Zoraptera

{{Short description|Order of insects}} {{Automatic taxobox | fossil_range = {{fossilrange|Albian|Present}} | image = Zorotypus from Los Bancos, Pichincha, Ecuador.jpg | image_caption = ''Zorotypus'' from Los Bancos, Pichincha, Ecuador | display_parents = 3 | taxon = Zoraptera | authority = Silvestri, 1913 | subdivision_ranks = Families | subdivision = * Zorotypidae * Spiralizoridae | diversity = 51 species }}

The insect order '''Zoraptera''', commonly known as '''angel insects''' and sometimes '''ground lice''',<ref>{{cite journal | last1=Matsumura | first1=Y. | last2=Lima | first2=S. P. | last3=Rafael | first3=J. A. | last4=Câmara | first4=J. T. | last5=Beutel | first5=R. G. | last6=Gorb | first6=S. N. | title=Distal leg structures of Zoraptera - did the loss of adhesive devices curb the chance of diversification? | journal=Arthropod Structure & Development | date=2022 | volume=68 | article-number=101164 | doi=10.1016/j.asd.2022.101164 | pmid=35468454 | url=https://www.sciencedirect.com/science/article/abs/pii/S1467803922000251 | url-access=subscription }}</ref> 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. Their mouthparts are adapted for chewing and are mostly found under bark, in dry wood or in leaf litter.<ref name="Rafael2008">{{cite journal | last1 = Rafael | first1 = JA | last2 = Godoi | first2 = FDP | last3 = Engel | first3 = MS | year = 2008 | title = A new species of Zorotypus from eastern Amazonia, Brazil (Zoraptera: Zorotypidae) | journal = Transactions of the Kansas Academy of Science | volume = 111 | issue = 3 & 4| pages = 193–202 | doi=10.1660/0022-8443-111.3.193| s2cid = 85821812 }}</ref> The order is found on most continents, but are absent in places like Canada, New Zealand, Australia and Europe.<ref>{{Cite journal |last1=Kaláb |first1=Oto |last2=Hoffmannova |first2=Johana |last3=Packova |first3=Gabriela |last4=Kočárková |first4=Ivona |last5=Kundrata |first5=Robin |last6=Kočárek |first6=Petr |date=2025 |title=Curated global occurrence dataset of the insect order Zoraptera |url=https://www.nature.com/articles/s41597-025-04696-4 |journal=Scientific Data |language=en |volume=12 |issue=1 |pages=360 |doi=10.1038/s41597-025-04696-4 |pmid=40021691 |pmc=11871360 }}</ref><ref>[https://www.nature.com/articles/s41597-025-04696-4/figures/1 Global distribution for Zoraptera]</ref> [[File:Zorotypus hirsutus Fig1 A.jpg|left|thumb|Winged fossil of ''Zorotypus hirsutus'' from the Late Cretaceous (Cenomanian) aged Burmese amber, around 99 million years old]]

== Description == left|thumb|''Zorotypus'' sp. The name Zoraptera, given by Filippo Silvestri in 1913,<ref name="Silvestri1913">{{cite journal | last1 = Silvestri | first1 = F. | year = 1913 | title = Descrizione di un nuovo ordine di insetti | url = https://archive.org/stream/bollettinodellab71913rscufo#page/193/mode/1up | journal = Bollettino del Laboratorio di Zoologia Generale e Agraria della R. Scuola Superiore d'Agricoltura in Portici | volume = 7 | pages = 193–209 }}</ref> 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, {{convert|3|mm|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.<ref name=":0">{{Cite book|title = The Insects: An Outline of Entomology|last1 = Gullan|year = 2005|last2 = Granston}}</ref> 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.<ref>{{cite journal | url=https://onlinelibrary.wiley.com/doi/pdf/10.1111/cla.12030 | doi=10.1111/cla.12030 | title=Evolution of attachment structures in the highly diverse <SCP>A</SCP>cercaria (<SCP>H</SCP>exapoda) | date=2014 | last1=Friedemann | first1=Katrin | last2=Spangenberg | first2=Rico | last3=Yoshizawa | first3=Kazunori | last4=Beutel | first4=Rolf G. | journal=Cladistics | volume=30 | issue=2 | pages=170–201 | pmid=34781597 }}</ref> 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 {{convert|7|mm|in}}, and the wings can be shed spontaneously. When observed, wings are paddle shaped and have simple venation.<ref name=":0" /> 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.<ref>{{Cite book |last=Engel |first=Michael S. |title=Innumerable Insects: The Story of the Most Diverse and Myriad Animals on Earth (Natural Histories) |date=October 9, 2018 |publisher=Union Square & Co. |isbn=978-1-4549-2323-7 |pages=}}</ref>

== 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.<ref name="Yoshizawa2007">{{cite journal| last1 = Yoshizawa | year = 2007| title = The Zoraptera problem: evidence for Zoraptera + Embiodea from the wing base | url =https://eprints.lib.hokudai.ac.jp/dspace/bitstream/2115/33766/1/SE32-2.pdf | journal = Systematic Entomology | volume = 32 | issue = 2| pages = 197–204 | doi=10.1111/j.1365-3113.2007.00379.x| bibcode = 2007SysEn..32..197Y| hdl = 2115/33766| s2cid = 53321436| hdl-access = free }}</ref><ref name="Yoshizawa2005">{{cite journal | last1 = Yoshizawa | first1 = K | last2 = Johnson | first2 = KP | year = 2005 | title = Aligned 18S for Zoraptera (Insecta): Phylogenetic position and molecular evolution | journal = Molecular Phylogenetics and Evolution | volume = 37 | issue = 2| pages = 572–580 | doi=10.1016/j.ympev.2005.05.008| pmid = 16005647 | bibcode = 2005MolPE..37..572Y | hdl = 2115/43133 | hdl-access = free }}</ref><ref name="Engel2002">{{cite journal | last1 = Engel | first1 = MS | last2 = Grimaldi | first2 = DA | year = 2002 | title = The first mesozoic Zoraptera (Insecta) | journal = American Museum Novitates | issue = 3362 | pages = 1–20 | doi=10.1206/0003-0082(2002)362<0001:tfmzi>2.0.co;2| citeseerx = 10.1.1.571.3443 | s2cid = 54764188 }}</ref><ref name="Ishiwata2011">{{cite journal | last1 = Ishiwata | first1 = K | last2 = Sasaki | first2 = G | last3 = Ogawa | first3 = J | last4 = Miyata | first4 = T | last5 = Su | first5 = Z-H | year = 2011 | title = Phylogenetic relationships among insect orders based on three nuclear protein-coding gene sequences | journal = Molecular Phylogenetics and Evolution | volume = 58 | issue = 2| pages = 169–180 | doi=10.1016/j.ympev.2010.11.001 | pmid=21075208| bibcode = 2011MolPE..58..169I }}</ref><ref>{{cite journal | last1 = Wang | first1 = X. | last2 = Engel | first2 = M.S. | last3 = Rafael | first3 = J.A. | last4 = Dang | first4 = K. | last5 = Wu | first5 = H. | last6 = Wang | first6 = Y. | last7 = Xie | first7 = Q. | last8 = Bu | first8 = W. | year = 2013 | title = A unique box in 28S rRNA is shared by the enigmatic insect order Zoraptera and Dictyoptera | journal = PLOS ONE | volume = 8 | issue = 1| article-number = e53679 | doi = 10.1371/journal.pone.0053679 | pmid=23301099 | pmc=3536744| bibcode = 2013PLoSO...853679W | doi-access = free }}</ref>

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:<ref>{{cite journal |last1=Wipfler|first1=Benjamin|last2=Letsch|first2=Harald|last3=Frandsen|first3=Paul B.|last4=Kapli|first4=Paschalia|last5=Mayer|first5=Christoph|last6=Bartel|first6=Daniela|last7=Buckley|first7=Thomas R.|last8=Donath|first8=Alexander|last9=Edgerly-Rooks|first9=Janice S.|last10=Fujita|first10=Mari|last11=Liu|first11=Shanlin |date=February 2019 |title=Evolutionary history of Polyneoptera and its implications for our understanding of early winged insects |journal=Proceedings of the National Academy of Sciences |volume=116 |issue=8 |pages=3024–3029 |doi=10.1073/pnas.1817794116 |pmid=30642969 |pmc=6386694 |bibcode=2019PNAS..116.3024W |doi-access=free }}</ref>

{{clade |label1=Polyneoptera |1={{clade |1={{clade |1='''Zoraptera''' (angel insects) 60px |2=Dermaptera (earwigs) 60px }} |2={{clade |1=Plecoptera (stoneflies) 70px |2={{clade |1=Orthoptera (grasshoppers, crickets, katydids) 50px |2={{clade |1={{clade |1={{clade |1=Grylloblattodea (ice crawlers) 70px |2=Mantophasmatodea (gladiators) 70px }} |2={{clade |1=Phasmatodea (stick insects) 70px |2=Embioptera (webspinners) 80px }} }} |label2=Dictyoptera |2={{clade |1=Mantodea (mantises) 80px |2=Blattodea (cockroaches and termites) 60px }} }} }} }} }} }}

=== 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.<ref name="Kocarek2020">{{cite journal | first1 = Petr | last1 = Kočárek | first2 = Ivona | last2 = Horká |first3 = Robin | last3 = Kundrata | date = 12 January 2020 | title = Molecular Phylogeny and Infraordinal Classification of Zoraptera (Insecta) | journal = Insects | volume = 11 | issue = 51 | page = 51 | doi = 10.3390/insects11010051 | pmid = 31940956 | doi-access = free | pmc = 7023341 }}</ref><ref>[http://zoraptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1218728 Zoraptera Species File (Version 5.0/5.0; retrieved 29 May 2021)]</ref><ref>{{Cite web |publisher=The Paleobiology Database |title=Order Zoraptera Silvestri 1913 |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=183857&is_real_user=1 |access-date= 26 November 2020}}</ref> There are eleven extinct species known as of 2017, many of the fossil species are known from Burmese amber.<ref>{{Cite journal|last1=Yin|first1=Ziwei|last2=Cai|first2=Chenyang|last3=Huang|first3=Diying|title=New zorapterans (Zoraptera) from Burmese amber suggest higher paleodiversity of the order in tropical forests|journal=Cretaceous Research|volume=84|pages=168–172|doi=10.1016/j.cretres.2017.11.028|year=2018|doi-access=free|bibcode=2018CrRes..84..168Y }}</ref> * Family Zorotypidae {{au|Silvestri, 1913}} ** Subfamily Zorotypinae {{au|Silvestri, 1913}} *** ''Zorotypus'' {{au|Silvestri, 1913}} — 7 spp. *** ''Usazoros'' {{au|Kukalova-Peck & Peck, 1993}} — 1 sp. ** Subfamily Spermozorinae {{au|Kočárek, Horká & Kundrata, 2020}} *** ''Spermozoros'' {{au|Kočárel, Horká & Kundrata, 2020}} — 6 spp. * Family Spiralizoridae {{au|Kočárek, Horká & Kundrata, 2020}} ** Subfamily Latinozorinae {{au|Kočárek, Horká & Kundrata, 2020}} *** ''Latinozoros'' {{au|Kukalova-Peck & Peck, 1993}} — 3 spp. ** Subfamily Spiralizorinae {{au|Kočárek, Horká & Kundrata, 2020}} *** ''Spiralizoros'' {{au|Kočárek, Horká & Kundrata, 2020}} — 12 spp. *** ''Centrozoros'' {{au|Kukalova-Peck & Peck, 1993}} (=''Meridozoros'' {{au|Kukalova-Peck & Peck, 1993}}; ''Floridazoros'' {{au|Kukalova-Peck & Peck, 1993}}) — 8 spp. *** ''Cordezoros'' {{au|Kočárek, Horká & Kundrata, 2020}} — 1 sp. *** ''Scapulizoros'' {{au|Kočárek, Horká & Kundrata, 2020}} — 1 sp. *** ''Brazilozoros'' {{au|Kukalova-Peck & Peck, 1993}} — 3 spp.

==== Incertae sedis ====

The following nine species are considered Zoraptera ''incertae sedis'':<ref name="Kocarek2020" /> * ''Zorotypus congensis'' {{au|van Ryn Tournel, 1971}} – Congo (Dem.Rep.) * ''Zorotypus javanicus'' {{au|Silvestri, 1913}} – Java * ''Zorotypus juninensis'' {{au|Engel, 2000}} (considered a synonym of ''Centrozoros hamiltoni''<ref>{{cite journal|last1=Kočárek|first1=P.|last2=Horka|first2=I.|year=2022|title=Identity of ''Zorotypus juninensis'' Engel, 2000, syn. nov. revealed: it is conspecific with ''Centrozoros hamiltoni'' (New, 1978) (Zoraptera, Spiralizoridae)|journal=Deutsche Entomologische Zeitschrift|volume=69|issue=1|pages=65–70|doi=10.3897/dez.69.83154|doi-access=free}}</ref>) – Peru * ''Zorotypus lawrencei'' New, 1995 – Christmas Island * ''Zorotypus leleupi'' {{au|Weidner, 1976}} – Galápagos Islands * ''Zorotypus longicercatus'' {{au|Caudell, 1927}} – Jamaica * ''Zorotypus newi'' {{au|(Chao & Chen, 2000)}} (=''Formosozoros newi'', is in actuality an immature earwig<ref>{{cite journal|last1=Kočárek|first1=P.|last2=Hu|first2=F.-S.|year=2023|title=An Immature Dermapteran Misidentified as an Adult Zorapteran: The Case of ''Formosozoros newi'' Chao & Chen, 2000|journal=Insects|volume=14|issue=1|page=53|doi=10.3390/insects14010053|doi-access=free|pmid=36661981 |pmc=9865158}}</ref>) – Taiwan * ''Zorotypus sechellensis'' {{au|Zompro, 2005}} – Seychelles * ''Zorotypus swezeyi'' {{au|Caudell, 1922}} – Hawaii

==== Extinct taxa ==== * ''Zorotypus'' Silvestri, 1913 ** Subgenus ''Zorotypus'' Silvestri, 1913 *** {{extinct}}''Zorotypus (Zorotypus) absonus'' Engel, 2008 – Dominican amber, Dominican Republic (Miocene) *** {{extinct}}''Zorotypus (Zorotypus) denticulatus'' Yin, Cai, & Huang, 2018 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Zorotypus) dilaticeps'' Yin, Cai, Huang, & Engel, 2018 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Zorotypus) goeleti'' Engel & Grimaldi, 2002 – Dominican amber, Dominican Republic (Miocene) *** {{extinct}}''Zorotypus (Zorotypus) mnemosyne'' Engel, 2008 – Dominican amber, Dominican Republic (Miocene) *** {{extinct}}''Zorotypus (Zorotypus) palaeus'' Poinar, 1988 – Dominican amber, Dominican Republic (Miocene) ** Subgenus {{extinct}}''Octozoros'' Engel, 2003 *** {{extinct}}''Zorotypus (Octozoros) acanthothorax'' Engel & Grimaldi, 2002 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) cenomanianus'' Yin, Cai, & Huang, 2018 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) hirsutus'' Mashimo, 2018 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) hudae'' (Kaddumi, 2005) – Jordanian amber, Jordan (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) hukawngi'' Chen & Su, 2019 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) nascimbenei'' Engel & Grimaldi, 2002 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) pecten'' Mashimo, 2019 – Burmese amber, Myanmar (Cretaceous) *** {{extinct}}''Zorotypus (Octozoros) pusillus'' Chen & Su, 2019 – Burmese amber, Myanmar (Cretaceous) ** {{extinct}}''Zorotypus cretatus'' Engel & Grimaldi, 2002 – Burmese amber, Myanmar (Cretaceous) ** {{extinct}}''Zorotypus oligophleps'' Liu, Zhang, Cai & Li, 2018 – Burmese amber, Myanmar (Cretaceous) ** {{extinct}}''Zorotypus robustus'' Liu, Zhang, Cai & Li, 2018 – Burmese amber, Myanmar (Cretaceous) * {{extinct}}''Xenozorotypus'' Engel & Grimaldi, 2002 ** {{extinct}}''Xenozorotypus burmiticus'' Engel & Grimaldi, 2002 – Burmese amber, Myanmar (Cretaceous)

== 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.<ref name="Choe1997">{{cite book | last=Choe | first=Jae C. | editor-last=Choe | editor-first=Jae C. | editor-last2=Crespi | editor-first2=Bernard J. | title=The Evolution of Mating Systems in Insects and Arachnids | chapter=The evolution of mating systems in the Zoraptera: mating variations and sexual conflicts | publisher=Cambridge University Press | location=Cambridge | isbn=978-0-511-72194-6 | doi=10.1017/cbo9780511721946.008 | pages=130–145| year=1997 }}</ref> Much of their time is spent grooming themselves or others.<ref>{{Cite book |title=Encyclopedia of Insects |publisher=Academic Press |year=2009 |isbn=978-0-12-374144-8 |edition=2nd |pages=Ch. 272}}</ref>

''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.<ref>[https://books.google.com/books?id=PYRFDrZs9QAC&dq=Zorotypus+gurneyi+hundred+30&pg=PA206 The Other Insect Societies]</ref>

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.<ref name="Choe233-237">{{cite journal | last=Choe | first=Jae C. | title=Sexual selection and mating system in ''Zorotypus gurneyi'' Choe (Insecta: Zoraptera) | journal=Behavioral Ecology and Sociobiology, II. Determinants and Dynamics of Dominance | volume=34 | issue=4 | year=1994 | issn=0340-5443 | doi=10.1007/bf00183473 | pages=233–237 | hdl=2027.42/46900 | s2cid=42298642 | url=https://deepblue.lib.umich.edu/bitstream/2027.42/46901/1/265_2004_Article_BF00183473.pdf | hdl-access=free }}</ref><ref name="Choe87-93">{{cite journal | last=Choel | first=Jae C. | title=Sexual selection and mating system in ''Zorotypus gurneyi'' Choe (Insecta: Zoraptera), I. Dominance hierarchy and mating success | journal=Behavioral Ecology and Sociobiology | volume=34 | issue=2 | year=1994 | issn=0340-5443 | doi=10.1007/bf00164179 | pages=87–93| bibcode=1994BEcoS..34...87C | hdl=2027.42/46900 | s2cid=9112078 | hdl-access=free }}</ref>

''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.<ref name="Choe1997" />

In ''Spermozoros impolitus'', copulation does not occur, but fertilization is accomplished instead by transfer of a spermatophore from the male to the female. This {{Convert|0.1|mm|in|adj=on}} spermatophore contains a single giant sperm cell, which unravels to about the same length as the female herself, {{Convert|3|mm|in}}. It is thought that this large sperm cell prevents fertilization by other males, by physically blocking the female's genital tract.<ref>{{cite journal | last1=Dallai | first1=R. | last2=Gottardo | first2=M. | last3=Mercati | first3=D. | last4=Machida | first4=R. | last5=Mashimo | first5=Y. | last6=Matsumura | first6=Y. | last7=Beutel | first7=R. G. | title=Divergent mating patterns and a unique mode of external sperm transfer in Zoraptera: an enigmatic group of pterygote insects | journal=Naturwissenschaften | volume=100 | issue=6 | date=12 May 2013 | issn=0028-1042 | doi=10.1007/s00114-013-1055-0 | pages=581–594| pmid=23666111| bibcode = 2013NW....100..581D| s2cid=16363067 | display-authors = 1}}</ref><ref>{{cite magazine |url=https://www.newscientist.com/article/mg21829195.600-the-tiny-insect-with-the-massive-sperm.html#.Uk_8o1CTg_Y |title=The tiny insect with the massive sperm |journal=New Scientist |date= 1 June 2013 |issue=2919 |page=17}}</ref>

=== 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.<ref>{{Cite journal|url = http://hbs.bishopmuseum.org/pubs-online/pdf/op91.pdf#page=35|title = The Zorotypidae of Fiji (Zoraptera)|last = Engel|first = Michael|date = 2007|journal = Bishop Museum Occasional Papers}}</ref>

== References == {{Reflist|2}}

== General references == * Costa JT 2006 Psocopera and Zoraptera. In: Costa JT The other Insect Societies. The Belknap Press of Harvard University Press, Cambridge, MA and London, UK pp 193–211 * {{cite book|last1=Grimaldi|first1=David|author-link1=David Grimaldi (entomologist)|last2=Engel|first2=Michael S.|author-link2=Michael S. Engel|title=Evolution of the Insects|url=https://books.google.com/books?id=odQmAAAAQBAJ&pg=PR4|date=16 May 2005|publisher=Cambridge University Press|isbn=978-1-107-26877-7| oclc=56057971}} * {{cite journal | last1 = Hubbard | first1 = Michael D. | year = 1990 | title = A Catalog of the Order Zoraptera (Insecta) | url = http://digitalcommons.unl.edu/insectamundi/393/ | journal = Insecta Mundi | volume = 4 | issue = 1–4 }} * {{Cite journal | last1 = Rafael | first1 = José Albertino | author-link2 = Michael S. Engel | last2 = Engel | first2 = Michael S. | year = 2006 | title = A new species of ''Zorotypus'' from central Amazonia, Brazil (Zoraptera: Zorotypidae) | url = http://digitallibrary.amnh.org/handle/2246/5806| journal = American Museum Novitates | issue = 3528 | pages = 1–11 | doi = 10.1206/0003-0082(2006)3528[1:ANSOZF]2.0.CO;2 | hdl = 2246/5806 | hdl-access = free | s2cid = 67852583 }} * {{cite journal |last=Kaddumi |first=Hani F. |year=2005 |title=Amber of Jordan, the Oldest Prehistoric Insects in Fossilized Resin. |journal=Publications of the Eternal River |publisher=Museum of Natural History |location=Amman. |page=168 |url=https://www.academia.edu/19770324 |oclc=235969870}}

== External links == * [http://tolweb.org/Zoraptera/8252 Tree of Life Zoraptera] * {{Cite web |url = http://zoraptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1218730 |website = Zorapotera Species File |title = Zorotypidae Silvestri, 1913 }} * {{Commons-inline}} * {{Wikispecies-inline}}

Category:Zoraptera Category:Insect orders Category:Extant Cretaceous first appearances Category:Taxa named by Filippo Silvestri Category:Polyneoptera