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Whitefly Biology

Whitefly Biology

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Annu. Rev. Entomol. Copyright

© 1991

1991. 36:431-57

by Annual Reviews Inc. All rights reserved

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.

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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.

Although such botanical associations pro­ vide some information concerning origin.36:431-457. The genera of the subfamily Aleurodicinae are almost entirely confined to the Neotropics. the following description offered by Mound & Halsey (131) is the best available and was undoubtedly accurate when published: Three of the largest whitefly genera. Bellitudo and Crenidorsum. Annu. Aleurotrachelus and Tetraleurodes. As a result. Aleurolonga. while always difficult. Pealius. Identifying whitefly distribution is troublesome. becomes particularly hard with a family that has not had the attention of entomologists for a long period of time. most of which are monophagous or oligophagous animals associated with woody perennial hosts. Many of the other genera. Nevertheless. Rev. For example. we have learned a great deal about the biology of whiteflies from work that concentrates on nonpest species. The genus Trialeurodes has most of its species in the New World as does Aleuroparadoxus. Dialeurodes chittendeni is thought to have a Himalayan origin because of its connection with rhododendrons. Addi- . Odontaleyrodes and Rhachisphora are also particularly common in the Oriental and Austro-Oriental regions. The process of making these determinations. Acaudaleyrodes. ORIGIN AND DISTRIBUTION The geographic origin of many whitefly species is largely speculation. Butani (26). have a more restricted distribution. drawing conclusions about the family based on what is known about a few species might not be appropriate.org by Universidad Nacional de Colombia on 02/05/12. are widely distributed in the Ethiopian and Oriental Regions. Thompson & Reinert (161). For personal use only. however. All are plant feeders with piercing. Dialeuropora and the largest genus. and Cock (45). Downloaded from www. such evidence should not be consid­ ered conclusive unless it is connected to additional evidence such as the presence of sibling species. whereas Orchamoplatus is apparently most common in the Pacific . Aleuroplatus. 1991. Aleurothrixus. Corbettia and Dialeurolonga are recorded only from Africa and Madagascar. Dialeurodes. Entomol. Aleurotubercularus. are more or less artificial assemblages of species with black pupal cases reported from many parts of the world.annualreviews. Bibliographies of the family include those by Trehan & Butani (163). Aleurolobus. Aleuropteridis.WHITEFLY BIOLOGY 433 corded in Aleyrodidae. which also originated there (115). Fortunately. and these determinations are becoming even more difficult as humans facili­ tate whitefly movement. Information is available concern­ ing > 25 others. sucking mouthparts. In contrast Africaleurodes. They are opisthognathus. and this is also true of a few genera of the Aleyrodinae such as Aleurocerus. PHYSICAL CHARACTERISTICS Whiteflies share many characteristics with other homopterans. Aleurocanthuys.

. except the egg. Finally. others are not.434 BYRNE & BELLOWS tionally. The dorsal placement of the anus in Aleyrodidae permits the effective handling of the honeydew that these phloem-feeding insects produce in copious amounts. Rev. and. This variance results from changes in the number of papillae (56. as columns. occur in this family although the marginal fringes in some species of Trialeurodes tend to have light yellow or orange tints (81). in Tetraleurodes spp. the lingula is cocked down into the liquid. wax can appear as a gelatinous mass. or as setae-like projections (81). Opaque white wax may be produced as a marginal fringe (e. In B.org by Universidad Nacional de Colombia on 02/05/12. The material is extruded from wax plates that consist of rows Annu. the wax forms tight curls of threads approximately 1 /-Lm in diameter (35). because it is viscous and associated with sooty mold fungi. Whitefly nymphs cannot walk away from their honeydew droplets. The number of these rays in some nymphs differs when the whiteflies are reared on different hosts (129). although it may reflect the color of the surrounding leaf surface. In psyllids the anus seems to h ave been displaced by enlarged genitalia ( 130). The transparent form usually appears as a very thin layer over the entire dorsal surface. when the lingula is released. One is clear or colorless. particularly for sessile nymphs. it extends to the eighth abdominal segment (87). can produce extracuticular waxes that cover the body. A dorsal anus is also found in psyllid adults (rarely in the nymphs) and it also functions to rid these insects of honeydew.36:431-457.annualreviews. dorsally as tufts (Aleurotrache­ Ius jelinekii and Aleuroplatus coronata). 151). the honeydew is . Waxes Whiteflies are distinctive in that all life stages. Transparent wax may also appear as a marginal fringe or as dorsal spikelike wax rays such as those found in Trialeurodes spp. For personal use only. In whitefly nymphs. but rather the depression into which the anus empties the contents of the digestive tract (honeydew). Other colors rarely.). floccosus and Aleurodicus dispersus). It is not the anus. The other is brilliant white. as plumes. vaporariorum. catapulted away. They flip their excreta away: the honeydew fills the vasiform orifice. adults of both sexes have four membranous wings. The vasiform orifice is generally located on the dorsum of the ninth abdominal segment of males. if ever. members of the family undergo incomplete metamorphosis (with certain com­ plications). Downloaded from www. tabaci and T. Wax in the adults takes on a different appearance. Honeydew can present problems. In females. 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).g. or as flocculent or woolly mats (A. 1991. Some of the waxes are similar in form to those found on coccids ( 1 22). Entomol. and comes in two colors (81).

which has an adult body length of> 2 mm and a wing expanse of > 3. For personal use only. tabaci at the lower end.. The wax of the two species consists primarily of triacylglycerols (65-75%) with a trace of wax esters. is expanded and opaque-white with dorsal and lateral waxy. and D. apolysis takes place. we believe that the word pupa has been so inculcated into whitefly literature that its replacement would only cause confusion. vapor­ ariorum. 92) was certain whiteflies had a pupal stage in the sense that this stage serves as a mold for some of the imaginal muscles. Fourth Nymphal Instar In the literature. Entomol. Females have two pairs of these wax plates. .5--4. Pupation has also never been reported to occur in other homopterous families. LIFE HISTORY Size Gill (81) reports that the physically largest genus is Aleurodicus spp.81).WHITEFLY BIOLOGY 435 of microtrichia found on the ventro-lateral abdominal surface. the fourth nymphal instar is commonly referred to as a pupa (20. The dimensions of most whitefly adults. in fact. when the pharate adult form is present. This form according to Gill (81) feeds and so clearly is not a pupa. Nechols & Tauber (134). J. Rev. and hydrocarbons (35). discussing T. however. Hinton (91. T. more closely approximate a range with B. At this point. however. Downloaded from www. The term implies that whiteflies exhibit a degree of holometabolism. The last stage. The next form. Although we do not consider Hinton's interpretation to be correct. R. vaporariorum in midrange. For functional purposes. distinct from what occurs in holometabolous families. has the red eyes and the yellow body pigment of the adult. At this stage.org by Universidad Nacional de Colombia on 02/05/12. Annu. spinelike processes. apolysis is complete and the adult cuticle is laid down. we would reserve the term pupa for the last. 1991. free fatty acids. males have four. What takes place during the whitefly's fourth nymphal instar is. 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. stated that the fourth nymphal instar has three morphologically distinct forms. he was unable to observe a distinct stadium that intervened between the last nymphal stage and the adult. alco­ hols.annualreviews. Nechols (personal communication) states. that even after careful histological examination.36:431-457. the transitional substage. Hind and forelegs distribute the particles over the wings and the rest of the body (except the eyes). 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).0 mm. The early fourth instar is flattened and transluscent. Each microtri­ chium is associated with a wax canal.

20 1 . S.annualreviews. Gameel (75) depicts 15 ovarioles in B.436 BYRNE & BELLOWS citri at the upper end (Table 1).20 ± :t ± ± . during which time the pedicel is filled with protoplasm. tabaci. Entomol.65 2.91 :t 0.07 0. usual for members of Aleyrodidae. 12 0. the protoplasm withdraws and the pedicel becomes a hollow tube.05 female Dialeurodes citri male female 1. In most species. Weber (174) observed that T. 144). The literature describes pedicels into stomata (58.05 0.04 0. but the eggs of Aleurocybotus occiduus lie on their side (144). Annu. occiduus 13 species that insert their (138. Bellows. Poinar (144) speculated that egg pedicles of A.06 1 . served as a guide for spermatozoa during fertilization. each ovariole contains one or more follicles and a germarium.99 :t ± 1. citri (T.85 :t 0. 86.05 0. very little of this material is present.06 2.03 0.06 2.org by Universidad Nacional de Colombia on 02/05/12. Rev. chittendeni (177) and Tetraleurodes acaciae (66). eggs assume an erect stance.03 0. 138.04 0. in addition to providing a means of attachment. For personal use only. This observa­ tion applies to late nymphal instars of D.13 :t 0.49 3. males are smaller (Table Oogenesis The internal reproductive system and oogenesis in whiteflies is similar to that of other homopterans. vaporariorum secretes a gluelike substance around the pedicel. 1991.01 ± ± 0. Sexual dimorphism in adult body dimension is 1). Pedicels inserted into interstitial spaces are associated with a much larger quantity of the glue. Weber postulated that Table 1 didae Body measurements (mm) of three species of Aleyro­ (n = 5) Body length Wing expanse Species Bemisia tabaci male female Trialeurodes vaporariorum male 0. When the pedicel is inserted into parenchyma cells.41 2. unpublished) as well as D. Quaintance & Baker (150) believed the pedicel. 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.06 0 . After fertilization. Eggs Whitefly eggs generally are pyriform or ovoid and possess a pedicel that is a peglike extension of the chorion. Downloaded from www.36:431-457.81 :t 0. The pedicel is either inserted into a slit made by the ovipositor in the leaf surface or into a stomatal opening.

tabaci from Arizona cotton laid no eggs at 14. For personal use only.7°C and 72 at 32. Fecundity increased fivefold when the insects were placed on a new leaf (25).5 at 33°C and to Aleurocanthus woglumi varied with the zero at 9°C. tabaci.WHITEFLY BIOLOGY 437 water passes osmotically across this colloidal mass and enters the egg through the pedicel. Ovipositional Habits Oviposition habits differ somewhat between species. Whiteflies such as B. Sudanese strains of early fall. the mean rate of oviposition was 252 eggs per female." Several other authors have also suggested that aleyrodids use the pedicel as a means of absorbing water into the egg. the mean was 204 the (DMT 22. and that it fell to 5. B. For the same species.36:431-457. During July and August when the daily maximum temperature (DMT) was 28. The eggs are laid indiscriminately.5°C. tabaci appear to be much more prolific than strains from other parts of the world (80).9°C. Dittrich et al (61) found that Sudanese whiteflies have a fecundity on cotton of 344 eggs.6 eggs per female (69). Poinar (144) used the fact that eggs of A. Rev. tabaci in Sudanese cotton fields to reveal a fecundity of 160. Azab et al (13) examined oviposition rates of B. In December and January (DMT 14.annualreviews. and associates the production of this material with a "cement gland. support the argument that the pedicel must absorb water from the plant. Burnett (25) reports that the fecundi­ ty of T.5 eggs. Gameel (75) reports a similar substance surrounding the egg pedicle of B.5. tabaci reared on cotton in Egypt in an open-air insectary. and Avidov (10) gives a fecundity value of approximately 50 eggs on eggplant.3 to 39. ranging from 8. The maximum number of eggs per female varied from 48 to 394.2°C (29). the mean rate was 61 eggs. 1991. vaporariorum at 18°C was 319. Downloaded from www. Oviposition rates also vary greatly in other species and are affected by environmental conditions and host plant.7°C). The fecundity of nymphal host plant. Husain & Trehan (99) report a fecundity 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. 3°C). Entomol. but produced 81 eggs at 26. Von Arx et al (168) calculated the fecundity of a Sudanese strain to be 127. Tetraleurodes stanjordi . We interpret Gameel's (75) results on B.4 eggs during 43 eggs on cotton in India.org by Universidad Nacional de Colombia on 02/05/12. Hinton (93) made the claim that the pedicel was involved in the transfer of water into the egg but offered no empirical evidence. occiduus dried up when removed from the leaf to Annu. During October and November. Citing Weber (174) and Wiggels­ worth (175). However. These results occurred under conditions of constant temperature and light. tabaci deposit a few eggs on the leaf upon which they emerge as adults and then move to newer growth. 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.

Walker also suggested that the possible presence of a probing deterrent might interfere with feeding. As the first-instar nymph of B. While the nymphs usually settle in a few hours. tabaci on young (5-leaf stage) vs mature (> 25-leaf stage) lettuce. Domenichini (62) suggests an additional segmentation or suture in legs of second. found 100% mortality on the mature lettuce and 58. usually on the same leaf upon which the egg was laid. although the divisions are indistinct. Downloaded from www. the process can take several days in cooler weather (10. Crawlers Annu. Crawler mortality has been attributed to several plant characteristics. tabaci begins to emerge. vaporariorum lays its eggs in a circular fashion on glabrous leaves. they insert their mouthparts into the phloem tissue and begin extracting sap.36:431-457. Some whiteflies oviposit their eggs in an arc or circle while their mouthparts remain inserted. the egg cracks at the apical end along a longitudinal line of dehiscence. personal observation). Some species possibly use the nymphal legs to assist in casting exuviae (81). .and third-instar nymphs appear to have only one segment (81). After settling. however.438 BYRNE & BELLOWS lays its eggs on both sides of the leaf (D. crawlers walk rather quickly over the leaf surface in search of an available minor vein. floccossus and Para­ leyrodes sp. 155). Following completion of development.1 % on the young lettuce.through fourth-instar nymphs. T. it bends in half until its front legs can clasp the leaf. Fourth-instar nymphs have distinct legs and antennae. woglumi lays its eggs in a spiral arrangement (86). Entomol. were found to move between plants (67). Rev. Nymphs of all nymphal stadia of the Aleurodicinae have three-segmented legs. A. it abandons that pattern on pubescent leaves (47). For personal use only. after which it walks away from the spent chorion. He attributed this mortality to the fact that the thick cuticle of mature leaves prevents penetration. com­ paring crawler survival of B.annualreviews. N. In warmer summer conditions. Crawlers (first-jnstar nymphs) of Aleyrodinae have functional walking legs (with three apparent segments) and antennae (with two apparent segments). Some crawlers. woglumi.org by Universidad Nacional de Colombia on 02/05/12. occiduus crawlers is slightly different because these eggs are laid on their side (144). Byrne. myricae on mature lemon leaves. was small (approximately 0. The portion of the population moving between host plants. in­ cluding cuticular thickness and nutritional factors. Walker (169) reported a high degree of crawler mortality for P. Legs and antennae of the second. also oviposit eggs in a circular fashion. Emergence behavior of A. although A. 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. 1991. Byrne & Draeger (34). such as those of A.3%) and the distances traveled were short « 30 mm).

Entomol. tabaci and T. Emergence patterns persisted under conditions of continuous light or darkness. 90% o f B. suggesting the presence of a circadian system. except for brief periods during molts. Gill (8 1 ) provides a recent review of the morphology of nymphs.and third-instar whitefly nymphs have an oval or elongate-oval body. No emergence was observed at temperatures below 1 7 ± 0. two thoracic and one caudal (R. Similar patterns of emergence were found for B. tabaci possesses dorsal setae and Siphoninus spp. While some members of the Trialeurodes spp. at 10°C and 1 6 .WHITEFLY BIOLOGY 439 If crawlers successfully reach the phloem of an appropriate host.7 h) ( 1 03). pupae. Whitefly nymphs have shallow breathing folds in the ventral body wall. Gill. Annu. Rev.annualreviews. proletella the teneral period was inversely related to temperature (57. NUTRITION AND EXCRETION Nutrition As far as is known. ec1osion of 50% of the total number of adults) and temperature. Middle Instars Most second. personal communication). whiteflies are phloem feeders. tabaci and T.3°e. possess dorsal papillae-B. particu- . floccosus was reported to eclose primarily between 0600 and 0900 hours in Hawaii (139).e. and adults for the family. Few emerged during hours of darkness.6°e and a photoperiod of 14: 1 0 LD. 4 h at 25°C) and was longer when the animals were reared on young leaves ( 1 6. they remain sessile until they reach the adult stage. a significant inverse correlation was found between the time of median emer­ gence (i. Downloaded from www.org by Universidad Nacional de Colombia on 02/05/12. . J. A. For personal use only. vaporariorum is approximately 4 h 10 min at 27°C (38).36:431-457. These form a passage to the spiracles and may assist in conduction of air. 1991. vaporariorum (38). Under a series of constant temperatures.4 h) than on older ones (5. The teneral period (between final ecdysis and first fight) for B. and the peak time of emergence was delayed when temperatures were fluctuated. Some nymphs may be circular or nearly heart-shaped. For A. The folds probably developed in response to the fact that nymphs have a flattened body and lie closely appressed to the leaf surface.5 ± 0. 8 h. Adult Eclosion Under a constant temperature of 29. have dorsal siphunculi­ most species have a relatively simple dorsum. A great deal of information is available about the nutritional requirements of other homopterans. This difference suggests that feeding takes place during the teneral period. tabaci adults emerged from their pupal cases between 0600 and 0930 hours (lights on occurred at 0600 hours) (94).

glutamine is a likely candidate. During the fall and spring. B. Entomol. During . ADULT CHARACTERISTICS Mating Behavior During summer months. Butler (27) stated that Aleurodes brassicae copulate even before their wings have dried and pigmentation is complete.04%) (37). tabaci feeding on the two hosts. 125. 1 26. tabaci feeding on poinsettia.440 BYRNE & BELLOWS Annu. sucrose. it was present in high concentrations in the honeydew. tabaci feeding on these host plants. 1 6 ± 0. Although trahalulose has been synthesized in the laboratory and appears to be created by certain microorganisms 02. After that time.36:431-457. approximately half were found at significantly lower levels in the honeydew produced by B. but avoid. tabaci females are attracted to. copulation takes place within 1 to 8 h following eelosion by B. This lack suggests that certain amino acids may be used to discharge nitrogenous compounds. Carbohydrates found in the phloem sap were common transport sugars and their constituents (e. Although several may be involved. a trisaccharide common in the honey­ dew of aphids ( 1 37). males are allowed to initiate courtship. 1991. and of the honeydew produced by B. tabaci from the pupal case. but until recently almost nothing was known about the needs of whiteflies. although xanthene occurred at a low level (0. The honeydew of whiteflies on both hosts also contained melezitose. Additionally. 52-54. Rev. Of the 1 4 or 1 5 amino acids found in the phloem sap. is relatively inexpensive energetically to produce (particularly in the presence of so much carbon). trahalulose (1-0-a-D-glucopyranosyl-D-fructofuranose) (37). and has a high nitrogen to carbon ratio. males within the first 10 hours following pupal eelosion. For personal use only. The pre­ dominant amino acid in honeydew was glutamine (> 50% of the total amino acid content). copulation takes place during the three days following eelosion (0). six amino acids not found in the phloem sap were found in the honeydew of whiteflies feeding on both hosts. glucose.annualreviews. Excretion No uric acid or hypoxanthine was found in the honeydew of B.g.org by Universidad Nacional de Colombia on 02/05/12. its production has never before been associated with members of the Insecta. 42. Byrne & Miller (37) analyzed the carbohydrate and amino acid contents of pumpkin and poinsettia phloem sap. The most noteworthy discovery was that of a disaccha­ ride. Downloaded from www. These pairings follow a complex mating behavior (117). 1 32). and fructose in poinsettia and these plus stachyose and raffinose in pumpkin) (37). This result indicates that these amino acids were metabolized either by the whiteflies or by the symbionts housed in their mycetocytes (96). larly aphids (7-9. 1 56).

tabaci with that of T.WHITEFLY BIOLOGY 441 Annu. whiteflies do not experience host alternation or seasonal sexual phases. Hargreaves (89) and Williams (176). The male then places his abdomen beneath the female at a 25° angle. and antennae of both sexes are held at a 24° angle to the horizontal axis of the head. vapor­ ariorum had two races. apparently reproduces by thelytoky and its popUlations consist entirely of females. tabaci. vapor­ ariorum (114. Polygyny and polyandry occur in T. they are active over only a few millimeters. tabaci (117). except in P. The male drums the flagellum of the female antennae with his antennae while moving his abdomen up and down in synchrony with antennal drumming. During phase IV . and another male doesn't interfere.and metathoracic legs. For personal use only. During this phase. A comparison of the sexual behavior of B. Unmated females. Other investigations indicated that T. bringing the aedeagus parallel to the longitudinal axis of the female body when it is inserted into the gonopore. reported that unfertilized eggs laid by virgin females gave rise exclusively to females.annualreviews.36:431-457. an American race showing arrhenotokous partheno­ genesis (which seems to be the more usual mode of reproduction) and an . Reproduction Most whiteflies reproduce by arrhenotoky. the female often rejects the male by flapping her wings or pushing the male away with her mesothoracic legs. England. antennuation ceases and the male raises the pair of wings closest to the female. If she doesn't move. Entomol. myricae. In phase II. myri­ cae. Li & Maschwitz (116) state that if pheromones are involved in the courtship behavior of B. Rev. Phase III involves pushing the female with the side of the male's body. Mated females may produce both males and females (XO and XX). one species. Copulation lasts from 125 to 265 s. When females accept males. vaporariorum courtship and that males can detect females from some distance (at least 5 cm). Downloaded from www. phase I . the terminal flap that covers the female gonophore is pulled open. Unlike aphids. The female terminates copUlation by prying the male free with her meso. This occurs in about 15% of the observed pairings. vapor­ ariorum (6) and B. vaporariorum in Merton. P. produce male offspring (XO). the male is situated parallel to the female. The abdomen of the male is bent upward at nearly a 90° angle and the aedeagus also is bent at a 90° angle. 1991. These authors (116) also state that pheromones are much more important during T. The female may flap her wings or push the male away to discourage him at this point. However.org by Universidad Nacional de Colombia on 02/05/12. The male claspers are open and the aedeagus protruded as he tries to clasp the female terminalia. the male encircles the female several times before placing a foretarsus or antenna on the edge of her wing. working with T. 116) reveals a great many similarities and some differences. he then moves parallel to the female.

vaporariorum do virgin females give rise to females (6). English race with thelytokous parthenogenesis (153). 127) had sex ratios of 2 females:l male. Many whiteflies are known from a s in gle host and so would be identified as monophagous. A lac k of information on host plants also applies to oligop hagous whiteflies. Downloaded from www. This may nevertheless follow a 1: 1 primary sex ratio because females live longer in the population as adults (130. The only . rather. the 11 chro mo som es divide. All eggs show two maturation divisions and undergo reduction. many of our ass um pt io ns concerning host plant associations are derived from published host lists. vaporariorum fem ales . Fertilized eggs develop with the diploid number into females. but these records are often for sp ecie s for which very few collections have been made (131). pseudoreduction results in only 11 chromo­ somes through the maturation divisions. Entomol. vaporariorum and B. however. The diploid chromosome number was the same with a normal meiosis through prophase. The American race was shown to have a haploid chromosome number of 11 and a diploid number of 22 (153). In sper­ matogenesis the haploid chromosome number is retained without further reduction. in the same way as in arrhenotokous strains (153).org by Universidad Nacional de Colombia on 02/05/12.36:431-457. HOST PLANT INTERACTIONS Few quantitative studies have been conducted on the degree of polyphagy among members of Aleyrodidae (e. polyphagy is pronounced in only a few whiteflies that feed on herbaceous plants. 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. which are haploid. After the second division. In t helytokous T. tabaci.annualreviews. T. however. up to a certain point. 164). 1991. For personal use only. Within the limitations imposed by such records. and 22 chromosomes are found in all segmentation nuclei and later on in the nymphal mitoses. so the assumption of oligophagy should also be viewed with caution.442 B YRNE & BELLOWS Annu. In no current cultures of T. Rev. Sex Ratios The ratio of male to female whitefly adults constantly changes throughout the course of the year. This information provides a po tentially biased view of host plant associations because more information usually is available for pest species. Adult field populations reported in two cases (83. At this stage. oogenesis occurs. 68).g. Unfertilized eggs give rise to males. polyphagous species are more likely to become pests. e.g. and oligophagy may be the most common host plant association (131). Most whiteflies are known primarily from woody angiosperms. The English race did not exhibit normal reduction and fertilization but rather a form of apomixis.

While color brings whiteflies into contact with the plant. in A. 1991. Walker ( 170) . In the dorsal (anterior) wall of the pharynx is the cribriform organ. Polyphagous species have a range of fitness on different hosts. tabaci (164). It is known only from Citrus spp. 1 78) demonstrated that leaf shape. a species widely reported as a pest of coconut in the Neotropics. They suggest that three of these have a chemosensory or mechano-chemosensory function. vaporariorum. woglumi reportedly is widely polyphagous in the Neotropics where it was introduced and where it had attained extraordinary population densities (60). A population of Aleurodicus cocois. The same is true for B. but this particular population is not found on coconut grown in the area (57b. The only report of a cue other than color is the finding that A. 69). proletella responds to the odor of crushed cabbage leaves (27). 65. vaporariorum while migrating responds initially to light with a wavelength of approximately 400 nm (that of the blue sky). structure.WHITEFLY BIOLOGY 443 Annu. 59). vaporariorum from Dioon sp. reproduction rates were higher on tomato than on Brassica spp.annualreviews. woglumi. A. examining the apex of the labium of adults for six species of whiteflies. and odor do not play a role in initial host finding for T. Rev. After a host is selected. 89). HOST PLANT SELECTION Once whiteflies enter an area containing suitable plant hosts.org by Universidad Nacional de Colombia on 02/05/12. Thus. In such situations. 83).. Woets and van Lenteren ( 1 65. 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. in T. one of each pair on either side of the labial groove. reportedly attacks and severely damages cashew in Brazil (57a. record of a whitefly from a gymnosperm is T. ( 1 04). which functions in tasting the phloem sap as it enters the mouth (86 . The mouth parts are typical for Homoptera. tabaci (22. entry into the leaf by the stylet bundle is a complicated process similar to that used by aphids. For personal use only. 99. 130). 1 28). 57b. as in B. Many populations that achieve high population densities may utilize plants as hosts that would not be utilized under lower population densities. Different populations of the same species may show different host ranges. in the Orient (44). but eventually responds to wavelengths of approx­ imately 550 nm (the green/yellow of plants). vaporariorum.36:431-457. and. found that each had seven pairs of sensilla . ( 1 3 1 ). Entomol. it does not correlate well with offspring survival ( 1 64). although the insects did respond to color. Walker & Gordh ( 1 72). Downloaded from www. most species respond to color as a cue to select landing sites for feeding and oviposition (48. survival and reproductive rates vary widely among several host plants (68. Coombe (48) discovered that T. 49. however.

short-range migration takes place regularly. the stylets of P. Flying strategies for these smaller insects are believed to be different. e. Annu. Byrne et al (32) found that whiteflies have relatively low wing loading values (. soft-bodied animals with their high surface-to-volume ratio. We do know that B. although pierced cells can sometimes be observed (169). tabaci occurs near ground level (below 10 cm) (39. myricae i s concentrated in the early morning and evening h ours (124). some decisions concerning host-plant qual ity are made before the leaf is fully penetrated. While even the migrators may be poor fliers.36:431-457.2 Hz) when compared to aphids.annualreviews. tabaci are routinely seen flying over fallow ground in extremely high numbers at least 150 m away from any vegetation (39. 1991. Flight of P.00532 gfcm2) and relatively high wingbeat frequencies (165. After penetration. much of the short-range move­ ment by B. It is probably appropriate to characterize them as poor fliers. 79). For personal use only.00174 to . Entomol. Apparently. 160). 169). Short-range migration is apparently all that whiteflies need once they are . Two morphs have been found to exist within popUlations of B. 123). 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.444 BYRNE & BELLOWS described how the labium is rubbed or tapped on the plant surface prior to insertion of the stylets. This was not true for insects we ighin g < 0. This finding was unexpected because many other animals (5. Byrne et al (32) found that more massive insects have significantly and positively correlated wing loading and wingbeat frequencies. Rev. Fli ght of B. whiteflies employ a "clap and fling" strategy (179). Nevertheless. a group that includes whiteflies and aphids.g. woglumi reportedly disperses over distances of up to 150 m (65. The question remains as to whether or not such whiteflies survive such a long journey. 79). Apparently. tabaci-a mi gr atory and a trivial-flying morph (36). Although it may occur. Most whitefly mouth parts enter the plant by piercing the epidermal cells (145. MIGRATION The literature offers no absolute evidence of long-range whitefly migration similar to that described for other homopterans (24.03 g. we assume that such journeys are rigorous for these small. 85) have high wing-beat frequencies to compensate for the smallness of their wings in relation to their body mass.org by Universidad Nacional de Colombia on 02/05/12. Downloaded from www. 38). indicating they do com­ pensate for hi gh wing loading by increasing wingbeat frequency. A. single individuals have reportedly traveled distances of up to 7 km (46). myricae for the most part follow an intercellular path. tabaci occurs during the morning and midday hours and has one peak (19.6 to 224.

B. and such species may develop and breed continually so long as temperature conditions permit [e. which appear to occur near lOoC for these two species. Adult whiteflies were routinely observed in the upper part of a cotton crop canopy. because they are poor fliers. however. preimaginal survival of B. tabaci varies inversely with relative humidity. 68. the vital rates reported for various whiteflies appear to vary widely. 104. These variations are attributable in part to the use of different populations of a particular species. More es­ pecially. They land on particular plants mostly by chance.annualreviews. Rev. fecundity. see 80. In the Near East (77) and India (100). . Downloaded from www. survival. Some generalizations can be made from such studies. DEMOGRAPHY AND POPULATION DYNAMICS The number of species for which life tables have been constructed is relatively limited (Table 2). Finally. 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. particularly regarding developmental threshold tempera­ tures. Siphoninus phillyreae (18). The direction of their flight is primarily dictated by the wind as they drift about in the manner of aerial plankton. it may be 2-80% in the range of 31-90% relative humidity (80).36:431-457. At this level. and fecundity are caused by rearing on different host plants (51. 164). Having left the original habitat. Many researchers report adult longevity. 77)]. 164). For personal use only. B. electing to stay on suitable hosts and moving away from those that are not (164).WHITEFLY BIOLOGY 445 Annu. vaporariorum and B. 1991. T.org by Universidad Nacional de Colombia on 02/05/12. Additional important differences in develop­ ment. 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. established in an area that has crop and weed hosts available all year. myricae overwinters as both adults and nymphs on avocado in Israel (158). tabaci have received the most attention in the literature regarding laboratory life tables (for reviews. 69. Most species are recorded from tropical or subtropical regions. while the majority of whiteflies migrating between habitats moved quickly to the ground after leaving the field (39). Voltinism and Overwintering Whiteflies generally appear to be multivoltine. and pre­ imaginal developmental and survival rates. P. different populations can have markedly different vital rates (80. with two to six yearly genera­ tions. Wherever these whiteflies are a serious problem. these wild and cultivated hosts grow in close proximity to one another. Entomol. 80. they have little control over what happens.g. they tumble along on the ground boundary level. 164). tabaci (51.

annualreviews. Rev. 1991.36:431-457. Entomol.2 < Silf 2332 16 10 17 40 17-22 Japan 3 108 Location India time (days) Adult Longevity Fecundity (eggs/female) Egg-adu lt survival gen.4 36. 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./yr 2 (days) (%) Density Reference 98 III 143 101 16 98 23 144 83 84 88 154 105 105 140 173 139 Guam 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 USSR 140 113 90 . Downloaded from www.org by Universidad Nacional de Colombia on 02/05/12. For personal use only.Table 2 Life history parameters for field populations of whiteflies Annu.4 53.

Annu. 1991.org by Universidad Nacional de Colombia on 02/05/12. Downloaded from www.36:431-457.annualreviews.J . Entomol. Aleyrodes jragariaeh Aleyrodes proletelld Asterobemisia carpinP Bemisia tabaci Norway France Italy India Israel Arizonak Florida Florida USSR USSR 3 73 15 2 20-35 30 0 17-65 2-5 71-8 2 10 102 99 10 58 2 206/lfl m 48/cm2 1O/cm2' 2 2/cm2' 127 180 29 Dialeurodes citri 41-333 149 3 II 11 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 3 2 159 98 177 2 98 109 14-20 2-3 120-150 1 46 :I: >-l tTl ><: c:I 0 t"'" 0 0 ><: � 6-7 25-30 45' 0-49' 110 158 fl 171 147 66 6/cm2 2 2-3 8 t -. Rev. For personal use only.

Adults overwinter. Unsprayed-22-71% parasitism. For personal use only. ("pupae") overwinter among fallen leaves. Laboratory study-values depend on temperature. g Overwinter h J Nymphs as nymphs. 1991. Following introduction of beetle and fungi. 'Laboratory study-35-49% survival on young and middle-aged leaves. Absence of natural enemies. Number Species Trialeurodes abutiloneus Trialellrodes laud' Trialellrodes lauri Trialeurodes vaporariorumw '64-96% egg hatch.). "60-100% parasitism q Cag e (Encarsia sp.36:431-457. Downloaded from www. 45% parasitism by EretmocerJls sp. study. b Sex ratio 2 females: 3. Greenhouse study. and Encarsia sp. m n °Nymphs of second and succeeding generations diapause. Year-round development. 'Emergence primarily between 0600-0900 h. t U v Authors suggest overwinter as adults. Iyr e time (days) 25-32 Adult Longevity (days) Fecundity (eggs/female) Egg-adult survival Location Arizonav USSR Sudan gen. Eretmocerus sp. 0% on old leaves. adults and nymphs overwinter on trees.4 days. and house study.lyr (0/0) Density Reference 28 220 21 21-30 13 9-50 100 5-319 69-93 113 71 25 I male. 'Winter nymphal survival on avocado leaves. 'Nine gen. Rev.Table 2 (Continued) Developmental Annu. overwinter as adults. 'Oviposition in February. k Laboratory I Hosted on Umbrella tree. d Sprayed--Iow natural enemy-caused mortality. overwinter in last nymphal stage. Entomol. preoviposition period in laboratory study.org by Universidad Nacional de Colombia on 02/05/12.annualreviews. W .

citri can vary widely for individuals from a single cohort of eggs from 48 to 333 days ( 1 27). Entomol. 107. citri) overwinter in temperate latitudes as nymphs on evergreen hosts (Table 2). densities of A. Developmental time for D. Whitefly introductions have often been followed by introductions of natural . 147 . 74.annualreviews. 2 1 . 112. spinijerus and D. causing leaf chlorosis. Most reports concern species whose populations increase dramatically on being introduced into regions.· WHITEFLY BIOLOGY 449 Annu. This general outcome indicates that whitefly populations have the potential for rapid. 15. 148.36:431-457.152). Some species are reported as univoltine. these are primarily known from temperate latitudes and evergreen hosts (Table 2). Other multivoltine species (e. High nymphal densities of 2�0/cm2 or up to several hundred per leaf are reported in such circumstances (Table 2). unpublished data). Downloaded from www. 50. 7 3 . Developmental times for multivoltine whiteflies vary usually with the season . and Asterobemisia carpini reportedly overwinters in the last nymphal stage on fallen leaves (102). for example . jelinekii). an introduced whitefly population reproduces so rapidly that its populations reach enormous densities. chittendeni and A. Overwintering A. Population Densities 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. Rev. 82). Bellows. were reduced from 5-10 nymphs per leaf to I nymph per 1000 leaves following the introduction of parasitic wasps (157). Some of these species overwinter as nymphs on foliage (e. Population Dynamics Studies of the population dynamics of whiteflies in natural settings are almost entirely lacking. Three species of Aleyrodes overwinter as adults (3.g. 1991. and ovarian develop­ ment simply takes place at a greatly reduced rate (4). 10 1 . 57. 118. however. 1 8 . For personal use only. 106.org by Universidad Nacional de Colombia on 02/05/12. populations often appear to increase unchecked except by the limitation of suitable foliage (31). but most species reported develop from egg to adult in from 25 to 50 days under field conditions (Table 2). No true refractory stage occurs.g. S. premature dehiscence. and plant death (1. A. In such settings. These differences may result from the action of natural enemies. ranging from 10 to less than lIcm2 (T. woglumi in Texas. defoliation. proletella adults experience a phase resembling ovarian diapause (3). Nymphal densities in more natural settings are generally much lower. D. leaf withering. perhaps exponential increase under favorable conditions of climate and host-plant availability. In nearly every case.

may be important. 1991. and competition among adults for oviposition sites may also occur [e.36:431-457. Climatic factors (temperature. enemies. internal regulatory processes. (abaci populations (17. The authors identified seven factors acting on the population between egg deposition and adult emergence: egg mortality.g. 5 5 . 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). wind. Downloaded from www. fungal disease of the nymphs. 45 . 95 . Long-term studies of the population dynamics of A. 1 55). rain. Several studies have examined B. These two species are polyphagous and therefore may not be typical of the family. Some information is available on the life history of approximately 25 other species. As a result. intraspecific competition among nym­ phal stages can be significant ( 1 35). Entomol.450 B YRNE & BELLOWS . predation .g.org by Universidad Nacional de Colombia on 02/05/12. realizing that as more research is conducted on this amazing group of insects our perception of their life history may change. B. the natural enemy introductions have resulted in substantial reductions in whitefl y population size (e. and Hemipterous predators (45 . 1 81). usually parasitic wasps in the families Aphelinidae and Platygaster­ idae and coccinellid predators. floccosus ( 1 35)] . 1 36. tabaci (10). Annu. 70. parasitism. reports of 2000 eggs/cm2 ( 1 35)]. The studies covered 1 7 generations. 1 67). rather than the action of natural enemies. relative humidity) also may play a role in certain populations [e. Pop­ ulations in cotton appear to increase nearly exponentially during the middle part of the growing season.annualreviews. Whitefly natural enemies have been considered in other reviews (e. jelinekii were con­ ducted in England (90. primarily B. Fungi have also been introduced in fewer cases.g. fallen fourth instar nymphs. In this population. crawler mortal­ ity. Analysis of these factors demonstrated density dependence (90). Neuroptera . and undoubtedly many more species are as yet undiscovered. CONCLUSION Much of what we know about aleyrodid biology comes from reports concern­ ing pest species. 1 66. For personal use only. Rev. The principal mortalities affecting the popUlations oc­ curred between adult emergence and egg deposition and during the fourth nymphal instar. and the maximum average density reached by the populations was approximately 5 nymphs per leaf. in this chapter we make statements about what is known currently about white­ flies. In populations that attain high densities . tabaci and T. l 3 1 ) . . 55. 1 In nearly every case. This atypical sp ecies is univoltine and was apparently introduced into England. A .g. This represents but a small fraction of the total number of described species. most frequently in crawler mortality. 157). [Other natural enemies recorded for the Aleyrodidae include preda­ cious mites. vaporariorum. 78. and unidentified deaths of each nymphal stage.

1981. The critical field photoperiod inducing ovarian diapause in the cabbage whitefly.annualreviews. L . Can. Biological control of citrus black­ fly. UK: Brit­ ish Crop Protection Council . Indian 1. . Benfatto. Thomas. For personal use only. 27:471-53 21. Linn. 190 . .. 17:483-87 18. Annu. Factors in resistance of peas to the pea aphid. Back. K. El­ Mirsawi. Proceeding o a Con­ f ference Held at Brighton. T. Auclair. Pak. 1021 pp. 1986. Inf. Econ. Appl. 1985 . H . England: Assoc. M . Perring. I. Physiol. 1963 . 1956. Arakawa. H. Z.org by Universidad Nacional de Colombia on 02/05/12. Azab. M. Entomol. 1. P. A comparative study on flight surface and aerodynamic parameters of insects. Jr. . and H. 1988. A. 1983. Biological control sought for ash whitefly. C. 2:448-49 15 . 1978. Cartier. 1965 . B . 9. Draeger and J. Exp. E. E. Exp . Khan. A. J . Y . D . J. Entomol. Patterns Dynamics of preimaginal populations of Bemisia tabaci (Homoptera: Aleyrodi­ dae) and Eretmocerus sp. Entomol. birds and bats. S. Kabashima. Ashby) in Sind and its control. (Hymenop­ tera: Aphelinidae) in southern California cotton. This is journal article number 7266 of the University of Arizona Agri­ cultural Experiment Station. Appl. 11:117-24 Ahmad. South. A. Bellows. Entomol. S. K . A. Agric. Relationships between whiteflies and whitefly-borne diseases. Remarques sur l'Aleurodes chelidonii. US Nat. Afr. DeVries for early review of this manuscript. Bioecology of the citrus whitefly and its integrated man­ agement. Feeding and excre­ thus woglumi. 44:4-6 19. Ara­ kawa. J. 1983. 6. 2025 November. B . L. C. S. J. Rev. . P. England. On the biology of Bemisia tabaci (Genn. The aleyrodids. Gill . Maltais. Synthesis of glucosyl­ 2. J. We also thank E. 17:225-28 20. . Lo mosca bianca lanosa (Aleurothrixusfloccosus [Mask. Bioi.]) si stadiffondendo negli agrumeti sici­ liani. En­ tomol. . 3. Abbas. J. R. 6:5-23 Abraham. Black fly o f Citrus (Aleurocan­ 1 2 . New record of Tetraleurodes similunaria Cor­ bett (Aleurodidae: Hemiptera) as a pest of lemon grass Cymbopogon flexuosus (Steud.WHITEFLY BIOLOGY ACKNOWLEDGMENTS 45 1 The authors are indebted to H. 8:439-90 Auclair. . 198 2 . C . Plant Prot.) (Homoptera: Aphididae). In 10th Int. J. Wellesbourne . S. Bordeaux 74:15 2-53 16. Plant Protection f Human Welf or are. B. Literature Cited Annu. Leddy. 1983. T. Costa. S. Paine. 2: 279-86 Auclair. 1 90 A new enemy of the 9. J. M. 28:117-3 2 17. Megahed. 1959. or 4 mealy-winged flies. 1990. . Entomol. 89:457-67 Avidov. Ekbom. A. 10: 243-49 Adams. 11. Workshop on f Pathogens Transmitted by White flies. 1 0. Entomol. Avigad. Jr.1 4 Adams. Calif. P. 8. Entomol. Aleyrodes pro­ letella (Homoptera: Aleyrodidae).). Sexual behaviour of the greenhouse whitefly (Trialeurodes vaporariorum): orienta­ tion and courtship. En­ tomol. S. J. E. . D . tion by the pea aphid. . 2 2: 270-78 Ahman. Arakawa. Vol. Oxford. Abstracts o the 1981 Int. Acyrthosiphon pisum (Harr. 1 9 5 7 . Bemis. II.36:431-457. The control of ovar­ ian development during adult diapause in the cabbage whitefly. Entom . 3. Entomol. Entomol. Meisenbacher. with references to other American species. 1955. A. T. Haque . BioI. Bird. Entomologist 3:3 1 3. Collins for technical assistance in its preparation. fructoses by the action of a yeast gluco­ sidase. Farrar. Latr. Amino acids. J. V. of California. M. A. S. Environ . . Bedford. Rev. K. Florida orange. 1959. 5. Downloaded from www. Bellows. Phys­ iol. This work was supported in part (T.) by grant BSR-8604546 from the US National Science Founda­ tion . N. Soc. S . T. . . 4. . 1988. M . in diel flight activity of Bemisia tabaci (Homoptera: A1eyrodidae) in cropping systems in southern California. L. H. Soc. Mus. Ktavim 7: 25-41 Avidzba. T. K . Bellows. Bull. 1984. C. 1981. Joy. Proc. J. Appl. Environ . Acyrthosiphon pisum (Harr. Aleyrodes pro­ letella. Agric. Bionomics of the tobacco whitefly (Bemisia tabaci Gen­ nad. A. Soc. . Biochemistry 73:573-87 13 . Aphid feeding and nutrition. E.) (Homoptera: Aphididae). 29:330-38 7. F. J. ) . E. 1971. . C. reared on different varieties of peas. Aleurocanthus woglumi (Ashby) (Homoptera: Aleyrodidae) in South Africa. Croydon. Congr. J. 1. Agric. M . Egypte 55:30 5-15 14. Bardie. G. 38: 23 187-90 2 2 . 1991. . G . D. D.) in Israel . Exp . 19 23 .

wing loading. L . Butler. L. 545 pp. 3 1 . Cohen. . Am. Butler. 4 1 . Bemisia tabaci (Homoptera: the banded-wing whitefly at different temperatures. 1986. 1986. Cheetham. 1990. 1. (Westwood) (Homoptera: Aleyrodidae) . Hoff­ man. 1990. 1. P. Hadley . G. Quantification of the greenhouse white­ N. .36:431-457. Bioi. D. Byrne. 1980. Coudriet . L. G . . . 44. D. D.org by Universidad Nacional de Colombia on 02/05/12.1 3 3 0 . Biochem. Byrne. G. T . On a severe attack by Siphoninus phillyreae Halliday supsb. Entomol. Russell. W. Byrne. Ann. Physial. . 1949. 43:2 1 5. D . wingbeat frequency and body mass iII homopterous insects. . T. whitefly with an 54 anotated bibliography . Pre­ liminary studies of the distribution of whiteflies (Bemisia tabaci) . N .1 8 2 4 . fluorescent dust to mark the insects. 35 27. A. Entomol. Agron. Soc. Wil­ son. Bemisia tabaci ment. 76:3 1 0. Bellows. All. . Physiol. J . 1985 . Beitr. 1990. Soc. P. J. Bioi. N . . S . 27:388-90 50. . 1986. F. Entomol. Water uptake from plant tissue by the egg pedicel of the green­ Trialeurodes vapor­ 38 48. M. Cult. 79:350- 1 978. E. 1 982. . ed. 1988. 220:21 3-20 tabaci-a literature survey on the cotton CAB Int. A. Geoponika 105:3-7 (in Greek) 5 1 . Emomol. Entomo!. D . . The effect of tempera­ ture on an insect host-parasite popula­ tion. R . Houck. 1 983. W . Berry. F. Butler. Visual behaviour vaporariorum. Rev. J. D . D. Carbohydrate and amino acid composi­ tion of phloem sap and honeydew pro­ duced by Bemisia tabaci. . N . Bull. Vari­ ation in developmental rate on different hosts and overwintering of the sweetpo- (Homoptera­ . USDA Tech. 1 990 . On the ecology of Aleurodes brassicae Walk (Hemiptera) . 35:379-94 25 . Entomal. G. C. Ecology 30: 1 1 3-34 26. B . 1989. S. Clausen. Clausen. A. 1 5 : iol. For personal use only. N . T. C. Jr. . N . Costa. Inst. Miller. Exp . D. Whiteflies in agricultural systems . S . Aphis migration and the efficiency of the trapping method. J . Econ. 83:487-93 Aleyrodidae). 1986. 1 938. N . Trialeurodes 7: 49. 1 987. 68: 1 1 93-95 34. . N . . D . Entomol. 1984. Introduced parasites and predators of arthropod pests and weeds: A world review. Can. 1. von Bretzel. USDA Handb. Draeger. . K. J. N . C . App!. 1 35 :9-23 33. 1. T. . 78. L . G. Cock. Coll man G. fly life cycle in a controlled environ­ ment. 58 pp 45. D. H . Soc. D. 1 3:267-76 36. A. composition and waxing behaviour. E. Cohen. . Byrne. P . 1 5(4):432- of the greenhouse whitefly. Burnett. B iologia de la mosca prieta de Ius citricos Aleuro­ canthus woglumi Ashby (Homoptera: Aleyrodidae) en el campo. 1967. D . Econ. Appl. 227-61 32. . Jf. Morphometric identification of wing polymorphism in Bemisia tabaci (Gen­ nadius) (Homoptera: Aleyrodidae). Environ . Be­ misia tabaci and Trialeurodes abuti­ fonea. 36:433-39 38. Ascot. J. Bemisia 43. . Prabhaker. The extraction and mechanism of a novel isomaltulose­ synthesizing enzyme from Erwinia rha­ (Homoptera: Aleurodidae): develop­ Aleyrodidae) on cotton: adult activity and cultivar oviposition preference. . UK:CAB 1 2 1 pp. . pp.452 BYRNE & BELLOWS 37 . Buchmann. K. A. Development of 60:877-78 29. Zool. Bioi.annualreviews. 1991. Environ. 1 975. 46. T . J. Parrella. Henneberry . N . Entomol. London 87:291-3 1 1 Similarity in flight activity rhythms in coexisting species of Aleyrodidae. L . Entomol. 1963. . N . A. Impact of trap design and placement when monitoring for the bandedwinged whitefly and the sweetpo­ tato whitefly . using Phytoparasitica 14: 1 52-53 47. C . Entomo!. Entomol. Costacos . N . . Draeger. Trans. P. ed. J. 3 1 :2 1 1 . 320. uvipusitiun and lungevity in rela­ tion to temperature. Downloaded from www. Butani. ticulate surface waxes of whiteflies: morphology. Meyerdirk. The citrus blackfly in Asia. E. 1. D . Henneberry. A. 1983 . 1 970. . D. K. von Sretzel . Am. Relationship between Exp. Span­ gler. Ga. Byrne. Deleted in proof 28. Trap. and the importa­ tion of its natural enemies into Tropical Ameria. P. Byrne . Ben-Joseph. 480. See Ref. E. 1 8 :429-32 35. Bibliography of Aleyrodidae II. Clayton. Entomo/. 1 948. M. E. Boscan de Martinez. . Effect of plant maturity on oviposition and nymphal mortality of Bemisia tabaci ariorum house whitefly. A. Soc. 1932. Insect Phys­ 23. M . P. S. 1988. D.1 9 3 9 . Byrne. Coombe. D . Control. N. Butler. R. T. . inaequalis Gautier on fruit trees and its control. Par­ (Homoptera: Aleyrodidae ) . C. M . S . 20:3 1 7- Annu. P . Cienc. Byrne. P . Fail­ ure of Bemisia tabaci to breed on cassa­ va plants in Brazil 243-5 1 Entomol. N . Deleted in proof 42. 300-4 40. . pontici. B yrne. Ann. Broadbent. Kisha­ ba. Berks.

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