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Zoraptera

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Zoraptera
Temporal range:Albian–Present
Zorotypusfrom Los Bancos, Pichincha, Ecuador
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Subclass: Pterygota
Infraclass: Neoptera
Cohort: Polyneoptera
Order: Zoraptera
Silvestri,1913
Families
Diversity
51 species

TheinsectorderZoraptera,commonly known asangel insects,contains small and soft bodied insects with two forms: winged with wings sheddable as intermites,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 inleaf litter.[1]

Winged fossil ofZorotypus hirsutusfrom theLate Cretaceous(Cenomanian) agedBurmese amber,around 99 million years old

Description[edit]

Zorotypussp.

The name Zoraptera, given byFilippo Silvestriin 1913,[2]is misnamed and potentially misleading: "zor" is Greek for pure and "aptera" means wingless. "Pure wingless" clearly does not fit the wingedalateforms, 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 resembletermitesin appearance and in their gregarious behavior. They are short and swollen in appearance. They belong to thehemimetabolousinsects. They possessmandibulatedbiting mouthparts, shortcerci(usually 1 segment only), and shortantennaewith 9 segments. The abdomen is segmented in 11 sections.[3]The maxillarypalpshave five segments, labial palps three, in both the most distal segment is enlarged. They have sixMalpighian tubules,and their abdominal ganglia have fused into two separate ganglionic complexes.[4]Immature nymphs resemble small adults. Each species showspolymorphism.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 havecompound eyesandocelli,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 simplevenation.[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[edit]

Phylogeny[edit]

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

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

Polyneoptera

Zoraptera (angel insects)

Dermaptera(earwigs)

Plecoptera(stoneflies)

Orthoptera(grasshoppers, crickets, katydids)

Grylloblattodea(ice crawlers)

Mantophasmatodea(gladiators)

Phasmatodea(stick insects)

Embioptera(webspinners)

Dictyoptera

Mantodea(mantises)

Blattodea(cockroaches and termites)

Classification[edit]

The Zoraptera are currently divided into twofamilies,foursubfamilies,ninegeneraand a total of 51species,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 fromBurmese amber.[15]

Incertae sedis[edit]

The following nine species are considered Zorapteraincertae sedis:[12]

Extinct taxa[edit]

Behavior and ecology[edit]

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

Centrozoros gurneyilives 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 ofUsazoros hubbardiare 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 barberilack 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]

InSpermozoros impolitus,copulation does not occur, but fertilization is accomplished instead by transfer of aspermatophorefrom 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[edit]

Zorapterans are thought to provide some important services to ecosystems. By consumingdetritus,such as dead arthropods, they assist indecompositionandnutrient cycling.[25]

References[edit]

  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.S2CID85821812.
  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. ^abGullan; 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.PMID34781597.
  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.ISBN978-1-4549-2323-7.{{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.S2CID53321436.
  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.PMID16005647.
  8. ^Engel, MS; Grimaldi, DA (2002). "The first mesozoic Zoraptera (Insecta)".American Museum Novitates(3362): 1–20.CiteSeerX10.1.1.571.3443.doi:10.1206/0003-0082(2002)362<0001:tfmzi>2.0.co;2.S2CID54764188.
  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.PMID21075208.
  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 Enigma tic insect order Zoraptera and Dictyoptera".PLOS ONE.8(1): e53679.Bibcode:2013PLoSO...853679W.doi:10.1371/journal.pone.0053679.PMC3536744.PMID23301099.
  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.PMC6386694.PMID30642969.
  12. ^abKočárek, Petr; Horká, Ivona; Kundrata, Robin (12 January 2020)."Molecular Phylogeny and Infraordinal Classification of Zoraptera (Insecta)".Insects.11(51): 51.doi:10.3390/insects11010051.PMC7023341.PMID31940956.
  13. ^Zoraptera Species File (Version 5.0/5.0; retrieved 29 May 2021)
  14. ^"Order Zoraptera Silvestri 1913".The Paleobiology Database.Retrieved26 November2020.
  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 ofZorotypus juninensisEngel, 2000, syn. nov. revealed: it is conspecific withCentrozoros 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 ofFormosozoros newiChao & Chen, 2000 ".Insects.14(1): 53.doi:10.3390/insects14010053.PMC9865158.
  18. ^abChoe, 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.ISBN978-0-511-72194-6.
  19. ^Encyclopedia of Insects(2nd ed.). Academic Press. 2009. pp. Ch. 272.ISBN978-0-12-374144-8.
  20. ^The Other Insect Societies
  21. ^Choe, Jae C. (1994)."Sexual selection and mating system inZorotypus gurneyiChoe (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.ISSN0340-5443.S2CID42298642.
  22. ^Choel, Jae C. (1994). "Sexual selection and mating system inZorotypus gurneyiChoe (Insecta: Zoraptera), I. Dominance hierarchy and mating success ".Behavioral Ecology and Sociobiology.34(2): 87–93.doi:10.1007/bf00164179.hdl:2027.42/46900.ISSN0340-5443.S2CID9112078.
  23. ^Dallai, R.; et al. (12 May 2013). "Divergent mating patterns and a unique mode of external sperm transfer in Zoraptera: an Enigma tic group of pterygote insects".Naturwissenschaften.100(6): 581–594.Bibcode:2013NW....100..581D.doi:10.1007/s00114-013-1055-0.ISSN0028-1042.PMID23666111.S2CID16363067.
  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[edit]

External links[edit]

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