Thrips
Thrips Temporal range:Permian– recent
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Winged and wingless forms | |
Scientific classification![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
(unranked): | Eumetabola |
(unranked): | Paraneoptera |
Order: | Thysanoptera Haliday,1836 |
Suborders & Families | |
Synonyms | |
Physopoda[1] |
Thrips(orderThysanoptera) are minute (mostly 1 mm (0.039 in) long or less), slenderinsectswith fringed wings and unique asymmetrical mouthparts.Entomologistshavedescribedapproximately 7,700 species. They fly only weakly and their feathery wings are unsuitable for conventional flight; instead, thrips exploit an unusual mechanism,clap and fling,to create lift using an unsteady circulation pattern with transientvorticesnear the wings.
Thrips are a functionally diverse group, with nearly half of the known species beingfungivorous.[2]A small proportion of species of thrips are seriouspestsof commercially important crops.[3]Some of these serve asvectorsfor over 20 viruses that causeplant disease,especially theTospoviruses.Many flower-dwelling species bring benefits as pollinators,[4]with some predatory thrips feeding on small insects ormites.[5]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. Due to their cryptic nature, thrips may aggregate in household objects such as furniture, bedding and computer monitors – in the latter case by forcing their way in between theLCDand its glass covering.[6] Their identification to species by standard morphological characteristics is often challenging.
Naming and etymology
[edit]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 BaronCharles De Geerdescribed two species in thegenusPhysapusin 1744, andLinnaeusin 1746 added a third species and named this group of insectsThrips.In 1836 the Irish entomologistAlexander Henry Halidaydescribed 41 species in 11 genera and proposed theordername of Thysanoptera. The first monograph on the group was published in 1895 byHeinrich Uzel,[7]who is regarded by Fedoret al.as the father of Thysanoptera studies.[8]
The generic andEnglishnamethripsis a direct transliteration of theAncient Greekwordθρίψ,thrips,meaning "woodworm".[9]Like some other animal-names (such assheep,deer,andmoose) in English the word "thrips" expresses both thesingular 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.[10][11][12]The older group name "physopoda" references the bladder-like tips to thetarsiof the legs. The name of the order,Thysanoptera,is constructed from the ancient Greek wordsθύσανος,thysanos,"tassel or fringe", andπτερόν,pteron,"wing", with reference to the insects' fringed wings.[13][14][15]
Morphology
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/8/8c/Thrips_physapus.jpg/170px-Thrips_physapus.jpg)
![](https://upload.wikimedia.org/wikipedia/commons/thumb/6/67/Leaf_suffering_from_thrips.jpg/170px-Leaf_suffering_from_thrips.jpg)
Thrips are smallhemimetabolicinsects with a distinctive cigar-shapedbody plan.They are elongated with transversely constricted bodies. They range in size from 0.5 to 14 mm (0.02 to 0.55 in) in length for the larger predatory thrips, but most thrips are about 1 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 twotarsalsegments with a bladder-like structure known as an "arolium" at the pretarsus. This structure can beevertedby means ofhemolymphpressure, enabling the insect to walk on vertical surfaces.[16][17]They have compound eyes consisting of a small number of ommatidia and three ocelli or simple eyes on the head.[18]
![](https://upload.wikimedia.org/wikipedia/commons/thumb/7/7b/Heliothrips_asymmetric_mouthparts.jpg/170px-Heliothrips_asymmetric_mouthparts.jpg)
Thrips have asymmetricalmouthpartsunique to the group. Unlike theHemiptera(true bugs), the rightmandibleof thrips is reduced and vestigial – and in some species completely absent.[19] 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.[20]
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 (seeterebra)ovipositoron 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.[18]
Evolution
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/6/6b/Parallelothrips_separatus.jpg/220px-Parallelothrips_separatus.jpg)
The earliest fossils of thrips date back to thePermianPermothrips longipennis,although it probably not a member of this group.[21]By theEarly Cretaceous,true thrips became much more abundant.[21]The extantfamilyMerothripidaemost resembles these ancestral Thysanoptera, and is probably basal to the order.[22]There are currently over six thousand species of thrips recognized, grouped into 777 extant and sixty fossil genera.[23]Some thrips were pollinators of theGinkgoalesas early as 110-105 Mya, in the Cretaceous.[24]Cenomanithrips primus,[25]Didymothrips abdominalisandParallelothrips separatusare known fromMyanmar amberof theCenomanianage.[26]
Phylogeny
[edit]Thrips are generally considered to be the sister group toHemiptera(bugs).[27]
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.[28][29]
Thysanoptera |
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Taxonomy
[edit]The following families are currently (2013) recognized:[29][30][18]
- SuborderTerebrantia
AdultFranklinothripsvespiformis(Aeolothripidae), a widely distributed tropical species
- AdiheterothripidaeShumsher, 1946(11 genera)
- AeolothripidaeUzel, 1895(29 genera) – banded thrips and broad-winged thrips
- FauriellidaePriesner,1949(four genera)
- †HemithripidaeBagnall,1923(one fossil genus,Hemithripswith 15 species)
- HeterothripidaeBagnall, 1912(seven genera, restricted to the New World)
- †Jezzinothripidaezur Strassen,1973(included by some authors inMerothripidae)
- †KarataothripidaeSharov, 1972(one fossil species,Karataothrips jurassicus)
- MelanthripidaeBagnall, 1913(six genera of flower feeders)
- MerothripidaeHood,1914(five genera, mostly Neotropical and feeding on dry-wood fungi) – large-legged thrips
- †Scudderothripidaezur Strassen, 1973(included by some authors inStenurothripidae)
- ThripidaeStephens,1829(292 genera in four subfamilies, flower living) – common thrips
- †TriassothripidaeGrimaldi & Shmakov, 2004(two fossil genera)
- UzelothripidaeHood, 1952(one species,Uzelothrips scabrosus)
- SuborderTubulifera
- PhlaeothripidaeUzel, 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.[31][32]
Biology
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/d/d9/Myrsine_howittiana.jpg/220px-Myrsine_howittiana.jpg)
Feeding
[edit]Thrips are believed to have descended from a fungus-feeding ancestor during the Mesozoic,[21]and many groups still feed upon and inadvertently redistribute fungal spores. These live among leaf litter or on dead wood and are important members of theecosystem,their diet often being supplemented withpollen.Other species are primitivelyeusocialand form plantgallsand still others are predatory on mites and other thrips.[14]Two species ofAulacothrips,A. tenuisandA. levinotus,have been found to be ectoparasites onaetalionidandmembracid plant-hoppersin Brazil.[33]Akainothrips francisiof Australia is a parasite within the colonies of another thrips speciesDunatothrips aneuraethat makes silken nests or domiciles onAcaciatrees.[34]A number of thrips in the subfamily Phlaeothripinae that specialize onAcaciahosts produce silk with which they glue together phyllodes to form domiciles inside which their semi-social colonies live.[35]
Mirothrips arbiterhas been found in paper wasp nests in Brazil. The eggs of the hosts includingMischocyttarus atramentarius,Mischocyttarus cassunungaandPolistes versicolorare eaten by the thrips.[36]Thrips, especially in the familyAeolothripidae,are also predators, and are considered beneficial in the management of pests like thecodling moths.[37]
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.[38][39]Besides feeding on plant tissues, thecommon blossom thripsfeeds 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.[40]
Pollination
[edit]Some flower-feeding thrips pollinate the flowers they are feeding on, and some authors suspect that they may have been among the first insects to evolve a pollinating relationship with their host plants.[41]Scirtothrips dorsaliscarries pollen of commercially importantchili peppers.[42][43][44]Darwin found that thrips could not be kept out by any netting when he conducted experiments by keeping away larger pollinators.[45]Thrips setipennisis the sole pollinator ofWilkiea huegeliana,a small, unisexual annually flowering tree or shrub in the rainforests of eastern Australia.T. setipennisserves as an obligate pollinator for other Australian rainforest plant species, includingMyrsine howittianaandM. variabilis.[46]The genusCycadothripsis a specialist pollinator ofcycads,the cones of which are adapted for pollination by small insects.[47]Thrips are likewise the primary pollinators of heathers in the familyEricaceae,[48]and play a significant role in the pollination ofpointleaf 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.[47]
Damage to plants
[edit]Thrips can cause damage during feeding.[49]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.[38][50]FamilyThripidaein particular is notorious for members with broad host ranges, and the majority of pest thrips come from this family.[51][52]For example,Thrips tabacidamages crops ofonions,potatoes,tobacco,andcotton.[39][53]
![](https://upload.wikimedia.org/wikipedia/commons/thumb/a/af/Australian_insects_%28Plate_XXXVII%29_%287268276524%29.jpg/170px-Australian_insects_%28Plate_XXXVII%29_%287268276524%29.jpg)
Some species of thrips creategalls,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.[54]A radiation of thrips species seems to have taken place onAcaciatrees in Australia; some of these species cause galls in thepetioles,sometimes fi xing two leaf stalks together, while other species live in every available crevice in the bark. InCasuarinain the same country, some species have invaded stems, creating long-lasting woody galls.[55]
Social behaviour
[edit]While poorly documented, chemical communication is believed to be important to the group.[56]Anal secretions are produced in the hindgut,[57]and released along the posterior setae as predator deterrents[57][58]In Australia, aggregations of malecommon blossom thripshave been observed on the petals ofHibiscus rosa-sinensisandGossypium hirsutum;females were attracted to these groups so it seems likely that the males were producingpheromones.[59]
In the phlaeothripids that feed on fungi, malescompeteto 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.[18]
Many thrips formgallson plants when feeding or laying their eggs. Some of the gall-formingPhlaeothripidae,such as generaKladothrips[60]andOncothrips,[61]formeusocialgroups similar toantcolonies, with reproductivequeensand nonreproductive soldier castes.[62][63][64]
Flight
[edit]Most insects create lift by the stiff-winged mechanism ofinsect flightwithsteady stateaerodynamics;this creates aleading edgevortexcontinuously as thewingmoves. The feathery wings of thrips, however, generate lift byclap and fling,a mechanism discovered by the Danish zoologistTorkel Weis-Foghin 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 theWagner effectwhich would otherwise counteract the growth of the circulation.[65][66][67][68]
- Clap and flingflight mechanism after Sane 2003
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Clap 1: wings close over back
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Clap 2: leading edges touch, wing rotates aroundleading edge,vortices form
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Clap 3: trailing edges close, vortices shed, wings close giving thrust
- Black circle and heavy line: wing (rachis and bristles); Black (curved) arrows: flow; Blue arrows: induced velocity; Orange arrows: net force on wing
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Fling 1: wings rotate around trailing edge to fling apart
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Fling 2: leading edge moves away, air rushes in, increasing lift
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Fling 3: new vortex forms at leading edge, trailing edge vortices cancel each other, perhaps helping flow to grow faster (Weis-Fogh 1973)
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 km of sea between Australia and South Island of New Zealand.[18]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.[69]
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 ameniscusby arching their bodies and working their way head-first and upwards along the water surface in order to escape.[70]
Life cycle
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/4/4f/Thrips_nymph.jpg/220px-Thrips_nymph.jpg)
Scale bar is 0.5 mm
Thrips lay extremely small eggs, about 0.2 mm long. Females of the suborderTerebrantiacut slits in plant tissue with their ovipositor, and insert their eggs, one per slit. Females of the suborderTubuliferalay their eggs singly or in small groups on the outside surfaces of plants.[71]Some thrips such asElaphothrips tuberculatusare known to be facultatively ovoviviparous, retaining the eggs internally and giving birth to male offspring.[72]Females in many species guard the eggs against cannibalism by other females as well as predators.[73]
Thrips arehemimetabolous,metamorphosing gradually to the adult form. The first twoinstars,called larvae or nymphs, are like small wingless adults (often confused withspringtails) 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 topupae:in these stages, the body's organs are reshaped, and wing-buds and genitalia are formed.[71]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.[74]The adult stage can be reached in around 8–15 days; adults can live for around 45 days.[75]Adults have both winged and wingless forms; in the grass thripsAnaphothrips 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.[76]Thrips can survive the winter as adults or through egg or pupaldiapause.[18]
Thrips arehaplodiploidwith haploid males (from unfertilised eggs, as inHymenoptera) and diploid females capable ofparthenogenesis(reproducing without fertilisation), many species usingarrhenotoky,a few usingthelytoky.[77]InPezothrips kellyanusfemales hatch from larger eggs than males, possibly because they are more likely to be fertilized.[78]The sex-determining bacterialendosymbiontWolbachiais a factor that affects the reproductive mode.[50][77][79]Several normally bisexual species have become established in the United States with only females present.[77][80]
Human impact
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/7/72/Tomato_with_Tomato_Spotted_Wilt_Virus.jpg/220px-Tomato_with_Tomato_Spotted_Wilt_Virus.jpg)
As pests
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/b/b2/Ponticulothrips_diospyrosi.jpg/220px-Ponticulothrips_diospyrosi.jpg)
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 astospoviruses.[81]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.[82]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 thetomato spotted wilt virusand 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.[83]Other viruses that they spread include the generaIlarvirus,(Alpha|Beta|Gamma)carmovirus,SobemovirusandMachlomovirus.[84]Their small size and predisposition towards enclosed places makes them difficult to detect byphytosanitary inspection,while their eggs, laid inside plant tissue, are well-protected from pesticide sprays.[75]When coupled with the increasingglobalizationof trade and the growth ofgreenhouseagriculture, thrips, unsurprisingly, are among the fastest growing group ofinvasive speciesin the world. Examples includeF. occidentalis,Thrips simplex,andThrips palmi.[85]
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 triticiandLimothrips 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.[86]
Management
[edit]![](https://upload.wikimedia.org/wikipedia/commons/thumb/d/d2/Asilidae_thrips.jpg/220px-Asilidae_thrips.jpg)
Thrips developresistancetoinsecticideseasily 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.[87]
Due to their small sizes and high rates of reproduction, thrips are difficult to control usingclassical 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 ofparasitoidHymenopteraparasitize eggs and larvae, theEulophidaeand theTrichogrammatidae.Other biocontrol agents of adults and larvae includeanthocorid bugsof genusOrius,andphytoseiidmites. Biological insecticides such as the fungiBeauveria bassianaandVerticillium lecaniican kill thrips at all life-cycle stages.[88]Insecticidal soapspray 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.[89]
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External links
[edit]![](https://upload.wikimedia.org/wikipedia/en/thumb/4/4a/Commons-logo.svg/30px-Commons-logo.svg.png)
![](https://upload.wikimedia.org/wikipedia/commons/thumb/d/df/Wikispecies-logo.svg/34px-Wikispecies-logo.svg.png)
- Thrips of the World checklist
- Thrips species wiki
- Thrips images from the "Pests and Diseases Image Library (PaDIL)" of AustraliaArchived2020-03-17 at theWayback Machine
- University of California Pest Management Guidelines for Thrips
- University of California Thrips Identification
- CISR: Center for Invasive Species Research Fact Sheets
- Thrips links on theUF/IFASFeatured Creatures Web site
- Frankliniella schultzei,common blossom thrips(Thripidae)
- Heliothrips haemorrhoidalis,greenhouse thrips(Thripidae)
- Scirtothrips dorsalis,chilli thrips(Thripidae)
- Selenothrips rubrocinctus,redbanded thrips(Thripidae)
- Thrips palmi,melon thrips(Thripidae)
- Thrips simplex,gladiolus thrips(Thripidae)