Zoraptera

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Zoraptera
Temporal range: Albian–Present
Zorotypus from Los Bancos, Pichincha, Ecuador.jpg
Zorotypus from Los Bancos, Pichincha, Ecuador
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Subclass: Pterygota
Infraclass: Neoptera
Cohort: Polyneoptera
Order: Zoraptera
Silvestri, 1913
Families
Diversity
51 species

The insect order Zoraptera, commonly known as angel insects, contains small and soft bodied insects with two forms: winged with wings sheddable as in termites, dark and with eyes (compound) and ocelli (simple); or wingless, pale and without eyes or ocelli. They have a characteristic nine-segmented beaded (moniliform) antenna. They have mouthparts adapted for chewing and are mostly found under bark, in dry wood or in leaf litter. [1]

Contents

Winged fossil of Zorotypus hirsutus from the Late Cretaceous (Cenomanian) aged Burmese amber, around 99 million years old Zorotypus hirsutus Fig1 A.jpg
Winged fossil of Zorotypus hirsutus from the Late Cretaceous (Cenomanian) aged Burmese amber, around 99 million years old

Description

Zorotypus sp. Zorotypus.jpg
Zorotypus sp.

The name Zoraptera, given by Filippo Silvestri in 1913, [2] is misnamed and potentially misleading: "zor" is Greek for pure and "aptera" means wingless. "Pure wingless" clearly does not fit the winged alate forms, which were discovered several years after the wingless forms had been described.

The members of this order are small insects, 3 millimetres (0.12 in) or less in length, that resemble termites in appearance and in their gregarious behavior. They are short and swollen in appearance. They belong to the hemimetabolous insects. They possess mandibulated biting mouthparts, short cerci (usually 1 segment only), and short antennae with 9 segments. The abdomen is segmented in 11 sections. [3] The maxillary palps have five segments, labial palps three, in both the most distal segment is enlarged. They have six Malpighian tubules, and their abdominal ganglia have fused into two separate ganglionic complexes. [4] Immature nymphs resemble small adults. Each species shows polymorphism. Most individuals are the apterous form or "morph", with no wings, no eyes, and no or little pigmentation. A few females and even fewer males are in the alate form with relatively large membranous wings that can be shed at a basal fracture line. Alates also have compound eyes and ocelli, and more pigmentation. This polymorphism can be observed already as two forms of nymphs. Wingspan can be up to 7 millimetres (0.28 in), and the wings can be shed spontaneously. When observed, wings are paddle shaped and have simple venation. [3] Under good conditions the blind and wingless form predominates, but if their surroundings become too tough, they produce offspring which develop into winged adults with eyes. These winged offspring are then able to disperse and establish new colonies in areas with more resources. Once established, future generations are once again born blind and wingless. [5]

Systematics

Phylogeny

The phylogenetic relationship of the order remains controversial and elusive. At present the best supported position based on morphological traits recognizes the Zoraptera as polyneopterous insects related to the webspinners of the order Embioptera. However, molecular analysis of 18s ribosomal DNA supports a close relationship with the superorder Dictyoptera. [6] [7] [8] [9] [10]

The following cladogram, based on the molecular phylogeny of Wipfler et al. 2019, places Zoraptera as the sister group of Dermaptera (earwigs); Zoraptera and Dermaptera together form the sister group of the remaining Polyneoptera: [11]

Polyneoptera

Zoraptera (angel insects) Zorotypus guineensis (white background).jpg

Dermaptera (earwigs) Earwig on white background.jpg

Plecoptera (stoneflies) Neoperla clymene hor.png

Orthoptera (grasshoppers, crickets, katydids) Gryllidae usda.png

Grylloblattodea (ice crawlers) Grylloblattidae (white background).jpg

Mantophasmatodea (gladiators) Mantophasma zephyra (white background).jpg

Phasmatodea (stick insects) Stick insect line diagram.png

Embioptera (webspinners) Embia major hor.png

Dictyoptera

Mantodea (mantises) Stagmomantis carolina usda hor.png

Blattodea (cockroaches and termites) Temnopteryx species Zebra Cockroach (white background).jpg

Classification

The Zoraptera are currently divided into two families, four subfamilies, nine genera and a total of 51 species, some of which have not been yet described. [12] [13] [14] There are eleven extinct species known as of 2017, many of the fossil species are known from Burmese amber. [15]

Incertae sedis

The following nine species are considered Zoraptera incertae sedis : [12]

Extinct taxa

Behavior and ecology

Zorapterans live in small colonies beneath rotting wood, lacking in mouthparts able to tunnel into wood, but feeding on fungal spores and detritus. These insects can also hunt smaller arthropods like mites and collembolans. [18] Much of their time is spent grooming themselves or others. [19]

Centrozoros gurneyi lives in colonies which range in size from a few dozen to several hundred individuals, but most often number about 30 individuals. The males are slightly larger than the females, and they fight for dominance. [20]

When two colonies of Usazoros hubbardi are brought together experimentally, there is no difference in behavior towards members of the new colony. Therefore, colonies in the wild might merge easily. Winged forms are rare. The males in most colonies establish a linear dominance hierarchy in which age or duration of colony membership is the prime factor determining dominance. Males appearing later in colonies are at the bottom of the hierarchy, regardless of their body size. By continually attacking other males, the dominant male monopolizes a harem of females. The members of this harem stay clumped together. There is a high correlation between rank and reproductive success of the males. [21] [22]

Latinozoros barberi lack such a dominance structure but display complex courtship behavior including nuptial feeding. The males possess a cephalic gland that opens in the middle of their head. During courtship they secrete a fluid from this gland and offer it to the female. Acceptance of this droplet by the female acts as behavioral releaser and immediately leads to copulation. [18]

In Spermozoros impolitus , copulation does not occur, but fertilization is accomplished instead by transfer of a spermatophore from the male to the female. This 0.1-millimetre (0.0039 in) spermatophore contains a single giant sperm cell, which unravels to about the same length as the female herself, 3 millimetres (0.12 in). It is thought that this large sperm cell prevents fertilization by other males, by physically blocking the female's genital tract. [23] [24]

Effects on ecosystem

Zorapterans are thought to provide some important services to ecosystems. By consuming detritus, such as dead arthropods, they assist in decomposition and nutrient cycling. [25]

Related Research Articles

<i>Zorotypus</i> Genus of angel insects

Zorotypus is a genus of angel insects in the family Zorotypidae.

<span class="mw-page-title-main">Embioptera</span> Order of insects

The order Embioptera, commonly known as webspinners or footspinners, are a small group of mostly tropical and subtropical insects, classified under the subclass Pterygota. The order has also been called Embiodea or Embiidina. More than 400 species in 11 families have been described, the oldest known fossils of the group being from the mid-Jurassic. Species are very similar in appearance, having long, flexible bodies, short legs, and only males having wings.

<span class="mw-page-title-main">Rhopalosomatidae</span> Family of insects

Rhopalosomatidae is a family of Hymenoptera containing about 68 extant species in four genera that are found worldwide. Three fossil genera are known.

Michael S. Engel, FLS, FRES is an American paleontologist and entomologist, notable for contributions to insect evolutionary biology and classification. In connection with his studies he has undertaken field expeditions in Central Asia, Asia Minor, the Levant, Arabia, eastern Africa, the high Arctic, and South and North America, and has published more than 925 papers in scientific journals.Some of Engel's research images were included in exhibitions on the aesthetic value of scientific imagery.

Spiralizoros silvestrii is a species of angel insect in the family Zorotypidae. It is found in Southern Asia.

Spermozoros impolitus is a species of insect in the order Zoraptera.

Spiralizoros cervicornis is a species of insect in the order Zoraptera. It was first found in Malaysia.

Spiralizoros magnicaudelli is a species of insect in the order Zoraptera. It was first found in Malaysia.

<i>Usazoros</i> Genus of insects

Usazoros hubbardi, commonly known as Hubbard's angel insect, is a species of insect in the order Zoraptera. It is native to the tropical and subtropical New World and has expanded its range into the eastern United States, where it lives in piles of sawdust, whereas in the hotter part of its range it lives under the bark of decomposing logs. It was named after the American entomologist Henry Guernsey Hubbard, who discovered the insect in the United States.

2019 in paleoentomology is a list of new fossil insect taxa that were described during the year 2019, as well as other significant discoveries and events related to paleoentomology that were scheduled to occur during the year.

Burmese amber is fossil resin dating to the early Late Cretaceous Cenomanian age recovered from deposits in the Hukawng Valley of northern Myanmar. It is known for being one of the most diverse Cretaceous age amber paleobiotas, containing rich arthropod fossils, along with uncommon vertebrate fossils and even rare marine inclusions. A mostly complete list of all taxa described up until 2018 can be found in Ross 2018; its supplement Ross 2019b covers most of 2019.

<span class="mw-page-title-main">Elcanidae</span> Extinct family of cricket-like animals

Elcanidae are an extinct family of Mesozoic and early Cenozoic orthopterans. Members of the family are distinguished by the presence of spurs on the distal part of the metatibia, unique among orthopterans, these have been suggested to have been used for controlling gliding, swimming aids, or for jumping on water. The group combines characteristics from both major groups of orthopterans, with long antennae and nymphal morphology similar to Ensifera, but with wing venation and adult morphology more similar to Caelifera. Elcanidae is part of Elcanoidea, which is thought to have diverged from living orthopterans by the beginning of the Permian, around 300 million years ago. The family also includes Permelcanidae, known from the Early-Late Permian. The relationship of Elcanoidea to Ensifera and Caelifera is currently unresolved. Elcanids are known from the Late Triassic to Paleocene of Eurasia, North and South America. Some members of the group exhibited aposematic coloration.

Burmese amber is fossil resin dating to the early Late Cretaceous Cenomanian age recovered from deposits in the Hukawng Valley of northern Myanmar. It is known for being one of the most diverse Cretaceous age amber paleobiotas, containing rich arthropod fossils, along with uncommon vertebrate fossils and even rare marine inclusions. A mostly complete list of all taxa described up until 2018 can be found in Ross 2018; its supplement Ross 2019b covers most of 2019.

Jouault, Corentin; Rosse-Guillevic, Simon. "A new genus of praeaulacid wasp from the mid-Cretaceous Kachin amber (Myanmar)". Annales de Paléontologie. 109 (1): 102599. doi:10.1016/j.annpal.2023.102599.

This paleoentomology list records new fossil insect taxa that were to be described during the year 2021, as well as notes other significant paleoentomology discoveries and events which occurred during that year.

Spiralizoros is a genus of zorapterans in the family Spiralizoridae. There are about eight described species in Spiralizoros. The species of this genus were transferred from the genus Zorotypus as a result of research published in 2020.

Spermozoros is a genus of angel insects in the family Zorotypidae. There are six described species in Spermozoros, transferred to Spermozoros from the genus Zorotypus as a result of research published in 2020.

This paleoentomology list records new fossil insect taxa that are to be described during the year 2022, as well as notes other significant paleoentomology discoveries and events which occurred during that year.

This list of 2023 in paleoentomology records new fossil insect taxa that are to be described during the year, as well as documents significant paleoentomology discoveries and events which occurred during that year.

This list of 2024 in paleoentomology records new fossil insect taxa that are to be described during the year, as well as documents significant paleoentomology discoveries and events which occurred during that year.

References

  1. ↑ Rafael, JA; Godoi, FDP; Engel, MS (2008). "A new species of Zorotypus from eastern Amazonia, Brazil (Zoraptera: Zorotypidae)". Transactions of the Kansas Academy of Science. 111 (3 & 4): 193–202. doi:10.1660/0022-8443-111.3.193. S2CID   85821812.
  2. ↑ Silvestri, F. (1913). "Descrizione di un nuovo ordine di insetti". Bollettino del Laboratorio di Zoologia Generale e Agraria della R. Scuola Superiore d'Agricoltura in Portici. 7: 193–209.
  3. 1 2 Gullan; Granston (2005). The Insects: An Outline of Entomology.
  4. ↑ Friedemann, Katrin; Spangenberg, Rico; Yoshizawa, Kazunori; Beutel, Rolf G. (2014). "Evolution of attachment structures in the highly diverse <SCP>A</SCP>cercaria (<SCP>H</SCP>exapoda)". Cladistics. 30 (2): 170–201. doi:10.1111/cla.12030. PMID   34781597.
  5. ↑ Engel, Michael S. (October 9, 2018). Innumerable Insects: The Story of the Most Diverse and Myriad Animals on Earth (Natural Histories). Union Square & Co. ISBN   978-1-4549-2323-7.{{cite book}}: CS1 maint: date and year (link)
  6. ↑ Yoshizawa (2007). "The Zoraptera problem: evidence for Zoraptera + Embiodea from the wing base" (PDF). Systematic Entomology. 32 (2): 197–204. doi:10.1111/j.1365-3113.2007.00379.x. hdl: 2115/33766 . S2CID   53321436.
  7. ↑ Yoshizawa, K; Johnson, KP (2005). "Aligned 18S for Zoraptera (Insecta): Phylogenetic position and molecular evolution". Molecular Phylogenetics and Evolution. 37 (2): 572–580. doi:10.1016/j.ympev.2005.05.008. hdl: 2115/43133 . PMID   16005647.
  8. ↑ Engel, MS; Grimaldi, DA (2002). "The first mesozoic Zoraptera (Insecta)". American Museum Novitates (3362): 1–20. CiteSeerX   10.1.1.571.3443 . doi:10.1206/0003-0082(2002)362<0001:tfmzi>2.0.co;2. S2CID   54764188.
  9. ↑ Ishiwata, K; Sasaki, G; Ogawa, J; Miyata, T; Su, Z-H (2011). "Phylogenetic relationships among insect orders based on three nuclear protein-coding gene sequences". Molecular Phylogenetics and Evolution. 58 (2): 169–180. doi:10.1016/j.ympev.2010.11.001. PMID   21075208.
  10. ↑ Wang, X.; Engel, M.S.; Rafael, J.A.; Dang, K.; Wu, H.; Wang, Y.; Xie, Q.; Bu, W. (2013). "A unique box in 28S rRNA is shared by the enigmatic insect order Zoraptera and Dictyoptera". PLOS ONE. 8 (1): e53679. Bibcode:2013PLoSO...853679W. doi: 10.1371/journal.pone.0053679 . PMC   3536744 . PMID   23301099.
  11. ↑ Wipfler, Benjamin; Letsch, Harald; Frandsen, Paul B.; Kapli, Paschalia; Mayer, Christoph; Bartel, Daniela; Buckley, Thomas R.; Donath, Alexander; Edgerly-Rooks, Janice S.; Fujita, Mari; Liu, Shanlin (February 2019). "Evolutionary history of Polyneoptera and its implications for our understanding of early winged insects". Proceedings of the National Academy of Sciences. 116 (8): 3024–3029. Bibcode:2019PNAS..116.3024W. doi: 10.1073/pnas.1817794116 . PMC   6386694 . PMID   30642969.
  12. 1 2 Kočárek, Petr; Horká, Ivona; Kundrata, Robin (12 January 2020). "Molecular Phylogeny and Infraordinal Classification of Zoraptera (Insecta)". Insects. 11 (51): 51. doi: 10.3390/insects11010051 . PMC   7023341 . PMID   31940956.
  13. ↑ Zoraptera Species File (Version 5.0/5.0; retrieved 29 May 2021)
  14. ↑ "Order Zoraptera Silvestri 1913". The Paleobiology Database . Retrieved 26 November 2020.
  15. ↑ Yin, Ziwei; Cai, Chenyang; Huang, Diying (2018). "New zorapterans (Zoraptera) from Burmese amber suggest higher paleodiversity of the order in tropical forests". Cretaceous Research. 84: 168–172. doi: 10.1016/j.cretres.2017.11.028 .
  16. ↑ Kočárek, P.; Horka, I. (2022). "Identity of Zorotypus juninensis Engel, 2000, syn. nov. revealed: it is conspecific with Centrozoros hamiltoni (New, 1978) (Zoraptera, Spiralizoridae)". Deutsche Entomologische Zeitschrift . 69 (1): 65–70. doi: 10.3897/dez.69.83154 .
  17. ↑ Kočárek, P.; Hu, F.-S. (2023). "An Immature Dermapteran Misidentified as an Adult Zorapteran: The Case of Formosozoros newi Chao & Chen, 2000". Insects. 14 (1): 53. doi: 10.3390/insects14010053 . PMC   9865158 .
  18. 1 2 Choe, Jae C. (1997). "The evolution of mating systems in the Zoraptera: mating variations and sexual conflicts". In Choe, Jae C.; Crespi, Bernard J. (eds.). The Evolution of Mating Systems in Insects and Arachnids. Cambridge: Cambridge University Press. pp. 130–145. doi:10.1017/cbo9780511721946.008. ISBN   978-0-511-72194-6.
  19. ↑ Encyclopedia of Insects (2nd ed.). Academic Press. 2009. pp. Ch. 272. ISBN   978-0-12-374144-8.
  20. ↑ The Other Insect Societies
  21. ↑ Choe, Jae C. (1994). "Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta: Zoraptera)" (PDF). Behavioral Ecology and Sociobiology, II. Determinants and Dynamics of Dominance. 34 (4): 233–237. doi:10.1007/bf00183473. hdl: 2027.42/46900 . ISSN   0340-5443. S2CID   42298642.
  22. ↑ Choel, Jae C. (1994). "Sexual selection and mating system in Zorotypus gurneyi Choe (Insecta : Zoraptera), I. Dominance hierarchy and mating success". Behavioral Ecology and Sociobiology. 34 (2): 87–93. doi:10.1007/bf00164179. hdl: 2027.42/46900 . ISSN   0340-5443. S2CID   9112078.
  23. ↑ Dallai, R.; et al. (12 May 2013). "Divergent mating patterns and a unique mode of external sperm transfer in Zoraptera: an enigmatic group of pterygote insects". Naturwissenschaften. 100 (6): 581–594. Bibcode:2013NW....100..581D. doi:10.1007/s00114-013-1055-0. ISSN   0028-1042. PMID   23666111. S2CID   16363067.
  24. ↑ "The tiny insect with the massive sperm". New Scientist. No. 2919. 1 June 2013. p. 17.
  25. ↑ Engel, Michael (2007). "The Zorotypidae of Fiji (Zoraptera)" (PDF). Bishop Museum Occasional Papers.

General references

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