The specific name, archaica, is derived from the Latin word archaicus (meaning "archaic"), used to mean that the species is an ancient, archaic beetle.[1]

Fossil locality and age

The single specimen used to describe Coleopsis archaica, ZfB 3315, comes from a small outcrop (exposed bedrock) south of the village of Grügelborn, approximately 6 kilometres (3.7 mi) north-northeast from the town of Sankt Wendel, in northeastern Saarland, Germany. The rocks belong to the Humberg Bed, located in the uppermost part of the Meisenheim Formation, which is part of the Rotliegend lithostratigraphic unit. Slightly older sediments from the same formation were dated to 297.0 ± 3.2 Ma, suggesting that the Coleopsis archaica fossil dates to the latest Asselian age or the earliest Sakmarian age of the Early Permian.[2][1]

Description

Coleopsis has a slender body with a total length of 7.8 millimetres (0.31 in) and a maximum width 2.5 millimetres (1⁄8 in), widest slightly anterior to (forward of) the posterior third of the body. The head is prognathous (having forward-facing mandibles) and has eyes that strongly protrude laterally. The antennae are not preserved in the fossil. The pronotum is unusually short and transverse relative to the rest of the body, being more than three times wide as it is long medially, and has flattened lateral processes (projections) with a coarsely serrated margin. Both the head and pronotum are covered with fine tubercules.

Unlike in extant beetles and most stem-group beetles except Tshekardocoleidae, the elytra (hardened forewings) of Coleopsis cover the body loosely, extend beyond the apex of the abdomen, and lack epipleura (outer margins) instead forming flattened lateral flanges. The loose fit of the elytra also means they do not form a tightly sealed subelytral space as in extant beetles. The elytra have several distinct veins spanning their length, as well as several less-distinct shorter veins on the posterior third of the body. "Window punctures" are present but only in a limited area of the elytra; this may either be an autapomorphy of Coleopsis or an artifact caused by the strong compression of the fossil. Window punctures are also present on the elytra of extant families Cupedidae and Ommatidae as well as some other stem-group beetle families such as Tshekardocoleidae and Permocupedidae.[1]

Taxonomy and systematics

Coleopsis was originally described by Kirejtshuk et al. (2014) as a member of the family Tshekardocoleidae within the superfamily Tshekardocoleoidea, itself within a broad concept of the suborder Archostemata. Later, Coleopsis was classified in its own family within Tshekardocoleidea by Kirejtshuk & Nel (2016). It was then placed in its own superfamily by Kirejtshuk (2020), proposed because it was claimed to have more remnants of the archedictyon in the wing venation of its elytra than that of tshekardocoleids.[3][1] In the classifications of Cai et al. (2022) and Bouchard et al. (2024), Coleopsis and Tshekardcoleidae among other stem-group beetles are assigned to the extinct suborder Alphacoleoptera (also known as Protocoleoptera),[4][5] though this is a paraphyletic group.[6][1]

A re-examination of Coleopsis using Reflectance Transformation Imaging by Schädel et al. (2022) found that it is a stem-group beetle, outside of a clade containing all beetles excluding Tshekardocoleidae. However, the authors could not determine if Coleopsis belonged to the Tshekardocoleidae or not, as they could not identify any synapomorphies (shared derived traits) uniting the two taxa and found that Tshekardocoleidae itself is possibly not a monophyletic group.[1]

The phylogenetic tree below shows the possible phylogenetic positions of Coleopsis, after Schädel et al. (2022):

Coleoptera (sensu lato)
 

†Coleopsis(?)

 
†Tshekardocoleidae

†Coleopsis(?)

 
 
 

†Permocupedidae

 
 

Metacoleoptera (all other beetles)

 
 
 

The family name was originally published in 2016 as "Coleopsidae" but Kirejtshuk (2020) claimed that this spelling was erroneous and proposed the spelling "Coleopseidae";[3] however, under the ICZN, Article 29.4, family-rank names originally published after 1999 are not subject to emendation because of incorrect spelling, so Coleopsidae would be maintained as the correct original spelling.[note 1]

References

  1. ↑ 1.0 1.1 1.2 1.3 1.4 1.5 1.6 Schädel, M.; Yavorskaya, M.; Beutel, R. (2022). "The earliest beetle †Coleopsis archaica (Insecta: Coleoptera) – morphological re-evaluation using Reflectance Transformation Imaging (RTI) and phylogenetic assessment". Arthropod Systematics & Phylogeny 80: 495-510. doi:10.3897/asp.80.e86582. 
  2. ↑ Cite error: Invalid <ref> tag; no text was provided for refs named Kirejtshuk2014
  3. ↑ 3.0 3.1 Kirejtshuk, A.G. (2020). "Taxonomic Review of Fossil Coleopterous Families (Insecta, Coleoptera). Suborder Archostemata: Superfamilies Coleopseoidea and Cupedoidea". Geosciences 10 (2): 73. doi:10.3390/geosciences10020073. 
  4. ↑ Cai, Chenyang; Tihelka, Erik; Giacomelli, Mattia; Lawrence, John F.; Ślipiński, Adam; Kundrata, Robin; Yamamoto, Shûhei; Thayer, Margaret K. et al. (March 2022). "Integrated phylogenomics and fossil data illuminate the evolution of beetles" (in en). Royal Society Open Science 9 (3). doi:10.1098/rsos.211771. ISSN 2054-5703. PMID 35345430. Bibcode: 2022RSOS....911771C. 
  5. ↑ Bouchard, Patrice; Bousquet, Yves; Davies, Anthony E.; Cai, Chenyang (2024-03-13). "On the nomenclatural status of type genera in Coleoptera (Insecta)". ZooKeys (1194): 1–981. doi:10.3897/zookeys.1194.106440. ISSN 1313-2970. PMID 38523865. PMC 10955229. Bibcode: 2024ZooK.1194....1B. https://zookeys.pensoft.net/article/106440/. 
  6. ↑ 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" (in en). Systematic Entomology 48: 69–96. doi:10.1111/syen.12562. ISSN 0307-6970. 

Notes

  1. ↑ ICZN Code 29.4. Acceptance of originally formed stem. If after 1999 a new family-group name is based on a generic name which is or ends in a Greek or Latin word or ends in a Greek or Latin suffix, but its derivation does not follow the grammatical procedures of Articles 29.3.1 or 29.3.2, its original spelling must be maintained as the correct original spelling

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