/thrips/, n., pl. thrips.
any of several minute insects of the order Thysanoptera, that have long, narrow wings fringed with hairs and that infest and feed on a wide variety of weeds and crop plants.
[1650-60; < NL < Gk thríps (sing.) woodworm]

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Any of some 5,000 species (order Thysanoptera) of tiny winged insects that feed chiefly on plants, inhabiting temperate and tropical areas worldwide.

Many species damage cultivated plants, by either sucking the sap or transmitting viral plant diseases. A few species are predators. Thrips reach a maximum length of about half an inch (15 mm). Most have two pairs of long, narrow, hair-fringed wings.

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▪ insect order

      any of approximately 5,000 species of insects that are among the smallest of the winged insects and are abundant in the tropical and temperate regions of the world. Thrips are economically important since some species transmit plant (plant disease) viruses. Feeding by thrips may reduce seed production, disfigure flowers and fruits, and damage plant leaves. In warm areas a number of them cause plant galls and leaf rolls. On the other hand, a few species prey on destructive mites and scale insects, and a number may aid in the pollination of flowers.

      Most thrips are 1.5 to 3 mm (0.06 to 0.12 inch) in length, with the smallest being about 0.6 mm (0.02 inch) and the largest about 15 mm (0.6 inch). Because of their minute size they can enter the smallest flowers or tiniest cracks in stems and bark. Their wings (wing), when present, are narrow and fringed, hence the name Thysanoptera (Latin for “fringed wings”). Despite their Lilliputian size, thrips are often elaborately ornamented with hexagonal designs, spines, or body flanges. The bristles (setae) of some are expanded at the tips or elongated, or the body wall may be expanded into prongs or forks. The larvae of thrips tend to be brightly coloured (red, orange, or yellow), whereas the adults range from whitish to brown or black. Adults have extensible bladders on the tarsi of the legs, as suggested by the common name bladder feet. Thrips is both singular and plural.

Natural history

Life cycle
      The egg stage is followed by two larval stages, during which all the immature feeding occurs. The development of thrips is unusual in that, although they belong to a group of insects that usually undergo simple metamorphosis, they have pupae resembling those that occur in insects that undergo complete metamorphosis. There may be one, two, or three pupal stages in which no feeding occurs.

      The primary method of reproduction among thrips is by bisexual union. The males frequently are shorter lived than females, and mating often takes place early in the life span, when the adults first appear. Reproduction by unfertilized eggs (parthenogenesis) has been known to occur occasionally.

      Thrips in the suborder Terebrantia always lay eggs, as do most thrips in the more advanced suborder Tubulifera. Hatching of eggs within the female (ovoviparity), occurs in some highly evolved members of the Tubulifera. Eggs are elongated, oval to kidney shaped, faintly to distinctly sculptured, and, as is the case in other minute insects, are proportionately large, often occupying two or three abdominal segments of the female before laying. Ordinarily, eggs of the Terebrantia are inserted into living plant tissue in a slit prepared by the sawlike ovipositor of the female. By contrast, eggs of the Tubulifera, whose females have no ovipositor, are pushed under bark, in or between flowers and leaf sheaths, or are glued onto leaf or bark surfaces. Eggs take from 3 to 16 days to hatch according to the species and conditions.

      Although similar in overall form to the adult, the larvae of thrips are smaller and simpler in structure. The long anal bristles characteristic of larvae may hold droplets of excreted liquid that are deposited on leaf surfaces. The presence of dark-coloured spots of excreta adjacent to light-coloured feeding zones are some typical signs of thrips damage.

      The first larval stage is short, lasting only a few days. The second may last up to many months when estivation or hibernation occurs. In the Terebrantia the second stage larva transforms into a prepupa, whereas in the Tubulifera an abbreviated form, the primipupa, is part of the sequence of development and is followed by the prepupal form. The prepupal stage lasts only a few hours to several days and gives rise to the final pupal stage, which may be of short or long duration. Pupation takes place in the ground in earthen cells (in the case of many advanced Terebrantia), in cocoons (in many primitive Terebrantia), or on a host plant (in many advanced Terebrantia and the Tubulifera). Some thrips, for example, the ear thrips, have only one generation per year, whereas others such as the onion thrips may have several generations each growing season.

      In regions having cold winters, thrips hibernate as adults or immature forms in detritus, hollow stems, or in the ground. Those that perish with the cold are replaced each year by spring migrants from warmer areas. In the tropics, many thrips are more active in the wet than in the dry seasons. Some thrips are sensitive to relative humidity. Although larvae are generally found in shady locations, on the underside of leaves or branches, adults occasionally occur on the upper side of leaves in bright light. Most thrips rest tightly against leaf veins or in crevices. They are primarily active during the daylight hours.

      Thrips are preyed upon by many insects (including other thrips), mites, birds, salamanders, and lizards. However, heavy rains, winds, and dust are probably as destructive to thrips as are predators.

Form and function

The larva
      In the larvae, the compound eyes are composed of a few facets, simple eyes (ocelli) are absent, antennal segments are fewer than in the adult, the middle portion of the body (thorax) is simple, the lower part of the leg (tarsus) is always one segmented (although also occasionally so in some adults), and the wings and genitalia are not developed. Larval bristles (setae) often differ in form, number, and arrangement from adult setae. Occasionally additional processes such as spines, combs, and plates are found on larvae.

The adult
      The head of a thrips is slightly skewed with the mouth cone directed posteriorly. Generally the antennae, which in most species are nine-segmented, are frequently reduced by fusion and project forward in front of the eyes. Of the mouthparts only the single left mandible is developed and the maxillae are modified into piercing stylets. Both the maxillae and the labium bear segmented sensory projections (palps).

      The middle portion of the body (thorax) bears the legs. The first part of the thorax (prothorax) of thrips bears the forelegs, the inner parts (coxae) of which may exhibit ridges that when rubbed with spurs from the lower parts (femurs) emit sounds, although these cannot be detected by the human ear without amplification. The remainder of the thorax (pterothorax) bears the wings as well as the mid and hind legs. The wings are usually fairly simple and straplike. At rest they are laid over the abdomen but never folded. Typically the wings have long posterior hairlike fringes with shorter fringes occurring on the leading edge.

      The abdomen is elongated and generally flattened dorso-ventrally, especially in the Tubulifera. There are 10 distinctive segments with a rudiment of the 11th. The first and eighth abdominal segments bear breathing holes (spiracles). Often in the Tubulifera, several setae on the back are S-shaped and hook into the wing fringes to hold the wings at rest. In the Terebrantia the terminal segment is rounded in the males and split ventrally up to the eighth segment in the females. In the Tubulifera the terminal segment is tubelike.

      The male genital organs are composed of paired appendages and an unpaired copulatory organ (aedeagus), all of which retract into the abdomen. The paired testes and accessory glands with their ducts occur in the posterior half of the abdomen. The female of the Terebrantia usually has an ovipositor composed of two pairs of sawlike blades. These are absent in the Tubulifera. There are eight egg sacs (ovarioles) and spherical, often pigmented, seminal receptacle for storage of sperm acquired during mating.

Evolution, paleontology, and classification
      The Thysanoptera are part of the Psocoptera or Corrodentia-Hemiptera phyletic line of insects. Their origin may have been correlated with the development of the left mandible as an efficient chisel for piercing pollen granules. Possibly during the period when ancestral types were pollen feeders early thrips proliferated into a distinct order, which has since become diverse. Because of their small size, the development of fringed wings seemingly was favoured to overcome disproportionate drag on the wings' upstroke.

      The earliest fossil, Permothrips longipennis, seems closest to the family Aeolothripidae, the most generalized and perhaps ancestral type. All extant families are known from fossils embedded in amber. The most advanced groups are the families Thripidae and Phlaeothripidae. The terebrantian families mostly retain their habit of feeding on vascular plants and pollen. Many members of the single extant tubiferan family Phlaeothripidae have adjusted to feeding in detritus.

Distinguishing taxonomic features
      Although no particular structure is absolutely indicative of a taxonomic level, many features are useful in separating the major groups of Thysanoptera. The form of the antennae and its appendages, the shape and structure of the forewing, abdominal characters including form of the ovipositor in females, and glandular and setal location are important characters for determining the familial relationships. At the generic and species level antennal details, mouthparts, and body sculpturing are useful taxonomic characters, along with colour and bristle arrangement.

Annotated classification
      The classification presented here is based on that of H. Priesner (1957, 1960).

Order Thysanoptera
 Insects averaging about 2 mm; unique in that the right mandible is atrophied, resulting in an asymmetrical mouthcone; wings, when present, bear long fringes. Extensible bladders on the tarsi of the adults.
      Suborder Terebrantia
 The 10th and last abdominal segment, rarely tubelike, always split ventrally, major anal setae arising from subapical region, never from separate platelets; with sawlike ovipositor in female; wings usually with fine hairs (micotrichia) and at least 1 longitudinal vein; larvae with intermediate antennal segments bearing rings and microsetae; pupae with antennal sheaths either extended forward of head or lying over head; eggs without polar thickenings. 2 fossil families, Permothripidae and an undetermined family are, respectively, from Permian and Jurassic periods. Present families include the following.

      Family Aeolothripidae
 Oligocene (Baltic amber) to present. Worldwide. Antennae 9-segmented; ovipositor may be upturned or straight; forewings broad and rounded at tips, surface with microtrichia and several longitudinal and cross veins; antennal sensors on intermediate segments in form of linear or circular disks.

      Family Merothripidae
 Oligocene (Baltic amber) to present. Worldwide. Antennae 8- or 9-segmented; ovipositor downturned, often weakly developed; forewings narrow, surface smooth; antennal sensors on intermediate segments disclike.

      Family Heterothripidae
 Cretaceous (Cedar Lake amber) to present. Western Hemisphere and India. Antennae nine-segmented; ovipositor downturned; forewings narrow, surface with microtrichia; antennal sensors on intermediate segments as flat discs or protruding peglike cones.

      Family Thripidae
 Oligocene (Baltic amber) to present. Worldwide. Antennae 6- to 9-segmented; ovipositor downturned, rarely weakly developed; forewings narrow, with microtrichia; antennal sensors on intermediate segments as simple or forked sense cones.

      Suborder Tubulifera
 The 10th abdominal segment tubelike, never split, major anal setae arising from separate plates adjacent to the tube; females without ovipositor; wings without longitudinal veins or fringe; larvae with antennal segments smooth not ringed; pupae with antennal sheaths hornlike or curved around (not over) head; eggs often with a tubercule on anterior pole.

      Family Phlaeothripidae
 This is the only family currently assigned to the suborder. Representatives occur throughout the world are known from Oligocene (Baltic amber) to the present.

Critical appraisal
      The subordinal disposition of the thrips is well established, but the ranking of family groups as given above is disputed by some who would raise the families to superfamily rank. The order was earlier (1838) designated the Physapoda by H. Burmeister, given in allusion to the swollen tarsi, the so-called bladder feet.

Lewis Judson Stannard, Jr.

Additional Reading
The literature on Thysanoptera is widely scattered throughout numerous entomological journals with occasional summaries given in monographs or lists. The monographs are predominantly faunal studies, though they often include regional revisions of some of the taxa. Only a few complete generic revisions have been published. Some of the latest, though not necessarily recent, faunal monographs are S.F. Bailey, “The Thrips of California, Part 1: Suborder Terebrantia,” Bull. Calif. Insect Surv., vol. 4, no. 5, pp. 143–220 (1957); H.E. Cott, Systematics of the Suborder Tubulifera (Thysanoptera) in California (1956); L.J. Stannard, “The Thrips, or Thysanoptera, of Illinois,” Bull. Ill. Nat. Hist. Surv., vol. 29, no. 4, pp. 215–552 (1968); G.D. Morison, “Thysanoptera of the London Area,” Lond. Nat., reprint no. 59 (1947–49). Two comprehensive catalogs and annotated synopses are C.F. Jacot-Guillarmod, “Catalogue of the Thysanoptera of the World (Part 1),” Ann. Cape Provincial Museums (Nat. Hist.) (1970); and L.J. Stannard, The Phylogeny and Classification of the North American Genera of the Suborder Tubulifera (Thysanoptera) (1957). Proposals for the classification of the thrips that have received general acceptance are those by H. Priesner, “Zur Vergleichender Morphologie des Endothorax der Thysanoptera,” Zool. Anz., vol. 159, no. 7–8, pp. 159–167 (1957), and “Das System der Tubulifera (Thysanoptera),” Anz. öst. Akad. Wiss., 13:283–296 (1960). Morphological treatments of the Thysanoptera are found in R.G. Davies, “Observations on the Morphology of the Head and Mouthparts in the Thysanoptera,” Proc. R. Ent. Soc. Lond. (A), vol. 33, no. 7–9, pp. 97–106 (1958); and P. Pesson, “Ordre des Thysanoptera,” in P. Grasse, Traité de zoologie, vol. 10, no. 2, pp. 1805–1869 (1951). A source of ecological information on thrips is that by L. Cederholm, Ecological Studies on Thysanoptera (1963).Lewis Judson Stannard, Jr.

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Universalium. 2010.

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