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Annu. Rev. Entomol. 1991.36:431-457. Downloaded from www.annualreviews.org by Universidad Nacional de Colombia on 02/05/12. For personal use only.
David N. Byrne
Department of Entomology, University of Arizona, Tucson, Arizona 85721
Thomas S. Bellows, Jr.
Department of Entomology, University of California, Riverside, California 92521
Aleyrodidae, Homoptera, life history, population dynamics, migration
The importance of whiteflies as economic pests seems to expand continually (3 1 ,78, 1 20). These homopteran insects damage crops by extracting large quantities of phloem sap, which can result in greater than 50% yield reduc tions ( 1 19). The honeydew excreted by these insects serves as a medium for sooty mold fungi (e.g. Capnodium spp.) that discolor parts of the plants used for food and fiber ( 14 1 , 1 42). Finally, a few species serve as vectors of several economically important viral plant pathogens ( 1 33). This article reviews characteristics of whiteflies that set them apart from other members of Stemorrhyncha and presents a review of whitefly literature to obtain a better understanding of their biology and assess the status of whitefly research. The family Aleyrodidae is considered to have two subfamilies. The Aleuro dicinae, endemic primarily to Central and South America, may be considered the more primitive taxa because of more complex wing venation (81, 131). However, the increased venation may be necessary because Aleurodicinae members are generally larger than whiteflies of the subfamily Aleyrodinae. This increased size [> 2.0 mm long (81)] likely requires greater wing support. The Aleyrodinae is larger in terms of number of species and is also more widespread.
0066-4 1 70/91/010 1 -0431 $02.00
BYRNE & BELLOWS
Enderlein (72) suggested a third subfamily, Udamoselinae on the basis of one specimen of a South American species, a male with a body length of 7 mm. Today the existence of the subfamily is thought to be dubious (131). HISTORICAL PERSPECTIVE Whiteflies are considered the tropical equivalent of aphids owing to their ordinal characteristics and their scarcity in temperate climates. Reaumur first described whiteflies in 1736, although he mistakenly placed Aleyrodes pro letella in Lepidoptera (64). In 1795, Latreille correctly placed them in Homoptera (64). The majority of the literature from the 19th century was taxonomic in nature and consisted primarily of descriptions of pupal charac teristics (e.g. 63, 121). Pupal cases are still generally of more value than other life stages when making taxonomic decisions. Maskell (121) reports that, from the time of the insects' first description in 1736 until 1895, little more than 50 noteworthy articles were published on them. Many of these papers were taxonomic treatments with little information about whitefly biology. Quaintaince (149) in 1900 solved many of the taxonomic problems con cerning the whiteflies of North America. Work by Bemis (20), who helped bring the total number of described North American species to 62, also did much to end taxonomic confusion. Trehan (162) and Mound & Halsey (131) played a large role in correcting global whitefly taxonomy. To date, however, little has been accomplished concerning whitefly systematics. The most comprehensive work available is Whitefly of the World (131); this very useful publication presents a list of genera and species but says little about systemat ic relationships. Data on whitefly cladistics appear to be unavailable. Information on basic biology has been even slower in developing, and early research was in part restricted to pest species. Back (14) was concerned about the presence of Aleurothrixus floccosus on citrus. Lloyd (119) published a report on the biology of Trialeurodes vaporariorum, but his primary intent was to describe greenhouse management strategies. Bemisia tabaci was first noticed on cotton in India in 1905 (97). Bionomic investigations did not begin in earnest, however, until several years later, when this whitefly was shown to seriously damage crops. After the tum of the century, more attention was focused on basic biology. Morrill & Back (127) examined the biology of Dialeurodes citri/olii and Dialeurodes citri. Hargreaves' (89) examination of T. vaporariorum in 1915 was another early report on whitefly bionomics and Garman & lewitt (76) studied whitefly biology in greenhouses. Because of concentration on pest species, particularly B. tabaci and T. vaporariorum, much of our data relate to a limited number of the > 1,200 known species. The most closely examined species are those that feed on a wide variety of herbaceous hosts. Polyphagy, however, is not usually re-
Annu. Rev. Entomol. 1991.36:431-457. Downloaded from www.annualreviews.org by Universidad Nacional de Colombia on 02/05/12. For personal use only.
Pealius. while always difficult. Aleurolonga. All are plant feeders with piercing. Nevertheless. Butani (26). Entomol. such evidence should not be consid ered conclusive unless it is connected to additional evidence such as the presence of sibling species. the following description offered by Mound & Halsey (131) is the best available and was undoubtedly accurate when published:
Three of the largest whitefly genera. For example. As a result.36:431-457. Aleurotrachelus and Tetraleurodes. Many of the other genera. and these determinations are becoming even more difficult as humans facili tate whitefly movement. Bellitudo and Crenidorsum. The genera of the subfamily Aleurodicinae are almost entirely confined to the Neotropics. and Cock (45). 1991.
Annu. Corbettia and Dialeurolonga are recorded only from Africa and Madagascar. have a more restricted distribution.
ORIGIN AND DISTRIBUTION
The geographic origin of many whitefly species is largely speculation. Dialeurodes. In contrast Africaleurodes. most of which are monophagous or oligophagous animals associated with woody perennial hosts. drawing conclusions about the family based on what is known about a few species might not be appropriate. however. The genus Trialeurodes has most of its species in the New World as does Aleuroparadoxus.
Whiteflies share many characteristics with other homopterans. Bibliographies of the family include those by Trehan & Butani (163).WHITEFLY BIOLOGY
corded in Aleyrodidae. Aleurothrixus. For personal use only. Acaudaleyrodes. Although such botanical associations pro vide some information concerning origin. are widely distributed in the Ethiopian and Oriental Regions. we have learned a great deal about the biology of whiteflies from work that concentrates on nonpest species. Fortunately. Aleuroplatus. Thompson & Reinert (161).org by Universidad Nacional de Colombia on 02/05/12. sucking mouthparts. The process of making these determinations. Identifying whitefly distribution is troublesome. Dialeuropora and the largest genus. and this is also true of a few genera of the Aleyrodinae such as Aleurocerus. Odontaleyrodes and Rhachisphora are also particularly common in the Oriental and Austro-Oriental regions. They are opisthognathus. Dialeurodes chittendeni is thought to have a Himalayan origin because of its connection with rhododendrons. Downloaded from www. Aleuropteridis. whereas Orchamoplatus is apparently most common in the Pacific . Aleurocanthuys. Information is available concern ing > 25 others. Aleurotubercularus. becomes particularly hard with a family that has not had the attention of entomologists for a long period of time. Addi-
. Aleurolobus.annualreviews. which also originated there (115). are more or less artificial assemblages of species with black pupal cases reported from many parts of the world. Rev.
Opaque white wax may be produced as a marginal fringe (e. The number of these rays in some nymphs differs when the whiteflies are reared on different hosts (129). or as setae-like projections (81). Rev. Vasiform Orifice One set of features that sets the Aleyrodidae apart from related familes is the presence of a vasiform orifice with its operculum and lingula (81). as plumes. The material is extruded from wax plates that consist of rows
Annu. the lingula is cocked down into the liquid. dorsally as tufts (Aleurotrache Ius jelinekii and Aleuroplatus coronata). vaporariorum. the honeydew is . and comes in two colors (81). if ever. 151). The dorsal placement of the anus in Aleyrodidae permits the effective handling of the honeydew that these phloem-feeding insects produce in copious amounts. In psyllids the anus seems to h ave been displaced by enlarged genitalia ( 130). It is not the anus. wax can appear as a gelatinous mass. The other is brilliant white. Entomol. although it may reflect the color of the surrounding leaf surface.org by Universidad Nacional de Colombia on 02/05/12. Some of the waxes are similar in form to those found on coccids ( 1 22).annualreviews. In whitefly nymphs. others are not. can produce extracuticular waxes that cover the body. Other colors rarely. 1991.
. They flip their excreta away: the honeydew fills the vasiform orifice. This variance results from changes in the number of papillae (56. or as flocculent or woolly mats (A. In females. Honeydew can present problems. because it is viscous and associated with sooty mold fungi. The transparent form usually appears as a very thin layer over the entire dorsal surface. it extends to the eighth abdominal segment (87).). when the lingula is released. floccosus and Aleurodicus dispersus). but rather the depression into which the anus empties the contents of the digestive tract (honeydew).434
BYRNE & BELLOWS
tionally. A dorsal anus is also found in psyllid adults (rarely in the nymphs) and it also functions to rid these insects of honeydew. catapulted away. occur in this family although the marginal fringes in some species of Trialeurodes tend to have light yellow or orange tints (81). For personal use only. Whitefly nymphs cannot walk away from their honeydew droplets. adults of both sexes have four membranous wings. Downloaded from www. as columns.36:431-457. the wax forms tight curls of threads approximately 1 /-Lm in diameter (35). in Tetraleurodes spp. The vasiform orifice is generally located on the dorsum of the ninth abdominal segment of males. except the egg. members of the family undergo incomplete metamorphosis (with certain com plications). In B. Finally. tabaci and T. Waxes Whiteflies are distinctive in that all life stages. One is clear or colorless. particularly for sessile nymphs. Wax in the adults takes on a different appearance.g. and. Transparent wax may also appear as a marginal fringe or as dorsal spikelike wax rays such as those found in Trialeurodes spp.
the fourth nymphal instar is commonly referred to as a pupa (20. Hinton (91. The dimensions of most whitefly adults. Hind and forelegs distribute the particles over the wings and the rest of the body (except the eyes). 1991. LIFE HISTORY Size Gill (81) reports that the physically largest genus is Aleurodicus spp. has the red eyes and the yellow body pigment of the adult. R. T. which has an adult body length of> 2 mm and a wing expanse of > 3. spinelike processes.36:431-457. free fatty acids. Downloaded from www. in fact. however. What takes place during the whitefly's fourth nymphal instar is. however. This form according to Gill (81) feeds and so clearly is not a pupa. is expanded and opaque-white with dorsal and lateral waxy. The term implies that whiteflies exhibit a degree of holometabolism. At this stage. he was unable to observe a distinct stadium that intervened between the last nymphal stage and the adult.81). stated that the fourth nymphal instar has three morphologically distinct forms. Rev. discussing T. For personal use only. and D. more closely approximate a range with B. that even after careful histological examination. Each microtri chium is associated with a wax canal. tabaci at the lower end.. The early fourth instar is flattened and transluscent.
Fourth Nymphal Instar In the literature. Nechols & Tauber (134). vaporariorum in midrange. The material is extruded as a continuous ribbon but is broken off as curly particles when the animal's hind tibiae pass over the plates (35). The next form. At this point. distinct from what occurs in holometabolous families. Although we do not consider Hinton's interpretation to be correct. Entomol.annualreviews. apolysis is complete and the adult cuticle is laid down.0 mm. we believe that the word pupa has been so inculcated into whitefly literature that its replacement would only cause confusion. alco hols. we would reserve the term pupa for the last. and hydrocarbons (35). Females have two pairs of these wax plates. J. Nechols (personal communication) states.
. nonfeeding portion of the last nymphal stadium found after apolysis has occurred and refer to the earlier portion of the last stadium as the fourth nymphal instar.5--4. Pupation has also never been reported to occur in other homopterous families.
Annu. vapor ariorum.WHITEFLY BIOLOGY
of microtrichia found on the ventro-lateral abdominal surface. For functional purposes. The last stage. males have four. 92) was certain whiteflies had a pupal stage in the sense that this stage serves as a mold for some of the imaginal muscles. the transitional substage.org by Universidad Nacional de Colombia on 02/05/12. The wax of the two species consists primarily of triacylglycerols (65-75%) with a trace of wax esters. when the pharate adult form is present. apolysis takes place.
0.13 :t 0. Weber
(174) observed that T.436
BYRNE & BELLOWS
citri at the upper end (Table 1). Poinar (144) speculated that egg pedicles of A. When the pedicel is inserted into parenchyma cells. the protoplasm withdraws and the pedicel becomes a hollow tube. In most species.07 0. The literature describes pedicels into stomata
(58.91 :t 0. Weber postulated that
Body measurements (mm) of three species of Aleyro
Body length Wing expanse
Bemisia tabaci male female
0. very little of this material is present. Pedicels inserted into interstitial spaces are associated with a much larger quantity of the glue.85 :t 0. S.99
1. 12 0.06
2.03 0. tabaci.20
. 138. in addition to providing a
means of attachment. 144).06 0 .org by Universidad Nacional de Colombia on 02/05/12.49 3. For personal use only.05 0. Bellows. Quaintance & Baker
(150) believed the pedicel. After fertilization. each
ovariole contains one or more follicles and a germarium.05
female Dialeurodes citri
1. 144) and four that utilize slits in the leaf epidermis
were inserted into the stomata because the epidermis of grasses and sedges contains large amounts of silica and lignin and would be difficult for females to penetrate. 1991. Entomol.03 0. Rev. Gameel
(75) depicts 15 ovarioles in B. occiduus
13 species that insert their (138. This observa tion applies to late nymphal instars of D. Sexual dimorphism in adult body dimension is 1).04
1 . Downloaded from www.annualreviews.36:431-457. males are smaller (Table
The internal reproductive system and oogenesis in whiteflies is similar to that of other homopterans.04 0. citri (T.
Whitefly eggs generally are pyriform or ovoid and possess a pedicel that is a peglike extension of the chorion. during which time the pedicel is filled with protoplasm.
Annu. 20 1 .06
2. vaporariorum secretes a gluelike substance
around the pedicel. served as a guide for spermatozoa during fertilization.
usual for members of Aleyrodidae. The
pedicel is either inserted into a slit made by the ovipositor in the leaf surface or into a stomatal opening. chittendeni (177) and Tetraleurodes acaciae (66). 86.41 2. but the eggs of
Aleurocybotus occiduus lie on their side (144). unpublished) as well as D. eggs assume an erect stance.
the mean rate of oviposition was 252 eggs per female. Dittrich et al (61) found that Sudanese whiteflies have a fecundity on cotton of 344 eggs. In December and January (DMT 14. We interpret Gameel's (75) results on
B. Hinton (93) made the claim that the pedicel was involved in the transfer of water into the egg but offered no empirical evidence. Citing Weber (174) and Wiggels worth (175).
support the argument that the pedicel must absorb water from the plant. Sudanese strains of
early fall. tabaci in Sudanese cotton fields to reveal a fecundity of 160. For the same species.
B. During October and November. Gameel (75) reports a similar substance surrounding the egg pedicle of
B. Recently Byrne et al (33) assayed eggs raised on plants irrigated with tritiated water and demonstrated that water extracted from plant tissue accounted for approximately 50% of the mass of a mature whitefly egg.36:431-457. vaporariorum at 18°C was 319.2°C (29).7°C).org by Universidad Nacional de Colombia on 02/05/12. the mean rate was 61 eggs. Rev.6 eggs per female (69).annualreviews. However. Poinar (144) used the fact that eggs of
A. Entomol. ranging from 8. Downloaded from www. but produced 81
eggs at 26. tabaci.WHITEFLY BIOLOGY
water passes osmotically across this colloidal mass and enters the egg through the pedicel." Several other authors have also suggested that aleyrodids use the pedicel as a means of absorbing water into the egg. and that it fell to 5. The eggs are laid indiscriminately. Tetraleurodes stanjordi
. The fecundity of nymphal host plant. Husain & Trehan (99) report a fecundity of
B. occiduus dried up when removed from the leaf to
Annu. tabaci deposit a few eggs on the leaf upon which they emerge as adults and then move to newer growth.5°C. tabaci appear to be much more prolific than strains
from other parts of the world (80). and Avidov (10) gives a fecundity value of
approximately 50 eggs on eggplant. The maximum number of eggs per female varied from 48 to 394. Von Arx et al (168) calculated the fecundity of a Sudanese strain to be 127.7°C and 72 at 32. Whiteflies such as B. Fecundity increased fivefold when the insects were placed on a new leaf (25).3 to 39. Oviposition rates also vary greatly in other species and are affected by environmental conditions and host plant. and associates the production of this material with a
"cement gland.9°C.4 eggs during
43 eggs on cotton in India. During July and August when the daily maximum temperature (DMT) was 28. 3°C). Burnett (25) reports that the fecundi ty of
T.5 at 33°C and to Aleurocanthus woglumi varied with the
zero at 9°C.
Oviposition habits differ somewhat between species. tabaci reared on cotton in
Egypt in an open-air insectary.5.5 eggs. 1991. Azab et al (13) examined oviposition rates of
B. tabaci from Arizona cotton laid no eggs at 14. These results occurred under conditions of constant temperature and light. the mean was 204 the (DMT 22. Parabemisia myricae limits oviposition on citrus to very young leaves and oviposits on the underside of the leaves as well as on the leaf margins. For personal use only.
crawlers walk rather quickly over the leaf surface in search of an available minor vein.36:431-457. T. Emergence behavior of
A. tabaci on young (5-leaf stage) vs mature (>
25-leaf stage) lettuce. it abandons that pattern on pubescent leaves (47). occiduus
crawlers is slightly different because these eggs are laid on their side (144). also oviposit eggs in a circular fashion.1 % on the young lettuce. As the first-instar nymph of B. Nymphs of all nymphal stadia of the Aleurodicinae have three-segmented legs. it bends in half until its front legs can clasp the leaf. after which it walks away from the spent chorion. For personal use only. After settling.annualreviews. found 100% mortality on the mature lettuce and 58. 155). They ascribed this difference to changes in the nutrition al quality of the plant because crawlers were equally successful in reaching the phloem tissue of both plant stages. floccossus and Para leyrodes sp. woglumi lays its eggs in a spiral arrangement (86). Domenichini (62) suggests an additional segmentation or suture in legs of second. woglumi. While the nymphs usually settle in a few hours. Crawlers (first-jnstar nymphs) of Aleyrodinae have functional walking legs (with three apparent segments) and antennae (with two apparent segments).
found to move between plants (67). Some whiteflies oviposit
their eggs in an arc or circle while their mouthparts remain inserted. Downloaded from www.
vaporariorum lays its eggs in a circular fashion on glabrous leaves. Walker also suggested that the possible presence of a probing deterrent might interfere with feeding. Byrne.438
BYRNE & BELLOWS
lays its eggs on both sides of the leaf (D.3%) and the distances traveled were short « 30 mm). The portion of the population moving between host plants.org by Universidad Nacional de Colombia on 02/05/12. com paring crawler survival of
B. myricae on mature lemon leaves. although the divisions are indistinct. Some crawlers.through fourth-instar nymphs. Crawler mortality has been attributed to several plant characteristics. Byrne & Draeger (34).
Annu. Rev. 1991.
. although A. Entomol. N.
Following completion of development. Some species possibly use the nymphal legs to assist in casting exuviae (81). usually on the same leaf upon which the egg was laid. was small (approximately 0. personal observation). the egg cracks at the apical end along a longitudinal line of dehiscence. they insert their mouthparts into the phloem tissue and begin extracting sap. in cluding cuticular thickness and nutritional factors. In warmer summer conditions. the process can take several days in cooler weather (10. however.and third-instar nymphs appear to have only one segment (81). He
attributed this mortality to the fact that the thick cuticle of mature leaves prevents penetration. such as those of
A. Fourth-instar nymphs have distinct legs and antennae. Legs and antennae of the second. Walker (169) reported a high degree of crawler mortality for
P. tabaci begins to emerge.
whiteflies are phloem feeders. Few emerged during hours of darkness. tabaci and T.3°e. Similar patterns of emergence were found for B.
Under a constant temperature of 29. While some members of the Trialeurodes spp. personal communication). tabaci adults emerged from their pupal cases between 0600 and 0930 hours (lights on occurred at 0600 hours) (94).e. tabaci possesses dorsal setae and Siphoninus spp. Entomol.
Annu. Gill (8 1 ) provides a recent review of the morphology of nymphs.7 h) ( 1 03). vaporariorum is approximately 4 h 10 min at 27°C (38).WHITEFLY BIOLOGY
If crawlers successfully reach the phloem of an appropriate host. For personal use only. suggesting the presence of a circadian system. A great deal of information is available about the nutritional requirements of other homopterans. vaporariorum (38).and third-instar whitefly nymphs have an oval or elongate-oval body. J. Emergence patterns persisted under conditions of continuous light or darkness. at 10°C and 1 6 . Some nymphs may be circular or nearly heart-shaped. and adults for the family. two thoracic and one caudal (R.org by Universidad Nacional de Colombia on 02/05/12. 8 h. These form a passage to the spiracles and may assist in conduction of air. ec1osion of 50% of the total number of adults) and temperature. The teneral period (between final ecdysis and first fight) for B. Downloaded from www. they remain sessile until they reach the adult stage. Under a series of constant temperatures. tabaci and T.
. No emergence was observed at temperatures below 1 7 ± 0. proletella the teneral period was inversely related to temperature (57. This difference suggests that feeding takes place during the teneral period.
Most second. 90% o f B. Gill.
NUTRITION AND EXCRETION Nutrition
As far as is known. pupae. 1991. A. 4 h at 25°C) and was longer when the animals were reared on young leaves ( 1 6. possess dorsal papillae-B. a significant inverse correlation was found between the time of median emer gence (i. For A.4 h) than on older ones (5. particu-
.36:431-457.6°e and a photoperiod of 14: 1 0 LD. have dorsal siphunculi most species have a relatively simple dorsum. except for brief periods during molts.5 ± 0. floccosus was reported to eclose primarily between 0600 and 0900 hours in Hawaii (139). and the peak time of emergence was delayed when temperatures were fluctuated. Whitefly nymphs have shallow breathing folds in the ventral body wall.annualreviews. The folds probably developed in response to the fact that nymphs have a flattened body and lie closely appressed to the leaf surface. Rev.
copulation takes place within 1 to 8 h following eelosion by B. tabaci feeding on the two hosts. tabaci from the pupal case. it was present in high concentrations in the honeydew. This result indicates that these amino acids were metabolized either by the whiteflies or by the symbionts housed in their mycetocytes (96).04%) (37). trahalulose (1-0-a-D-glucopyranosyl-D-fructofuranose) (37). approximately half were found at significantly lower levels in the honeydew produced by B.440
BYRNE & BELLOWS
Annu. males within the first 10 hours following pupal eelosion. 125. 1991. The honeydew of whiteflies on both hosts also contained melezitose. The most noteworthy discovery was that of a disaccha ride.annualreviews. Rev. For personal use only.36:431-457. glucose. Although several may be involved. ADULT CHARACTERISTICS
During summer months. These pairings follow a complex mating behavior (117). is relatively inexpensive energetically to produce (particularly in the presence of so much carbon). Downloaded from www. and of the honeydew produced by B. Entomol. males are allowed to initiate courtship. copulation takes place during the three days following eelosion (0). Carbohydrates found in the phloem sap were common transport sugars and their constituents (e. During
. and fructose in poinsettia and these plus stachyose and raffinose in pumpkin) (37). The pre dominant amino acid in honeydew was glutamine (> 50% of the total amino acid content). This lack suggests that certain amino acids may be used to discharge nitrogenous compounds. 1 32). its production has never before been associated with members of the Insecta. Of the 1 4 or 1 5 amino acids found in the phloem sap. Excretion No uric acid or hypoxanthine was found in the honeydew of B. tabaci females are attracted to. although xanthene occurred at a low level (0. 42. B. 1 56). Additionally. six amino acids not found in the phloem sap were found in the honeydew of whiteflies feeding on both hosts. tabaci feeding on poinsettia. glutamine is a likely candidate. Although trahalulose has been synthesized in the laboratory and appears to be created by certain microorganisms 02. During the fall and spring.org by Universidad Nacional de Colombia on 02/05/12.g. Byrne & Miller (37) analyzed the carbohydrate and amino acid contents of pumpkin and poinsettia phloem sap. 1 6 ± 0. but avoid. After that time. 1 26. tabaci feeding on these host plants. 52-54. a trisaccharide common in the honey dew of aphids ( 1 37). and has a high nitrogen to carbon ratio. but until recently almost nothing was known about the needs of whiteflies. Butler (27) stated that Aleurodes brassicae copulate even before their wings have dried and pigmentation is complete. sucrose.
larly aphids (7-9.
These authors (116) also state that pheromones are much more important during T. However. Downloaded from www. tabaci (117). England.annualreviews. an American race showing arrhenotokous partheno genesis (which seems to be the more usual mode of reproduction) and an
. he then moves parallel to the female.and metathoracic legs. Hargreaves (89) and Williams (176). The female may flap her wings or push the male away to discourage him at this point. tabaci. Entomol. and another male doesn't interfere. vapor ariorum (6) and B. Mated females may produce both males and females (XO and XX). antennuation ceases and the male raises the pair of wings closest to the female. A comparison of the sexual behavior of B. The female terminates copUlation by prying the male free with her meso. Li & Maschwitz (116) state that if pheromones are involved in the courtship behavior of B. the female often rejects the male by flapping her wings or pushing the male away with her mesothoracic legs. Copulation lasts from 125 to 265 s. Rev. vaporariorum courtship and that males can detect females from some distance (at least 5 cm). working with T. 116) reveals a great many similarities and some differences. reported that unfertilized eggs laid by virgin females gave rise exclusively to females. they are active over only a few millimeters. This occurs in about 15% of the observed pairings. vapor ariorum had two races. The abdomen of the male is bent upward at nearly a 90° angle and the aedeagus also is bent at a 90° angle. The male claspers are open and the aedeagus protruded as he tries to clasp the female terminalia. one species.36:431-457. The male drums the flagellum of the female antennae with his antennae while moving his abdomen up and down in synchrony with antennal drumming. tabaci with that of T. produce male offspring (XO). For personal use only. myricae. myri cae.
phase I . Unlike aphids.org by Universidad Nacional de Colombia on 02/05/12. During this phase.WHITEFLY BIOLOGY
Annu. the terminal flap that covers the female gonophore is pulled open. In phase II. and antennae of both sexes are held at a 24° angle to the horizontal axis of the head. When females accept males.
Most whiteflies reproduce by arrhenotoky. except in P. vaporariorum in Merton. Other investigations indicated that T. whiteflies do not experience host alternation or seasonal sexual phases. If she doesn't move. Unmated females. vapor ariorum (114. apparently reproduces by thelytoky and its popUlations consist entirely of females. bringing the aedeagus parallel to the longitudinal axis of the female body when it is inserted into the gonopore. the male encircles the female several times before placing a foretarsus or antenna on the edge of her wing. 1991. During phase IV . the male is situated parallel to the female. P. The male then places his abdomen beneath the female at a 25° angle. Polygyny and polyandry occur in T. Phase III involves pushing the female with the side of the male's body.
and 22 chromosomes are found in all segmentation nuclei and later on in the nymphal mitoses. polyphagous species are more likely to become pests. Most whiteflies are known primarily from woody angiosperms. vaporariorum fem ales . pseudoreduction results in only 11 chromo somes through the maturation divisions. In no current cultures of T. Downloaded from www. the 11 chro mo som es divide. Entomol. up to a certain point. in the same way as in arrhenotokous strains (153). which are haploid. The American race was shown to have a haploid chromosome number of 11 and a diploid number of 22 (153). e. Fertilized eggs develop with the diploid number into females.org by Universidad Nacional de Colombia on 02/05/12.
The ratio of male to female whitefly adults constantly changes throughout the course of the year. so the assumption of oligophagy should also be viewed with caution.annualreviews. but these records are often for sp ecie s for which very few collections have been made (131).36:431-457.
English race with thelytokous parthenogenesis (153). tabaci. polyphagy is pronounced in only a few whiteflies that feed on herbaceous plants.
HOST PLANT INTERACTIONS
Few quantitative studies have been conducted on the degree of polyphagy among members of Aleyrodidae (e. 68). vaporariorum and B. T. vaporariorum do virgin females give rise to females (6). A lac k of information on host plants also applies to oligop hagous whiteflies. The only
B YRNE & BELLOWS
Annu.g. however. 1991. All eggs show two maturation divisions and undergo reduction. Unfertilized eggs give rise to males. The English race did not exhibit normal reduction and fertilization but rather a form of apomixis.g. many of our ass um pt io ns concerning host plant associations are derived from published host lists. Adult field populations reported in two cases (83. In t helytokous T. and oligophagy may be the most common host plant association (131). 127) had sex ratios of 2 females:l male. This information provides a po tentially biased view of host plant associations because more information usually is available for pest species. however. Rev. Within the limitations imposed by such records. For personal use only. This may nevertheless follow a 1: 1 primary sex ratio because females live longer in the population as adults (130. At this stage. Many whiteflies are known from a s in gle host and so would be identified as monophagous. oogenesis occurs. 164). Such censor ing of data may be responsible for the fact that the majority of the records are for pest species that appear to be polyphagous. The diploid chromosome number was the same with a normal meiosis through prophase. After the second division. rather. In sper matogenesis the haploid chromosome number is retained without further reduction.
Rev. Thus. The mouth parts are typical for Homoptera. It is known only from Citrus spp. They suggest that three of these have a chemosensory or mechano-chemosensory function. ( 1 3 1 ). Coombe (48) discovered that T. Entomol. 130). Woets and van Lenteren ( 1 65. 1991.. The only report of a cue other than color is the finding that A. entry into the leaf by the stylet bundle is a complicated process similar to that used by aphids. 99. 1 28).org by Universidad Nacional de Colombia on 02/05/12. 57b. as in B. Different populations of the same species may show different host ranges. A. which functions in tasting the phloem sap as it enters the mouth (86 . vaporariorum while migrating responds initially to light with a wavelength of approximately 400 nm (that of the blue sky). In such situations. 59). tabaci (164). survival and reproductive rates vary widely among several host plants (68. 49. Downloaded from www. 89). For personal use only. and odor do not play a role in initial host finding for T. in A.36:431-457. although the insects did respond to color. proletella responds to the odor of crushed cabbage leaves (27). one of each pair on either side of the labial groove. vaporariorum. found that each had seven pairs of sensilla . The same is true for B. reportedly attacks and severely damages cashew in Brazil (57a. the action of natural enemies or other mortality factors in the species' native range may sufficiently reduce the fitness of populations on marginal hosts such that the whiteflies rarely use these hosts. examining the apex of the labium of adults for six species of whiteflies. Many populations that achieve high population densities may utilize plants as hosts that would not be utilized under lower population densities. In the dorsal (anterior) wall of the pharynx is the cribriform organ. structure. a species widely reported as a pest of coconut in the Neotropics.WHITEFLY BIOLOGY
Annu. and. While color brings whiteflies into contact with the plant. 65. however. Polyphagous species have a range of fitness on different hosts. in T. tabaci (22. ( 1 04). Walker ( 170)
. most species respond to color as a cue to select landing sites for feeding and oviposition (48. Walker & Gordh ( 1 72).
HOST PLANT SELECTION
Once whiteflies enter an area containing suitable plant hosts. woglumi. vaporariorum from Dioon sp. vaporariorum. woglumi reportedly is widely polyphagous in the Neotropics where it was introduced and where it had attained extraordinary population densities (60). it does not correlate well with offspring survival ( 1 64). A population of Aleurodicus cocois. but eventually responds to wavelengths of approx imately 550 nm (the green/yellow of plants). 1 78) demonstrated that leaf shape.annualreviews. reproduction rates were higher on tomato than on Brassica spp. but this particular population is not found on coconut grown in the area (57b.
record of a whitefly from a gymnosperm is T. After a host is selected. 83). in the Orient (44). 69).
although pierced cells can sometimes be observed
(169). Rev. Entomol. some decisions concerning host-plant
qual ity are made before the leaf is fully penetrated. It is probably appropriate to characterize them as poor fliers.
Short-range migration is apparently all that whiteflies need once they are
. While even the migrators may be poor fliers. myricae for the most part follow an intercellular path.444
BYRNE & BELLOWS
described how the labium is rubbed or tapped on the plant surface prior to insertion of the stylets.6 to 224. Most whitefly mouth parts enter the plant by piercing the epidermal cells (145. soft-bodied animals with their high surface-to-volume ratio. After penetration. Movement of more than a few hundred meters is likely assisted by humans.
When comparing data on whiteflies and aphids to data for other insects. myricae i s concentrated in the early morning and evening h ours (124). For personal use only.
123). Nevertheless. Fli ght of B. 79). Flight of P. a group that includes whiteflies and aphids. 1991.00532 gfcm2) and relatively high wingbeat frequencies (165. Apparently.annualreviews. Downloaded from www. short-range migration takes place regularly. whiteflies employ a "clap and fling" strategy (179). A. Two morphs have been found to exist within popUlations of B. Byrne et al (32) found that whiteflies have relatively low wing loading values (. 160). 38). woglumi reportedly disperses over distances of up to 150 m (65. 85) have high wing-beat frequencies to
compensate for the smallness of their wings in relation to their body mass.03 g. Apparently. e. tabaci occurs near ground level (below 10 cm) (39. indicating they do com pensate for hi gh wing loading by increasing wingbeat frequency. We do know that B. Although it may
occur. much of the short-range move ment by B. single individuals have reportedly traveled distances of up to 7 km (46).
Annu. 169). This was not true for insects we ighin g < 0. tabaci are routinely seen flying over fallow ground in extremely high numbers at least 150 m away from any vegetation (39.g. The question remains as to whether or not such whiteflies survive such a long journey. Flying strategies for these smaller insects are believed to be different.36:431-457.
The literature offers no absolute evidence of long-range whitefly migration
similar to that described for other homopterans (24. the stylets of P. 79).2 Hz) when compared to aphids.00174 to . Byrne et al (32) found that more massive insects have significantly and positively correlated wing loading and wingbeat frequencies. we assume that such journeys are rigorous for these small. This finding was unexpected because many other animals (5. tabaci occurs during the morning and midday hours and has one peak (19. tabaci-a mi gr atory and a trivial-flying morph (36).org by Universidad Nacional de Colombia on 02/05/12.
These variations are attributable in part to the use of different populations of a particular species.
Voltinism and Overwintering
Whiteflies generally appear to be multivoltine. 104. Downloaded from www. B. Adult whiteflies were routinely observed in the upper part of a cotton crop canopy. B. 80. it may be 2-80% in the range of 31-90% relative humidity (80). Finally. preimaginal survival of B. They land on particular plants mostly by chance. and such species may develop and breed continually so long as temperature conditions permit [e. Additional important differences in develop ment. Wherever these whiteflies are a serious problem.36:431-457. 1991.
. particularly regarding developmental threshold tempera tures. they tumble along on the ground boundary level. T. More es pecially. and pre imaginal developmental and survival rates. P. For personal use only. 69. which appear to occur near lOoC for these two species. 164). they have little control over what happens. and fecundity are caused by rearing on different host plants (51. Gerling & Horowitz (79) and Byrne & Houck (36) surmised that subsets of whitefly populations leave their original habitat in response to deteriorating conditions in search of better feeding or oviposition sites. different populations can have markedly different vital rates (80. survival.org by Universidad Nacional de Colombia on 02/05/12. 164). tabaci (51.
established in an area that has crop and weed hosts available all year. these wild and cultivated hosts grow in close proximity to one another. with two to six yearly genera tions. tabaci have received the most attention in the literature regarding laboratory life tables (for reviews. 77)]. The direction of their flight is primarily dictated by the wind as they drift about in the manner of aerial plankton. tabaci populations overwinter on a variety of cultivated and wild vegetation such as vegetables and cheeseweed and move to such spring hosts as potato and cultivated sunflower. Rev. because they are poor fliers. At this level. Having left the original habitat. however.g. fecundity. Entomol. Siphoninus phillyreae (18). electing to stay on suitable hosts and moving away from those that are not (164). the vital rates reported for various whiteflies appear to vary widely. tabaci varies inversely with relative humidity. In the Near East (77) and India (100).annualreviews.WHITEFLY BIOLOGY
Annu. while the majority of whiteflies migrating between habitats moved quickly to the ground after leaving the field (39).
DEMOGRAPHY AND POPULATION DYNAMICS
The number of species for which life tables have been constructed is relatively limited (Table 2). Most species are recorded from tropical or subtropical regions. Some generalizations can be made from such studies. see 80. vaporariorum and B. Many researchers report adult longevity. 68. myricae overwinters as both adults and nymphs on avocado in Israel (158). 164).
2 < Silf 2332 16 10 17 40 17-22 Japan 3 108 Location India time (days) Adult
Egg-adu lt survival
gen.4 36.annualreviews. For personal use only.4 53. Entomol.
Developmental Number Species Aleurocanthus husaini Aleurocanthus spinosus Aleurocanthus spiniferus' Aleurocanthus woglumi Japan 4 5-6 5 3 4 54-103 36 -50 36 50 c 9 28-35 50-929/lf 330-425/1fd 7011fO 6 6 30 27. Downloaded from www.Table 2
Life history parameters for field populations
143 101 16 98 23 144 83 84 88 154 105 105 140 173 139
Cuba South Africa India Venezuela
Aleurocybotus occidus Aleurodicus cocois anacardib Aleurodicus pimentae Aleurolobus barodensis
California Brazil Jamaica India India Pakistan Pakistan
Aleurothrixus flocossus Aleurotrachelus jelinekUg
Spain Oman Hawaiif
. 1991.org by Universidad Nacional de Colombia on 02/05/12. Rev.
For personal use only.Annu. Entomol. 1991.org by Universidad Nacional de Colombia on 02/05/12.
Aleyrodes jragariaeh Aleyrodes proletelld Asterobemisia carpinP Bemisia tabaci
Norway France Italy India Israel Arizonak Florida Florida
2 20-35 30 0 17-65 2-5 71-8 2 10
10 58 2 206/lfl m 48/cm2 1O/cm2' 2 2/cm2' 127 180
Dialeurodes chittendeni Dialeurolonga elongata Neomaskellia andropogonisP Neomaskellia bergjjq Parabemisia myricae Siphoninus phillyreaet Tetraleurodes acaciaeu Tetraleurodes semilunaris
France India Britain India Pakis tan India Japan Israel California Egypt Florida
98 109 14-20 2-3
120-150 1 46
:I: >-l tTl ><: c:I 0 t"'" 0 0 ><:
25-30 45' 0-49'
147 66 6/cm2 2
t -.annualreviews.36:431-457. Downloaded from www.J
time (days) 25-32
Adult Longevity (days)
Location Arizonav USSR Sudan
°Nymphs of second and succeeding generations diapause. 'Nine gen.
ratio 2 females:
Number Species Trialeurodes abutiloneus Trialellrodes laud' Trialellrodes lauri Trialeurodes vaporariorumw
'64-96% egg hatch.
Greenhouse study. 0% on old leaves. ("pupae") overwinter among fallen leaves.
k Laboratory I
Hosted on Umbrella tree.lyr
21 21-30 13 9-50 100 5-319 69-93
I male. Absence of natural enemies.4 days. Laboratory study-values depend on temperature. overwinter in last nymphal stage. Eretmocerus sp.
h J Nymphs
as nymphs. 'Laboratory study-35-49% survival on young and middle-aged leaves. 1991. and Encarsia
'Oviposition in February. "60-100% parasitism
Annu. d Sprayed--Iow natural enemy-caused mortality. adults and nymphs overwinter on trees.
t U v
Authors suggest overwinter as adults.
Adults overwinter. preoviposition period in laboratory study. For personal use only. Downloaded from www. Entomol.annualreviews. Following introduction of beetle and fungi. study. overwinter as adults. 45% parasitism by
EretmocerJls sp. Rev.
'Emergence primarily between 0600-0900
'Winter nymphal survival on avocado leaves.org by Universidad Nacional de Colombia on 02/05/12.).
densities of A. These differences may result from the action of natural enemies. however. 7 3 .36:431-457. S. unpublished data). premature dehiscence. A. populations often appear to increase unchecked except by the limitation of suitable foliage (31). Three species of Aleyrodes overwinter as adults (3. were reduced from 5-10 nymphs per leaf to I nymph per 1000 leaves following the introduction of parasitic wasps (157). citri can vary widely for individuals from a single cohort of eggs from 48 to 333 days ( 1 27).152).annualreviews. citri) overwinter in temperate latitudes as nymphs on evergreen hosts (Table 2). 50. spinijerus and D. Whitefly introductions have often been followed by introductions of natural
. Developmental time for D. 148. defoliation. Overwintering A. and Asterobemisia carpini reportedly overwinters in the last nymphal stage on fallen leaves (102). 112.
Studies of the population dynamics of whiteflies in natural settings are almost entirely lacking. and ovarian develop ment simply takes place at a greatly reduced rate (4). Entomol. For personal use only. D. these are primarily known from temperate latitudes and evergreen hosts (Table 2).g. causing leaf chlorosis. and plant death (1. In such settings. jelinekii). This general outcome indicates that whitefly populations have the potential for rapid. but most species reported develop from egg to adult in from 25 to 50 days under field conditions (Table 2). 82). leaf withering. Nymphal densities in more natural settings are generally much lower. 147 . Developmental times for multivoltine whiteflies vary usually with the season . 1991.
Many records of population densities are probably not characteristic of white fly populations in natural settings and generally refer to whitefly species introduced into new areas lacking natural enemies or to popUlations under pesticide treatment. chittendeni and A. 106.g. an introduced whitefly population reproduces so rapidly that its populations reach enormous densities. 2 1 . Some of these species overwinter as nymphs on foliage (e. In nearly every case.org by Universidad Nacional de Colombia on 02/05/12. No true refractory stage occurs. proletella adults experience a phase resembling ovarian diapause (3). 15. 1 8 . Most reports concern species whose populations increase dramatically on being introduced into regions. woglumi in Texas. Bellows. ranging from 10 to less than lIcm2 (T. Some species are reported as univoltine.
Other multivoltine species (e.·
Annu. Downloaded from www. 107. perhaps exponential increase under favorable conditions of climate and host-plant availability. 74. 57. High nymphal densities of 2�0/cm2 or up to several hundred per leaf are reported in such circumstances (Table 2). 10 1 . for example . Rev. 118.
78. parasitism. Downloaded from www. Several studies have examined B. In this population. 45 . the natural enemy introductions have resulted in substantial reductions in whitefl y population size (e. may be important. 1 66. In populations that attain high densities . Pop ulations in cotton appear to increase nearly exponentially during the middle part of the growing season.
.g. 95 . 70. tabaci and T. primarily B. internal regulatory processes. most frequently in crawler mortality. and competition among adults for oviposition sites may also occur [e. and unidentified deaths of each nymphal stage. This may in part result from the adaptation of the population to cotton as a host ( 1 64) and in part from the suppression of natural-enemy activity by insecticide application ( 17). l 3 1 ) . Long-term studies of the population dynamics of A.g.g. This atypical sp ecies is univoltine and was apparently introduced into England. in this chapter we make statements about what is known currently about white flies. and Hemipterous predators (45 . rain. jelinekii were con ducted in England (90. 1 81). wind. Whitefly natural enemies have been considered in other reviews (e. 157).annualreviews. B. 55. fallen fourth instar nymphs.org by Universidad Nacional de Colombia on 02/05/12. tabaci (10). As a result. This represents but a small fraction of the total number of described species. 5 5 . These two species are polyphagous and therefore may not be typical of the family. 1991. rather than the action of natural enemies. and undoubtedly many more species are as yet undiscovered. relative humidity) also may play a role in certain populations [e. For personal use only. vaporariorum. Analysis of these factors demonstrated density dependence (90). floccosus ( 1 35)] . crawler mortal ity. 1 36. usually parasitic wasps in the families Aphelinidae and Platygaster idae and coccinellid predators. Neuroptera .450
B YRNE & BELLOWS .g. Climatic factors (temperature. Some information is available on the life history of approximately 25 other species. The authors identified seven factors acting on the population between egg deposition and adult emergence: egg mortality. reports of 2000 eggs/cm2 ( 1 35)]. 1 55). The studies covered 1 7 generations. Rev.
enemies. and the maximum average density reached by the populations was approximately 5 nymphs per leaf. Entomol. fungal disease of the nymphs. [Other natural enemies recorded for the Aleyrodidae include preda cious mites.
Much of what we know about aleyrodid biology comes from reports concern ing pest species.36:431-457. predation .
Annu. realizing that as more research is conducted on this amazing group of insects our perception of their life history may change. (abaci populations (17. 1 67). intraspecific competition among nym phal stages can be significant ( 1 35). The principal mortalities affecting the popUlations oc curred between adult emergence and egg deposition and during the fourth nymphal instar. A . Fungi have also been introduced in fewer cases. 1 In nearly every case.
B . C. A. 1984. Bellows. K . with references to other American species. J . 1 9 5 7 . 1965 . Pak. Bemis. 1985 . Benfatto. S . Environ . Azab. . Entom . Lo mosca bianca lanosa (Aleurothrixusfloccosus [Mask. Bioecology of the citrus whitefly and its integrated man agement. birds and bats. of California. 10: 243-49 Adams. Bardie. . 1983. . Bellows. Plant Prot. . E.) by grant BSR-8604546 from the US National Science Founda tion . 1955. Leddy. Agric. or 4 mealy-winged flies. J. 1963 . Oxford. Remarques sur l'Aleurodes chelidonii. . Soc.]) si stadiffondendo negli agrumeti sici liani. Ara kawa. J. 1021 pp. 1983.
44:4-6 19. 17:225-28 20. Avigad. E. Bull. Agric.
3. M .annualreviews. Exp . 1988. Kabashima.
1 0. Z. Entomol. II. 1991. Afr. Jr. 2025 November. . S. P. Aleurocanthus woglumi (Ashby) (Homoptera: Aleyrodidae) in South Africa. D . El Mirsawi.org by Universidad Nacional de Colombia on 02/05/12. H . G .
4. Perring. US Nat. En tomol. Abstracts o the 1981 Int.. England. 1 90 A new enemy of the 9. and H. Relationships between whiteflies and whitefly-borne diseases.) (Homoptera: Aphididae). Acyrthosiphon pisum (Harr. Paine. Entomol. Soc.
11. On the biology of Bemisia tabaci (Genn.
Annu. Exp . Aphid feeding and nutrition.
2:448-49 15 . T. Thomas. T. K. V. Entomol. M. 1981. 89:457-67 Avidov. J.
8. 1971. A. Bedford. Ktavim 7: 25-41 Avidzba. K . A. B. A. 2 2: 270-78 Ahman.) in Israel . Factors in resistance of peas to the pea aphid. Rev.
9. G. 1983. S. Croydon. Entomol. S. Back. 1981. England: Assoc. D. Gill . . Costa. 1956. A. Soc. Entomol. The aleyrodids. Entomologist 3:3 1 3. 11:117-24 Ahmad. 1990. Calif. South. J. A. Sexual behaviour of the greenhouse whitefly (Trialeurodes vaporariorum): orienta tion and courtship.
5. . Bionomics of the tobacco whitefly (Bemisia tabaci Gen nad. Bioi. Aleyrodes pro letella. Joy. For personal use only. Mus. I. Florida orange. . Agric. Linn. T. D. 6:5-23 Abraham. Entomol. 2: 279-86 Auclair. P. J. 3. Meisenbacher.). . S. 38: 23 187-90 2 2 .
tion by the pea aphid. Aleyrodes pro letella (Homoptera: Aleyrodidae). Latr. Indian 1. B .
fructoses by the action of a yeast gluco sidase. Appl. . DeVries for early review of this manuscript. J. Y . F. J. H. S. 1959. J. Bellows. Arakawa. Khan. Auclair. Plant Protection f Human Welf or are. T. Farrar. UK: Brit ish Crop Protection Council . 19 23 . 17:483-87 18. 27:471-53 21. 1988. Econ.WHITEFLY BIOLOGY
The authors are indebted to H. Draeger and J. L. Workshop on f Pathogens Transmitted by White flies.1 4 Adams. Appl. Inf. New record of Tetraleurodes similunaria Cor bett (Aleurodidae: Hemiptera) as a pest of lemon grass Cymbopogon flexuosus (Steud. Megahed. 28:117-3 2 17. Synthesis of glucosyl
2.) (Homoptera: Aphididae). Entomol. Maltais. Amino acids. Downloaded from www. 1986. Biochemistry 73:573-87 13 .
29:330-38 7. Patterns
Dynamics of preimaginal populations of Bemisia tabaci (Homoptera: Aleyrodi dae) and Eretmocerus sp. S. Bordeaux 74:15 2-53 16.
6. Abbas. 198 2 . . C. C . Proceeding o a Con f ference Held at Brighton. Feeding and excre
thus woglumi. J. 1. Jr. Physiol. Entomol.
1. Arakawa. Haque . Proc. E. R. . E. In 10th Int. C. S. M . L . Can. Phys iol. 190 . Bird. Biological control sought for ash whitefly. The control of ovar ian development during adult diapause in the cabbage whitefly.
in diel flight activity of Bemisia tabaci (Homoptera: A1eyrodidae) in cropping systems in southern California. A comparative study on flight surface and aerodynamic parameters of insects. . K. (Hymenop tera: Aphelinidae) in southern California cotton. Acyrthosiphon pisum (Harr. 1959. Ashby) in Sind and its control. T. J. P. . The critical field photoperiod inducing ovarian diapause in the cabbage whitefly. Congr. Environ . Exp. D . J. Annu. C. BioI. We also thank E. Cartier. L. This is journal article number 7266 of the University of Arizona Agri cultural Experiment Station. Entomol. J. M. 8:439-90 Auclair. Collins for technical assistance in its preparation. H. reared on different varieties of peas. This work was supported in part (T. A. . Black fly o f Citrus (Aleurocan 1 2 . Appl. A. M . ) . Rev. . 1978. Egypte 55:30 5-15 14. N. Biological control of citrus black fly. E. Ekbom.36:431-457. Wellesbourne
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. . E. B . 1984. 88:784-88 76. Calif. E. Fjelddalen. Entomol. Stainton. Development o f a new primary pest of cotton in the Sudan: Bemisia tabaci (Gennadius) in cotton fields in Israel. Steinberg. V . F . P. V. Am.
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