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{{Short description|Order of insects}}
{{For|the type genus|Thrips (genus)}}
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{{Automatic taxobox
|name              = Thrips
|fossil_range      = {{fossilrange |299 |0}}<small>[[Permian]] – recent</small>
|image             = Thysanoptera.jpg
|image_caption     = Winged and wingless forms
|taxon             = Thysanoptera
|authority         = [[Alexander Henry Haliday |Haliday]], 1836
|display_parents   = 2
|subdivision_ranks = Suborders & Families
|subdivision       = [[Terebrantia]]
: [[Adiheterothripidae]]
: [[Aeolothripidae]]<!-- Zootaxa 864: 1–16 (2005) -->
: [[Fauriellidae]]
: † [[Hemithripidae]]
: [[Heterothripidae]]
: † [[Jezzinothripidae]]
: † [[Karataothripidae]]
: [[Melanthripidae]]
: [[Merothripidae]]
: † [[Scudderothripidae]]
: [[Stenurothripidae]]
: [[Thripidae]]
: † [[Triassothripidae]]
: [[Uzelothripidae]]
[[Tubulifera]]
: [[Phlaeothripidae]]
: † [[Rohrthripidae]]
|synonyms =Physopoda<ref name="Fedor2010">{{cite journal |last1=Fedor |first1=Peter J. |last2=Doricova |first2=Martina |last3=Prokop |first3=Pavol |last4=Mound |first4=Laurence A. |title=Heinrich Uzel, the father of Thysanoptera studies |journal=Zootaxa |year=2010 |volume=2645 |pages=55–63 |url=http://www.mapress.com/zootaxa/2010/f/zt02645p063.pdf |doi=10.11646/zootaxa.2645.1.3 |doi-access=free}}</ref>
}}
'''Thrips''' ([[Order (biology) |order]] '''Thysanoptera''') are minute (mostly {{convert |1 |mm |abbr=on}} long or less), slender [[insect]]s with fringed wings and unique asymmetrical mouthparts. [[Entomologists]] have [[species description |described]] approximately 7,700 species. They fly only weakly and their feathery wings are unsuitable for conventional flight; instead, thrips exploit an unusual mechanism, [[#Flight |clap and fling]], to create lift using an unsteady circulation pattern with transient [[vortices]] near the wings.
Thrips are a functionally diverse group; many of the known species are [[fungivorous]]. A small proportion of the species are serious [[Pest (organism) |pest]]s of commercially important crops. Some of these serve as [[Disease vector |vectors]] for over 20 viruses that cause [[plant disease]], especially the [[Tospoviruses]]. Many flower-dwelling species bring benefits as pollinators, with some predatory thrips feeding on small insects or [[mite]]s. In the right conditions, such as in greenhouses, invasive species can exponentially increase in population size and form large swarms because of a lack of natural predators coupled with their ability to reproduce asexually, making them destructive to crops. Their identification to species by standard morphological characteristics is often challenging.
==Naming and etymology ==
The first recorded mention of thrips dates from the 17th century, and a sketch was made by Philippo Bonanni, a Catholic priest, in 1691. Swedish entomologist Baron [[Charles De Geer]] described two species in the [[genus]] ''Physapus'' in 1744, and [[Linnaeus]] in 1746 added a third species and named this group of insects ''Thrips''. In 1836 the Irish entomologist [[Alexander Henry Haliday]] described 41 species in 11 genera and proposed the [[Order (biology) |order]] name of Thysanoptera. The first monograph on the group was published in 1895 by [[Heinrich Uzel]],<ref>{{cite book |title=Monografie řádu Thysanoptera |publisher=Hradec Králové |year=1895 |last=Uzel |first=Jindrich |url=https://archive.org/details/monografieraduth00uzel}}</ref> who is regarded by Fedor ''et al.'' as the father of Thysanoptera studies.<ref>{{cite journal |last1=Fedor |first1=Peter J. |last2=Doricova |first2=Martina |last3=Prokop |first3=Pavol |last4=Mound |first4=Laurence A. |title=Heinrich Uzel, the father of Thysanoptera studies |journal=Zootaxa |year=2010 |volume=2645 |pages=55–63 |url=http://www.mapress.com/zootaxa/2010/f/zt02645p063.pdf |doi=10.11646/zootaxa.2645.1.3 |doi-access=free |quote =Jindřich (Heinrich) Uzel, a Czech phytopathologist and entomologist, published in 1895 a monograph on the Order Thysanoptera that provided the basis for almost all subsequent work on this group of insects.}}</ref>
The generic and [[English language |English]] name ''thrips'' is a direct transliteration of the [[Ancient Greek]] word {{lang |grc |[[wikt:θρίψ |θρίψ]]}}, ''thrips'', meaning "[[woodworm]]".<ref>{{LSJ |qri/y |θρίψ |ref}}.</ref> Like some other animal-names (such as ''sheep'', ''deer'', and ''moose'') in English the word "thrips" expresses both the [[Grammatical number |singular and plural]], so there may be many thrips or a single thrips. Other common names for thrips include thunderflies, thunderbugs, storm flies, thunderblights, storm bugs, corn fleas, corn flies, corn lice, freckle bugs, harvest bugs, and physopods.<ref name="Kobro 2011">{{cite journal |last=Kobro |first=Sverre |title=Checklist of Nordic Thysanoptera |journal=Norwegian Journal of Entomology |volume=58 |pages=21–26 |year=2011 |url=http://www.entomologi.no/journals/nje/2011-1/pdf/nje-vol58-no1-kobro.pdf |access-date=October 25, 2014}}</ref><ref>{{cite book |last=Kirk |first=W.D.J. |year=1996 |title=Thrips: Naturalists' Handbooks 25 |publisher=The Richmond Publishing Company}}</ref><ref>{{cite book |last1=Marren |first1=Peter |last2=Mabey |first2=Richard |author2-link=Richard Mabey |title=Bugs Britannica |url=https://books.google.com/books?id=Ah62bUZLDOwC |year=2010 |publisher=Chatto & Windus |isbn=978-0-7011-8180-2 |page=141}}</ref> The older group name "physopoda" references the bladder-like tips to the [[Tarsus (skeleton) |tarsi]] of the legs. The name of the order, ''Thysanoptera'', is constructed from the ancient Greek words {{lang |grc |[[wikt:θύσανος |θύσανος]]}}, ''thysanos'', "tassel or fringe", and {{lang |grc |[[wikt:πτερόν |πτερόν]]}}, ''pteron'', "wing", with reference to the insects' fringed wings.<ref>{{cite web |title=Thysanoptera |url=https://www.merriam-webster.com/dictionary/Thysanoptera |publisher=Merriam-Webster |access-date=15 February 2017}}</ref><ref name=Tipping>{{cite book |last=Tipping |first=C. |editor=Capinera, John L. |title=Encyclopedia of Entomology |url=https://books.google.com/books?id=i9ITMiiohVQC&pg=PA3769 |year=2008 |publisher=Springer Science & Business Media |isbn=978-1-4020-6242-1 |pages=3769–3771}}</ref><ref>{{LSJ |qu/sanos |θύσανος}}, {{LSJ |ptero/n |πτερόν |shortref}}.
</ref>
==Morphology ==
[[File:Thrips physapus.jpg|thumb|upright|left|Typical Tubulifera thrips: the feathery wings are unsuitable for the [[leading edge]] [[vortex]] flight of most other insects, but support [[clap and fling]] flight.]]
Thrips are small [[hemimetaboly |hemimetabolic]] insects with a distinctive cigar-shaped [[body plan]].<ref name=uga/> They are elongated with transversely constricted bodies. They range in size from {{convert |0.5 |to |14 |mm |in |2 |abbr=on}} in length for the larger predatory thrips, but most thrips are about 1&nbsp;mm in length. Flight-capable thrips have two similar, strap-like pairs of wings with a fringe of bristles. The wings are folded back over the body at rest. Their legs usually end in two [[Arthropod leg |tarsal]] segments with a bladder-like structure known as an "arolium" at the pretarsus. This structure can be [[wikt:eversion |everted]] by means of [[hemolymph]] pressure, enabling the insect to walk on vertical surfaces.<ref name=Gillott>{{cite book |last=Gillott |first=Cedric |year=2005 |title=Entomology |url=https://archive.org/details/entomology00cgil |url-access=limited |publisher=Springer |page=[https://archive.org/details/entomology00cgil/page/n244 234] |isbn=978-0-306-44967-3}}</ref><ref>{{cite journal  |year=1971 |title=Functional morphology of the thysanopteran pretarsus |journal=Canadian Journal of Zoology |volume=49 |issue=1 |pages=91–108 |doi=10.1139/z71-014 |pmid=5543183 |last1=Heming |first1=B. S. }}</ref> They have compound eyes consisting of a small number of ommatidia and three ocelli or simple eyes on the head.<ref name=ReshCarde />
[[File:Heliothrips asymmetric mouthparts.jpg|thumb|upright |Asymmetric mouthparts of ''Heliothrips'']]
Thrips have asymmetrical [[Insect mouthparts |mouthparts]] unique to the group. Unlike the [[Hemiptera]] (true bugs), the right [[mandible (arthropod) |mandible]] of thrips is reduced and vestigial – and in some species completely absent.<ref>{{cite book |last1=Childers |first1=C.C. |last2=Achor |first2=D.S. |year=1989 |chapter=Structure of the mouthparts of Frankliniella bispinosa (Morgan) (Thysanoptera: Thripidae). |editor1=Parker, B.L. |editor2=Skinner, M. |editor3=Lewis, T. |title=Towards Understanding Thysanoptera. Proceedings of the International Conference on Thrips |publisher=USDA Technical Report NE-147 |place=Radnor, PA}}</ref>
The left mandible is used briefly to cut into the food plant; saliva is injected and the maxillary stylets, which form a tube, are then inserted and the semi-digested food pumped from ruptured cells. This process leaves cells destroyed or collapsed, and a distinctive silvery or bronze scarring on the surfaces of the stems or leaves where the thrips have fed.<ref>{{cite journal |last1=Chisholm |first1=I.F. |last2=Lewis |first2=T. |title=A new look at thrips (Thysanoptera) mouthparts, their action and effects of feeding on plant tissue |journal=Bulletin of Entomological Research |year=2009 |volume=74 |issue=4 |pages=663–675 |doi=10.1017/S0007485300014048}}</ref> The mouthparts of thrips have been described as “rasping-sucking”,<ref name="ncsu">{{cite web |title=Thrips: Biology and Rose Pests |url=https://entomology.ces.ncsu.edu/thrips-2/ |website=North Carlolina State University |access-date=14 October 2024}}</ref> “punching and sucking”,<ref name=uga>{{cite web |title=Biology and Management of Thrips Affecting the Production Nursery and Landscape |url=https://extension.uga.edu/publications/detail.html?number=C1158 |website=University of Georgia |access-date=14 October 2024 |language=en}}</ref> or, simply just a specific type of “piercing-sucking” mouthparts.<ref>{{cite web |title=Thrips (Thysanoptera: Thripidae) |url=https://agsci.colostate.edu/wp-content/uploads/sites/115/2013/01/Thrips.pdf |website=Colorado State Universiy}}</ref>
Thysanoptera is divided into two suborders, Terebrantia and Tubulifera; these can be distinguished by morphological, behavioral, and developmental characteristics. Tubulifera consists of a single family, [[Phlaeothripidae]]; members can be identified by their characteristic tube-shaped apical abdominal segment, egg-laying atop the surface of leaves, and three "pupal" stages. In the Phlaeothripidae, the males are often larger than females and a range of sizes may be found within a population. The largest recorded phlaeothripid species is about 14 mm long. Females of the eight families of the Terebrantia all possess the eponymous saw-like (see [[wikt:terebra |terebra]]) [[ovipositor]] on the anteapical abdominal segment, lay eggs singly within plant tissue, and have two "pupal" stages. In most Terebrantia, the males are smaller than females. The family Uzelothripidae has a single species and it is unique in having a whip-like terminal antennal segment.<ref name=ReshCarde/>
==Evolution ==
[[File:Parallelothrips separatus.jpg|thumb |right |''[[Parallelothrips |Parallelothrips separatus]]'', [[Myanmar amber]] ]]
The earliest fossils of thrips date back to the [[Permian]] ''[[Permothrips |Permothrips longipennis]]'', although it probably not a member of this group.<ref name=Grimaldietal2004/> By the [[Early Cretaceous]], true thrips became much more abundant.<ref name=Grimaldietal2004/> The extant [[Family (biology) |family]] [[Merothripidae]] most resembles these ancestral Thysanoptera, and is probably basal to the order.<ref name=Mound1997>{{cite book |last1=Mound |first1=L.A. |authorlink1=Laurence Alfred Mound |editor-last=Lewis |editor-first=T. |title=Biological diversity |publisher=CAB International |chapter=Thrips as Crop Pests |year=1997 |pages=197–215 }}</ref> There are currently over six thousand species of thrips recognized, grouped into 777 extant and sixty fossil genera.<ref>{{cite web |url=http://thrips.info/wiki/Main_Page |title=Thrips Wiki |access-date=19 February 2017}}</ref> Some thrips were pollinators of the [[Ginkgoales]] as early as 110-105 Mya, in the Cretaceous.<ref>{{cite journal |last=Peñalver |first=Enrique |last2=Labandeira |first2=Conrad C. |last3=Barrón |first3=Eduardo |last4=Delclòs |first4=Xavier |last5=Nel |first5=Patricia |last6=Nel |first6=André |last7=Tafforeau |first7=Paul |last8=Soriano |first8=Carmen |display-authors=3 |title=Thrips pollination of Mesozoic gymnosperms |journal=Proceedings of the National Academy of Sciences |publisher=Proceedings of the National Academy of Sciences |volume=109 |issue=22 |date=21 May 2012 |doi=10.1073/pnas.1120499109 |doi-access=free |pages=8623–8628|pmc=3365147 }}</ref> ''[[Cenomanithrips |Cenomanithrips primus]]'',<ref name="Tong2019">{{cite journal |last=Tong |first=Tingting |last2=Shih |first2=Chungkun |last3=Ren |first3=Dong |title=A new genus and species of Stenurothripidae (Insecta: Thysanoptera: Terebrantia) from mid-Cretaceous Myanmar amber |journal=Cretaceous Research |publisher=Elsevier BV |volume=100 |year=2019 |issn=0195-6671 |doi=10.1016/j.cretres.2019.03.005 |pages=184–191}}</ref> ''[[Didymothrips |Didymothrips abdominalis]]'' and ''[[Parallelothrips |Parallelothrips separatus]]'' are known from [[Myanmar amber]] of the [[Cenomanian]] age.<ref name="Guo2024">{{cite journal |last=Guo |first=Dawei |last2=Engel |first2=Michael S. |last3=Shih |first3=Chungkun |last4=Ren |first4=Dong |title=New stenurothripid thrips from mid-Cretaceous Kachin amber (Thysanoptera, Stenurothripidae) |journal=ZooKeys |publisher=Pensoft Publishers |volume=1192 |date=21 February 2024 |doi=10.3897/zookeys.1192.117754 |doi-access=free |pages=197–212|pmc=10902785 }}</ref>
===Phylogeny===
Thrips are generally considered to be the sister group to [[Hemiptera]] (bugs).<ref name="Li">{{cite journal |last1=Li |first1=Hu |display-authors=etal |title=Higher-level phylogeny of paraneopteran insects inferred from mitochondrial genome sequences |journal=Scientific Reports |year=2015 |volume=5 |pages=8527 |doi=10.1038/srep08527 |pmid=25704094 |pmc=4336943 |bibcode=2015NatSR...5E8527L}}</ref> The phylogeny of thrips families has been little studied. A preliminary analysis in 2013 of 37 species using 3 genes, as well as a phylogeny based on ribosomal DNA and three proteins in 2012, supports the monophyly of the two suborders, Tubulifera and Terebrantia. In Terebrantia, Melanothripidae may be sister to all other families, but other relationships remain unclear. In Tubulifera, the Phlaeothripidae and its subfamily Idolothripinae are monophyletic. The two largest thrips subfamilies, Phlaeothripinae and Thripinae, are paraphyletic and need further work to determine their structure. The internal relationships from these analyses are shown in the cladogram.<ref name=Terry2013>{{cite journal |last1=Terry |first1=Mark |last2=Whiting |first2=Michael |title=Evolution of Thrips (Thysanoptera) Phylogenetic Patterns and Mitochondrial Genome Evolution |journal=Journal of Undergraduate Research |year =2013 }}</ref><ref name=Buckman2012>{{cite journal |last1=Buckman |first1=Rebecca S. |last2=Mound |first2=Laurence A. |authorlink2=Laurence Alfred Mound |last3=Whiting |first3=Michael F. |title=Phylogeny of thrips (Insecta: Thysanoptera) based on five molecular loci |journal=Systematic Entomology |year=2012 |volume=38 |issue=1 |pages=123–133 |doi=10.1111/j.1365-3113.2012.00650.x |s2cid=84909610 }}</ref>
{{clade
|label1='''Thysanoptera'''
|1={{clade
|label1=[[Terebrantia]]
|1={{clade
|1=[[Melanthripidae]]
|2=other Terebrantia
}}
|label2=[[Tubulifera]]
|2=[[Phlaeothripidae]]
}}
}}
===Taxonomy===
The following families are (2013) recognized:<ref name=Buckman2012/><ref>{{cite journal |last=Mound |first=L.A. |authorlink=Laurence Alfred Mound |year=2011 |url=http://www.mapress.com/zootaxa/2011/f/zt03148p202.pdf |title=Order Thysanoptera Haliday, 1836 in Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness |pages=201–202 |journal=Zootaxa |volume=3148 |doi=10.11646/zootaxa.3148.1.38 }}</ref><ref name=ReshCarde />
* Suborder [[Terebrantia]] [[File:Frankadult.jpg|thumb|upright |Adult ''[[Franklinothrips]] vespiformis'' ([[Aeolothripidae]]), a widely distributed tropical species]]
:* [[Adiheterothripidae]] {{small |Shumsher, 1946}} (11 genera)
:* [[Aeolothripidae]] {{small |Uzel, 1895}} (29 genera) – banded thrips and broad-winged thrips
:* [[Fauriellidae]] {{small |[[Hermann Priesner |Priesner]], 1949}} (four genera)
:* †[[Hemithripidae]] {{small |[[Richard Siddoway Bagnall |Bagnall]], 1923}} (one fossil genus, ''[[Hemithrips]]'' with 15 species)
:* [[Heterothripidae]] {{small |Bagnall, 1912}} (seven genera, restricted to the New World)
:* †[[Jezzinothripidae]] {{small |[[Otto zur Strassen |zur Strassen]], 1973}} (included by some authors in [[Merothripidae]])
:* †[[Karataothripidae]] {{small |Sharov, 1972}} (one fossil species, ''[[Karataothrips jurassicus]]'')
:* [[Melanthripidae]] {{small |Bagnall, 1913}} (six genera of flower feeders)
:* [[Merothripidae]] {{small |[[J. Douglas Hood |Hood]], 1914}} (five genera, mostly Neotropical and feeding on dry-wood fungi) – large-legged thrips
:* †[[Scudderothripidae]] {{small |zur Strassen, 1973}} (included by some authors in [[Stenurothripidae]])
:* [[Thripidae]] {{small |[[James Francis Stephens |Stephens]], 1829}} (292 genera in four subfamilies, flower living) – common thrips
:* †[[Triassothripidae]] {{small |Grimaldi & Shmakov, 2004}} (two fossil genera)
:* [[Uzelothripidae]] {{small |Hood, 1952}} (one species, ''[[Uzelothrips scabrosus]]'')
* Suborder [[Tubulifera]]
:* [[Phlaeothripidae]] {{small |Uzel, 1895}} (447 genera in two subfamilies, fungal hyphae and spore feeders)
The identification of thrips to species is challenging as types are maintained as slide preparations of varying quality over time. There is also considerable variability leading to many species being misidentified. Molecular sequence based approaches have increasingly been applied to their identification.<ref>{{cite journal |doi=10.1653/024.096.0250 |journal=Florida Entomologist |volume=96 |issue=2 |pages=318–322 |year=2013 |title=Homologies and Host-Plant Specificity: Recurrent Problems in the Study of Thrips |first=Laurence A. |last=Mound |authorlink=Laurence Alfred Mound |doi-access=free}}</ref><ref>{{cite journal |journal=J. Econ. Entomol. |volume=99 |issue=5 |pages=1813–1819 |year=2006 |title=Molecular Identification Key for Pest Species of Scirtothrips (Thysanoptera: Thripidae) |first1=Paul F. |last1=Rugman-Jones |first2=Mark S. |last2=Hoddle |first3=Laurence A. |last3=Mound |first4=Richard |last4=Stouthamer |pmid=17066817 |doi=10.1093/jee/99.5.1813 |url=https://www.researchgate.net/publication/6729061}}</ref>
==Biology==
[[File:Myrsine howittiana.jpg|thumb |The Australian rainforest shrub ''[[Myrsine howittiana |Myrsine (Rapanea) howittiana]]'' is pollinated by ''[[Thrips setipennis]]''.]]
===Feeding===
Thrips are believed to have descended from a fungus-feeding ancestor during the Mesozoic,<ref name=Grimaldietal2004>{{cite journal |last1=Grimaldi |first1=D. |author2=Shmakov, A. |author3=Fraser, N. |title=Mesozoic Thrips and Early Evolution of the Order Thysanoptera (Insecta) |journal=Journal of Paleontology |year=2004 |jstor=4094919 |volume=78 |issue=5 |pages=941–952 |doi=10.1666/0022-3360(2004)078<0941:mtaeeo>2.0.co;2 |s2cid=85901347 }}</ref> and many groups still feed upon and inadvertently redistribute fungal spores.<ref>{{cite journal |url=https://doi.org/10.1146/annurev.ento.51.110104.151044 |doi=10.1146/annurev.ento.51.110104.151044 |title=Invasion Biology of Thrips |year=2006 |last1=Morse |first1=Joseph G. |last2=Hoddle |first2=Mark S. |journal=Annual Review of Entomology |volume=51 |pages=67–89 |pmid=16332204 }}</ref> These live among leaf litter or on dead wood and are important members of the [[ecosystem]], their diet often being supplemented with [[pollen]]. Other species are primitively [[eusocial]] and form plant [[gall]]s and still others are predatory on mites and other thrips.<ref name=Tipping/><ref>{{cite journal |url=https://doi.org/10.1002/ps.7176 |doi=10.1002/ps.7176 |title=Thysanoptera as predators: Their diversity and significance as biological control agents |year=2022 |last1=Wang |first1=Zhaohong |last2=Mound |first2=Laurence A. |last3=Hussain |first3=Mubasher |last4=Arthurs |first4=Steven P. |last5=Mao |first5=Runqian |journal=Pest Management Science |volume=78 |issue=12 |pages=5057–5070 |pmid=36087293 |s2cid=252181342 }}</ref> Two species of ''Aulacothrips'', ''A. tenuis'' and ''A. levinotus'', have been found to be ectoparasites on [[Aetalionidae |aetalionid]] and [[Membracidae |membracid plant-hoppers]] in Brazil.<ref>{{cite journal |doi=10.1007/s11230-014-9526-z |pmid=25274260 |title=Two new ectoparasitic species of Aulacothrips Hood, 1952 (Thysanoptera: Heterothripidae) associated with ant-tended treehoppers (Hemiptera) |journal=Systematic Parasitology |volume=89 |issue=3 |pages=271–8 |year=2014 |last1=Cavalleri |first1=Adriano |last2=Kaminski |first2=Lucas A. |s2cid=403014 }}</ref> ''Akainothrips francisi'' of Australia is a parasite within the colonies of another thrips species ''Dunatothrips aneurae'' that makes silken nests or domiciles on ''Acacia'' trees.<ref>{{Cite journal |last1=Gilbert |first1=James D. J. |last2=Mound |first2=Laurence A. |last3=Simpson |first3=Stephen J. |date=2012 |title=Biology of a new species of socially parasitic thrips (Thysanoptera: Phlaeothripidae) inside Dunatothrips nests, with evolutionary implications for inquilinism in thrips: A NEW INQUILINE THRIPS |journal=Biological Journal of the Linnean Society |language=en |volume=107 |issue=1 |pages=112–122 |doi=10.1111/j.1095-8312.2012.01928.x |doi-access=free }}</ref> A number of thrips in the subfamily Phlaeothripinae that specialize on ''Acacia'' hosts produce silk with which they glue together phyllodes to form domiciles inside which their semi-social colonies live.<ref>{{Cite journal |last1=Gilbert |first1=James D. J. |last2=Simpson |first2=Stephen J. |date=2013 |title=Natural history and behaviour of Dunatothrips aneurae Mound (Thysanoptera: Phlaeothripidae), a phyllode-gluing thrips with facultative pleometrosis: Natural History of Dunatothrips Aneurae |journal=Biological Journal of the Linnean Society |language=en |volume=109 |issue=4 |pages=802–816 |doi=10.1111/bij.12100 |doi-access=free }}</ref>
''Mirothrips arbiter'' has been found in paper wasp nests in Brazil. The eggs of the hosts including ''Mischocyttarus atramentarius'', ''Mischocyttarus cassununga'' and ''Polistes versicolor'' are eaten by the thrips.<ref>{{cite journal |title=Egg predation within the nests of social wasps: a new genus and species of Phlaeothripidae, and evolutionary consequences of Thysanoptera invasive behaviour |first1=Adriano |last1=Cavalleri |first2=André R. |last2=De Souza |first3=Fábio |last3=Prezoto |first4=Laurence A. |last4=Mound |journal=Biological Journal of the Linnean Society |year=2013 |volume=109 |issue=2 |pages=332–341 |doi=10.1111/bij.12057 |doi-access=free }}</ref> Thrips, especially in the family [[Aeolothripidae]], are also predators, and are considered beneficial in the management of pests like the [[codling moth]]s.<ref>{{Cite book |title=The Biology of the Codling Moth as the Basis for Its Control |last=Tadic |first=M. |publisher=Univerzitet U Beogradu |year=1957}}</ref>
Most research has focused on thrips species that feed on economically significant crops. Some species are predatory, but most of them feed on pollen and the chloroplasts harvested from the outer layer of plant epidermal and mesophyll cells. They prefer tender parts of the plant, such as buds, flowers and new leaves.<ref name=Kirk1995>{{cite book |last=Kirk |first=W.D.J. |year=1995 |title=Feeding behavior and nutritional requirements |pages=21–29 |editor1=Parker, B.L. |editor2=Skinner, M. |editor3=Lewis, T. |work=Thrips Biology and Management |publisher=Plenum Press}}</ref><ref name=NCSUOnionThrips>{{cite web |title=Onion Thrips |url=http://ipm.ncsu.edu/AG136/thrips9.html |publisher=NCSU |access-date=23 February 2017}}</ref> Besides feeding on plant tissues, the [[common blossom thrips]] feeds on pollen grains and on the eggs of mites. When the larva supplements its diet in this way, its development time and mortality is reduced, and adult females that consume mite eggs increase their fecundity and longevity.<ref>{{cite journal |last1=Milne |first1=M. |last2=Walter |first2=G.H. |year=1997 |title=The significance of prey in the diet of the phytophagous thrips, ''Frankliniella schultzei'' |journal=Ecological Entomology |volume=22 |issue=1 |pages=74–81 |doi=10.1046/j.1365-2311.1997.00034.x |bibcode=1997EcoEn..22...74M |s2cid=221682518 }}</ref>
===Pollination ===
[[File:Leaf curling in coffee caused by hoplandothrips.JPG|thumb |[[Coffea |Coffee tree]] leaves rolled up by ''[[Hoplandrothrips]]'' ([[Phlaeothripidae]]) damage]]
Some flower-feeding thrips pollinate the flowers they are feeding on,<ref>{{cite journal |last1=Terry |first1=I. |last2=Walter |first2=G.H. |last3=Moore |first3=C. |last4=Roemer |first4=R. |last5=Hull |first5=C. |title=Odor-Mediated Push-Pull Pollination in Cycads |journal=Science |date=2007 |volume=318 |issue=5847 |page=70 |doi=10.1126/science.1145147 |pmid=17916726 |bibcode=2007Sci...318...70T |s2cid=24147411 |doi-access=free }}</ref> and some authors suspect that they may have been among the first insects to evolve a pollinating relationship with their host plants.<ref>{{cite journal |last=Terry |first=I. |year =2001 |title=Thrips: the primeval pollinators? |journal=Thrips and Tospoviruses: Proceedings of the 7th Annual Symposium on Thysanoptera |pages=157–162}}</ref> Amber fossils of ''Gymnopollisthrips'' from the [[Early Cretaceous]] show them to be coated in ''[[Cycadopites]]''-like pollen.<ref>{{Cite journal |last=Peñalver |first=Enrique |last2=Labandeira |first2=Conrad C. |last3=Barrón |first3=Eduardo |last4=Delclòs |first4=Xavier |last5=Nel |first5=Patricia |last6=Nel |first6=André |last7=Tafforeau |first7=Paul |last8=Soriano |first8=Carmen |date=2012 |title=Thrips pollination of Mesozoic gymnosperms |url=https://pnas.org/doi/full/10.1073/pnas.1120499109 |journal=Proceedings of the National Academy of Sciences |language=en |volume=109 |issue=22 |pages=8623–8628 |doi=10.1073/pnas.1120499109 |issn=0027-8424 |pmc=3365147 |pmid=22615414}}</ref> ''[[Scirtothrips dorsalis]]'' carries pollen of commercially important [[chili pepper]]s.<ref>{{cite journal |last=Sakai |first=S. |title=Thrips pollination of androdioecious ''Castilla elastica'' (Moraceae) in a seasonal tropical forest |journal=American Journal of Botany |volume=88 |issue=9 |pages=1527–1534 |year=2001 |pmid=21669685 |jstor=3558396 |doi=10.2307/3558396 |doi-access=free }}</ref><ref>{{cite journal |last1=Saxena |first1=P. |last2=Vijayaraghavan |first2=M.R. |last3=Sarbhoy |first3=R.K. |last4=Raizada |first4=U. |year=1996 |title=Pollination and gene flow in chillies with ''Scirtothrips dorsalis'' as pollen vectors |journal=Phytomorphology |volume=46 |pages=317–327 |title-link=Scirtothrips dorsalis}}</ref><ref>{{cite journal |first1=Dawn |last1=Frame |year=2003 |title=Generalist flowers, biodiversity and florivory: implications for angiosperm origins |journal=Taxon |volume=52 |issue=4 |pages=681–5 |jstor=3647343 |doi=10.2307/3647343}}</ref> Darwin found that thrips could not be kept out by any netting when he conducted experiments by keeping away larger pollinators.<ref>{{cite book |last=Darwin |first=Charles |author-link=Charles Darwin |year=1892 |title=The effects of cross and self fertilization in the vegetable kingdom |publisher=D. Appleton & Company |url=https://archive.org/stream/effectsofcrossse00darw_0#page/n23/mode/2up/ |page=11}}</ref>''Thrips setipennis'' is the sole pollinator of ''[[Wilkiea huegeliana]]'', a small, unisexual annually flowering tree or shrub in the rainforests of eastern Australia. ''T. setipennis'' serves as an obligate pollinator for other Australian rainforest plant species, including ''[[Myrsine howittiana]]'' and ''[[Myrsine variabilis |M. variabilis]].''<ref>{{cite journal |title=Thrips (Thysanoptera) pollination in Australian subtropical rainforests, with particular reference to pollination of ''Wilkiea huegeliana'' (Monimiaceae) |journal=Journal of Natural History |year=2001 |pages=1–21 |volume=35 |issue=1 |doi=10.1080/002229301447853 |first1=G.A. |last1=Williams |first2=P. |last2=Adam |first3=L.A. |last3=Mound |bibcode=2001JNatH..35....1W |s2cid=216092358 |authorlink3=Laurence Alfred Mound}}</ref> The genus ''[[Cycadothrips]]'' is a specialist pollinator of [[cycad]]s, which are normally wind pollinated but some species of ''[[Macrozamia]]'' are able to attract thrips to male cones at some times and repel them so that they move to female cones.<ref name=Eliyahu>{{cite journal |last1=Eliyahu |first1=Dorit |last2=McCall |first2=Andrew C. |last3=Lauck |first3=Marina |last4=Trakhtenbrot |first4=Ana |last5=Bronstein |first5=Judith L. |year=2015 |title=Minute pollinators: The role of thrips (Thysanoptera) as pollinators of pointleaf manzanita, ''Arctostaphylos pungens'' (Ericaceae) |journal=Journal of Pollination Ecology |volume=16 |pages=64–71 |pmc=4509684 |pmid=26207155 |doi=10.26786/1920-7603(2015)10}}</ref> Thrips are likewise the primary pollinators of heathers in the family ''[[Ericaceae]]'',<ref>{{cite journal |last1=García-Fayos |first1=Patricio |last2=Goldarazena |first2=Arturo |year=2008 |title=The role of thrips in pollination of ''Arctostaphyllos uva-ursi'' |journal=International Journal of Plant Sciences |volume=169 |issue=6 |pages=776–781 |doi=10.1086/588068 |s2cid=58888285 }}</ref> and play a significant role in the pollination of [[pointleaf manzanita]]. Electron microscopy has shown thrips carrying pollen grains adhering to their backs, and their fringed wings are perfectly capable of allowing them to fly from plant to plant.<ref name=Eliyahu/>
===Damage to plants ===
Thrips can cause damage during feeding.<ref>{{cite book |last=Childers |first=C.C. |year=1997 |title=Feeding and oviposition injuries to plants |work=Thrips as Crop Pests |editor=Lewis, T. |pages=505–538 |publisher=CAB International}}</ref> This impact may fall across a broad selection of prey items, as there is considerable breadth in host affinity across the order, and even within a species, varying degrees of fidelity to a host.<ref name=Kirk1995/><ref name=Mound2005>{{cite journal |last=Mound |first=L.A. |authorlink=Laurence Alfred Mound |year=2005 |title=Thysanoptera: diversity and interactions |journal=Annual Review of Entomology |volume=50 |pages=247–269 |doi=10.1146/annurev.ento.49.061802.123318 |pmid=15355240 }}</ref> Family [[Thripidae]] in particular is notorious for members with broad host ranges, and the majority of pest thrips come from this family.<ref>{{cite journal |last=Bailey |first=S.F. |year=1940 |title=The distribution of injurious thrips in the United States |journal=Journal of Economic Entomology |volume=33 |issue=1 |pages=133–136 |doi=10.1093/jee/33.1.133}}</ref><ref>{{cite journal |last=Ananthakrishnan |first=T.N. |author-link=T. N. Ananthakrishnan |year=1993 |title=Bionomics of Thrips |journal=Annual Review of Entomology |volume=38 |pages=71–92 |doi=10.1146/annurev.en.38.010193.000443}}</ref><ref>{{cite journal |doi=10.3390/insects13010061 |doi-access=free |title=Problems with the Concept of "Pest" among the Diversity of Pestiferous Thrips |year=2022 |last1=Mound |first1=Laurence A. |last2=Wang |first2=Zhaohong |last3=Lima |first3=Élison F. B. |last4=Marullo |first4=Rita |journal=Insects |volume=13 |issue=1 |page=61 |pmid=35055903 |pmc=8780980 }}</ref> For example, ''[[Thrips tabaci]]'' damages crops of [[onion]]s, [[potato]]es, [[tobacco]], and [[cotton]].<ref name=NCSUOnionThrips/><ref name=CABI>{{cite web |url=http://www.cabi.org/isc/datasheet/53746 |title=''Thrips tabaci'' (onion thrips) |work=Invasive Species Compendium |publisher=CABI |access-date=24 February 2017}}</ref>
[[File:Australian insects (Plate XXXVII) (7268276524).jpg|thumb|upright |[[Eusociality |Eusocial]] colonies of ''[[Kladothrips]]'' live in [[gall]]s induced on ''[[Acacia]]'' trees.]]
Some species of thrips create [[gall]]s, almost always in leaf tissue. These may occur as curls, rolls or folds, or as alterations to the expansion of tissues causing distortion to leaf blades. More complex examples cause rosettes, pouches and horns. Most of these species occur in the tropics and sub-tropics, and the structures of the galls are diagnostic of the species involved.<ref>{{cite journal |last1=Jorge |first1=Nina Castro |author2=Cavalleri, Adriano |author3=Bedetti, Cibele Souza |author4=Isaias, Rosy Mary Dos Santos |year=2016 |title=A new leaf-galling ''Holopothrips'' (Thysanoptera: Phlaeothripidae) and the structural alterations on ''Myrcia retorta'' (Myrtaceae) |journal=Zootaxa |volume=4200 |issue=1 |pages=174–180 |doi=10.11646/zootaxa.4200.1.8 |pmid=27988645 |issn=1175-5334 }}</ref> A radiation of thrips species seems to have taken place on ''[[Acacia]]'' trees in Australia; some of these species cause galls in the [[Petiole (botany) |petioles]], sometimes fixing two leaf stalks together, while other species live in every available crevice in the bark. In ''[[Casuarina]]'' in the same country, some species have invaded stems, creating long-lasting woody galls.<ref>{{cite journal |last=Mound |first=Laurence A. |authorlink=Laurence Alfred Mound |year=2014 |title=Austral Thysanoptera: 100 years of progress |journal=Australian Journal of Entomology |volume=53 |issue=1 |pages=18–25 |doi=10.1111/aen.12054 |s2cid=85793869 |doi-access=free }}</ref>
=== Social behaviour ===
While poorly documented, chemical communication is believed to be important to the group.<ref>{{cite journal |last=Blum |first=M.S. |year=1991 |title=Towards understanding Thysanoptera: Chemical ecology of the Thysanoptera |pages=95–108 |editor1=Parker, B.L. |editor2=Skinner, M. |editor3=Lewis, T. |journal=Proceedings of the International Conference on Thrips. USDA Technical Report NE-147}}</ref> Anal secretions are produced in the hindgut,<ref name=":0">{{Cite journal |last1=Howard |first1=Dennis F. |last2=Blum |first2=Murray S. |last3=Fales |first3=Henry M. |year=1983 |title=Defense in Thrips: Forbidding Fruitiness of a Lactone |journal=Science |language=en |volume=220 |issue=4594 |pages=335–336 |doi=10.1126/science.220.4594.335 |issn=0036-8075 |pmid=17732921 |bibcode=1983Sci...220..335H |s2cid=24856539}}</ref> and released along the posterior setae as predator deterrents<ref name=":0" /><ref>{{cite journal |last1=Tschuch |first1=G. |last2=Lindemann |first2=P. |last3=Moritz |first3=G. |year=2002 |title=Chemical defence in thrips |pages=277–278 |editor1=Mound, L.A. |editor2=Marullo, R. |journal=Thrips and Tospoviruses: Proceedings of the 7th International Symposium on Thysanoptera }}</ref> In Australia, aggregations of male [[common blossom thrips]] have been observed on the petals of ''[[Hibiscus rosa-sinensis]]'' and ''[[Gossypium hirsutum]]''; females were attracted to these groups so it seems likely that the males were producing [[pheromone]]s.<ref>{{cite journal |last1=Milne |first1=M. |last2=Walter |first2=G.H. |last3=Milne |first3=J.R. |year=2002 |title=Mating Aggregations and Mating Success in the Flower Thrips, ''Frankliniella schultzei'' (Thysanoptera: Thripidae), and a Possible Role for Pheromones |journal=Journal of Insect Behavior |volume=15 |issue=3 |pages=351–368 |doi=10.1023/A:1016265109231 |s2cid=23545048}}</ref>
In the phlaeothripids that feed on fungi, males [[competition (biology) |compete]] to protect and mate with females, and then defend the egg-mass. Males fight by flicking their rivals away with their abdomen, and may kill with their foretarsal teeth. Small males may sneak in to mate while the larger males are busy fighting. In the Merothripidae and in the Aeolothripidae, males are again polymorphic with large and small forms, and probably also compete for mates, so the strategy may well be ancestral among the Thysanoptera.<ref name="ReshCarde">{{cite book |editor1=Resh, Vincent H. |editor2=Cardé, Ring T. |title=Encyclopedia of Insects |year=2003 |publisher=Academic Press |isbn=978-0-12-586990-4 |pages=[https://archive.org/details/encyclopediaofin00bada/page/999 999–1003] |chapter=Thysanoptera |last=Mound |first=L.A. |chapter-url-access=registration |chapter-url=https://archive.org/details/encyclopediaofin00bada |url=https://archive.org/details/encyclopediaofin00bada/page/999 }}</ref>
Many thrips form [[gall]]s on plants when feeding or laying their eggs. Some of the gall-forming [[Phlaeothripidae]], such as genera ''[[Kladothrips]]''<ref>{{cite journal |last1=Kranz |first1=B.D. |author2=Schwarz, M.P. |author3=Mound, L.A. |authorlink3=Laurence Alfred Mound |author4=Crespi, B.J. |author-link4=Bernard Crespi |year=1999 |title=Social biology and sex ratios of the eusocial gall-inducing thrips ''Kladothrips hamiltoni'' |journal=Ecological Entomology |volume=24 |issue=4 |pages=432–442 |doi=10.1046/j.1365-2311.1999.00207.x |bibcode=1999EcoEn..24..432K |s2cid=83180900}}</ref> and ''[[Oncothrips]]'',<ref>{{cite journal |first1=Brenda D. |last1=Kranz |first2=Michael P. |last2=Schwarz |first3=Taryn E. |last3=Wills |first4=Thomas W. |last4=Chapman |first5=David C. |last5=Morris |first6=Bernard J. |last6=Crespi |year=2001 |title=A fully reproductive fighting morph in a soldier clade of gall-inducing thrips (''Oncothrips morrisi'') |journal=Behavioral Ecology and Sociobiology |volume=50 |issue=2 |pages=151–161 |jstor=4601948 |doi=10.1007/s002650100347 |s2cid=38152512 }}</ref> form [[eusocial]] groups similar to [[ant]] colonies, with reproductive [[Queen ant |queens]] and nonreproductive soldier castes.<ref>{{cite book |last1=Crespi |first1=B.J. |author2=Mound, L.A. |authorlink2=Laurence Alfred Mound |year=1997 |chapter-url=https://books.google.com/books?id=qkHHO_SmM_kC&pg=PA166 |chapter=Ecology and evolution of social behaviour among Australian gall thrips and their allies |editor1=Choe, J.C. |editor2=Crespi, B.J. |title=The evolution of social behaviour of insects and arachnids |publisher=Cambridge University Press |pages=166–180 |isbn=978-0-521-58977-2}}</ref><ref>{{cite journal |last1=Chapman |first1=T.W. |last2=Crespi |first2=B.J. |year=1998 |title=High relatedness and inbreeding in two species of haplodiploid eusocial thrips (Insecta: Thysanoptera) revealed by microsatellite analysis |journal=Behavioral Ecology and Sociobiology |volume=43 |issue=4 |pages=301–306 |doi=10.1007/s002650050495 |jstor=4601521 |s2cid=32909187 }}</ref><ref>{{cite journal |first1=Brenda D. |last1=Kranz |first2=Michael P. |last2=Schwarz |first3=David C. |last3=Morris |first4=Bernard J. |last4=Crespi |year=2002 |title=Life history of ''Kladothrips ellobus'' and ''Oncothrips rodwayi'': insight into the origin and loss of soldiers in gall-inducing thrips |journal=Ecological Entomology |volume=27 |issue=1 |pages=49–57 |doi=10.1046/j.1365-2311.2002.0380a.x |bibcode=2002EcoEn..27...49K |s2cid=85097661 }}</ref>
===Flight===
{{See|Insect flight#Clap and fling}}
Most insects create lift by the stiff-winged mechanism of [[insect flight]] with [[steady state]] [[aerodynamics]]; this creates a [[leading edge]] [[vortex]] continuously as the [[insect wing |wing]] moves. The feathery wings of thrips, however, generate lift by [[clap and fling]], a mechanism discovered by the Danish zoologist [[Torkel Weis-Fogh]] in 1973. In the clap part of the cycle, the wings approach each other over the insect's back, creating a circulation of air which sets up vortices and generates useful forces on the wings. The leading edges of the wings touch, and the wings rotate around their leading edges, bringing them together in the "clap". The wings close, expelling air from between them, giving more useful thrust. The wings rotate around their trailing edges to begin the "fling", creating useful forces. The leading edges move apart, making air rush in between them and setting up new vortices, generating more force on the wings. The trailing edge vortices, however, cancel each other out with opposing flows. Weis-Fogh suggested that this cancellation might help the circulation of air to grow more rapidly, by shutting down the [[Wagner effect]] which would otherwise counteract the growth of the circulation.<ref>{{cite journal |last=Weis-Fogh |first=Torkel |author-link=Torkel Weis-Fogh |title=Quick estimates of flight fitness in hovering animals, including novel mechanisms for lift production |journal=Journal of Experimental Biology |volume=59 |pages=169–230 |year=1973 |doi=10.1242/jeb.59.1.169 |url=http://jeb.biologists.org/content/59/1/169.abstract}}</ref><ref name=Sane>{{cite journal |url=http://www.cs.washington.edu/homes/diorio/MURI2003/Publications/sane_review.pdf |title=The aerodynamics of insect flight |last=Sane |first=Sanjay P. |journal=The Journal of Experimental Biology |year=2003 |volume=206 |pages=4191–4208 |doi=10.1242/jeb.00663 |pmid=14581590 |issue=23 |s2cid=17453426 |doi-access=free }}</ref><ref name="physics">{{cite journal |url=http://www.physics.emory.edu/~weeks/lab/papers/jwang-arf05.pdf |title=Dissecting Insect Flight |last=Wang |first=Z. Jane |author-link=Z. Jane Wang |doi=10.1146/annurev.fluid.36.050802.121940 |year=2005 |bibcode=2005AnRFM..37..183W |journal=Annual Review of Fluid Mechanics |volume=37 |issue=1 |pages=183–210 }}</ref><ref>{{cite journal |last=Lighthill |first=M.J. |author-link=M. J. Lighthill |title=On the Weis-Fogh mechanism of lift generation |journal=Journal of Fluid Mechanics |volume=60 |pages=1–17 |year=1973 |doi=10.1017/s0022112073000017 |bibcode=1973JFM....60....1L |s2cid=123051925 }}</ref>
<gallery class="center" mode="nolines" widths="180px" caption="[[Clap and fling]] flight mechanism after Sane 2003">
File:Clap and Fling 1- clap 1.svg |Clap 1: wings close over back
File:Clap and Fling 2- clap 2.svg |Clap 2: leading edges touch, wing rotates around [[leading edge]], vortices form
File:Clap and Fling 3 - clap 3.svg |Clap 3: trailing edges close, vortices shed, wings close giving thrust
</gallery>
<gallery class="center" mode="nolines" widths="180px" caption="Black circle and heavy line: wing (rachis and bristles); Black (curved) arrows: flow; Blue arrows: induced velocity; Orange arrows: net force on wing">
File:Clap and Fling 4- fling 1.svg |Fling 1: wings rotate around trailing edge to fling apart
File:Clap and Fling 5- fling 2.svg |Fling 2: leading edge moves away, air rushes in, increasing lift
File:Clap and Fling 6- fling 3.svg |Fling 3: new vortex forms at leading edge, trailing edge vortices cancel each other, perhaps helping flow to grow faster (Weis-Fogh 1973)
</gallery>
Apart from active flight, thrips, even wingless ones, can also be picked up by winds and transferred long distances. During warm and humid weather, adults may climb to the tips of plants to leap and catch air current. Wind-aided dispersal of species has been recorded over 1600&nbsp;km of sea between Australia and South Island of New Zealand.<ref name=ReshCarde /> It has been suggested that some bird species may also be involved in the dispersal of thrips. Thrips are picked up along with grass in the nests of birds and can be transported by the birds.<ref>{{Cite journal |last1=Fedor |first1=Peter |last2=Doričová |first2=Martina |last3=Dubovský |first3=Michal |last4=Kisel'ák |first4=Jozef |last5=Zvarík |first5=Milan |date=2019-08-15 |title=Cereal pests among nest parasites – the story of barley thrips, Limothrips denticornis Haliday (Thysanoptera: Thripidae) |url=https://journal.fi/entomolfennica/article/view/84532 |journal=Entomologica Fennica |volume=21 |issue=4 |pages=221–231 |doi=10.33338/ef.84532 |s2cid=82549305 |issn=2489-4966 |doi-access=free }}</ref>
A hazard of flight for very small insects such as thrips is the possibility of being trapped by water. Thrips have non-wetting bodies and have the ability to ascend a [[Meniscus (liquid) |meniscus]] by arching their bodies and working their way head-first and upwards along the water surface in order to escape.<ref>{{cite journal |doi=10.1098/rsbl.2016.0279 |pmid=27624795 |year=2016 |title=Meniscus ascent by thrips (Thysanoptera) |journal=Biology Letters |volume=12 |issue=9 |page=20160279 |pmc=5046919 |last1=Ortega-Jiménez |first1=Victor Manuel |last2=Arriaga-Ramirez |first2=Sarahi |last3=Dudley |first3=Robert }}</ref>
===Life cycle ===
[[File:Thrips nymph.jpg|thumb |Thrips nymph.<br/>Scale bar is 0.5&nbsp;mm]]
Thrips lay extremely small eggs, about 0.2&nbsp;mm long. Females of the suborder [[Terebrantia]] cut slits in plant tissue with their ovipositor, and insert their eggs, one per slit. Females of the suborder [[Tubulifera]] lay their eggs singly or in small groups on the outside surfaces of plants.<ref name=GullanCranston>{{cite book |last1=Gullan |first1=P.J. |last2=Cranston |first2=P.S. |year=2010 |title=The Insects: An Outline of Entomology |url=https://archive.org/details/insectsoutlineen00pjgu |url-access=limited |publisher=Wiley |edition=4th |isbn=978-1-118-84615-5 |page=[https://archive.org/details/insectsoutlineen00pjgu/page/n535 511]}}</ref> Some thrips such as ''Elaphothrips tuberculatus'' are known to be facultatively ovoviviparous, retaining the eggs internally and giving birth to male offspring.<ref>{{Cite journal |last=Crespi |first=Bernard J. |date=1989 |title=Facultative viviparity in a thrips |url=https://www.nature.com/articles/337357a0 |journal=Nature |language=en |volume=337 |issue=6205 |pages=357–358 |doi=10.1038/337357a0 |issn=0028-0836}}</ref> Females in many species guard the eggs against cannibalism by other females as well as predators.<ref>{{Cite journal |last=Crespi |first=Bernard J. |date=1990 |title=Subsociality and female reproductive success in a mycophagous thrips: An observational and experimental analysis |url=http://link.springer.com/10.1007/BF01049195 |journal=Journal of Insect Behavior |language=en |volume=3 |issue=1 |pages=61–74 |doi=10.1007/BF01049195 |issn=0892-7553}}</ref>
Thrips are [[hemimetabolous]], metamorphosing gradually to the adult form. The first two [[instar]]s, called larvae or nymphs, are like small wingless adults (often confused with [[springtail]]s) without genitalia; these feed on plant tissue. In the Terebrantia, the third and fourth instars, and in the Tubulifera also a fifth instar, are non-feeding resting stages similar to [[pupa]]e: in these stages, the body's organs are reshaped, and wing-buds and genitalia are formed.<ref name=GullanCranston/> The larvae of some species produce silk from the terminal abdominal segment which is used to line the cell or form a cocoon within which they pupate.<ref>{{Cite journal |last1=Conti |first1=Barbara |last2=Berti |first2=Francesco |last3=Mercati |first3=David |last4=Giusti |first4=Fabiola |last5=Dallai |first5=Romano |date=2009 |title=The ultrastructure of malpighian tubules and the chemical composition of the cocoon of Aeolothrips intermedius Bagnall (Thysanoptera) |url=https://onlinelibrary.wiley.com/doi/10.1002/jmor.10793 |journal=Journal of Morphology |volume=271 |issue=2 |language=en |pages=244–254 |doi=10.1002/jmor.10793 |pmid=19725134 |s2cid=40887930 }}</ref> The adult stage can be reached in around 8–15 days; adults can live for around 45 days.<ref name=UMassAmherst/> Adults have both winged and wingless forms; in the grass thrips ''Anaphothrips obscurus'', for example, the winged form makes up 90% of the population in spring (in temperate zones), while the wingless form makes up 98% of the population late in the summer.<ref name=CapineraPests>{{cite book |last=Capinera |first=John L. |title=Handbook of Vegetable Pests |url=https://books.google.com/books?id=l_0KOVxv-tYC&pg=PA538 |year=2001 |publisher=Gulf |isbn=978-0-12-158861-8 |page=538}}</ref> Thrips can survive the winter as adults or through egg or pupal [[diapause]].<ref name=ReshCarde />
==Human impact==
[[File:Tomato with Tomato Spotted Wilt Virus.jpg|thumb |A [[tomato]] infected with the thrips-borne [[Tospovirus]], [[tomato spotted wilt virus]]]]
===As pests===
[[File:Ponticulothrips diospyrosi.jpg|thumb |''Ponticulothrips diospyrosi'' on finger |alt=|left]]
Many thrips are pests of commercial crops due to the damage they cause by feeding on developing flowers or vegetables, causing discoloration, deformities, and reduced marketability of the crop. Some thrips serve as vectors for plant diseases, such as [[tospoviruses]].<ref>{{cite journal |last=Nault |first=L.R. |year=1997 |title=Arthropod transmission of plant viruses: a new synthesis |journal=Annals of the Entomological Society of America |volume=90 |issue=5 |pages=521–541 |doi=10.1093/aesa/90.5.521}}</ref> Over 20 plant-infecting viruses are known to be transmitted by thrips, but perversely, less than a dozen of the described species are known to vector tospoviruses.<ref>{{cite journal |last=Mound |first=L.A. |date=2001 |title=So many thrips – so few tospoviruses? |journal=Thrips and Tospoviruses: Proceedings of the 7th International Symposium on Thysanoptera |pages=15–18}}</ref> These enveloped viruses are considered among some of the most damaging of emerging plant pathogens around the world, with those vector species having an outsized impact on human agriculture. Virus members include the [[tomato spotted wilt virus]] and the impatiens necrotic spot viruses. The western flower thrips, ''[[Frankliniella occidentalis]]'', has spread until it now has a worldwide distribution, and is the primary vector of plant diseases caused by tospoviruses.<ref>{{cite journal |doi=10.1146/annurev.ento.51.110104.151044 |pmid=16332204 |title=Invasion Biology of Thrips |journal=Annual Review of Entomology |volume=51 |pages=67–89 |year=2006 |last1=Morse |first1=Joseph G. |last2=Hoddle |first2=Mark S. |s2cid=14430622 }}</ref> Other viruses that they spread include the genera ''[[Ilarvirus]]'', ''[[Carmovirus |(Alpha |Beta |Gamma)carmovirus]]'', ''[[Sobemovirus]]'' and ''[[Machlomovirus]].''<ref>{{Cite journal |last=Jones |first=David R |date=2005 |title=Plant Viruses Transmitted by Thrips |url=http://link.springer.com/10.1007/s10658-005-2334-1 |journal=European Journal of Plant Pathology |language=en |volume=113 |issue=2 |pages=119–157 |doi=10.1007/s10658-005-2334-1 |bibcode=2005EJPP..113..119J |s2cid=6412752 |issn=0929-1873}}</ref> Their small size and predisposition towards enclosed places makes them difficult to detect by [[phytosanitary inspection]], while their eggs, laid inside plant tissue, are well-protected from pesticide sprays.<ref name="UMassAmherst">{{cite web |last1=Smith |first1=Tina M. |title=Western Flower Thrips, Management and Tospoviruses |url=https://ag.umass.edu/greenhouse-floriculture/fact-sheets/western-flower-thrips-management-tospoviruses |publisher=University of Massachusetts Amherst |access-date=21 February 2017 |date=2015}}</ref> When coupled with the increasing [[globalization]] of trade and the growth of [[greenhouse]] agriculture, thrips, unsurprisingly, are among the fastest growing group of [[invasive species]] in the world. Examples include ''F. occidentalis'', ''[[Thrips simplex]]'', and ''[[Thrips palmi]]''.<ref name=Carlton>{{cite book |last=Carlton |first=James |title=Invasive Species: Vectors And Management Strategies |url=https://books.google.com/books?id=li6hUvpnp2sC&pg=PA54 |year=2003 |publisher=Island Press |isbn=978-1-61091-153-5 |pages=54–55}}</ref>
Flower-feeding thrips are routinely attracted to bright floral colors (including white, blue, and especially yellow), and will land and attempt to feed. It is not uncommon for some species (e.g., ''[[Frankliniella tritici]]'' and ''[[Limothrips cerealium]]'') to "bite" humans under such circumstances. Although no species feed on blood and no known animal disease is transmitted by thrips, some skin irritation has been described.<ref>{{cite journal |last=Childers |first=C.C. |author2=Beshear, R.J. |author3=Frantz, G. |author4=Nelms, M. |date=2005 |title=A review of thrips species biting man including records in Florida and Georgia between 1986–1997 |journal=Florida Entomologist |volume=88 |issue=4 |pages=447–451 |doi=10.1653/0015-4040(2005)88[447:AROTSB]2.0.CO;2 |doi-access=free }}</ref>
===Management===
[[File:Asilidae_thrips.jpg|thumb |A robberfly preying on thrips]]
Thrips develop [[pesticide resistance |resistance]] to [[insecticide]]s easily and there is constant research on how to control them. This makes thrips ideal as models for testing the effectiveness of new pesticides and methods.<ref>{{Cite journal |last1=Kivett |first1=Jessica M. |last2=Cloyd |first2=Raymond A. |last3=Bello |first3=Nora M. |year=2015 |title=Insecticide rotation programs with entomopathogenic organisms for suppression of western flower thrips (Thysanoptera: Thripidae) adult populations under greenhouse conditions |journal=Journal of Economic Entomology |volume=108 |issue=4 |pages=1936–1946 |doi=10.1093/jee/tov155 |issn=0022-0493 |pmid=26470338 |s2cid=205163917 }}</ref>
Due to their small sizes and high rates of reproduction, thrips are difficult to control using [[Biological pest control |classical biological control]]. Suitable predators must be small and slender enough to penetrate the crevices where thrips hide while feeding, and they must also prey extensively on eggs and larvae to be effective. Only two families of [[parasitoid]] [[Hymenoptera]] parasitize eggs and larvae, the [[Eulophidae]] and the [[Trichogrammatidae]]. Other biocontrol agents of adults and larvae include [[Anthocoridae |anthocorid bugs]] of genus ''[[Orius]]'', and [[phytoseiid]] mites. Biological insecticides such as the fungi ''[[Beauveria bassiana]]'' and ''[[Verticillium lecanii]]'' can kill thrips at all life-cycle stages.<ref name="Hoddle">{{cite web |last1=Hoddle |first1=Mark |title=Western flower thrips in greenhouses: a review of its biological control and other methods |url=http://biocontrol.ucr.edu/wft.html#Biological%20Control%20Approaches |publisher=University of California, Riverside |access-date=21 February 2017}}</ref> [[Insecticidal soap]] spray is effective against thrips. It is commercially available or can be made of certain types of household soap. Scientists in Japan report that significant reductions in larva and adult melon thrips occur when plants are illuminated with red light.<ref>{{Cite journal |title=Efficacy of red LED irradiation for controlling ''Thrips palmi'' in greenhouse melon cultivation |journal=Japanese Journal of Applied Entomology and Zoology |year=2015 |pages=1–6 |volume=59 |issue=1 |first1=Yusuke |last1=Katai |first2=Ryusuke |last2=Ishikawa |first3=Makoto |last3=Doi |first4=Shinichi |last4=Masui |doi=10.1303/jjaez.2015.1 |doi-access=free }}</ref>
==References==
{{Reflist |30em}}
==External links==
{{Commons category |Thysanoptera}}
{{Wikispecies |Thysanoptera}}
* [http://www.ento.csiro.au/thysanoptera/worldthrips.html Thrips of the World checklist]
* [http://thrips.info/wiki/Main_Page Thrips species wiki]
* [http://www.padil.gov.au/pests-and-diseases/Search?queryText1=&queryType1=All&sortType=ScientificName&viewType=Details&pageSize=10&page=1&taxonomy=Thysanoptera Thrips images from the "Pests and Diseases Image Library (PaDIL)" of Australia] {{Webarchive |url=https://web.archive.org/web/20200317161043/http://www.padil.gov.au/pests-and-diseases/search?queryText1=&queryType1=All&sortType=ScientificName&viewType=Details&pageSize=10&page=1&taxonomy=Thysanoptera |date=2020-03-17 }}
* [http://www.ipm.ucdavis.edu/PMG/PESTNOTES/pn7429.html University of California Pest Management Guidelines for Thrips]
* [http://keys.lucidcentral.org/keys/v3/thrips_of_california/Thrips_of_California.html University of California Thrips Identification]
{{Orders of Insects}}
{{Thysanoptera}}
{{Taxonbar |from=Q185628}}
{{Authority control}}
[[Category:Thrips |Thrips ]]
[[Category:Agricultural pest insects]]
[[Category:Extant Permian first appearances]]
<!-- [[Category:Pollinator insects]] "category:Thrips" is already listed here -->
[[Category:Taxa named by Alexander Henry Haliday]]
[[Category:Condylognatha]]