Bock 1983; De Bruijn &Guthrie 1982; Githunguri,. Ndong'a .... a aftel' wnlis J. C. (1973 ) (Eighth Ed ition revised by H. K. Airy Shaw) A ...... Dis!. and Bemisia tabaci vere carried out on cotton near Ludhiana in ...... 17th September to 17th October. ...... camed out between mid-September and earl\' :-':o\'ember, the inseCls.
BEMISIA TABACI
A LITERATURE SURVEY
on the
cotton whitefly
with an
annotated bibliography
BEMISIA TABACI
A LITERATURE SURVEY
on the cotton whitefly
with an annotated bibliography
Editor: M.J.W. Cock
IIBC
Published on behalf of the
FOOD AND AGRICULTURE ORGANIZATION OF THE
UNITED NATIONS
in association with the
COMMONWEALTH SCIENCE COUNCIL
INTER-AMERICAN INSTITUTE FOR COOPERATION IN AGRICULTURE
by the
INTERNATIONAL INSTITUTE OF BIOLOGICAL CONTROL
1986, reprinted 1992
BEMISIA TABACI
A LITERATURE SURVEY
on the cotton whitefly
with an annotated b:ibliography
Editor: M.J .w. Cock
lIBC
Published on behalf of the
FOOD AND AGRICULTURE ORGANIZATION OF THE·
UNITED NATIONS
!
•
~ki::"
\
in association with the COMMONWEALTH SCIENCE COUNCIL
INTER-AMERICAN INSTITUTE FOR COOPERATION IN AGRICULTURE
by the
INTERNATlONAL INSTITUTE OF BIOLOGICAL CONTROL
1986, reprinted 1992
This document was prepared and published by:
International Institute of Biological Control,
Silwood Park,
Buckhurst Road,
Ascot,
Berles. SL5 7TA,
UK.
ISBN 0 85198 575 0
@)Food and Agriculture Organization of the United Nations, 1986, 1992 The copyright in this book is vested in the Food and Agriculture Organization of the United Nations. Applications for permission to issue the book in other languages are welcomed, especially from publishers wishing to produce editions in French and Spanish. Such applications should be addressed to: Director, Publications Division, Food and Agriculture Organization of the United Nations, Via delle Terme di Caracalla, Rome, Italy. The information contained in this publication is based on published, and occasionally unpublished, research by many contributing experts. While every effort has been made to ensure accuracy, it is not possible for the International Institute of Biological Control or C' A' B International to assume responsibility for the statements contained herein, nor does the mention of any method or product supplier constitute a recommendation thereof. Furthermore, the designations employed and the presentation of the material in this document do not imply the expression of any opinion whatsoever on the part of the International Institute of Biological Control or C' A' B International concerning the legal status of any country, territory, city or area or of its authorities, or concerning the delimitation of its frontiers or boundaries.
Hi
INTRODUCTION 1.
TAXONCMY AND BIOLOGY • A. Lopez-Avila
2.
DISTRIBUTION • • . . . . . • • . . . . . . . • . . . . C.A.B International Institute of Entomology
13
3.
HOST PLANTS . . . . . • A.H. Greathead
17
4.
NATURAL ENEMIES . . . • . • • • • . . • . . . . . . . . . . . • . . . A. Lopez-Avila
27
5.
POPULATION ECOLOGY . . M.J.W. Cock
.....................•.
37
6.
TRANSMISSION OF DISEASES . . . . • . . . . . . . . . . . . . . . . . . A.A. Brunt
43
7.
ECONŒ1IC DAMAGE . . . • • . . . . . . . . . . . . . . . . . . . . . . A. Lopez-Avila & M.J.W. Cock
51
8.
OVERVIEW OF CHEMICAL CONTROL G.A .. Matthews
. . . . . . . . • . . . . . . .
55
9.
arHER CONTROL METHODS M.J.W. Cock
• • . • . . . . . . . . . . . . . . . . . . .
59
10.
POSSIBILITIES FOR CLASSrCAL BIOLOGICAL CONTROL. . . . . . . . . . . . M.J.W. Cock
63
11.
ANNOTATED BIBLIOGRAPHY . . . • . . . . . . . . . . . . . . . . . . . .
73
3
This work was prepared and published by the C.A.B International Institute of Biological Control (ClBC) on a contract from the Food and Agriculture Organization of the United Nations (FAO). lt provides a near complete bibliography of the published works on the cotton whitefly, Bemisia tabaci (Gennadius), with in most cases an abstract for each title. The cut off date for the literature search is early 1986 in the CABI abstract journals, but sorne additional more recent references have been added. Based upon this bibliography, an overview has been prepared to summarize the important research results to date, and try to indicate where future research effort may be most usefully deployed. The preparation of this volume has been very much a team effort. In alphabetical order: Mrs A.J. Girling pasted up the bibliography; Mr D.J. Girling (CIEC Infonnation Officer) obtained many of the references, did much of the legwork and initial planning, organized the checking of insect names in the C.A.B International Institute of Entomology (CIE) eard index and by CIE taxonOOlists, and co-ordinated the printingj Mrs H.A. Girling input aIl the mpterial cnto word processor with commendable accuracy; Mrs A.H. Greathead was responsible for much of the detail checking in both the bibliography and overview; Dr D.J. Greathead (Director, CIBC) played a supervisory role and reviewed aIl sections of the overviewj Mr C.J. Hamilton (CIE) earried out· the initial database searchesj Mr J.M.B. Harley (CIE) organized the checking of plant names by Ms J. Hickman and Mrs S. Niedbalaj Mr K. Hudson (C.A.B International Mycological Institute) checked many of the plant pathology references j Dr N.W. Hussey reviewed a chapter j Mrs S. Niedbala (CIE) prepared the distribution mapj Mr B. Stride photocopied much materialj Dr J.K. Waage (then Imperial College, now CIBC) reviewed a chapter. My thanks go to aIl these and the authors of the overview chapters.
M.J.W. Cock August 1986
3 Qlapter 1
Taxonaay and biology
A. Lopez-Avila
Instituto Colombia
Colanbiano Agropecuario
(ICA),
A.A.
151123
Eldorado
Bogotâ,
Presently at: Imperial College, Silwood Park, Ascot, Berks SL5 7PY, U. K. 1. 1. Tazoncmy
Bemisia tabaci (Gennadius) belongs to the sub-family Aleyrodinae, family Aleyrodidae, superfamily Aleyrodoidea, which is placed either in the sub-order Homoptera of the order Hemiptera (Richards & Davies 1977; Woodward, Evans & Eastop 1970) or in the sub-order Sternorrhyncha, order Homoptera (Borror & Delong 1964). Mound & Halsey (1978) provide a catalogue of the Aleyrodidae (over 1,100 species) with much information on host plants and natural enemies. While there are keys for classification of the aleyrodids at the generic and subfamilial level based on adult morphology (Hidalgo ~ aL 1975), the majority of whitefly species cannot be identified by morphological characters of the adult. genera and species are defined on the structure of the fourth nymphal instar, the so-called "pupal case" (Mound & Halsey 1978) . Unfortunately polyphagous whi tefly species, such as Trialeurodes vaporariorum (Westwood) and~ tabaci, vary in the appearance of their pupal cases depending on the form of the host plant cuticle on which they develop (Mound 1963; Russell 1948). Mound (1963) raised offspring of a single virgin female of~. tabaci on different species of plants, and demonstrated that two' of these forros, from tobacco and cassava, differ significantly in shape as weIl as size. He suggested that these variants are induced phenotypically by morphological characters of the host plant leaves, such as cuticle irregularity and hairiness. It is essential to understand the significance of these morphological forros, particularly where whitefly species are vectors of diseases between crops or between crop and reservoir. B. tabaci was first described as Aleurodes tabaci from tobacco in Greece (Gennadius 1899), but host-correlated morphological variation and host-plant diversity has led to a large number of synonyms. Takahashi (1936) and Russell (1958) synonymi zed several described species of Bemisia with B. tabaci, and Mound & Halsey (1978) list the synonyms given in Table 1.1. - Hidalgo et al. (1975) listed Bemisia hancocki Corbett as a synonym of ~. tabaci but they can be separated using the key given by Mound (1965) in his study of aleyrodids from West Africa which recognizes only these two species of Bemisia from area. Lebn (pers. comm. cited in Hidalgo et al. 1975) suggests that B. tabaci may be indigenous to Africa, but Mound (1965; 1983) suggestS-that it originated in the Oriental Region where related species occur, and that it was introduced and dispersed from India into America and Africa by man.
The
1.2. Biology and morphology
Many studies on the life cycle and morphology of ~ tabaci have been carried out under different climatic conditions, on different host plants (particularly cotton), and under different names: Avidov (1956); Azab, Megahed & EI-Mirsawi (1970a; 197üb; 1972); Corbett (1935); EI-Helaly, EI-Shazli & EI-Gayer (1971a; 1971b); Husain & Trehan (1933); Misra & Lamba (1929); Priesner & Hosny (1934). This chapter reviews these and also reflects the results of recent studies by the author.
Bemisia tabaci
4 . Table 1.1
Synonyms of Bemisia tabaci (Gennadius 1899) Species
Type locality
achyranthes Singh 1931 bahiana Bondar 1928 costa-limai Bandar 1928 emiliae Corbett 1926 gOldingi Corbett 1935 gossyPiperda Misra & Lamba 1929 gossypiperda var. mosaicivectura Ghesquière 1934 hibisci Takahashi 1933 inconspicua Quaintance 1900 10ngispina Priesner & Hosny 1934 lonicerae Takahashi 1957 manihotis Frappa 1938 minima Danzig 1964 miniscula Danzig 1964 nigeriensis Corbett 1935 rhodesiaensis Corbett 1935 signata Bandar 1928 vayssierei Frappa 1939
India Brazil Brazil Sri Lanka Nigeria India, Pakistan zaire Taiwan Florida EgyPt Japan Madagascar USSR (Georgia) USSR (Georgia) Nigeria Nigeria Brazil Madagascar
The immature stages of whiteflies are usually called larvae and most workers on J!.: tabaci employ this term, but because of the incomplete metamorphosis which this insect undergoes, it seems to be more appropriate to call them nymphs, which is the term used here. Azab, Megahed & EI-Mirsawi (1970a; 1972), EI-Helaly, EI-Shazli & EI-Gayer (1971a; 1971b) and Sharaf & Batta (1985) recognized four immature stages of ~ tabaci between the egg and the adul t emergence. They call the first three larval instars and the fourth the pupal stage. Since sorne morphological changes take place in the late fourth instar, and also because of the differences in behaviour of parasitoids towards these stages, four nymphal instars and a pupal stage are recognized here. 1.2.1. 'nle egg
The egg (Figure 1.1) is oval, sub-elliptical in shape, tapering towards its distal end and is provided at one side of its base with a stalk which serves to attach it to the plant. The dimensions (mean ~ SE) are: length: 0.211 ~ 0.005 mm; breadth at the broadest part: 0.096 ~ 0.002 mm; length of stalk: 0.024 ~ 0.003 mm. Eggs are almost always laid on the lower surface of the leaves, sometimes in small circular or semicircular groups produced by the female rotating about her feeding stylets whilst laying eggs (a characteristic of the subfamily Aleyrodinae (Mound 1983)). The newly-Iaid egg has a shiny chorion, which is smooth, whitish-yellow and covered with a mealy powder from the wings of the female. Before hatching it becornes light brown, the distal tip turns dark brown with two small red spots which correspond to the eyes of the nymph, and two yellow spots appear on the base of the egg which are the mycetomes of the nymph. 80th the eyes and the mycetornes are easily visible through the chorion. Several workers have stated that the incubation period varies mainly with the temperature and relative humidity (RH). Avidov (1956) found that eggs are not affected by the air humidity as they are attached to the leaf; but it has also been found that when the leaf withers and dies, the eggs collapse even when they are about to hatch.
Taxonomy
Px
5
biology
a-
d_
1.1
1.2
Figure
1.1 Bemisia tabaci egg ready to hatch: a, longitudinal hatching line; b, nymph eye; c, nymph mycetome; d, egg stalle
Figure
1.2 Bemisia tabaci first nymphal instar: D.V. dorsal view; V.V. ventral view; a, cephalic marginal seta; b, cephalic-dorsal microseta; c, eye; d, thoracic marginal seta; e, 1st abdaninal-dorsal microseta; f, abdaninal-marginal seta; g, 8th abdominal-dorsal microseta; h, vasiform orifice; i, caudal seta; j, anterocephalic ventral microseta; k, cephalic-ventral microseta; l, antenna; m, fore-leg; n, spiracle; 0, mandibular and maxillary stylets; p, meso-leg; q, hind-leg; r, abdaninal-ventral microseta; s, operculum; t, lingula. Azab, Megahed & El-Mirsawi (1972), rearing~ tabaci on sweet potato under field conditions in Egypt, found that the incubation period varied from 3 to 29 days for the periods June-September and December-January with daily mean temperature (DMT) of 28.4·C and 14.3"C respectively. In their analysis they found a negative and highly significant correlation between DMT and the incubation periode They also found a positive effect of RH on the incubation periode Butler, Henneberry & Clayton (1983) found that the duration of the egg stage varied from 22.5 days at 16.7 Oc to 7.6 days at 25° C and 5.0 days at 32.5°C, but eggs failed to hatch at 36.0°c. The author' s unpublished observations show that under constant conditions of 25°C, RH 75%, light: dark (L:D) 16:8 h, the duration of the egg stage was 6.14 days on bean plants, 6.37 days on tobacco plants, 6.9 days on lantana, 7.30 days on tomato and 7.67 days on cotton. Azab, Megahed & El-Mirsawi (1972) described the hatching process as follows: at the time of hatching, a crack appears proximally on the curved side of the egg and the proximal end of the nymph emerges. The emerging nymph bends itself towards the leaf surface until it can grip it; the legs, as they emerge, are used to push away from the egg-shell. This process is assisted by bending and alternate contraction and expansion of the abdomen. The empty egg-sheH is left on the leaf surface, maintaining its upright position and shape.
Bemisia tabaci
6 1.2.2. F.lrst
tlJliillbal
instar
The first nymphal instar is also called the crawler, because of its
habit of crawling on the leaf surface frOID its eclosion until it finds a suitable place to settle down and start feeding. It is oval in shape and measures 0.261 :!:. 0.001 DID long and 0.144 :!:. 0.010 mm broad. The dorsal surface is convex and the ventral surface fiat. The colour is whitish-green and on each side of the abdominal cavity there is a yellow spot, the mycetome, which is quite apparent through the integLment. In the morphological descriptions given by Azab, Megahed & El-Mirsawi (1910a) and El-Helaly, El-Shazli & EI-Gayer (1911b), there are sixteen pairs of marginal setae of variable lengths. There are three pairs in the cephalic region, five pairs in the thoracic region, and eight pairs in the abdominal reglon. The anal pair ls the longest. Dorsally, there are three pairs of microsetae: the first is cephalic, the second is on abdominal segment 1 and the third on abdominal segment 8. Ventrally there are also three pairs of microsetae. The small eyes are on each side of the cephalic region, situated antero-ventrally and contracted in the middle, or completely divided, and are apparent as amall inconspicuous red spots. The antennae are three-segmented and end in a fine spine. Legs on the first instar are well developed and four-segmented. The coxa is small and thick and bears one spine on the inner side; the trochanter i8 inconspicuous; the femur is longer than the coxa and as long as the tibia which is slender and bears a large and curved spine in the Middle of its outer side. The tarsus is unisegmented and ends in two small claws. The mouth parts are situated at the level of the front pair of legs and comprise two pairs of stylets which correspond to the mandibles and the maxillae. The vasiform orifice (a large dorsal opening on the last abdominal segment characteristic of whiteflies) is semicircular and elongated; the lingula (a process within the vasiform orifice on which the honey-dew collects) is finger-shaped, has two setae and projects beyond the operculum (Figure 1.2).
The duration of the first and other nymphal instars varies and is correlated with the temperature as was stated by Azab, Megahed & El-Mîrsawi (1972). They found that the correlation between the DMT and the duration of this instar is negative and an increase of 1°C was associated with a decrease of 0.30 day. Under their study conditions, which were close to ambient conditions, they found that the duration of the first instar varied from 2 ta 6 days. Sharaf & Batta (1985) stated that the duration of the first instar at 25°C was 2.8 :!:. 0.2 days, and 9.0 :!:. 0.7 days at 14°C. 1.2.3. SecoM nympal instar
Th e body, as in the first instar, is oval in shape measuring 0.365 + 0.026 mm long and 0.218 :!:. 0.012 mm at the broadest part (at the level of th; thorax) . The margin is crenulated and the marginal setae present in the first instar are missing except the antero-lateral, postero-lateral and caudal pairs. Azab, Megahed & EI-Mirsawi (1970a) mentioned only two pairs of marginal setae in the second instar. There are also three pairs of dorsal microsetae, the first are cephalic the second are situated on the first abdominal segment and the third on the 8th abdominal segment on each side of, and anterior to, the vasiform orifice. EI-Helaly, EI-Shazli & EI-Gayer (1971b) stated that there is one more pair of dorsal microsetae on the mesothorax, and two pairs of ventral microsetae, the first cephalic, situated near the base of the mouth parts, and the second on abdominal segment 8, situated at the same level as the dorsal pair. There are three pairs of spiracles, two pairs on the thorax and the third on abdominal segment 8. The eyes are small, inconspicuous, and not divided as in the first instar. The antennae are atrophied, two-segmented and directed backwards. The legs are also atrophied, two-segmented, conical, without setae, and end in a disc-like sucker. The mouth parts are more developed than in the first instar. The vasiform orifice is triangular, the operculum i3 semicircular
Taxonomy
& biology
7
and the lingula is long, thick, and anned with two praninent setae. The colour of the body i5 greenish-yellow and the mycetomes which are visible through the integument are orange-yellow. Azab, Megahed & El-Mirsawi (1972) found the duration of this instar to vary from one to five days. Sharaf & Butler (1985) reported 7.0 : 0.4 days at 14·C, and 2.8! 0.1 days at 25·C. 1.2.Il. 1hird DJIIIPbal instar The body is oval, elongate, measuring 0.489 ~ 0.022 !!ID long, 0.295 ~ 0.018 mm broad, with a crenulated margin. The third instar bears the same setae as the second instar. There is a slight constriction of the margin at the cephalic region. The eyes are circular, small and inconspicuous. The antennae are a trophied and directed towards the median line of the body; they look like hooks, due to the setaceous third segment bent back on the second. The mouth parts, as in the previous instars, are situated at the level of the front pair of legs and consist of four long stylets, but in the base there is a trapezoidal structure which is more elongate in this instar than in the two preceding ones. The legs are unsegmented, atrophied and end in disc-like suckers. The vasiform orifice is triangular, and the lingula is long and projects beyond the operculum. The colour of the third instar i5 greenish-yellow· and as in the second instar the mycetanes are orange-yellow and conspicuous. Azab, Megahed & El-Mirsawi (1972) found that the duration of the third instar varied from 2 to 7 days under their study conditions. They also found a negative correlation between the temperature and the duration of this instar with a variation of 0.38 day pel" 1"C. 1.2.5. Fourth nympbal instar Two entities are readily recognizable in the development of ~ tabaci between the third moult and adult emergence. They are referred to here as the fourth nymphal instar and the pupal stage. The description given here for the fourth instar is ccxnparable ta those given by previous workers for the early, or freshly formed, pupa. The fourth nymphal instar is large, elliptical in shape with the cephalic region semicircular, thin and flat with the margin crenulated and sometimes deeply indented due to the leaf hairs. It is 0.662 ~ 0.023 mm long and 0.440 :t. 0.003 mm broad. The marginal setae are as for the 2nd and 3rd instars, the anal pair being the most conspicuous. The dorsal setae are quite conspicuous, but vary in number from one to seven. t-bund (1963) stated that when the insect develops on glabrous leaves, it has fewer setae than when it develops on hirsute leaves. When a full set of seven pairs of setae is present, its distribution is: two pairs on the cephalic region , two pairs in the thoracic region and three pairs on the abdaninal region. In this instar, there is a series of dorsal and bilateral structures including tubercules, pores, porettes and ridges which become more evident in the pupal stage. The eyes are still small in size and apparent as two ama11, inconspicuous, red spots. The legs are short curved and unsegmented. The vasifonn orifice is triangular and elongated. The lingula, with a pair of setae, projects beyond the operculum towards the caudal margin. The colour of the four th nymphal instar is greenish-yellow and the mycetomes are elongated and yellow. The author' s unpublished observations show that under constant conditions pf 25·C, RH 75%, L:D 16:8 h, the fourth nymphal instar lasted 3.8 days on lantana, 3.4 days on beans, 2.2 days on tobacco, 2.1 days on cotton and 2.0 days on tcmato.
Bemisia tabaci
8
1.2.6. Pupa
There 1s no mou! t between the fourth nymphal instar and the pupal stage, as recognized here, but sane morpho1ogical characters are quite different. The body is elliptical in shape with the cephalic region semicircular. The dorsal surface is convex, and the thoracic and abdominal segments apparent. It ia 0.700 mm long and 0.376 :t 0.022 lIIn broad at the broadeat part, the mesothorax. The marginal and dorsal setae are as for the fourth nymphal instar, but the marginal ones are inconspicuous except the anal pair which ia the most obvious of aU. When the full set of seven pairs of dorsal setae is present they are arranged as in Figure 1.3: two pairs on the cephalic region, two pairs on the thoracic region and three pairs on the abdominal region. Ventrally, there is a pair of microsetae on abdominal segment 8. The effect of the degree of pubescence of the host plant on the number and distribution of dorsal setae in the pupal stage has been studied in detail by Mound (1963), Azab, Megahed & El-Mirsawi (1970b) and Harakly (1974). Pores and porettes are present in the pupal stage but they are visible on1y with careful staining of the pupa1 cases. The inverted T-shaped fracture 1ine of ecdysis is visible. Thoracic tracheal fo1ds are present in the pupa and extend between the first pair of spiracles and the body margin. There are four pairs of spiracles: two thoracic and two abdanina1. The vasiform orifice is triangular as in the fourth instar and the lingula extends beyond the operculum towards the caudal furrow, which appears only in the pupal stage. The surface of the lingula is covered with minute microspines and is armed distally with a pair of praninent setae. The anal (caudal) pair of setae is more than half the length of the caudal furrow (Mound 1965) (Figures 1.3, 1.4). The eyes look like two red spots constricted in the Middle and are qui te conspicuous, which is a distinctive character of the pupal stage. The body colour changes from greenish in the fourth instar to yellowish in this stage and the mycetanes becane less apparent. Pruthi & samuel (1941) and Mound (1963) stated that there is a sexual dimorphism among pupae of ~ tabaci; the sexes can be distinguished by size, wi th females produced from large pupae and males from small pupae. However Azab, Megahed & El-Mirsawi (1970a) found that male and fanale pupae were of a similar size. The duration of the pupal stage in the author's work was 4.5 days on tobacco, 4.4 days on beans, 2.4 days on tomato, 2.3 days on lantana and 1.7 days on cotton.
1.2.7. the adult Azab, Megahed & EI-Mirsawi (1970a) and Gupta (1972) describe and discuss the external morphology of the adul t male and fanale of~. tabaci. The newly emerged adult is soft, and whitish-yellow in colour, but after a few hours the colour changes to canpletely white, due to the deposition of wax on the body and wings. The body measures from vertex to tip of the genitalia 0.96 mm in the fanale and 0.82 mm in the male, and from vertex to tip of the wings 1.30 mm for females and 1.15 mm for males. The head is more or less conical, broadest at the level of the antennae and narrow towards the lDOuth parts. The antennae are long, fHiform and seven-segmented. The mouth parts are of the typical piercing and sucking type. The compound eyes are red and comprize one optical mass which is crossed on the surface by a triangular strip-like cuticular projection which is covered with white wax. The adult has two pairs of wings covered with white waxy powder; their venation is reduced.. The legs are slender, the hind pair longest. The abdomen is spindle--shaped with the vasiform orifice dorsal, near the apex. The male is generally sma11er and more slender than the female, and they also differ in the external genita1ia. The ovipositor consists of two pairs of sharply pointed lobes and, when at rest, is bent dorsally towards the
Taxonomy
9
& biology o. v.l v.v.
1.3
1.4
Figure 1.3 Bemisia tabaci pupal stage: D.Y. dorsal view,
Y.Y. ventral view: a, 1st cephalic setaj b, eyej c, 2nd cephalic setaj d, 1st thoracic spiraclej e, 1st thoracic setaj f, 2nd thoracic spiraclej g, inverted T-shaped line of ecdysis j h, 2nd thoracic setaj i, 1st abdominal setaj j, abdominal segmentationj k, 3rd-4th abdominal setaj l,8th abdominal setaj m, lingulaj n, caudal f'urroWj 0, caudal (anal) setaj p, thoracic-tracheal foldj q, fore-Iegj r,meso-Iegj s, hind-legj t, mandibular and maxillary stylets j U, lst abdominal spiraclej v, 8th abdominal-ventral microsetaj v, 8th abdominal spiracle.
Figure 1.4 Bemisia tabaci vasiform orifice: l,8th abdominal setaj m, lingulaj n, caudal
furrowj 0, caudal (anal) setaj v, 8th abdominal ventral microsetaj v, spiraclej x, vasiform orificej y, operculum. vasiform orifice. The male genitalia, consisting of an aedeagus and a pair of curved claspers, are pemanently extended. From data obtained over one year in Egypt, Azab, Megahed & El-Mirsawi (1972) found that in the male, longevity varied from 2 to 17 days, while in females i t varied from 8 to 60 days. Butler, Henneberry & Clayton (1983) found that at 26.7 and 32.2·C males lived an average of 7.6 and 11.7 days and females an average of 8.0 and 10.4 days respectively. Sharaf & Batta (1985) stated that at 25"C females live from 1 to 29 days (mean 14.8) while males live from 2 to 19 days (mean 8.9). Under standard laboratory conditions (25 ·C, RH 60%, L:D 16: 8), the author found that the longevity varied from 5 to 15 days (mean 8.66) for males and 5 to 32 days (mean 19.75) for females.
Bemisia tabaci
10
1.3. Lif'e cycle 1.3. 1. Duration Russell (1975) observed that, in different studies, the reported life cycle of B. tabaci varies considerably and is correlated with the climatic and host plant conditions. Azab, Megahed & El-Mirsawi (1971) stated that the duration of the life cycle varied from 14 to 75 days under field conditions in Egypt. The life cycle was shortest in summer (June-September) at 14-20 days (mean 16.4 OOys) and longest in winter (December-March) at 74-75 days (mean 74.6 days). In the author's unpublished studies, the time from oviposition to adult emergence was 21.5 days on bean, 22.4 days on tobacco, 23.0 days on cotton, 23.5 days on tomato and 25.35 days on lantana. Similar results were obtained by Coudriet et al. (1985). They found that the time required for B. tabaci to complete development from egg ta adult at 26.7 + 1°C was influenced by the host plant on which it was confined. Development was completed in 30% less time on lettuce, cucumber, aubergine, and squash than on broccoli or carrot (Table 1.2).
Table 1.2 Development of Bemisia tabaci from egg to adult on different hosts at 26.7 + 1°C (after Coudriet et al. 1985). Development time (days) Host
Garrot (Daucus carota)
Broccoli (Brassica oleracea)
Tomate' (Lycopersicon esculentum)
Flax (Linum usitatissimum)
Guar (eyamopsis tetragonoloba)
Pepper (Gapsicum annuum)
Guayule (Parthenium argentatum)
Cantaloupe (Cucumis Melo)
Watermelon (Ci trullu~atus)
Bean (Phaseolus vulgaris)
Cotton (Gossypium hirsutum)
Alfalfa (Medicago sativa)
Squash (Cucurbita maxima)
Aubergine (Solanum melongena)
Cucumber (Cucumis sativus)
Lettuce (Lactuca sativa)
Sweet patato (Ipomoea batatas)
Mean
SD
Range
29.8 29.7 27.3 23.9 23.9 23.4 23.2 22.3 22.3 21.8 21.7 21.4 21.3 20.9 20.6 19.4 18.6
2.2 1.6 1.0 1.8 1.2 1.8 1.6 1.1 1.8 2.6 1.9 0.8 0.7 1.0 1.4 1.9 1. 1
27-38 28-36 22-28 21-28 22-28 21-26 21-25 21-27 19-30 19-29 20-27 20-23 20-23 19-24 19-29 17-25 16-27
1.3.2. Fecundity and preoYiposition period Sharaf & Batta (1985) stated that the fecundity of B. tabaci was 76.0 and 56.4 at 25°C and 14"c respectively. They also found a preoviposition period of 3.6 and 4.9 days for the two temperatures. In the author's unpublished studies it was found that at 25· C, J!.: tabaci start ovipositing during the first 24 hours after emergence. Husain & Trehan (1933) found that in captivity a maximum of 119 eggs was laid in 18 days by a single female. The average number of eggs laid during a 24 hour period was 6 and 8 in two different years and the maximum number was 16. Azab, Megahed & El-Mirsawi (1972) stated that the number of eggs per female varied from 48 to 394 with a Mean of 161. Butler et al. (1983) report that the average number of eggs per female at 26.7 and 32.2"C was 81 and 72
Taxonomy & biology
11
respectivèly. Of the eggs laid at 26.7"C, 68% hatched and of those laid at 32.2°C, 75% hatched. Gerling, Horowitz & Baumgartner (in Israel 1985) report that there are indications that repeated insecticide applications have created an insecticide-resistant, highly fecund (300 eggs/female) strain of ~. tabaci in Sudan.
1.3.3. PartheDogenesis and sex ratio Virgin females of ~ tabaci lay eggs which give rise only to males ( Azab, Megahed & El-Mirsawi 1972 j Husain & Trehan 1933; Mound 1983 j Sharaf & Batta 1985). Sharaf & Batta (1985) found that a decrease in temperature from 25 to 14·C caused a rE'lllarkable increase in the number of adult females. The sex ratios were 1:1.8 and 1:3.1 (male:female) respectively at those temperatures. In the author' s studies, the sex ratio in samples taken fran cages where B. tabaci had been reared at 25·C on bean plants for more than ten generations was 1:2.15 (male:female)
1.3.4. Generations per year The observations of Husain ( 1931 ) , Avidov ( 1956 ) , Azab, Megahed & El-Mirsawi (1972) are in agreement that the number of generations of B. tabaci varies between 11 and 15 per year, under tropical field conditions.
12
Bemisia tabaci
13 OJapter 2
Distribution
C.A.B Intematiooal Institute of Entœology
56 Queen's Gate, London SW7 5JR, U.K. Bemisia tabaci ls widespread throughout the tropics, with the apparent exception of equatorial South America. The distribution map (Figure 2.1) was prepared to update the earlier CIE (1971) distribution map. It is based upon published records and specimens in the British Museum (BotNH) as fol1ows: EUROPE Cyprus France Greece Italy Portugal Sicily Spain Turkey [United KingdoroJ
Cyprus 1977; 1979; 1980; 1981 BMNH (1981, Provence) Mound & Halsey 1978 Iaccarino 1981; Patti & Rapisarda 1981; Viggiani & Battaglia 1983 Gentry 1965 Rapisarda & Patti 1983; Russo 1942 Mound & Halsey 1978 Dincer 1979; 1984; ~engonca 1982; Tunç et al. 1983; Turhan et al. 1983; Uygun & Ozgur 1980 Th e record of Mound & Halsey (1978) bas been omi tted as this was based on an accidentaI introduction into Kent and no permanent colony has been established there.
USSR
Azerbaijan SSR Georgian SSR
Moskovetz 1941 Danzig 1964
Angola Pagalu cameroun Cape Verde Is. Central African Republic Chad
Mound & Halsey 1978 Mound & Halsey 1978 Mound & Halsey 1978 Lobo-Lima & Klein-Koch 1981 Mound & Halsey 1978 Bink 1973; Bink-Moenen 1983; Lourens, van der Laan & Brader 1972 Matile-Ferrero 1978; Nkouka, Onore & Fabres 1981 Abdel Fattah, Sharaf & El Sebae 1983; Awadallah, Tawfik & Shalaby 1980 (Giza); El Nawawy et al. 1983; Hafez et al. 1983a; 1983b; 1983c (Upper & Lower Egypt); Kalil, Watson & Guirguis 1983 (Kafr El-Shiekh); Shaheen 1983 (Ismailia); Shanab & Awad-Allah 1982 Gentry 1965; Mound & Halsey 1978 (Eritraea) King 1980; Mound & Halsey 1978 Forsyth 1966; Mound & Halsey 1978 Delattre 1961; Mound & Halsey 1978 Bock 1983; De Bruijn & Guthrie 1982; Githunguri, Ndong'a & Ammadalo 1984; Seif 1981 Gentry 1965; Viggiani 1982 Appert 1967; Paulson & Kumashiro 1985
AFRICA
Congo Egypt
Ethiopia Gambia Ghana Ivory Coast Kenya Libya Madagascar
Bemisia tabaci
14 Malawi
Mauritius M:>rocco Mozambique Nigeria Réunion Rwanda Senegal Sierra Leone Somalia South Africa Sudan
Tanzania Togo Tunisia Uganda Zaire Zimbabwe
M:>und & Halsey 1978; Sauti 1982; 1984 Mound & Halsey 1978 Brader 1981 lI1NH (Lourenço Marques); Mound & Halsey 1978 Van Rheenen 1973; Vetten & Allen 1983 Russell & Etienne 1985 Mulindangabo & Birandano 1984 Thewys, Herve & Larroque 1979 M:>und & Halsey 1978 Gastellani, Nur & Mohamed 1982; Skaf 1968 Hill 1968 (Transvaal); 1969 Bindra 1983; Bindra & Rahman 1983; Dittrich & Ernst 1983; Dittrich, Hasan & Ernst 1985; He ijne & Peregrine 1984; Khalifa & Gameel 1983; Kisha 1984; Siddig 1984; von Arx, Baumgartner & Delucchi 1983; 1984; etc. Mound & Halsey 1978 Dengel 1981; Reckhaus 1979 EMNH (1980, Hammamet) Mound & Halsey 1978 Mound & Halsey 1978 BMNH (1935, Umtali; 1953, Salisbury; 1980, Chisumbanje)
ASIA Afghanistan Burma Cllina India
Indonesia Iran Iraq Israel
Japan Jordan Kuwait Lebanon Malaysia
Gentry 1965 EMNH (1984, Rangoon); Crowe 1985
Chou 1949 (Chekiang, Fukien, Kwangtung, Shensi, Szechwan); Mound & Halsey 1978 Anon. 1963 (Bihar, Andhra Pradesh); BMNH (1965, Madhya Pradesh); Butani & Jotwani 1983 (Delhi); Chavan 1983 (West Bengal); Chhabra & Kooner 1981 (Punjab); Gangwar & Sachan 1981 (Meghalaya); Gupta & Singh 1983 (Uttar Pradesh); LaI & Pillai 1982 (Kerala); Muniyappa 1983 (Karnataka); Murugesan & Chelliah 1981 (Tamil Nadu); Patel & Patel 1966 (Gujarat); Patil & Pokharker 1979 (Maharashta); Satyavir 1983 (Rajasthan); Sinha & Chakrabarti 1982 (Haryana) ; Yein 1981 (Assam) BMNH (1985, Sulawesi); Martin 1985 (Java); Mound & Halsey 1978 (Sumatra) Habibi 1975; Mansef & Kashkooli 1978 El-Serwiy, EI-Haidari & Saad 1984; Rizk & Ahmed 1981 Berlinger & Dahan 1983; Berlinger, Dahan & Cohen 1983; Gerling 1983; Gerling & Horowitz 1984; Horowitz, Podoler & Gerling 1984; Israel 1983; 1984; 1985; Kobatake, Osaki & Inouye 1981 (Honshu); Takahashi 1955 (Shikoku) Ohnesorge 1981; Ohnesorge, Sharaf & Allawi 1980; Sharaf 1982; 1984; Sharaf, Al-Musa & Nazer 1984a; 1984b BMNH (1976, Jabriyah)
Gentry 1965
Corbett 1935 (Malaya); Martin 1985 (Sarawak);
Parker, Booth & Bellotti 1978 (Selangor) BMNH (1976, Rumais, Nizwar)
15
Distribution Pakistan Philippines Saudi Arabia South Yemen Sri Lanka Syria Taiwan Thailand
Baloch & Soanro 1980 (Sind Province); Habib & Mohyuddin 1981 (Rawalpindi); Shafee & Rizri 1982 (Charsadda) Deang 1969; Paulson & Kl.IIIB.Shiro 1985
Abu Yaman 1971; BMNH (1968, Riyadh) Mound & Halsey 1978 (Aden) Shivanathan 1983
FAO 1966
Cllang 1969; Chou 1949
Mabbett 1979; 1980; 1983; Nachapong & Mabbett
1979; Wangboonkong 1981
AUSTRALASIA AND PACIFIC lSI.ANOO
Australia Caroline Is. Fiji Hawaii Mariana Is. Papua New Guinea Solomon Is. Tuvalu Western Samoa
EMNH (1962, Western Australia; 1981, Queensland;
1982, New South Wales); Mound & Halsey 1978
Mound & Halsey 1978
Paulson & Kumashiro 1985
Paulson &Kumashiro 1985 (Oahu) Paulson & Kumashiro 1985
Martin 1985 (Buso, Lae, Lasaranga Is. , New Britain) IMNH (1977 , Guadalcanal) EMNH (1976, FunafUti, Makin, S. Tarawa) EMNH (1977, Upolu, Manua)
AMERICA, IIJRTH
USA Arizona California District of Columbia Florida Georgia Maryland Texas Mexico
Butler & Henneberry 1984; Butler, Henneberry Natwick 1985; Butler & Wilson 1984
Butler & Henneberry 1984; Butler, Henneberry Natwick 1985; Butler & Wilson 1984; Duffus Fleck 1982; Natwick & Zalom 1984
Russell 1975
Russell 1975
Russell 1975
Russell 1975
Russell 1975
Espinosa 1972; Leon 1976 (Chiapas); Leon Sifuentes 1973
&
&
&
&
AMERICA, e»n'RAL AND CARIBBEAN
Barbados Costa Rica Cuba Dominican Republic El Salvador Guatemala Honduras Jamaica Nicaragua Panama Puerto Rico
Bennett & Alam 1985; BMNH (1920, 1921); Paulson & Kumashiro 1985
BMNH (1983, N. Heredia Province)
Blanco & Bencomo 1978; 1981
IMNH (1979, San Juan)
Granillo, Anaya & Diaz 1974; Meiners et al. 1973
Gamez 1971
Howell 1978
BMNH (1965, Kingston); Paulson & Kumashiro 1985
Falcon 1971; Laboucheix 1973
BMNH (1983, Darien Province); Gamez 1971
Bird & Sanchez 1971; Bird, Sanchez & Vakili 1973
Bemisia tabaci
16 JMKRICA, :DJ'I1I
Argentina Brazil
Colombia Venezuela
Figure
IMNH (1943, Olaco) j Paulson & Kumashiro 1985
Barradas & Chagas 1982 (sao Paulo); Barreto, Silva & Teixeira 1980 (Rio Grande do Sul); Bortoli & Giaccmini 1981; d' Araûjo e Silva et al. 1968 (Bahia, Santa Catarina, Sao Paulo); Santis 1981 Hohmann, SChoonhoven & cardona 1980; Posada et al. 1976 Guagliami 1967
2.1 The presently recorded distribution of Bemisia tabaci
17
a.apter 3 Host plants
A.B. Greatbead.
C.A.B International Institute of Biological Control, Silwood Park, Ascot, Berks SL5 7PY, U.K.
3. 1. Introductioo The starting point for compilation of this list of plants on which Bem!sia tabaci bas been recorded was the list in Mound & Halsey (1978). Those which they attributed to BM(NH) , i.e. in the collection of the British Museum (Natural History), have here been attributed ta Mound & Halsey (1978). More names were added as the l iterature search proceeded. Plant names have been checked against standard texts and Index Kewensis, and those names not traced have been marked *. No subspecies or varieties have been included in the liste Table 3.1 Ban1cing of faailies of Be-isia tabaci hast plants Family
No of species
No recorded by Mound & Halsey
No of species in family a
(1978 )
Leguminosae Canpositae Malvaceae Solanaceae Euphorbiaceae Convolvulaceae Cucurbitaceae Labiatae Verbenaceae Cruciferae Amaranthaceae Rosaceae Moraceae Oleaceae Gramineae Capparidaceae Chenopodiaceae Tiliaceae Umbelliferae 3 families 9 families 15 families 28 families
96 56 35 33
56 33 25 27 24 13 12 9 11 8 7 6
~
20 17 16 16 15 12 12 10
"other" (3) "other" (2) 5
8
8 7 6
6
6 5
4
3
3
5
2 1
11 30
"ether" (1) "other" (4) "ether" (1) other families other families other families other families
12,000 13,000 1,000 2,000 2,000 1,650 640 3,500 3,000 3,200 850 2,000 1,400 600 10,000 650 400 450 2,850
a aft el' wnlis J. C. ( 1973 ) (Eighth Ed i tion revised by H. K. Airy Shaw) A dictionary of the flowering plants and ferns. London, U.K.; Cambridge University Press. 1245 + !xvi pp.
Bemisia tabaci
18
The list shows the very wide range of host plants of B. tabaci and
contains 506 plant names representlng 14 families, an increase over the list in M:>und & Halsey (1918) of
SaDe 200 names and 12 families. In Table 3.1 the numbers of host plants in different families are given. The ranking of host plant families is similar to that in Mound &: Halsey (1978).
3.2.
List of BEwis1a tabaci bost plant records
Adhatoda vasica (Mound & Halsey 1978), Asystasia gangetica (Cohie 1969), Ruellia patula (El Khidir 1965; Gameel 1972), Ruellia prostrata (Azab, Megahed &: El-Mirsawi 1911; Gameel 1912; Pruthi &: Samuel 1942) Acer macrophyllum (Penny 1922) Aceraceae Achyranthes aspera (Gameel 1912; Pruthi &: Samuel 1942; Singh 1931), Amaranthaceae Amaranthus blitum (Cllang 1969), Amaranthus gangeticus (Azab, Megahed &: El-Mirsawi 1971; Gameel 1972; Pruthi &: Samuel 1942), Amaranthus graecizans (Gameel 1912) , Amaranthus hybridus (Chagas, Barradas & Vicente 1981), Amaranthus spinosus (Azab, Megahed &: El-Mirsawi 1971; Chang 1969; Gameel 1912; Pruthi &: Samuel 1942), Amaranthus viridis (Azab, Megahed &: El-Mirsawi 1911; Gameel 1972; Pruthi &: Samuel 1942), Amaranthus sp. (Avidov 1956; Tunç et al. 1983), Celosia cristata (Azab, Megahed & El-Mirsawi 1911; Chang 1969; Pruthi & Samuel 1942), Celosia plumosa (Mound & Halsey 1978), Digera arvensis (:alternifolia) (CIBC Pakistan Station 1983; El Khidir 1965; Gameel 1972), Gomphrena globosa (Chang 1969) Anacardiaeeae Odina aspleniifolia (Misra & Lamba 1929) Annona muricata (Cohie 1966a), Annona retieulata (Cohie 1966a), Annona Annonaeeae sguamosa (Cohie 1966a; Mound &: Halsey 1978), Fissistigma oldhamii (Mound &: Halsey 1978) Anthurium andraeanum (Herold 1967), Coloeasia antiguorum (Mound &: Araeeae Halsey 1978) Aristolochiaeeae Aristolochia braeteolata (El Khidir 1965; Gameel 1972), Aristolochia eyrnbifera (=labiosa) (Mound &: Halsey 1978), Aristoloehia punjabensis (CIBC Pakistan Station 1983) Leptadenia heterophylla (El Khidir 1965), Pergularia extensa (Mound &: Asclepiadaceae Halsey 1978) Impatiens balsarnina (Chang 1969) Balsarninaeeae Spathodea earnpanulata (=nilotiea) (El Khidir 1965; Gameel 1972) Bignoniaeeae Bornbaeopsis glabra (Cohie 1966b), Ceiba sp. (Cohie 1966a) Bornbaeaceae Boraginaceae Heliotropium europaeurn (CIBC Pakistan Station 1983), Heliotropium ovalifolium (Garneel 1972), Heliotropiurn sudanieurn (Gameel 1972) Cannabidaceae Cannabis sativa (Azab, Megahed &: El-Mirsawi 1971; Gameel 1972; Pruthi &: Samuel 1942) Capparidaeeae Boseia senegalensis (Cohie 1969), Cadaba rotundifolia (Mound &: Halsey 1978), Capparis sp. (Azab, Megahed &: El-Mirsawi 1971), Cleome ehelidonii (Azab, Megahed &: El-Mirsawi 1971; Pruthi &: Samuel 1942), Cleome gynandra (El Khidir 1965; Garneel 1972), Cleome viseosa (Misra &: Lamba 1929; Pruthi &: Samuel 1942), Cleorne sp. (Costa 1980) Caprifoliaceae Lonieera japoniea (Kobatake, Osaki &: Inouye 1981; Mound &: Halsey 1978; Takahashi 1957) Cariea papaya (Singh 1969) Caricaceae Caryophyllaceae Dianthus chinensis (Chang 1969), Saponaria vaeearia (Verma 1974e) Beta vulgaris (Duffus &: Flock 1982), Chenopodium album (Azab, Megahed Chenopodiaceae &: El-Mirsawi 1971; Chang 1969 j Pruthi &: Samuel 1942; Tunç et al. 1983), Chenopodium murale (Gameel 1972), Chenopodium sp. (Avidov 1956), Coehlospermurn planehoni (Cohie 1969), Spinaeia oleraeea (Chang
1969)
Chrysobalanaceae Chrysobalanus orbieularis (Cohie 1968)
Cistaceae Cistus salvifolius (Danzig 1964a; Tunç et al. 1983)
Commelinaceae Commelina benghalensis (Azab, Megahed &: El-Mirsawi 1971; Gameel 1972;
Pruthi &: Samuel 1942)
Cornpositae Aeanthospermum hispidurn (Mariappan &: Narayanasamy 1972) , Ageraturn eonyzoides (Garneel 1972; Prutbi &: Samuel 1942; Takahashi 194Gb), Acanthaceae
Host plants
Convolvulaceae
Cruciferae
19
Aspilia africana (Cohic 1969), Aspilia sp. (Mound & Halsey 1978), Aster tataricus (Danzig 1966) , Aster sp. (Chang 1969) , Bidens bipinnata (Chang 1969), Bidens pilosa (Chagas, Barradas & Vicente 1981 ) , Blainvillea rhomboidea (Chagas, Barradas & Vicente 1981) , Blumea neilgherrensis (Wilson & PottY 1972), calendula officinalis (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & samuel 1942), Carthamus oxyacantha (Azab, Megahed & El-Mirsawi 1971), Carthamus tinctorius (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Centaurea africana (Mimeur 1946), Centaurea sp. (Danzig 1964a) , Chromolaena odorata (van der Laan 1940) , Olrysanthemum argenteum (~engonca 1975) , Chrysanthemum coronarium (Chang 1969), Chrysanthemun indictun (Olang 1969) , O1rysanthemtun sinense (Corbett 1935b), Chrysanthemum sp. (Tunç et al. 1983), Cirsium arvense (Tunç et al. 1983) , Conyza aegyptiaca (Mound & Halsey 1978) , Coreopsis tinctoria (Mound & Halsey 1978), Cosmos bipinnatus (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Eclipta erecta (=alba, prostrata) (Azab, Megahed & El-Mirsawi. 1971; El Khidir 1965; Gameel 1972), Emilia javanica (=coccinea) (Cohie 1969), Emilia sonchifolia ( Corbett 1926; Mound & Halsey 1978), Erigeron linifolius (=Conyza crispus) (Avidov 1956; Tunç et al. 1983), Eupatorium cannabinum (Danzig 1964b), Eupatoritm chinense (Osaki & Inouye 1978), Galinsoga parviflora (01agas, Barradas & Vicente 1981), Gerbera jamesonii (01ang 1969) , Helianthus annuus (Chang 1969; Mound & Halsey 1978; Young 1944) , Helianthus debilis (El Khidir 1965) , Helianthus tuberosus (Mound & Halsey 1978), Inula viscosa (Avidov 1956), Lactuca saUva (Chang 1969), Lactuca scariola (Avidov 1956), Launaea asplenifolia (Pruthi & Samuel 1942), Parthenium argentatum (Coudriet et al. 1985), Pseudelephantopus spiralis (Mound & Halsey 1978), Serratula quinquefolia (Danzig 1964b), Sonchus arvensis (Azab, Megahed & El-Mirsawi. 1971), Sonchus cornutus (Azab, Megahed & El-Mirsawi. 1971), Sonchus oleraceus (Azab, Megahed & El-Mirsawi 1971; Gameel 1972), Sonchus spp. (Mound & Halsey 1978; Tunç et al. 1983), Tagetes erecta (CIBC Pakistan Station 1983), Vernonia anthelmintica (Azab, Megahed & EI-Mirsawi 1971; Pruthi & Samuel 1942), Vernonia cinerea (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Vernonia sp. (Mound 1963; Mound & Halsey 1978), Vicoa vestita (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Xanthium pungens (Chagas, Barradas & Vicente 1981), Xanthium strumarium (=brasilicum) (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Zinnia angustifolia (:;>engonca 1975) , Zinnia elegans (Azab, Megahed & EI-Mirsawi 1971; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942) Convolvulus arvensis (Azab, Megahed & EI-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942; Tunç et al. 1983); Convolvulus sp. (Avidov 1956), Ipomoea aguatica (=reptans) (Azab, Megahed & El-Mirsawi 1971; Gameel 1972), Ipomoea batatas (Avidov 1956; Chang 1969; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942; Quaintance 1900), Ipomoea blepharosepala (Gameel 1972), Ipomoea cairica (David & Subramaniam 1976), Ipomoea cardiosepala (Azab, Megahed & El-Mirsawi 1971; Chagas, Barradas & Vicente 1981), Ipomoea carnea (CIBC Pakistan Station 1983), Ipomoea cordofana (Azab, Megahed & EI-Mirsawi 1971; Chagas, Barradas & Vicente 1981; Gameel 1972), Ipomoea hederacea (Azab, Megahed & EI-Mirsawi 1971), Ipomoea iovolucrata (Mayné & Ghesquière 1934), Ipomoea 011 (Chang 1969), Ipomoea palmata (Mound & Halsey 1978), Ipomoea purga (Azab, Megahed & EI-Mirsawi 1971), Ipomoea purpurea (Mound & Halsey 1978), Ipomoea quioguefolia (Chagas, Barradas & Vicente 1981), Ipomoea rubrocaerulea (Avidov 1956), Ipomoea sagittata (Mimeur 1946), Jacguemontia tamnifolia (Chagas, Barradas & Vicente 1981), 'furbina corymbosa (=mollissima) (Varma 1963) Brassica alboglabra (Chang 1969), Brassica campestris (Misra & Lamba 1929; Pruthi & Samuel 1942), Brassica caulorapa (Misra & Lamba 1929),
20
Cucurbitaceae
Ericaceae lliphorbiaceae
Fagaceae Flacourtiaceae Geraniaceae
Bemisia tabaci
Brassica chinensis (Chang 1969), Brassica juncea (Azab, Megahed & El-Mirsawi 1971; Chang 1969; Gameel 1972), Brassica napus (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Brassica oleracea (Gameel 1972; Misra & Lamba 1929; Mound & Halsey 1978; Pruthi & samuel 1942), Brassica rapa (Chang 1969; Pruthi & Samuel 1942), Capsella bursa-pastoris (Tunç et al. 1983), Eruca sativa (El Khidir 1965; Gameel 1972), NasturtillIl officinale (CIBC Pakistan Station 1983), Raphanus sativus (Azab, Megahed & El-Mirsawi 1971; Chang 1969; Gameel 1972; Pruthi & Samuel 1942), Raphanus sp. (Tunç et al. 1983) , Sinapis arvensis (Tunç et al. 1983) , ZilIa myagroides (Azab, Megahed & El-Mirsawi 1971) Benincasa hispida (01ang 1969), Citrullus colocynthis (Misra & Lamba 1929), Citrullus lanatus (Avidov 1956; Azab, Megahed & El-Mirsawi 1971; Chang 1969; crnc Pakistan Station 1983), Coccinia indica (Azab, Megahed & El-Mirsawi 1971), CucllIlis melo (=maderaspatanus) (Avidov 1956; Chang 1969; Gameel 1972; Misra & Lamba 1929; Pruthi & Samuel 1942), Cucumis sativus (Avidov 1956; Chang 1969; El Khidir 1965; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942; Tunç et al. 1983), Cucurbita maxima (Coudriet et al. 1985), Cucurbita moschata (Chang 1969), Cucurbita pepo (Avidov 1956; Corbett 1935b; Tunç et al. 1983), Ecballium elaterium (Avidov 1956), Lagenaria siceraria (Crnc Pakistan Station 1983), Lagenaria vulgaris (Azab, Megahed & EI-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Luffa acutangula (Azab, Megahed & EI-Mirsawi 1971; Gameel 1972; Mound & Halsey 1978), Luffa aegyptiaca (=cylindrica) (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Gameel 1972; Pruthi & Samuel 1942), Momordica charantia (Azab, Megahed & EI-Mirsawi 1971), Trichosanthes anguina (Corbett 1935b; Pruthi & Samuel 1942), Trichosanthes dioica (Gameel 1972; Misra & Lamba 1929) Arbutus menziesii (Bemis 1904) Acalypha hispida (01ang 1969; Mound & Halsey 1978), Acalypha indica (David & Subramaniam 1976), Bridelia ferruginea (Cohic 1969), Chamaesyce hypericifolia (Chagas, Barradas & Vicente 1981), Chrozophora tinctoria (Avidov 1956), Croton lobatus (Kim et al. 1986), Croton sparsiflorus (Varna 1963), Euphorbia aegyptiaca (El Khidir 1965; Gameel 1972), Euphorbia convolvuloides (Mound & Halsey 1978), Euphorbia falcata (CrnC Pakistan Station 1983), Euphorbia geniculata ( =prunifolia) ( Costa & Bennett 1950), Euphorbia helioscopia (CrnC Pakistan Station 1983) , Euphorbia heterophylla (Azab, Megahed & EI-Mirsawi 1971; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Euphorbia hirtella (Bondar 1928; Mound & Halsey 1978), Euphorbia hypericifolia (Azab, Megahed & EI-Mirsawi 1971; Pruthi & Samuel 1942) , Euphorbia peplus (Tunç et al. 1983) , Euphorbia pilulifera (=hirta) (Azab, Megahed & EI-Mirsawi 1971; Mayné & Ghesqui~re 1929; Mound & Halsey 1978; Pruthi & Samuel 1942), Euphorbia prostrata (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Euphorbia pulcherrima (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Mound & Halsey 1978) , Jatropha curcas (Cobic 1966a) , Jatropha gossypifolia (Bird 1957), Jatropha multifida (Mayné & Ghesquière 1934), Macaranga tanarius (Chang 1969; Mound & Halsey 1978), Manihot esculenta (=utilissima) (Cohie 1969; Gameel 1972; Mound 1963; Mound & Halsey 1978), Manihot glaziovii (Gohic 1966a; Mound & Halsey 1978) , Manihot sp. (Corbett 1935a; Mound & Halsey 1978), Mercurialis annua (Tunç et al. 1983), Microcoeca mercurialis* (Nair & Menon 1978), Phyllanthus maderaspatensis (Gameel 1972), Phyllanthus niruri (El Khidir 1965; Gameel 1972; Mound & Halsey 1978), Rieinus eommunis (Chang 1969; El Khidir 1965), Trewia nudiflora (Mayné & Ghesquière 1929) Quereus agrifolia (Bemis 1904), Quercus densiflora (Bemis 1904) Rawsonia lucida (Mound & Halsey 1978) Erodium sp. (Tunç ~ al. 1983), Pelargonium odoratissimum (Azab, Megahed & El-Mirsawi 1971), Pelargonium sp. (McWhorter 1957)
Host plants
Gramineae
Grossulariaceae Guttiferae Labiatae
Lauraeeae Legtuninosae
21
Coix lacryma-jobi, (Cohie 1969), Wodon daetylon (Azab, Megahed & El-Mirsawi 1971), Hemarthria canpressus (=faseieulata) (Avidov 1956), Oplismenus burmannii (Azab, Megahed & El4tirsawi 1971; Pruthi & samuel 1942), Oryza sativa (David & Subramaniam 1976), Pennisettun amerieam.Jll (=Penieillaria spicata) (Avidov 1956), Saeehartun officinarum (David & Subramaniam 1976), Zea mays (Avidov 1956) Ribes cynosbati (Ganez-Menor 1968), Ribes gracile. (Ganez-Menor 1968), Ribes grossularia (Gomez-Menor 1968) Psorospennl.ll1 corymbiferum (Cohic 1969) Anisomeles ovata (Azab, Megahed & El-Mirsawi 1971; Pruthi & samuel 1942), Elsholtzia cristata (=patrini) (Danzig 1964a; Mound & Halsey 1978), Lamium purpureum (Danzig 1964a; Mound & Halsey 1978), Lamium sp. (Tunç et al. 1983), Leucas ciliata (Vanna 1963), Leucas stelligera (Varma 1963), Mentha lOngifolia (CIBC Pakistan Station 1983), Mentha sativa (Azab, Megahed & El-Mirsawi 1971), Mentha sylvestris (Tunç et al. 1983), Nepeta ruderalis (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Ocimum basilicum (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), OCiml.ll1 gracile (El Khidir 1965; Gameel 1972), Oeimum sanetum (Azab, Megahed & EI-Mirsawi 1971; Pruthi & Samuel 1942), Origanum sp. (Habib & Farag 1971), Salvia splendens (Chang 1969), Stachys spp. (Tunç et al. 1983) Persea gratissima (Cohie 1968), Umbellularia ealiforniea (Bemis 1904) Acacia nilotica (CIBC Pakistan Station 1983), Acacia sp • (Rao 1958), Albizia lebbeck (CIBC Pakistan Station 1983), Arachis hyPogaea (Chang 1969; Cohie 1969; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Astragalus sinieus (Olang 1969), Bauhinia purpurea (Olang 1969; Mound & Halsey 1978), Bauhinia raeemosa (Mound & Halsey 1978), Bauhinia tomentosa* (Cohie 1968), Bauhinia variegata (El Khidir 1965; Gameel 1972), Butea frondosa (Rao 1958), Caesalpinia puleherrima (Cohic 1969; Mound & Halsey 1978), Cajanus cajan (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Cajanus indicus (Avidov 1956; Azab, Megahed & El-Mirsawi 1971), Calopogonium sp. (Mound & Halsey 1978), Canavalia ensiformis (Takahashi 1941), Cassia fistula (CIBC Pakistan Station 1983), Cassia javanica (Cohie 1966b), Cassia senna (Gameel 1972), Cassia sp. (Mound & Halsey 1978), Centrosema pubescens (Olang 1969; Mound 1963), Cen trosema sp. (Muniyappa, Reddy & Shivashankar 1976; Nene 1972; Williams, Grewal & Amin 1968), Cicer arietinum (Azab, Megahed & El-Mirsawi 1971; Gameel 1972), Clitoria ternatea (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Crotalaria juncea (Avidov 1956; Azab, Megahed & El-Mirsawi 1971; Chang 1969; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Crotalaria pycnostachya (Gameel 1972), Crotalaria saltiana (El Khidir 1965; Gameel 1972), Crotalaria sp. (Mound 1963; Mound & Halsey 1978), Cyamopsis psoralioides (Azab, Megahed & El-Mirsawi 1971; Mound & Halsey 1978), Cyamopsis tetragonoloba (Rao, Ragunathan & Joshi 1983), Dalbergia sissoo (Rao 1958), Desmodium intortum (Chang 1969), Desmodium latifolium (=lasiocarpum) (Cohie 1966b), Desmoditun triquetrum (Mound & Halsey 1978), Dolichos biflorus (Mound & Halsey 1978), Doliehos lablab (Gameel 1972; Mound 1963; Mound & Halsey 1978; Pruthi & Samuel 1942), Erythrina indica (Cohie 1966a), Erythrina suberosa (CIBC Pakistan Station 1983), Glycine hispida (Pruthi & Samuel 1942), Glycine javanica (Chang 1969), Glycine koidzumii (Chang 1969), Glycine max (Azab, Megahed & El-Mirsawi 1971; Kobatake, Osaki & Inouye 1981), Glycine sp. (Takahashi 1956), Indigofera hirsuta (Muniyappa & Reddy 1979), Indigofera sp. (Mound 1963), Lathyrus articulatus (Azab, Megahed & El-Mirsawi 1971), Lens esculenta (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Lespedeza bicolor (Chang 1969), Lotus arabicus (Azab, Megahed & El-Mirsawi 1971), Lupinus hartwegii (Verma 1974a), Lupinus perennis (Chang 1969) , Macrotyloma uniflorum (Muniyappa 1983) , Medicago
Bemisla tabaci
22 dentieùlata· (Rahman
1940) ,
Medieago
hispida
(Azab,
Megahed
&
EI-Mirsaw1 1971), Med1cago sat1va (Av1dov 1956; Azab, Megahed &
Linaceae
Loganiaceae Lythraceae Malvaceae
El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942; Tunç et al. 1983), Melilotus indiea (=parviflora) (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Millettia drastiea (Cohie 1966b), Mucuna cochinchinensis (David & Subramaniam 1976), Mucuna sp. (Mound & Halsey 1978), Parkinsonia aculeata (Azab, Megahed & El-Mirsawi. 1971), Phaseolus atropurpureus (Chang 1969), Phaseolus calcaratus (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Phaseolus lathyroides (Bird, Sanchez & Vakili 1973; Goodman 1977b), Phaseolus lOngepedunculatus (Bird & Maramorosch 1975; Flores & Silbersc1'Jnidt 1966), Phaseolus lunatus (Avidov 1956), Phaseolus mediatus (Azab, Megahed & El-Mirsawi. 1971), Phaseolus vulgaris (Avidov 1956; Chang 1969; Gameel 1972; Gomez~enor 1953; Mound & Halsey 1978; Pruthi & Samuel 1942), Piliostigma thonningii (Gohic 1966 b) , Pisum arvense (Gameel 1972; Pruthi & Samuel 1942), Pisum sativum (Avidov 1956; Azab, Megahed & El-Mirsawi. 1971; Chang 1969; Gameel 1972; Pruthi & Samuel 1942), Platysepalum vanderysti (Gohie 1966b) , Prosopis stephaniana (Avidov 1956), Psoralea bituminosa (Danzig 1964a), Pterocarpus erinaceus (Gohic 1969), Pueraria sp. (Mound & Halsey 1978), Rhynchosia memnonia (Azab, Megahed & El-Mirsawi. 1911; Gameel 1972), Rhynehosia minima (Bird 1962; Chagas, Barradas & Vicente 1981), Sesbania macrocarpa (Avidov 1956), sesbania sesban (CIBC Pakistan Station 1983), Stylosanthes gracilis (01ang 1969), Tephrosia appollinea (El Khidir 1965; Gameel 1972), Teramnus uncinatus (Muniyappa, Reddy & Shivashankar 1976), Trifolium alexandrinum (Avidov 1956 ; Azab, Megahed & El ~irsawi 1971; Pruthi & Samuel 1942), Vicia dasycarpa (01ang 1969), Vicia faba (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Gameel 1972), Vigna aconitifolia (CIBC Pakistan Station 1983; Vil' 1984), Vigna catjang (Gameel 1972; Pruthi & Samuel 1942), Vigna luteola (Barradas & Chagas 1982), Vigna MungO (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942; Rathi & Nene 1975), Vigna radiata (Gameel 1972; Nene 1973a; Pruthi & Samuel 1942; Srivastava & Singh 1916), Vigna repens (01ang 1969), Vigna sesguipedalis (Avidov 1956), Vigna sinensis (Avidov 1956; Chang 1969; Mayné & Gesquière 1934), Vigna unguiculata (Srivastava & Singh 1976), Vigna sp. (Chang 1969), Voandzeia subterranea (Munniyappa, Reddy lx Shivashankar 1976) Linum usitatissimum (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Gameel 1912; Pruthi & Samuel 1942), Reinwardtia trigyna (Mound lx Halsey 1918) Gen. indet. (Gohic 1966b) Lawsonia alba (Mound & Halsey 1978) Abutilon avicennae (Chang 1969), Abutilon bidentatum (CISC Pakistan Station 1983), Abutilon figarianum (=graveolens) (Azab, Megahed lx El-Mirsawi 1911), Abutilon glaucum (=pannosum) (Azab, Megahed lx El-Mirsawi 1971; Gameel 1972), Abutilon grandiflorum (Gerling 1984), Abutilon indicum (CISC Pakistan Station 1983), Abutilon zanzibaricum (Azab, Megahed & EI-Mirsawi 1971), Abutilon sp. (Avidov 1956; Mound lx Halsey 1918), Althaea cannabina (Azab, Megahed & EI-Mirsawi 1971), Althaea rosea (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Mound lx Halsey 1918; Pruthi & Samuel 1942), Gossypium arboreum (David lx Subramaniam 1976), Gossypium barbadense (Mound 1963; Mound & Halsey 1978), Gossypium herbaceum (Avidov 1956; Corbett 1935b; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Gossypium hirsutum (Chang 1969; Mound 1963; Mound & Halsey 1978), Gossypium sp. (Corbett 1935aj Mound & Halsey 1978), Hibiscus cannabinus (Chang 1969; Gameel 1972; Mound 1963; Pruthi & Samuel 1942), Hibiscus esculentus (Chang 1969; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942; Quaintance 1900), Hibiscus mutabilis (=sinensis) (Azab, Megahed & EI-Mirsawi 1971; CISC Pakistan Station 1983), Hibiscus rosa-sinensis (Avidov
Host plants
Menispermaceae Moraceae
Moringaceae Musaceae Myrtaceae Nyctaginaceae Oleaceae
Oxalidaceae Passifloraceae Pedaliaceae
Periplocaceae Plantaginaceae Polygonaceae Portulacaceae Proteaceae Punicaceae Ranunculaceae Rhamnaceae
Rosaceae
23 1956;' Chang 1969; Gameel 1972; Hound& Halsey 1978; Pruthi & Samuel 1942; Takahashi 1933), Hibiscus sabdariffa (Azab, Megahed & El-Mirsawi 1911; Chang 1969; Gameel 1912), Hibiscus temifoliolus (Azab, Megahed & El-Mirsawi 1971; Gameel 1972), Hibiscus sp. (Avidov 1956), Halva sylvestris (Azab, Megahed & El-Mi.rsawi 1971), Halva sp. (Tunç et al. 1983), Malvastrum coromandelianun (CIBC Pakistan Station 1983), Malvaviscus arboreus (Hound & Halsey 1978), Sida alba (El Khidir 1965; Gameel 1972), Sida asperifolia* (Hound & Halsey 1978), Sida carpinifolia (Chagas, Barradas & Vicente 1981), Sida cordifolia (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Sida grewioides (CIBC Pakistan Station 1983), Sida rhOOlbifolia (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Sida veronicaefolia (=hunilis) (Gameel 1972), Urena lobata (Chang 1969; Gameel 1972; Mound & Halsey 1978), Wissadula amplissima (Schuster 1964) Stephania japonica (Takahashi 1955) Broussonetia papyrifera (=Morus australis) (CIBC Pakistan Station 1983; Mound & Halsey 1978), Ficus carica (Tunç et al. 1983), Ficus palmata (CIBC Pakistan Station 1983), Ficus religiosa (CIBC Pakistan Station 1983), Ficus sycomorus (Habib & Farag 1971), Ficus sp. (Mound & Halsey 1978), Maclura aurantiaca (Tunç et al. 1983), Morus alba (CIBC Pakistan Station 1983), Morus indica (Raychaudhuri, O1atterjee & Dhar 1961), Morus sp. (Tunç et al. 1983) Moringa pterygosperma (=oleifera) (CIBC Pakistan Station 1983; Rao 1958) Musa sapientum (CIBC Pakistan Station 1983), Musa sp. (Habib & Farag 1971) &!gania sp. (Rao 1958), Psidium guajava (Chang 1969; Gameel 1972; M::>und & Halsey 1918; Freisner & Hosny 1934) Boerhavia repens (=diffusa) (Azab, Megahed & EI-Mirsawi 1971; El Khidir 1965; Gameel 1972) Forsythia suspensa (Chang 1969) , Jasminum humile (CmC Pakistan Station 1983), Jasminum officinale (CIBC Pakistan Station 1983), Jasminllll pubigerum (CIBC Pakistan Station 1983), Jasminum sambac (Wilson 1972), Jasminum sp. (M::>und & Halsey 1918), Ligustrum vulgare (Tunç et al. 1983), Olea europaea (Mimeur 1946) Oxalis corniculata (Azab, Megahed & El-Mirsawi 1971; Chang 1969; Gameel 1972; Pruthi & Samuel 1942) Barteria bagshawi· (M::>und & Halsey 1978), Passiflora edulis (Ondieki 1975) Sesamum indicum (Azab, Megahed & El-Mirsawi 1971; Chang 1969; Gameel 1972; Fruthi & Samuel 1942), Sesamum orientale (Brunt this volume), Sesamum sp. (Mound & Halsey 1978) Periploca graeca (Gomez~enor 1968) Plantago sp. (Tunç et aL 1983) Antigonon leptopus (CISC Pakistan Station 1983), Polygonum persicaria (Tunç et al. 1983), Rumex spp. (Tunç et al. 1983) Portulaca oleracea (Avidov 1956) Helicia ochinchinensis* (Chang 1969) Punica granatum (Mimeur 1946) ligusticifolia (Bernis 1904) , Rammculus langsdorfii Cl ematis (=japonicus) (Chang 1969), Ranunculus muricatus (Tunç et al. 1983)
Rhamnus californica (Bernis 1904) , Rhamnus crocea (Bemis 1904) ,
Ziziphus mauri ti ana (CIBC Pakistan Station 1983) , Ziziphus
spina-christi (Mound & Halsey 1978)
Fragaria vesca (~engonca 1915), Heteromeles arbutifolia (Bernis 1904),
Potentilla sp. (Tunç et al. 1983), Pyrus calleryana (Azab, Megahed &
El-Mirsawi 1971), pus communis (Avidov 1956; Mimeur 1946), Pyrus
malus (Avidov 1956), J7rus mamorensis (Mimeur 1946), Rosa centrifolia
(Azab, Megahed & EI-Mirsawi 1971; Gameel 1972; Fruthi & Samuel 1942), Rosa gallica (=bourbonia) (Bhargava & Joshi 1962; Sastry 1966), Rosa
24
Bemisia tabaci indica (=chinensis) (CIBC Pakistan Station 1983), Rosa sp. (Avidov
1956; Cohie 1969; Tun9 et al. 1983), Rubus fruticosus (Tunç et al. 1983) Gardenia Jasminoides (CIBC Pakistan Station 1983), ~rinda tinctoria Rubiaceae (David & Subramaniam 1976) Citrus sp. (Avidov 1956; Frappa 1939), Huta sp. (Habib & Farag 1971) Rutaceae Salix sp. (Tunç et al. 1983) - Salicaceae Scrophulariaceae capraria biflora (Gomez~enor 1968), Scoparia dulcis (Azab, Megahed & El-Mirsawi 1971 ; Gameel 1972; Pruthi & Sarouel 1942), Veronica sp . (~und & Balsey 1978) capsicum annuum (Avidov 1956; Chang 1969; Cohic 1966a; Gameel 1972 j Solanaceae Hound & Halsey 1978; Pruthi & Samuel 1942), Capsicum frutescens (Shivanathan 1983), cestrum nocturnum (~und & Balsey 1978), Datura alba (Azab, Hegahed & El-Mirsawi 1971), Datura fastuosa (David & Subramaniam 1976) , Datura metel (El Khidir 1965; Gameel 1972) , Datura stramonium (Azab, Hegahed & El-Mirsawi 1971; Chang 1969; Gameel 1972; Pruthi & Samuel 1942; Tunç et al. 1983), Datura suaveolens (=gardneri) (Azab, Hegahed & El-Mirsawi 1971), Datura sp. (Gcxnez-Menor 1968), Hyoscyamus niger (Pruthi & Samuel 1942), Lycium chinense (01ang 1969), Lycopersicon esculentum (Avidov 1956; Chang 1969; El Khidir 1965; Gameel 1972; Mound & Halsey 1978; Pruthi & Samuel 1942), Lycopersicon pimpinellifolium (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Nicandra physalodes (Azab, Megahed & EI-Mirsawi 1971; Mound & Halsey 1978; Pruthi & Samuel 1942), Nicotiana glauca (Bandar 1928; Fruthi & Samuel 1942), Nicotiana glutinosa (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Gameel 1972; Pruthi & Samuel 1942), Nicotiana plumbaginifolia (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Nicotiana rustica (Azab, Megahed & El-Mirsawi 1971; Gameel 1972; Pruthi & Samuel 1942), Nicotiana tabacum (Avidov 1956; Chang 1969; Cohic 1969; Gameel 1972; Gennadius 1889; ~und & Halsey 1978; Pruthi & Samuel 1942), Petunia hybrida (~engonca 1975), Petunia violacea (=phoenicea) (Pruthi & Samuel 1942), Petunia sp. (ClBC Pakistan Station 1983), Physalis angulata (Azab, Megahed & EI-Mirsawi 1971; Pruthi & Samuel 1942), Physalis floridana (Chang 1969), Physalis minima (Mound & Halsey 1978), Physalis peruviana (Avidov 1956; Azab, Megahed & El-Mîrsawi 1971; Chang 1969; Misra & Lamba 1929; Pruthi & Samuel 1942), Solanum dubium (Azab, Megahed & EI-Mirsawi 1971; Gameel 1972), Solanum melongena (Avidov 1956; Chang 1969; Gameel 1972; Kobatake, Osaki & lnouye 1981; Mound & Halsey 1978; Pruthi & Samuel 1942; Takahashi 1956; Tunç et al. 1983), Solanum nigrum (Avidov 1956; Azab, Megahed & El-Mirsawi 1971; Chang 1969; Gameel 1972; Fruthi & Samuel 1942; Tunç et al. 1983), Solanum tuberosum (Avidov 1956; Chang 1969; Gameel 1972; Misra & Lamba 1929; Mound & Halsey 1978; Pruthi & Samuel 1942; Tunç et al. 1983), Solanum verbascifolium (Azab, Megahed & El-Mirsawi 1971; Pruthi & Samuel 1942), Solanum xanthocarpum (Misra & Lamba 1929), Withania sœmifera (Azab, Megahed & EI~irsawi 1971) Glossostemon bruguieri (Azab, Megahed & El-Mirsawi 1971), Guazuma Sterculiaceae tomentosa (David & Subramaniam 1976) , Theobroma cacao (Mayné & Ghesqui~re 1934) Daphne gnidium (Gomez-Menor 1954) Thymeliaceae Corchorus acutangulus (Azab, Megahed & EI-Mirsawi 1971; Pruthi & Tiliaceae Samuel 1942), Corchorus aestuans (Gameel 1972), Corchorus capsularis (Azab, Megahed & EI-Mirsawi 1971; Chang 1969; Gameel 1972; Pruthi &
Samuel 1942), Corchorus olitorius (Gameel 1972; Mound & Halsey 1978),
Corchorus trilocularis (Avidov 1956; Misra & Lamba 1929), Grewia
asiatica (ClBC Pakistan Station 1983)
Trema guineensis (Cohic 1969), Ulmus campestris (Tunç et al. 1983)
Ulmaceae Apium graveolens (Chang 1969), caucalis latifolia (Tunç et al. 1983),
Umbelliferae Coriandrum sativum (Azab, Megahed & El-Mirsawi 1971; Gameel 1972;
Pruthi & Samuel 1942), Daucus carota (Chang 1969), Eryngium sp. (Tunç
et al. 1983)
Host plants
Urticaceae
Verbenaceae
Violaceae Zingiberaceae Zygophyllaceae
25 Urtica urens (Tunç et al. 1983), Villebrunnea frutescens (=Boehmeria
frutescens) (Takahashi 1934)
Call1carpa sp. (Takahashi 1955), Clerodendrum infortunatum (Misra &
Lamba 1929; Pruthi & Samuel 1942), Clerodendrum splendens (Motmd &
Halsey 1978), Clerodendrum villosum (Corbett 1935b), Duranta plumieri
(Pruthi & Samuel 1942), IX1ranta repens (Azab, Megahed & El-Mirsawi
1971), Holmskioldla sanguinea (Mound & Halsey 1978), Lantana camara
(Avidov 1956; El Khidir 1965; Gameel 1972; Motmd & Balsey 1978),
Llppia gemminata (Misra & Lamba 1929), Nyctanthes arbor-tristis (Misra
& Lamba 1929), Verbena bonariensis (01agas, Barradas & Vicente 1981),
Verbena officinalis (Tunç et al. 1983), Verbena sp. (Avidov 1956),
Vitex agnus-castus (Azab, Megahed & El-Mirsawi 1971; Gameel 1972),
V1tex keniensis (Mound 1963; Mound & Halsey 1978), Vitex negundo (CIBC
Pakistan Station 1983)
Viola tricolor (~engonca 1975), Viola sp. (Tunç et al. 1983)
Elettaria cardamanum (Vanna & Gapoon 1958)
Tribulus terrestris (El Khidir 1965; Gameel 1972)
26
Bem1s1a tabac1
27 QJapter JI
llabra! eDeIIies
A. Lopez-Avila
Instituto Colombia
Colombiano
Agropecuario
(ICA),
A.A.
151123
Eldorado Bogotâ,
Presently at: Imperial College, Silwood Park, Ascot, Berks SL5 7PY, U.K. JI. 1. 'nie recorded species The following catalogue of natural enemy records is an update of the The natural enanies are sunmary provided by Greathead & Bennett (1981). listed in Tables 4.1 (parasitoids) and 4.2 (predators). Most of these records are from cotton and almost all the parasitoids belong to the family Aphelinidae (Hymenoptera). The odd records of an encyrtid and a ceraphronid are probably errora, 50 that aIl true records of parasitoids are Aphelinidae. The principal genera are Encarsia (= Prospaltella) and Eretmocerus. The genera Prospaltella and Aspidiotiphagus were recently synonymized with Table 4.1
Hymenoptera parasitoids recorded from Bemisia tabaci
Parasitoid species Aphelinidae Aphelosoma sp. Encarsia sp. (= Prospaltella sp.)
Locality
References
Pakistan
Ahmad & Muzaffar 1977 Azab, Megahed & El-Mirsawi 1970c; Khalifa & EI-Khidir 1965 Elmosa 1979; Sharaf 1982; 1984 Ahmad & Muzaffar 1977 Abdelrahman 1986; Cowland 1934; Fulmek 1943; Shires, Murray & Sadig 1983 Elmosa 1979 Elmosa 1979 Ahmad & Muzaffar 1947 Santis 1981 Santis 1981 Gerling & Rivnay 1984 LaI 1980; 1983; Nair & Nambiar 1983 cmc Pakistan Station 1983 Gerling 1967 Elmosa 1979 Gerling 1972; 1984; Gerling, Metro & Horowitz 1980; Horowitz, Motro & Gerling 1980; Wool, Gerling & Cohen 1984 Viggiani & Mazzone 1980 cmc Pakistan Station 1983 Abdelrahman 1986; Gameel 1969 Gerling 1967 Viggiani 1982 Shafee & Rizvi 1982
Egypt
Jordan Pakistan Sudan
E. E. E. E. E. E.
aspidioticola (Mercet) sp. ?aurantii (Howard) bemisiae de SantisC bicolor de Santis deserti Gerling & Rivnay shafeei Hayat (=~. flava Shafee) E. formosa Gahan
E. lutea (Masi)
E. meritoria Gahan E. mineoi Viggiani E. mohyuddini Shafee & Rizvi
Turkey Turkey Pakistan Brazil Brazil Israel Iodia Pakistan California Jordan Israel
Italy Pakistan Sudan California
Libya Pakistan
Bemisia tabaci
28 E. partenopea Masi
Egypt
Morocco
India
Kenya, Malawi,
Zimbabwe Pakistan a Hawaii Pakistan
E. smithi (Silvestri) E. sublutea (Silvestri)
E. sp. nr. tricolor Foerster Eretmocerus sp.
Egypt
Italy Pakistan Sudan Pakistan California
E. aligarhensis Khan & Shafee E. californicus Howard E. corni Haldanan
Egypt
India Pakistan E. "diversiciliatus Silvestri"d
Egypt
Sudan California
E. haldanani Howard
& Zimbabwe Egypt
Sudan Syria Turkey USSR India
Gerling 1985 EI-Helaly, El-Shazli & EI-Gayar 1971a; Hafez et al. 1983a; 1983bj 1983c; Tawfik et al. 1983 Hayat 1972; Pruthi 1941; Pruthi & Samuel 1942; Samuel 1950 Avidov 1956; Gerling 1972; 1984; Gerling, Motro & Horowitz 1980; Horowitz, Metro & Gerling 1980; Wool, Gerling & Cohen 1984 Tremblay 1959 Elmosa 1979; Sharaf 1982; 1984; Sharaf & Batta 1985 Ahmad & Muzaffar 1977; CIBC Pakistan station 1983 Abdelrahman 1986; Gameel 1969 Greathead & Bennett 1981 Elmosa 1979 Mound & Halsey 1978 Pruthi & Samuel 1942; Samuel 1950
zaire
Mound & Halsey 1978
India Israel
Italy Jordan Pakistan
Pteroptrix bemisiae Masi e b
Aphanogmus fumipennis Thomson
a b
c
d
e
In lit. as
Gerling 1985 Mohyuddin pers. comm. 1986 Gerling 1985 CIBC Pakistan Station 1983 Azab, Megahed & El-Mirsawi 19700; El-Helaly, El-Shazli & El-Gayar 1971a Viggiani & Battaglia 1983 Ahmad & Muzaffar 1977 Joyce 1955a; Shires, Murray & Sadig 1983 CIBC Pakistan Station 1983 Gerling 1966 Priesner & Hosny 1940 Hayat 1972 CIBC Pakistan Station 1983 Khalifa & El-Khidir 1965 Abdelrahman 1986; Cowland 1934 ; Fulmek 1943; Gerling 1966; 1967; Natwick & Zalom 1984
Kenya, Malawi,
E. mundus Mercet
(= masii Silvestri)
cerapbronidae
Priesner & Hosny 1940 Mimeur 1946 Pruthi & Samuel 1942; Samuel 1950
I.
transvena (Timberlake)
Probably an error as this is normally a parasitoid of cecidomyiid larvae, although once reared from a thrips (Dessart & Bournier 1971) Encarsia bemisiae de Santis is a homonym of -.I: (Prospaltella) bemisiae (Ishii) j however, until a replacement name is published, this name will be used Possible misidentification; needs confirmation We have been unable to trace this name
29
Natural enemies Table Il.2 Predators recorded attacking Bemisia tabaci Locality
References
Morocco India
Brazil India Morocco Morocco Israel Pakistan Sudan
Mimeur 1946 Nasir 1947; Rahman 1940 El-Helaly, El-Shazli & El Gayar 1971a Thomas 1932; Husain & Trehan 1933 Link & Costa 1981 Nasir 1947 Mimeur 1946 Mimeur 1946 Israel 1985 CIBC Pakistan Station 1983 Abdelrahman 1986
Sudan
Abdelrahman 1986
India Pakistan India Pakistan Pakistan Brazil Brazil Brazil Pakistan Pakistan India
Rahman 1940; Thomas 1932 CIBC Pakistan Station 1983 Husain & Trehan 1933 CIBC Pakistan Station 1983 CIBC Pakistan Station 1983 Link & Costa 1981 Link & Costa 1981 Link & Costa 1981 CIBC Pakistan Station 1983 CISC Pakistan Station 1983 Rahman 1940
Amblyseius aleyrodis El Badry
Sudan
A. chilenensis Dosse A. limonicus Garman & McGregor A. rubini (Swirski & Amitai) A. swirskii Athias-Henriot
EUseius hibisci (Chant)
Israel Israel Israel Israel California Israel India Israel Sudan Israel Sudan
El Badry 1967; 1968; Gameel 1971 Swirski, Amitai & Dorzia 1970 Swirski & Dorzia 1968 Teich 1966 Teich 1966 Meyerdick & Coudriet 1985 Swirski, Amitai & Dorzia 1970 Gupta & Chaudhry 1972 Swirski, Amitai & Dorzia 1970 El Badry 1967 Swirski & Dorzia 1969 El Badry 1967; Gameel 1971
Egypt
Soliman et al. 1976
Predator species IIeuroptera: QIrysopidae
Anisochrysa flavifrons (Brauer) Brinckochrysa scelestes (Banks) a Olrysopa (s • ...b) sp(p).
Egypt
India
f... cymbele Banks
f... flava (Scopoli)
C. fonnosa Brauer
Chrysoperla carnea (Stephens)
8eIIli.ptera: Anthoooridae
Grius albidipennis (Reuter) Coleoptera: Coccinellidae
Brumoides suturalis (Fabricius) Brumus sp.
Catana parcesetosa (Sicard)
Coccinella septempunctata Linnaeus
Coleomegilla maculata (DeGeer)
Cycloneda sanguinea (Linnaeus)
Eriopis connexa (Germar)
Harmonia dimidiata (Fabricius)
Menochilus sexmaculatus (Fabricius)
Scymnus sp.
Acarina: Pbytoseiidae
Polyphagotarsonemus latus (Banks) b Typhlodromus athiasae Porath & Swirski 1. medanicus El Badry 1. occidentalis Nesbitt T. sudanicus El Badry Acari.na: Stigmaeidae
Agistemus exsertus Gonzalez
a
b
Chrysopa has now been divided into several generaj there is nothing ta indicate that these records refer to Chrysopa (~. ~.) as now defined This is most probably an errar as
~.
latus is a phytophagous species.
Bemisia tabaci
30
Encarsia; under the old definition Encarsia and ProspalteIIa were exclusively parasitoids of whiteflies, whl1e As pidiotiphagus were associated with daspid1d scales. Eretmocerus spp. are also exclusively parasitoids of whiteflies. Species of these genera are often host specifie (Gerling 1912). The recorded host ranges of the parasitoids of Bemisia tabaci are given in Table 4.3. As yet studies have not been carried out to check whether 'species' with wide host ranges are not really complexes of morphologically identical sibling species with specifie host preferences. Since the males of Encarsia spp. are hyperparasitoids on Encarsia females (Gerling 1961) and those of E. lutea have also been reared from moth eggs on infested cotton, albeit with rates of parasitism of only 4-6% (Stoner & Butler 1965), assessment of the potential of this genus as control agents is complicated (Williams 1911). Table
'.3 Alternative
Parasitoid species
hosts for parasitoids recorded from Bemisia tabaci ether hosts
References
Dynaspidiotus abietis (Schrank)a
At least 15 spp. of Diaspididae
Thompson 1950 (Spain) Herting 1912; Thompson 1950; etc. Fulmek 1943 (USA) Thompson 1950 (Hawaii) Thompson 1950 (Hawaii) Shafee 1973 (India) CABI (Pakistan) Mound & Halsey 1918 t-bund & Halsey 1978 Gerling 1966; 1961 (California) Thompson 1950 (England) Ferriere 1965 Gerling 1967 (California) Extensive literature Gameel 1969 (Sudan); Rosen 1966 (Israel) Gameel 1969 (Sudan)
Encarsia spp.
E. aspidioticola E. aurantii E. bicolor E. shafeei E. fonnosa
E. lutea
Aleuroplatus coronata (Quaintance)
Hemiberlesia rapax (Comstock)a
!L. lataniae (Signoret)a
"A scale insect"
Aleurolobus barodensis (Maskell)
Aleyrodes lonicerae Walker
A. proletella (Linnaeus) !. spiraeoides Quaintance Dialeurodes chittendeni Laing D. citri (Ashmead)
TriaieUrodes abutiloneus (Haldeman)
I.. vaporariorum (Westwood)
Acaudaleyrodes citri
(Priesner & Hosny) Aleurolobus niloticus Priesner & Hosny Asterobemisia carpini (Koch) Heliothis zea (Boddie) [eggs]c Pealius setosus Danzig Trialeurodes abutiloneus
I.. vaporariorum Trichoplusia ni (Hübner) [eggs]c
E. meritoria
Aleyrodes spiraeoides
Trialeurodes abutiloneus
I.. floridensis (Quaintance)
E. partenopea
Aleyrodes proletella (Linnaeus)
Asterobemisia carpini (Koch)
!. paveli (Zahradnik)
CABl Stoner & Butler 1965 (Arizona) Danzig 1964b (USSR) Stoner & Butler 1965 (Arizona) CABl (Japan); 3toner & Butler 1965 (Arizona) Stoner & Butler 1965 (Arizona) Gerling 1966; 1967; Oatman 1910 (California) Gerling 1967 (California) Thompson 1950 (USA) CABl (Egypt, ltaly); Thompson 1950 (England) CABl Mound & Haisey 1978
Natural enemies
31
Bulgarialeurodes coteaii (Maakell) Siphoninus immaculatus (Heeger) ~. phillyreae (Haliday)
Trialeurodes vaporariorum
E. smithi
Aleurocanthua citriperdus Quaintance & Baker A. spiniferus (Quaintance) A. woglumi Ashby .. Bemls~a
d
E. sublutea E. transvena
sp. b Melanaphis sacchari (Zehntner) Singhius hibisci (Kotinsky) Trialeurodes vaporariorum
E. sp. nr. tricolor
Aleurocanthus woglumi Aleuroclava sp. Aleurolobus sp.
!. niloticus Priesner & Hosny Dialeurodes citri (Ashmead)
CABl (Turkmenia, USSR) Mound & Halsey 1978 CABl (Bulgaria, Italy); Mentzelos 1967 (Greece); Thcmpson 1950 (ltaly); Tremblay 1969 CABl (Italy); ThOOlpson 1950 (UK); Timofeeva 1963 (USSR) Fulmek 1943 (Sri Lanka, Cl1ina, Malaya , lndonesia) CABl (China, Japan); ThOOlpson 1950 (China, Japan) Flanders 1969 (Mexico introduced); Thompson 1950 (China, Sri Lanka) ThOOlpson 1950 (Scmalia)
Thompson 1950 (Hawaii)
Thompson 1950 (Hawaii)
CABI (Hawaii); Thompson 1950
(Hawaii) CIBC Pakistan Station 1983 (Pakistan) CIBC Pakistan Station 1983 (Pakistan) CIBC Pakistan Station 1983 (Pakistan) CIBC Pakistan Station 1983 (Pakistan) CIBC Pakistan Station 1983 (Pakistan)
Eretmocerus spp.
E. califomicus
E. corni
Aleyrodes spiraeoides Aleurothrixus floccosus (Maskell) Tetralicia sp. Trialeurodes abutiloneus I.. vaporariorum Aleyrodes sp. Dialeuronga fici David & Subramanian Lepidosaphes beckii (Newman)a Pealius quereus (Signoret) Singhius hibisci Siphoninus phillyreae Tetraleurodes comi (Haldeman) Trialeurodes packardi (Morrill)
I.. vaporariorum E. "diversiciliatus" Acaudaleyrodes citri Aleyrodes spiraeoides E. haldemani Aleyrodes spiraeoides Aleurobus niloticus Aleurothrixus floccosus
Gerling 1966 (California)
Thompson 1950 (Puerto Rico)
Gerling 1966 (California)
Gerling 1966 (California)
Gerling 1966 (California)
Fulmek 1943 (Italy)
CABI (India)
Thompson 1950 (Paraguay)
Mound & Halsey 1978
Thompson 1950 (Hawaii)
Mentzelos 1967 (Greece);
Tremblay 1969 Fulmek 1943 (USA) Thompson 1950 (USA); CABl (New York) Thompson 1950 (Chile) Rosen 1966 (Israel) Rosen 1966 (Israel) Gerling 1967; Oatman 1970 (Califomia) Hayat 1972 (India) CABI (California); Thompson 1950 (USA)
Bemisia tabaci Trialeurodes abutiloneus
.I.. vaparariorum Aleuroplatus cadabae Priesner & Hosny Aleyrodes sp. Asterobemisia paveli Bemisia ovata Neomaskellia bergii (Signoret)
E. mundus
a b
c
CABl (Louisiana); Dysart 1966 (Illinois) i Gerling 1967 (California) CABl (Japan) Gameel 1969 (Sudan) Fulmek 1943 (ltaly, Spain)
Mound & Halsey 1978
Mound & Halsey 1978 (USSR)
CABl (India)
Diaspididae Aphididae Noctuidae
d This could weIl be B. tabaci e
CABl refers to the CABI database searched The older records of predators are of Chrysopidae and Coccinellidae, which are polyphagous, but it has not been shown whether whitefly are preferred or incidental prey, but some at least can be recognized as usually feeding on other prey (Table 4.4). The predatory mites are mostly more recently recorded and many have yet to be evaluated. One pathogen has been recorded attacking B. tabaci: a fungus, Paecilomyces farinosus (Dickson & Fries) Brown- & Smith, in India (Balakrishnan & Nene 1980; Nene 1973). P. farinosus is recorded from a wide range of arthropod hosts.. Relatively few natural enemies of other Bemisia spp. have been recorded, and these are summarized in Table 4.5.
table _.4 Prey of predators recorded attacking Bemisia tabaci Predator
Prey preference
Neuroptera: OJrysopidae
Chrysopa and related spp.
Highly polyphagous
Hemiptera: Antbocoridae
Orius albidipennis
Polyphagous
Coleoptera: Coccinellidae
Brumoides suturalis Coccinella septempunctata Coleomegilla maculata Cycloneda sanguinea Eriopus connexa Harmonia dimidiata Menochilus sexmaculatus
Polyphagous Polyphagous, usually associated with aphids Highly polyphagous, often associated wi th aphids Polyphagous, usually associated with aphids Polyphagous Aphids Highly polyphagous
33
Na tural enemies
Acarina: Pbytose1idae
A. chilenensis A. limonicus A. rubini
A. swirskii Typhlodranus athiasae T. occidentalis
Polyphagous Phytophagous mites; also recorded attacking a scale (~Mlrtry 1963) Field records on Parabemisia myricae (Kuwana) in Israel (Wysoki & Cohen 1983); eriophyiid mites. Highly polyphagous in labo tests Field records on whiteflies, scale insects and mealybugs. Highly polyphagous , including pollen, in labo tests Highly polyphagous in lab. tests Usually associated with phytophagous mites
Acarina: stiglaeidae
Agistemus exsertus
Phytophagous mites
Table Il.5 Natural enemies of Bemisia spp. other than Bemisia tabaci and Parabemisia myricae
Host
Natural enemy
Locality
Reference
Bemisia sp(p).
Encarsia nigricephala Dozier
Puerto Rico
Colanbia
Scmalia
Herting 1972 Posada &Garcia 1976 Fulmek 1947
Eretmocerus sp.
Egypt
Priesner & Hosny 1934
Encarsia strenua (Silvestri)
China,
Herting 1972 Malaya, Java,
Sumatra
Eretmocerus mundus Mercet
USSR
Nikolskaya & Jasnosh 1968
Encarsia bemisiae (Ishii)a Encarsia sp. not bemisiae
Japan
Jap~
Eretmocerus sp.
Japanb
Herting 1972 Rose, Debach & Woolley 1981 Rose, Debach & Woolley 1981
~.
afer -B. (Priesner & Hosny) ~.
giffardi (Kotinsky)
B. ovata
(GouX)
Parabem1sia mwricae (Kuwana)
a b
Not to be confused
sublutea (Silvestri)
with~.
bemisiae de Santis (cf. Table 4.1)
Successfully introduced ioto California to control suggest these species have adapted to B. tabaci
-!.=
myricae; there are no reports to
Detailed biological information is only available for the following two species, but as studies in California and Israel, and those of the author, are completed more detail will become available for other species.
Bemisia tabaci
34
'.2. Encarsia lutea. (Masi 1910) (ApbeliDidae) Taxooc-J and diBtribution Encarsia lutea was described as Prospaltella lutea from two female specimens reared t'ran a "cochenille" on Cistus salvifolius collected at Portici, Italy. Mercet (cited in Ferrière 1965) described the male from one specimen collected in Spain. It has since been redescribed by Ganez-Menor (1944), Ferrière (1965) and Viggiani & Mazzone (1980). ~ lutea is the type-species of a group of Encarsia spp. characterized by a peculiar structure of the male antennae (Viggiani & Mazzone 1980). Bionaai.cs
Some aspects of the biology and ecology of E. lutea have been studied. Gerling, Foltyn & Horowitz (in Israel 1982, p. 293) and Foltyn & Gerling (in Israel 1984, p. 142) found that ~ lutea preferred the third and fourth instar of J!.= tabaci for ovipositionj the first and second instars were not used under normal circumstances. Developmental duration of females at 25 c C was 17 days and males developed 2 days more quickly than females. Theil" observations also showed that ~ lutea will lay more than once on the same host even when other unparasitized host are available. Gerling, Metro & Horowitz (1980) found that ~ lutea and Eretmocerus mundus were the only important natural enemies of J!.= tabaci in cotton fields on the coastal plain of Israel, but they also found that the percentage of parasitism did not rise with increases in the population of the whitefly. Horowitz, Metro & Gerling (1981) made similar observations during the summers of 1977-78. 11.3. Eret.>cerus lIUldus Hercet 1931 (Aphelinidae) Taxonaay and distribution
Eretmocerus m.mdus was described from specimens from Italy and Spain reared on a species of Aleurodes infesting aubergine. Ferrière (1965) synonymizes ~ corni Masi with ~ mundus and gives the distribution as: Algeria, Italy, Spain, and the whole Mediterranean Region. Viggiani & Battaglia (1983) report that that ~. mundus is a very active parasitoid on ~. tabaci in the Palaearctic Region. It has also has been reported in Egypt, Greece, Illinois (USA), and Sudan and places given in Table 4.1. Bionanics
Foltyn & Gerling (1985), Gameel (1969), Hafez et al. (1983bj 1983c), Sharaf (1982), Sharaf & Batta (1985) and Tawfik et al. (1983) have studied aspects of the biology of E. mundus. The eggs can be laid under all four nymphal instars, but not the pupaj second or third instar nymphs are preferred. When ovipositing, the female stands at an angle of 90· to the host, with wings raised and inserts the ovipositor under the whitefly nymph. The egg is laid close to the insertion point of the whitefly's proboscis into the leaf. After oviposition, the female apparently marks the host while drumming on it with her hind legs. She distinguishes already parasitized hosts from unparasitized ones and refrains frem laying undel' the former. Discrimination is accomplished after antennal drumming only. Hence, in most cases only one egg is deposited pel' hosto In the few cases when two eggs were deposited, only the second egg hatched and accordingly in aIl cases one parasitoid adult emerged from each host pupa. Under laboratory conditions at 18°C, a mated female deposits about 15 eggs throughout its life, whereas at 30·C, a mated female deposits 48 eggs and an unmated female 42 eggs. At 30·C, the total developmental period takes 18 days, whereas i t takes 20.5 days at 25°C, 26 days at 19'C and 35 days at 12 c C. 80th males and females can copulate within a few hours of emergence. Parthenogenetic reproduction gives both males and females. The percentage of females in progeny of mated females averaged 81.4%, whereas female progeny of virgin females averaged only 36%. Adult parasitoids live for a maximum of 5 days at ambient conditions (mean tempo 25.6°C) when provided wit~ water and honey.
Natural enemies
35
,The egg hatches only under the fourth instar or the pupa. After hatching, the first instar Parasitoid larva bores into the host and develops internally. The parasitized nymph becanes swollen, and the pupa becanes brownish and is smaller than unparasitized ones. Pupation is within the host pupa and the adult wasp emerges through a semi-circular hole at the antero-dorsal part of the host.
36
Bemisia tabaci
37 Q1apter 5
Population
Ecol~
M.J.V. Cook
C.A.B International Institute of Biological Control, Silwood Park, Ascot, Berks SL5 7PY, U.K.
5.1. Introduction The population dynamics of whiteflies as a group has been little studied. With the exception of the studies on Bemisia tabaci discussed below, the only intensive study is a long term one carried out on viburnum whitefly, Aleurotrachelus jelinekii (Frauenfeld), at Silwood Park, Ascot, The population England which was initiated by T.R.E. Southwood in 1962. dynamics of this whitefly on three isolated bushes has been followed for 18 years in order to understand the role of different mortality and biological factors in the population fluctuations (Hassell, Southwood & Reader in press; Reader 1981; Reader & Southwood 1984; Southwood" & Reader 1976). The population dynamics of ~ tabaci with multiple overlapping generations, many host plants and cl:1mates are clearly far more complex. 5.2. Saapling Methods The underpinning of any population ecology study is the sampling programme used to assess the incidence of a pest and its natural enemies. A general text on sampling (e.g. Kogan & Herzog 1980; Morisita 1977; Southwood 1978) should be consulted, and sampling for absolute population estimates and relative monitoring estimates need to be distinguished. The following methods appear in the literature on B. tabaci. 5.2.1. Sticky traps Yellow sticky traps have been quite widely used to monitor flying adults. Gerling & Horowitz (1984) review their use. Studies in Galifornia (Butler, Henneberry & Natwick 1985) have shown a good correlation with other methods. Studies on the attractiveness of light of different wavelengths to ad ults of ~ tabaci (EI-Helaly, Rawash & Ibrahim 1981 a; 1981 b; Mound 1962 ) show why the use of yellow traps is effective. 5.2.2. &letion samplers Dvacs or modified vacuum cleaners have been used, particularly in California (Butler, Henneberry & Natwick 1985; Natwick & zalom 1984) to monitor the incidence of adult whitefly on cotton. 5.2.3. Coœts per plant tmit The distribution of whitefly on plants is far from random. Adults and eggs mostly occur on the young leaves, and the sessile early stages mature with the leaves so that pupae tend to be on the older leaves (Ohnesorge, Sharaf & Allawi 1980). Hence, adults and nymphs must be sampled separately (Mabbett 1983). Adults should be sampled on a per plant basis (Mabbett 1983). Rangaraju & Chenula (1980) describe the use of a bell jar to sample adults from small plants such as Vigna radiataj the jar is placed over the plant which is then tapped, and the adults fly up and settle on the inside of the bell jar where they can be counted. Nymphs should be sampled on a per leaf or per defined portion of leaf basis, but because of the non-random distribution on the plant the choiee of leaf or leaf portion must be earefully defined. For example, early studies in the Gezira, Sudan, sample 10 cm2. from the central area of 100 three-week-old leaves (Cowland 1933).
Bemisia tabaci
38
.. Recent developments in m.merical and sequential sampling theory (e.g. Iwao 1977; Kuno 1977; Sheperd 1980) have been applied to B. tabaci on cotton (von Arx, Baumgartner & Delucchi 1984) and sampling strategies for defined levels of accuracy developed. For monitoring purposes to implement IPM decision rules, such strategies considerably reduce sampling costs, while for population studies they can be used to generate data of predetermined accuracy.
5.3. Seasonal patterns Because B. tabaci is so widespread and polyphagous subtropical areas, it is impossible to generalize about population development, although in the next sections some upon the effects of specific factors. Phenology reported areas is as follows:
in tropical and the patterns of comments are made for some specifie
~
The population on tomato is low over winter, and increases during dry periods leading to outbreaks between April and November with a peak during August - Dctober. Late sowings in April were severely affected while early sowings in March were not (Khalifa & EI-Khidir 1965; Shaheen 1983).
Kerala, Iodia LaI (1981) records a peak on cassava June - August, and Lal & Pillai (1982) record peaks in January, March and June. Punjab, Iodia 1!..: tabaci overwinters on weeds such as Convolvulus and Euphorbia or Brassica crops. In the early spring i t moves onto Hibiscus esculentus, cucurbits and ratoon cotton, then onto the germinating cotton crop in April. It reaches a peak of infestation on cotton in July - August and declines in September - October, before moving to patata, on which peaks occur in September and December, and its overwinter hosts (Husain & Trehan 1933; Husai.n, Trehan & Verna 1936; Sidhu & Dhawan 1981). Iraq On the autumn crop of cucumber population peaks occur in October (EI-Serwij, El-Haidari & Saad 1984). Nigeria Breeding is continuous on cassava, but populations of adults are severely decreased by the effects of heavy rain (Golding 1936). South Africa Over winter survival is on ratoon tobacco. The population build-up is slow in September - November, but increases December - February before declining again (Hill 1968). Gezira, Sudan Until quite recently, B. tabaci was an early season pest. Cowland (1933) reported population peaks on cotton in Dctober, November and December. Bedford (1936) found there is a peak in September on early cotton and in November on late cotton. B. tabaci is numerous on Ipomoea cordofana in Augus t (Bedford 1940) and moves from there onta cotton; in a year of pOOl' growth of...!...: cordofana, there was little problem with ~ tabaci on cotton (Bedford 1937). Towards the end of the year, the population on cotton declines and that on Dolichos lablab increases (Bedford 1940). Since the 1970s multiple applications of broad spectrLml insecticides have changed the situation. Abdelrahman (1986) presents a useful summary for the Gezira. Cotton is planted in late July to early August. B. tabaci populations build up on weeds which appear in July following rains. In
Population ecology
39
September- 'the- whitefly "startto disperse "and colonize cotton, and mass movement occurs in October and November when the weeds dry up. Late rains slow this process. In unsprayed cotton, B. tabaci reaches a population peak in November, whereas in sprayed cotton it continues until late December. During March to June, !h. tabaci can be found on green host plants in gardens, vegetable areas and beside standing water. In July rains restart the sequence on newly sprouted weeds. Turkey Peaks of outbreaks on cotton occur June - August (Baloch & Soomra 1980).
5.Ja. Factors infiuencing populations
5.4.1.
~tion ~ tabaci ls able to maintain itself in flight for quite long periods and is therefore dispersed principally by the action of wind. The initiation of out breaks under suitable conditions is largely defined by the density of ~ tabaci in the air (Joyce 1983) which is a function of emigration fran other infestations. Dùe to wind dispersal, it tends to be more abundant at the edges of fields - in Egypt Khalifa & EI-Khidir (1965) report that it is mos~ frequent on the north edge of cotton fields in summer when the wind is from the south, and on the south edge in winter when the wind is frcm the north.
5.11.2. Kmgration By the time adults of B. tabaci anerge from the pupa, the host plant leaf is generally in pool' condition (Ohnesorge, Sharaf & Allawi 1980) and the adults fly to locate suitable young leaves. Most movanent within a crop is at a low level and is to locate fresh oviposition sites (Gerling & Horowitz 1984). When the crop as a whole begins to deteriorate/senesce, or is harvested, then large nlmbers of adults will leave the plant, fly upwards and be carried by the wind to colonize new sites downwind from the original infestation. These founder fanales will then initiate new infestations. Hence emigration May be on a large scale at certain times, but immigration is usually on a small scale and it is the progeny of these colonizers which develop the new infestation; Le. there is not a mass movement from one crop to another (as with locusts) but a dispersal colonization-population growth-outbreak sequence. 5. Il.3. Climate The statement that rainfa 11 substantially reduces ~ tabaci populations occurs repeatedly in the literature - Khalifa & El-Khidir (1965), for example, suggest that more than 10 mm (0.4") leads to substantial drops in population. Several workers have monitored climatic factors such as temperature, rainfall, relative humidity (RH) and attempted to correlate these with the pattern of attack by B. tabaci. Favourable factors suggested include high temperature 1 low rainfall (Trehan 1944) , period of no/low rainfa11 (Khalifa & El-Khidir 1965), temperature and humidity (Seif 1981), low humidity and high temperature (Cowland 1933), high rainfall, high humidity and a stable maximum temperature (laI 1981) , high maximum temperature (LaI & Pillai 1982), mild winters (Johnson et al. 1982), low humidity (Shanab &: Awad-Allah (1982), adequate humidity Œusain, Trehan & Verna 1936), although there is little indication of how such factors may favour population growth.
5.4.11. RaturaI enemies In Chapter 4 the natural enemies attacking ~ tabaci were reviewed. There is, however, very little information on the importance of these natural
enemies and most of this is anecdotal.
Thus, in the Punjab natural enemies
were said to be important in curtailing outbreaks, but less so than rain (Thomas 1932) or were unimportant (Husain &: Trehan 1933) i in Nigeria the only
40
Bemisia tabaci natural enemies noted were spiders (Golding 1936); in Sudan the concensus is
that natural enemies are caumon, but not effective (Bedford 1936; Cowland 1935) or act tao late in the crop season (Abdelrahman 1986); in Kerala, India, Encarsia flava parasitized 6-21% 1!.: tabaci and 'appeared' to prevent large outbreaks. Several authors have suggested that the use of pyrethroid insecticides has had adverse effects on the natural enemies of B. tabaci leading ta increased outbreaks (cf. Chapter 8). The only detailed studies on the role of natural enemies (parasitoids) in the population dynamics of..!!.: tabaci have been carried out on cotton in Israel (Gerling 1984; Gerling, Matro & Horowitz 1980; Horowitz, Podoler & Girling 1984). By manipulating the infestation with cages, the Israeli workers were able to set up synchronized generations of whitefly on parts of the cotton bushes which they then monitored to accumulate life table data. Carrying out key factor analysis (as described by Varley, Gradwell & Hassell 1973) showed that the natural enemies, principally Eretmocerus mundus and Encarsia lutea, attacked quite a large number of whitefly but were not the key mortality factor. To what extent they MaY still play a regulatory role (cf. Hassell 1985) is left open. When the overwintering populations on a variety of plants in Israel were monitored (Gerling 1984), i t became apparent that populations of bath..!!.: tabaci and its t';oK) parasitoids fluctuated markedly between sample dates, between leaves of the same plant, between plants, between plant species and between localities. Clearly, dealing with average parasitism or spot sampIes of parasitism is not meaningful. The highest parasitism usually occurred on plants which maintained a population of B. tabaci for long periods (e.g. Lantana camara) , suggesting that the parasitoids play more of a regulating role in the long term, more stable situation than the short term, unstable, crop situation. Similar detailed studies are now badly needed for other areas in the range of ~. tabaci. There is no information as to how natural enemies emigrate and immigrate betwe~n host colonies of B. tabaci. Emigration MaY occur later than for the whitefly as development within the host can continue after the host plant is senescing and even as the majority of adult whitefly disperse, their progeny Conversely, the colonizing..!!.: remain and are available for parasitism. tabaci merely have to locate a suitable hast plant; parasitoids have to find a suitable host plant with a population of..!!.: tabaci. Hence parasitoids probably disperse later than their hosts and are less successful at colonizing, so that a significant generation gap MaY develop such that the parasitoids are unable to control the developing outbreak. 5.4.5. Interactions vith hast plants The variation in degree of infestation of plant has already been referred to in the last section and the earlier section on sampling. In developing a sampling programme for cotton, von Arx, Baumgartner & Delucchi (1984) found that the whitefly concentrate on a single main stem leaf, the location of which varied depending upon plant growth and cotton variety. In coolel" seasons, the rate of development of the whitefly decreases, so that even though eggs are laid on young leaves, the sedentary whitefly fails to complete development before the leaf senesces, leading to heavy mortality (Ohnesorge, Sharaf & Allawi 1981). As long as a host plant remains in good condition, MOSt whitefly will remain on it and increase in numbers. Von Arx, Baumgartner & Delucchi (1984) have suggested that there is a density dependent migration effect, but offer no evidence in support; this aspect merits further investigation. Once the plant starts to senesce, or it is destroyed during or after harvest, the adult whitefly will disperse or emigrate. 5.5. Understanding/Prediction The life table analysis study of Horowitz, Podoler & Gerling (1984) has already been referred to (Section 5.4.4.). The largest mortalities were
Population ecology
41
those of crawlera which' disappeared', probably principally for climatic reasons, but also due to the action of predators, and perhaps the effects of plant hairiness. They found that the mortality of instar l nymphs between the crawler settling and the first moult was the key mortality factor which contributed most to the pattern of generational IOOrtality. This mortality was again attributed largely to climatic factors, but it often acted in an opposite manner to the crawler mortality - i.e. if many crawlers disappear, Most of those that settle survive, but if Many crawlers settle, fewer survive. Mortality of the other nymphal stages and the pupa was attributed almost entirely to the action of parasitoids. The pattern of mortalities shown by their data suggests the parasitoids may act in a delayed density dependent manner which could provide a stabilizing, reguiating effect upon the population in the long terme In developing predictive models, the effects of climate upon developnent may be critical and recent studies in California using environment monitoring devices are relevant. Natwick & Zalam (1984) found upper and lower development thresholds for ~ tabaci of 90·F (32·C) and SO.S·F (10·C). Measuring the heat unit acclDulations in the field they found 601.3 and 600.8 day degrees F accumulated between successive population peaks. Von Arx, Batlllgartner & Delucchi (1983b) have developed a simulation model of the growth of cotton and the population dynamics of Bemisia tabaci in the Sudan. This is based upon earlier models of the growth of cotton, and their own detailed laboratory studies on development rates, mortality and fecundity in response to temperature (Von Arx, Baumgartner & Delucchi 1983a). The model uses field-collected temperature data to predict populations accurately over two seasons. As the role of natural enemies was incorporated in descriptive, rather than mechanistic, equations, the model cannot he used to explore the role of the natural enemies in the populations. Emigration of adult whitefly, included as a density dependent effect, was used to fit the model to the observed data. Comparing the model' s predictions with field data, the relative importance of the different components was examined by making predictions with successive components excludedj this showed that migra tion and host plant effects were important, but natural enemies were note From the model the authors conclude that irrigation should be interrupted as soon as possible to prevent further vegetative growth (which would produce further whitefly population growth) and crop sanitation at the end of the season can reduce the number of whitefly which survive to the next season.
42
Bemisia tabaci
43 Qlapter 6
Transmssioo of diseases
A.A. Brœt Glasshouse Crops Research Institute, Littlehampton, W. SUssex BN17 6LP, U.K.
6. 1. Introductioo A virus-like disease of Eupatorium chinense var. simplicifolium, now known to be transmissible by Bemisia tabaci and possibly induced by tobacco leaf curl virus, was first described in 752 A.D. by Empress Koken of Japan in a poem (No. 4268, Vol. 19) included in a classical anthology entitled "Manyoshi" . It 1s thus possibly the earliest record of a virus-induced plant disease (Inouye & Osaki 1981). Viruses transmitted by ~ tabaci are now known to cause ManY severe diseases in tropical crops. Gassava mosaic and abutilon mosaic (= abutilon infectious variegation) viruses have been recognized for almost a century (Baur 1906; Warburg 1894), tobacco leaf curl virus for 75 years (Peters & Schwartz 1912) and others such as those causing euphorbia mosaic and tomato golden mosaic have caused concern for weIl over 50 years (Costa & Bennett 1950; Debrot, Herold & Dao 1963). Whitefly transmitted viruses and as yet uncharacterized virus-like agents are still often prevalent and very damaging pathogens throughout tropical areas (Bird & Maramorosch 1975; 1978; Bird et al. 1975; Costa 1975; Muniyappa 1980; Muniappa & Veeresh 1984; Nene 1972), but occur also in sub-tropical and temperate regions (Costa 1965; Duffus & Flock 1982; Loebenstein & Harpaz 1960) .
6.2. Viroses Nineteen viruses are known to be transmitted by B. tabaci (Table 6.1). One of these (tomato yellow leaf curl) , although ithas been tentatively designated a distinct virus from cytopathological studies, has yet to be further characterized and compared with morphologically similar viruses. With the exception of horse gram yellow mosaic, lettuce infectious yellows, squash leaf curl, tobacco leaf curl, tomate leaf curl and tomato yellow leaf curl, the viruses are mechanically transmissible and moderately stable in vitro. Thus, Most are still infective in sap after 10 min at 50-60·C, after dilution to 10-.2 to 10- 5 and after 21-48 br at 18·C (Bird et al. 1975; Brown & Nelson 1984; 1986; Brunt & Kenten 1973; Cohen & Nitzany 1960; Costa 1976; Costa & Garvalho 1960; Galvez & Gastano 1975; Hollings et al. 1976; Kim et al. 1986; Ni tzany 1970). AlI except cowpea mild mottle, cucumber yellow vein, lettuce infectious yellows and sweet potato mild mottle viruses are members of the geminivirus group, a taxon first recognized by the International Committee for the Taxonomy of Viruses in 1978 (Harrison et al. 1917; Matthews 1979). Geminiviruses have individual particles which measure c. 18 x 20 nm but which often occur in vitro in pairs with overall dimensions of 20 x 30 nm. The geminiviruses such as African cassava mosaic, bean golden mosaic and tomato golden mosaic viruses have been shown to contain a single coat protein (polypeptide mol. wt. 28-34 x 10~) and c. 20% by weight of ss-DNA which occurs predominantly as two covalently closed circular Molecules which are indistinguishable in size (each containing c. 2,600 nucleotides) but differing in nucleotide sequence and genetic content (Bisaro et al. 1982; Goodman 1977a; 1977b; Haber et al. 1981; Hamilton, Bisaro & Buck 1982; Hamilton et al. 1983; Harrison-et-aI. 1977; Reismann et al. 1979). The two Molecules of African cassava mosaic virus ss-DNA have been completely sequenced (Stanley & Gay 1983); both are needed for infection (Stanley 1983),
Table 6.1
~ ~
Viruses transmitted by Bemisia tabaci
Virus
Main host(s)
Occurrence
References
Abutilon mosaic
Abutilon spp.
Tropical America
Bean golden mosaic
Phaseolus lunatus P. longepedunculatus P. vulgaris Manihot esculenta
Tropical & sub-tropical America Africa India
Abouzid &: Jeske 1986; Baur 1906; Kitajima &: Costa 1974 Bird & Maramorosch 1975; Bird et al. 1972; Flores & Silberschmidt 1966; GameZ-1971
Cassava mosaic Cotton leaf crumple Cowpea miId mottle
Cucumber yellows virus Euphorbia mosaic Horse gram yellow mosaic
Harrison et al. 1977; Warburg 1894 Muniyappa & Veeresh 1984 USA Gossypium hirsutum Brown & Nelson 1984 India, Ivory Coast Dubern & Dolet 1981; Iizuka et al. 1984; Muniyappa !rachis hypogaea &: Veeresh 1984 -----Iwaki et al. 1982 Thailand Glycine max Lycopersicon esculentum Nigeria Brunt & Phillips 1981 Phaseolus vulgaris Brazil Costa, Gaspara &: Vega 1983 Vigna unguiculata Africa, Asia Brunt &: Kenton 1973 Cucumis sativus Israel Cohen & Nitzany 1960; Nitzany 1970 Euphorbia geniculata Tropical America Costa &: Bennett 1950 Arachis hypogaea India
Muniyappa et al. in press; Rao, Rao & Reddy 1980 India
Muniyappa, Reddy &: Shivashankar 1976; Muniyappa et Cajanus cajan al. in press; Nene 1972; Williams, Grewal &: Amin 1968 Muniyappa, Reddy &: Shivashankar 1976; Muniyappa et Centrosema sp. India al. in press; Williams, Grewal &: Amin 1968 India Muniyappa & Reddy 1979 Indigofera hirsuta M.m iyappa, Reddy &: Shivashankar 1975; Muniyappa et India Macrotyloma uniflorum al. in press; Williams, Grewal &: Amin 1968 Muniyappa, Reddy &: Shivashankar 1976; Muniyappa et India Glycine max al. in press; Ramakrishnan et al. 1972; Singh, Sandhu &: Mavi 1972 Muniyappa, Reddy &: Shivashankar 1976 Macroptilium lathyroides India Capoor &Varma 1948; Muniyappa, Reddy &: India Phaseolus lunatus Shivashankar 1976; Muniyappa et al. in press Muniyappa et al. in press India Phaseolus vUlgaris
r
1-'. CIl 1-"
Il»
~
~ .....
Teramnus uncinatus Vigna acontifolia
India Indla
~mungo
India India
Vigna radiata Voandzeia subterranea Jatropha mosaic* Lettuce infectious yellows
Melon leaf curl Mungbean yellow mosaic Squash leaf curl
Sweet potato mild mottle Tobacco leaf curl
Tomato golden mosaic Tomato yellow mosaic Tomato yellow leaf curl* Watermelon curly mottle
Jatropha gossypifolia J. multifida Croton lobatus Beta vulgaris Citrullus lanatus Cucumis melo Cucurbita maxima C. moschata Lactuca sativa Cucumis melo Vigna radiata Citrullus lanatus Cucumis Melo Cucurbita maxima C. moschata Ipomoea batatas Capsicum annuum ?Eupatorium chinense Lonicera japonica Lycopersicon esculentum Nicotiana tabacum Lycopersicon esculentum Lycopersicon esculentum Lycopersicon esculentum Cltrullus lanatus
Indla Caribbean
Muniyappa, Reddy & Shivashankal" 1976 Muniyappa, Reddy & Shivashankar 1976; Vil" 1984; Williams, Gl"ewal & Amin 1968 Chhabra & Koonel" 1981; Muniyappa et al. in press Gupta & Singh 1983; Muniyappa, Reddy& Shivashankar 1976; Muniyappa et al. in press; Nal"iani 1960 -----Muniyappa, Reddy & Shivashankar 1976; Muniyappa et al. in press Bird & Sanchez 1911; Cook 1981; Kim et al. 1986
USA
Duffus & Flock 1982; Duffus, Larsen & Liu 1986
USA Asia USA
Duffus & Johns 1985 Honda et al. 1983; Thongmeearkom et al. 1981 Cohen et al. 1983; Flock & -----Mayhew 1981
';/
~ ~ .....
CI) CI)
1-'.
o
::l
oH) 0 1-" CI)
CD
~ CD
CI)
Africa Hollings, Stone & Bock 1916; Hollings et al. 1976 Tropical, sub Mishra, Raychaudhuri & Jha 1963; Osaki & Inouye tropical and some 1978; Osaki, Kobatake & Inoue 1919; Pal & Tandon temperate regions 1931; Vasudeva & Sam Raj 1948 Tropical America Tropical America Middle East Arizona (USA)
Costa 1916; Matyis 1975; Stein, Coutts & Buck 1983 Lastra & Gil 1981 Cohen & Nitzany 1966; Russo, Cohen & Martelli 1980 Brown & Nelson 1986
* Designation as viruses based on ultrastructural studies
~
Vl
46
Bemisia tabaci ,andalthough' they have acoumon sequence .of .c. 200 nucleotides, they are otherwise different. The other four viruses have filamentous particles. Those of cowpea mild mottle virus are mostly c. 13 x 650 nm and contain c. 95% protein (polypeptide mol. wt. 33 x 10 3 ) and 5% ss-RNA (mol. wt. 2.7 x 10') (Brunt & Kenten 1973; Iizuki et al. 1984; Iwaki et al. 1982). Particles of cucumber yellow vein, sweet potato mild mottle and lettuce infectious yellows viruses mostly measure c. 740-800, 850-900 and 1,800-2,000 nm, respectively (Duffus, Larsen & Liu 1986; Hollings et al. 1976; sela et al. 1980). Although the nucleic acids of the latter two viruses have yet to be characterized that of cucumber yellow vein virus is reported to contain DNA (Sela et al. 1980). The geminiviruses that have been investigated occur within the nuclei of phloem and, occasionally, adjacent parenchyma cells. In such locations the particles occur in ribbon-like and/or paracrystalline aggregates. Infected nuclei also contain spherical or ovoid inclusions which, in section, are seen as so-called "fibrillar rings" (Goodman 1981; Horvat & Verhoyen 1981; Kim & Flores 1919; Kim & Fulton 1984; Kim, Shock & Goodman 1918; Kim et al. 1986; Lastra & Gil 1981; Osaki & Inouye 1918; Osaki, Kobatake & Inoue 1979; Russo, Cohen & Martelli 1980; Thongmeearkom et al. 1981). Viruses with filamentous particles that are transmitted by ~ tabaci induce inclusions which, by contrast, occur only within the cytoplasm of aIl cell types. Cowpea mild mottle virus induces the formation of brush-like inclusions which are possibly aggregates of virus particles (Brunt, Atkey & Woods 1983; Thongmeearkon et al. 1984). Sweet potato mild mottle virus induces cytoplasmic cylindrical inclusions ("pin-wheels") which are essentially similar to those induced by potyviruses (Hollings, Stone & Bock 1916). The inclusions associated with infection by lettuce infectious yellows virus are similar to those of closteroviruses (Houk & Hoefert 1983). Inclusions, consisting of aggregated virus particles, and fibrillar bodies induced by geminiviruses in nuclei of phloem cells are readily detectable by light microscopy (Christie et al. 1986). African cassava mosaic, bean golden mosaic and tomato golden mosaic virus es are serologically related (Sequeira & Harrison 1982; Stein, Coutts & Buck 1983). These rela tionships were confirmed by immunosorbent electron microscopy which, in addition, showed that the three viruses are also closely related to euphorbia mosaic and squash leaf curl virus (Roberts, Robinson & Harrison 1984). Spot nucleic acid hybridization tests using probes for African cassava mosa;i.c virus DNA 1 confirmed the results of serological tests; thus, imperfect nucleotide sequences have been found between African cassava mosaic and bean golden mosaic, tomato golden mosaic, tobacco leaf curl, tomato leaf curl and tomato yellow leaf curl. There is, however, no relationship between whitefly- and plant hopper-transmitted geminiviruses (Roberts, Robinson & Harrison 1984).
6.3. YinJs-llke Agents Many plant species are affected by diseases which, although of unknown etiology, are thought to be induced by virus-like agents transmissible by B. tabaci. Some examples of such diseases are given in Table 6.2. Further intensive investigations are required to determine whether sorne species are merely hosts of previously described viruses, or infected by hitherto undescribed viruses or other pathogens. Although some of the virus-like agents are known to have wide host ranges, affected species are for convenience listed separately in Table 6.2. 6._. 'nie Yectors Of the 1,156 species of whiteflies reported to occur world-wide (Mound & Halsey 1978), only three, Bemisia tabaci, Trialeurodes vaporariorum (Westwood) and L abutiloneus (Haldeman), are known to be vectors of plant viruses. Of these, ~. tabaci is by far the Most important; thus, it is known
41
Transmission of diseases Table 6.2 Sorne hosts of virus-like diseases transmitted by Bemisia tabaci
Species
OCcurrence
References
Pakistan
Ahmad 1979
Venezuela
Herold 1967
India
Verma & Singh 1973
Brazil
Costa 1980
lndia
Thomas &
India
Verma 1974c
Sri Lanka lOOia lndia
Gadd & Loos 1941
Wilson & Potty 1972
Srivastava et al. 1977
Taiwan Puerto Rico Puerto Rico India
Liao et al. 1979
Bird & Sanchez 1971
Bird et al. 1975
Vanna. 1963
India
Varma 1963
India lndia lndia Puerto Rico India
Chenulu & Phatek 1965
Vanna 1963
Capoor 1939; Radhakrishnan Nair &
Wilson 1970
Bird 1957
Nair & Menon 1978
USA
McWhorter 1957
India lndia India
Varma 1963
Vanna 1963
Verma 1974b
lndia India Brazil Thailand USSR India Puerto Rico Nigeria Puerto Rico Pakistan Nigeria India Pakistan
Rao, Ragunathan & Joshi 1985
Capoor & Vanna 1950
Costa 1975
Iwaki et al. 1983
Reifman & Polivanova 1969
Verma 1974a
Bird et al. 1972
Williams 1975
Bird 1962; Bird et al. 1975
Ahmed & Harwood 1974
Anno-Nyako et al. 1983
Nene 1972
Ahmad 1978b
India Sudan Nigeria
Singh & Misra 1971
Jones & Mason 1930
Golding 1930
Allarantha.ceae
Digera arvensis Araceae
Anthurilln andraeanum Bal san' naceae
Impatiens balsamina Capparidaceae Cleome sp. Caricaceae Carica papaya
Krishmaswami 1939
CaryopbJ'llaceae
Saponaria vaccaria C20 nymphs/leaf), two as medium (5-20 nymphs/leaf) and the rest as low. Only two parasitoids were reared, but levels of parasitism were generally high.
64
Bemisia tabaci Almost aU samples contained Eretmocerus
..mus.
The rate of parasitism
shows an upward trend in response ta increased whitefly density (Table 10. 1) . Encarsla sublutea occurred in an samples, but the rate of Parasitism falls in meditm infestations and was low in the high infestation. This lack of a density dependent effect suggests~ sublutea may not be very effective at curtailing an infestation, but at low whitefly population densities could play a useful role in regulating the population. Gerling (1985) states that ~ sublutea is synonymous with ~ transvena in Hawaii and also occurs in Japan. In aU three areas it also parasitizes Trialeurodes vaporariorum.
10.2.3. Sudan
Abdelrahman (1986) summarizes the present situation in the Sudan Gezira, based upon parallel studies in sprayed and unsprayed cotton. Four parasitoids attack Jh tabaci: Eretmocerus "diversiciliatus", ~ mundus, Encarsia lutea and another Encarsia sp., but the first of these is of no importance. In treated fields ParaSitism never exceeds 44%, but in untreated fields a Peak of 77% occurred 3-4 weeks after the peak incidence of adult whitefly. In the absence of spraying, parasitism curtails the Jh tabaci population earlier in the season, so that as in the time prior to extensive spraying, it is an early- to mid-season pest. The application of broad spectrum pesticides from the air prevents the parasitoids and predators (principaUy O1rsysoperla carnea) frOID multiplying, until the crop canopy closes over, and leads to the delayed peak in ~ tabaci population. Insecticides are applied early in the season to control Heliothis armigera and Empoasca lybica, pointing to the need for an integrated crop protection approach. 10.2. JI. Egypt and Israel In Israel, Encarsia lutea and Eretmocerus mundus are the only significant parasitoids and they do not provide adequate control (Gerling, Motro & Horowitz 1980). Comments on their incidence are given in Section 4.2. In Egypt the same parasitoids occur and Azab, Megahed & EI-Mirsawi (1970c) report parasitized pupae to be present throughout the year on Lantana camara, but scarce frOID February to June. The maximum parasitism occurs in December (62%). In October parasitism was high on Euphorbia, but low on sweet potato, tomato and cauliflower. 10.2.5. Pakistan In surveys carried out by the CIBC Pakistan station) between 1979 and 1982, natural enemies were plants. The results are given in some detail here available (CIBC Pakistan Station 1983), yet provide assess the potential of biological control.
Station (now PARC-eIBC assessed upon a range of as they are not readily important background to
Encarsia mohyuddini was recently described from Charsadda, Pakistan (Shafee & Rizri 1982) where it parasitized 3% of ~. tabaci on Morus alba. It was not recorded frcxn B. tabaci on other host plants in Pakistan and hence is of little potential for biological control.
crnc Pakistan Station (1983) reported Encarsia sp. Dr. tricolor to parasitize B. tabaci on Morus alba at Multan from May ta December (1-9% parasi tismfa.nd on Lantana camara at Rawalpindi throughout the year (2-11 % parasitism). Although they did not record it from ~ tabaci on other host plants, they did record it from several other hosts as shown in Table 10.2. Thus, this species has been recorded from a variety of hosts and host plants. However, it has not been found to attack~. tabaci on cotton or other crops, and hence is unlikely to be useful for biological control on these.
65
Biological control
Table 10.1 Parasitism of Bemisia tabaci in Eastern Africa (after Gerling 1985)
Host plant
Country
Cassava
Kenya
Cotton Sweet potato Unknown sp.
Malawi Malawi Malawi Malawi
Degree of Infestation
% Eretmocerus mundus
% Encarsia sublutea
high medium lCM low low low low
50 a 17 19 30 19 17 16
very low a 20 27 51 30 24 39
a In one collection Eretmocerus mundus was common and Encarsia sublutea very rare, and in the other, the opposite was true.
Table 10.2 Parasitism by Encarsia sp. nr. tricolor in Pakistan (after CIBC Pakistan Station 1983) Host
Food plant
Locality
Season Parasitism
Aleurocanthus wqglumi Aleuroclava sp. Aleurolobus sp. A. niloticus
Citrus sinensis Morus alba - Ziziphus mauritiana Acacia nilotica Dalbergia sissoo Morus alba - Lantana camara Citrus sinensis
Multan Rawalpindi Multan Rawalpindi Rawalpindi Multan Rawalpindi Mardan
x-i x-iii i-viii vii-x v-viii v-xii i-xii i
Bemisia tabaci Dialeurodes citri
Table 10.3 Parasitism by Encarsia sp.
'c'
2-8% 3-7% 5-18% 4-16% 1-3% 1-9% 2-11% 2%
in Pakistan (after CIBC Pakistan Station 1983)
Host
Food plant
Bemisia tabaci
Aristolochia punjabensis Rawalpindi xii-iii Gardenia jasminoides Rawalpindi vii-ix GossyPium herbaceum Rawalpindi v-x Helianthus annuus Rawalpindi vi-viii Hibiscus rosa-sinensis Peshawar xi Rawalpindi vii Morus alba Multan x-iv --Rawalpindi iii-v Nasturtium sp. Sida grewioides Multan x Multan ~ acontifolia ix Multan v ~.~ Dalbergia sissoo Charsedda ix
3-25% 1-7% 1-5% 1-3% 2% 2% 2-12 3-14% 2% 3% 1% 12
Dalbergia sissoo
9
B. hancocki
Locality
Season
Parasitism
Aleurolobus niloticus
Charsedda
ix
66
Bemisia tabaci Encarsla sp. wC' was reared principally t'rom ~ tabaci, on a range of
host plants excluding cotton (Table 10.3).
Although parasitism rates are
low, this species is consistently associated with ~ tabaci and potentially useful for biological control. Encarsia sp. 'E' was only reared from B. tabaci. Q.lite high levels of parasitism (>10%) were recorded on Morus alba, Hibiscus mutabilis, Lantana camara, Rosa indica and Sesbania sesban, but it was much less coumon on other hosts (Table 10.4). At Hultan parasitism was low August to October, and at Rawalpindi June to August. Apparently host specifie, this species is potentially useful for the biological control of~. tabaci elsewhere. Although it has been reared t'rom Aleurocanthus woglumi and !. husaini on citrus (Sialkot, Rawalpindi) , and commonly (27% parasitism) fran Bemisia hancocki on Dalbergia sissoo in June (Lahore), Encarsia lutea is more consistently associated with ~ tabaci (Table 10.5). Parasitism on cotton was low, compared ta that reported in other countries, and this is attributed principally to excessive use of pesticides. EDcarsia sublutea was listed in CIBC Pakistan Station (1983) as Encarsia (= Prospaltella) sp.
(A) (A.I. Mohyuddin pers. COllIn. 1986) and was reared only from..!!.: tabaci. lt was generally uncommon on aU hostplants except Hibiscus rosa-sinensis and Lantana camara (Table 10.6) and hence is unlikely to be useful for biological control. Aleurolobus barodensis Maskall on sugar cane was regularly parasitized by Encarsia shafeei, but ~ tabaci was also occasionally used. Thus collections t'rom Albizia lebbeck at Multan (10% parasitism) and from Morus alba at Charsadda yielded this species. As it is only casually associated with ~. tabaci, it would net be a usefuI biological control agent. Encarsia (= Prœpaltella) sp. 'c' should not be confused with Encarsia sp. 'e 1 above. lt was reared occasionally from..!!.: tabaci (Table 10.7) and once from Acaudaleyr'Odes citri on Bauhinia variegata in small numbers. It is not likely to be useful for biological control as it is only occasionally associated with B. tabaci and not common. Eret.>cerus aligarbensis was MOst commonly and consistently reared from tabaci, although other hosts are also attacked (Table 4.3). At Rawalpindi it was active throughout the year on a succession of host plants, of which Solanum melongena, cotton, Gardenia jasminoides, soyabean, Lantana camara and Aristolochia punjabensis were Most consistently used. On cotton at Rawalpindi and Multan, parasitism increased from May to a peak of 13% in August-September before dropping off. Incidences on these and other host plants are sunmarized in Table 10.8. Sorne brief biology studies (?host plant) showed that 43% instar l, 19% instar II and 2% instar III were parasitized, and development took 23, 21 and 19 days on each instar respectively. The range of alternative hosts, suitable host plants and moderately high parasitism of..!!.: tabaci on several plants, including cotton, suggest this parasitoid could usefully be introduced into areas where it does net occur.
~.
The widespread parasitoid, Eretmocerus cami, was reared from..!!.: tabaci and 1!..: hancocki, but would seem to be casually associated with..!!.: tabaci and of little potential for biological control. ErebDocerus .mus is the parasitoid Most coumonly associated with ~ tabaci in Pakistan, although it uses other hosts as weIl (Table 4.3). It is active throughout the year and, as for ~. aligarhensis, can be found on a succession of host plants (Table 10.9).
67
Biological control
Table 10.JI Parasitism of Bemisia tabaci by Encarsia sp.
'E' in Pakistan (after CIBC
Pakistan Station 1983) Host plant
Locality
Season
Cucumis sativus Gardenia jasminoides GossyPium spp.
Sialkot Rawalpindi Faisalabad, Multan & Rawalpindi Rawalpindi Multan
vi viii-xi
2 1-2
v-x
1-2
iv-vii ,x-xii i-xii
1-10 1-16
Rawalpindi
i-xii
0.5-10
xii..,.iii
2-7
i-xii i-xii xi i-xii i-xii v
0.5-20 2-23 4 1-18 0.5-20 2
Helianthus annuus Hibiscus mutabilis
Hibiscus rosa-sinensis Lantana camara Morus alba - Ricinus communis Rosa indica Sesbania sesban ~~
Charsadda, Multan & Peshawar Rawalpindi Multan Sahiwal Multan Multan Khanewal
Parasitism rate Peaks Range (%)
vii(10%) xii(13%) v-vi( 11-16%) xii(10%) vi(5%)
xi-iv(>1 0%) iii-iv(>20%) vi-vii(>15% ) ii-iv(>15%)
Table 10.5 Parasitism of Bemisia tabaci by Encarsia lutea in Pakistan Cafter CIBC Pakistan Station 1983) Host plant
Locality
Achyranthes aspera Sialkot Althaea rosea Rawalpindi Cucurbita ~ Rawalpindi Helianthus annuus Rawalpindi Hibiscus rosa-sinensis Peshawar Lantana camara Peshawar Solanum melongena Rawalpindi Sialkot Rosa indica Rawalpindi Peshawar
Season x-xi vii vi v,x-xii ii xi-iii viii v ix xi
% Parasitism 1-2 8
3 3, 6-11 4 0-10 4 7 5 6
68
Bemisia tabaci
Table 10.6 Parasitism of BEmisia tabaci by Encarsia sublutea in Pakistan (after
cmc
Pakistan Station 1983)
Host plant
Locality
Multan Faisalabad Sialkot Faisalabad Multan Rawalpindi Helianthus annuus Rawalpindi Multan Hibiscus mutabilis Hibiscus rosa-sinensis Multan Peshawar Sialkot Rawalpindi Hibiscus sabdariffa Lantana camara Kohat Peshawar Rawalpindi Sialkot Morus alba Charsadda -Albizia lebbeck Cucumis melo Cuctlllis sativus Gossypium spp.
Season
%Parasitism
xi-xii v vi vi-ix x v-x xi-xii xi-xii xii xi ii xii xi iii xi xii-ii xi
4 1 2
1-3 3 1-2 1-2 1-9 7 1 8 4 6 6 9 6-10 5
Table 10.7 Parasitism of Bemisia tabaci by Encarsia (=Prospaltella) sp. (after CISC Pakistan Station 1983) Host plant
Locality
5eason
Althaea rosea Gossypium spp.
Multan Rawalpindi, Faisalabad Faisalabad Multan Faisalabad Sahiwal Khanewal
vii v-x
Helianthus annuus Sida grewioides Solanum melogena ~ aconitifolius
!.
~
x xi-xii x x v
'c'
in Pakistan
% Parasitism 2
1-3 1
4-7 1 4 2
69
Biological control
Table 10.8 Parasitism of Bemisia tabaci by Eretmocerus aligarhensis in Pakistan (after crac Pakistan Station 1983) Host plant
Locality
Season
Albizia lebbeck Aristolochia punjabensis CuClllÙ.S Melo Cucumis sativus Gardenia jasminoides Glycine max GossyPium spp.
Multan Rawalpindi Faisalabad Sialkot Rawalpindi Rawalpindi Faisalabad , Multan & Rawalpindi Faisalabad Rawalpindi Mlltan Multan Peshawar Kohat Peshawar Rawalpindi Faisalabad Sahiwal Rawalpindi Charsadda Faisalabad Rawalpindi Mlltan Khanewal
xi-vi x-ii
3-13 6-10
v
4
vi viii-xi viii-xi
4 3-15 6-10
ix(15%) x(10%)
v-xi
2-10
ixC>10%)
x vi
6 7
v
6
x-iii ix xi xii-ii x-ii ix xi ix
4-8
i v
4 4
iii-x ix
1-5 13 6
Helianthus annuus Hibiscus esculentus Hibiscus mutabilis Hibiscus rosa-sinensis Lantana camara Luffa acutangula Ricinus communis Rosa indica Solanum melongena Solanllll nigrum ~mungo
v
Parasitism rates Range (%) Peak
xii(10%)
1 4
2-3 6-12 5 3 10
xii-i(> 10%) xii-i(>10%)
viii(5% )
Table 10.9 Parasitism of Bemisia tabaci by Eretmocerus mundus in Pakistan Cafter Pakistan Station 1983) Host plant
Locality
Season
Parasitism rate Range (%) Peak
Aristolochia punjabensis Cucumis Melo Cucumis sativus Cucurbita ~ Gardenia jaminoides Glycine max
Rawalpindi Faisalabad Sialkot Rawalpindi Rawalpindi Faisalabad, Rawalpindi Faisalabad, Multan & Rawalpindi Rawalpindi Multan Rawalpindi Rawalpindi Rawalpindi Rawalpindi Faisalabad Rawalpindi
xi-ii
5-12 8
GossyPium spp. Helianthus annuus Hibiscus esculentus Hibiscus rosa-sinensis Hibiscus sabdariffa Lantana camara Rosa indica Solanum ~elongena
v
crac
xii( 12%)
4 vi vi, x-xii 5, 2-4 viii-xi 6-18
ix(18%)
viii-xi
6-11
x(11%)
iv-xi
2-20
vi-ix(>15%)
vi, x-xii 5, 2-4 v 3 10 xii 4 xii 7-14 x-ii ix 5 x 3 iii-x 1-6
xii-iiC>10%)
viii-ixC>5% )
70
Bemisia tabaci
10.2.6. ID1ia
In northern Iodia, Samuel (1951) Incarsia mtbi, F.ret:.x!erus lIB.Sii and report the relative contribution of parasitism (Table 10.10), beyond that ~'
fOWld B. tabaci to be attacked by ~ bœd siae. He did not these three species to the total bemisiae seemed the IDOst coumon.
Table 10.10 Parasitism of Bemisia tabaci in Iodia, October to March, 1940-42 (after Samuel 1951)
Host plant
Ageratum conyzoides Capsicum annuum Crotalaria juncea Duranta p1l.IDieri Glycine max Gossypium herbaceum Nicotiana tabacum Oplismenus burmannii Sesamum indicum ~catjang
Mean % parasitism 4.1 4. 1 2.8
4.4 5.0
39.7 1.2
9.2 4.0 4.0
Range of
J parasitism 2.6-6.1 3.5-7.3 1. 5-4. 6 3.0-6.0
3.8-7.7 22-73 0-2.3 5.9-12.8 2.0-5.9 2.5-5.8
Parasitism occurs principally in the winter months, October to March, and the rate of parasitism increases steadily during this periode Parasitism is strikingly high on cotton, but ooly moderate on the other host plants sampled. Further studies on the incidence and biology of these parasitoids would he worthwhile as they are potentially useful as biological control agents. 10.2.1. Java
In their review of the crop pests of Iodonesia, Kalshoven & van der Vech t ( 1950) make passing mention of the natural enemies of ~ tabaci! listing two parasitic wasps (a yellow species attacking 10-53% of pupae and a black one attacking 1.5-5%) and a small Scymnus sp. (Coccinellidae). The yellow (? Encarsia sp.) and black parasitoids may weIl consist of more than two species 1 and further studies are needed. 10.3.
Possible introductions At this time there are no candidate natural enemies which could he introduced elsewhere in the confident expectation of achieving an effective biological control of Bemisia tabaci. However, as outlined above, we can now recognise several parasitoids coomonly found a t tacking B. tabaci to an apparently significant degree (Table 10.11), and these canIbe considered for introduction into areas where they do not occur. Introductions of these common parasitoids to areas where they do not occur are already starting. Thus, three Encarsia spp. and one Eretmocerus s p. from Pakistan are in cul ture in California (Meyerdirk pers. comm. 1986), at least two exotic Encarsia spp. are in culture in Israel (Gerling pers. comm. 1986), and releases in these countries and Fast Africa are under consideration.
71
Biological control
Table 10.11 The more important parasitoids of Bemi.sia tabaci Locality
Parasitoids
California
Eretmocerus haldanani
East Africa
a Encarsia sublutea Eretmocerus mundus
Sudan, Eastern Med.iterranean
Encarsia lutea Eretmocerus mundus
Pakistan
Encarsia sp. 'c' Encarsia sp. 'E' Eretmocerus aligarhensis Eretmocerus mundus
Northern India
Pteropteryx bemisiae
a
10. JI.
Also present in Pakistan but unimportant.
Discussion The low diversity of parasitoids attacking ~ tabaci in Africa and the Eastern Mediterranean compared to Pakistan is striking and supports the suggestion (Section 1.1) that the origin of J!: tabaci lies in Asia. Accordingly any strategy for biological control of ~ tabaci must include surveys in South-east Asia. The recent survey in Pakistan led ta the discovery of several undescribed species in the taxonomically difficult genus Encarsia (including Prospaltella). Already sorne of these are being cultured for release in other countries, but as yet cannot be related to the coded names in eISe Pakistan Station (1983). Further research for biological control must include an element for taxonomie research to clarify and describe these, and other yet to be diseovered, new species.
Greathead & Bennett's (1981) conclusions about what was needed to develop the potential for classical biological control of B. tabaei still stand: "In addition to minimizing pesticide application to create environment where natural enemies can thrive, studies are needed to:
an
(a) provide life tables [from different regions] to evaluate the action of existing natural enemies and indicate which additional species would be Most desirablej Cb)
conduct comparative studies on cotton and alternative hosts, particularly crops and weeds, growing nearby to evaluate the importance of these other plants as sources of whitefly infestation and of parasitoidsj
(c) carry out surveys of whiteflies in climatically suitable areas in Asia on cotton and other major hasts to search for additional effective natural enemies which MaY be active in the crop and its surroundings j
72
Bemisia tabaci (d)
elucidate the determinants of host and host plant preference of suitable natural enem1es, bearing in mind that formaI taxonomy may conceal sibling species or strains which are more host specifie than their names suggest;
(e)
arrange colonization introductions."
and
post-colonization
monitoring
of
any
73 .Im1otated B1bllograpb;y
Introduct1œ
The bibliography which follows provides a near complete coverage of the publications which refer to Bemisia tabaci. It also includes all papers referred to in the overview chapters - not all of which concem ~ tabaci. AH the citations have been arranged in a standard format. . The titles are either given in the original language, or in English with an explanation in square brackets appended. Journal titles follow the World List of SCientific Periodicals. Where possible, abstracts copied from, or based on, those of the Review of Applied Entanology (Series A) have been included. Those fran the early years have been copied directIy, as have those from the later years, but fol' the intennOOiate period the published format cannot be adapted to that used here and where appropriate these have been re-keyed. Fol' the abstracts copied from the early period of the Review of AppliOO Entanology this has 100 to some anomalies. Thus the scientific names used are those valid at the time, which in the case of B. tabaci means many references to its synonYmS . (cf. Table 1.1.), of which B. gossyPiperda is the most cOlDlDon. Similarly, some of these abstracts are cl'Oss-referred to earlier abstracts (by Review of Applied Entomology volume and page or abstract number) , but these earlier abstracts cannot be located without referring back to the Review of Applied Entomo10gy. Names or phrases in square brackets in a tit1e or abstract have been added during abstracting and do not appear in the original. Where scientific names in the abstracts W8re altered fol' the Review of Applied Entomology, the original names used by authors follow (in brackets) the names considerOO val id at the time. The common names of chemicals follows Review of Applied Entanology usage; a list appears in Volume 68, pp. 7-10. Rewsletter
There is now a news1etter dOOicated to B. tabaci, of which the first two issues have appeared. It is edited by Professor D. Gerling (Tel Aviv University) and Professor B. Ohnesorge, Institut fÜr Phytornedizin, Universitat Aohenheim, Postfach 700562, D-7000 Stuttgart 70, Federal Republic of Germany (address fol' correspondence). The newsletter lists recent publications, relevant meetings and researeh activities, and at present is a t tempting an economic assessment of the ~ tabaci pest problem by means of questionnnaires.
Bemisia tabaci
74 1 AIIOEL FATIAII, S.A,S'i SIIARAF, LH.r.; EL SEIlAE, A. (1983) Perfot'lllaJ\ee of flueythrinate (Cybolt) against a vide speetl'Ulll of agrieultural pests in Egypt. Arab Journal of Plant Proteetion 1 (2), 74-78. (In Arabie vith English Slllllllary.) Field trials earried out in Egypt shOlled that flueythrinate (Cybolt), gave exeellent control at applieation rates of 54-107 g a.L/ha of _isia tabac! on soyabeans. 2 ABDEL-SAllEED, G.A.; ABDEL-SALAH, A.H.; ASSEH, H.A.; AHIN, S.M. (1972) Control of the pests of snake-eucUlIber (CucWllis IIelo L. var flexuosus L.) and cucWllber (C. sativus L.) in Arab Republfcof Egypt. îïïdian Journal of ricultural SciëiiCe"s 42, 95-99. Against Bemisia ta ci on cucumber, OIIethoate and fot'llOthion gave DOrtalitie:s0I96.7Saïïd 90.25%, respeetively. 3 ABOEL-SALAH, A.H.; ASSEH, H.A.; ABOEL-SRAIIEED, G.A. (1971) Experimental studies on tomato pests. 1. Effect of seme pesticides on touto pests in the U.A.R. Zeitschrift für Angewandte Ent0lll01ogie 69, 55-59. TYo tests vere carried out near Alexandria, Egypt, to evaluate the effeetlveness of earbaryl (Sevin), triclùorphon (Oipterex), dillethoate (Roxion). fot1llOthion (Anthio), fenthion (Lebaycid), earbophenothion (Tri thion), phosalone (Zolone) , azinphos-llll!thyl (Guthion), IIIalathion and omethoate (Folilllat) in sprays for the control of 8emisia tabaci and other pests on tOllatO. Omethoate, carbophenothion-;-Ienthion, dimethoate, fot'llOthion and phosalone gave effeetive control. AlI the treated plants outyie1ded untreated ones, but only those sprayed vith carbaryl did so significantly. 4 ABDEL-SALAH, A.H.; ASSEH, H.A.; ABDEL-SllAllEED, G.H.; lIAHMAD, S.M. i RAGAB, F. Y. (1972) Chemiea1 control of SOOK! squash pests in U.A.R. Zeitschrift fUr Angevandte Entomologie 70, 169-174. ExperillE!nts vere conductëd in Egypt to evaluate granular insecticides in comparison vi th emulsion sprays for the control of pests of squash, inc1uding 8emisia tabaci. 5 ABOEL-SALAH, A.H.; ASSEH, H.A.; IIAHIIAD, S.H.; ElO, G.B. (1972) Studies on potato pests in U.A.R. II. SUsceptibility of same potato varieties to insect infestation in the field and in the storage. Zei tschrift für Angevandte Entomologie 70, 76-a2. Four experiments vere conducted in Egypt to ascertain the susceptibility of the potato varieties Alpha (A), Alpha (B), Alpha (E), Aran Banner, Claudia, Gevont 'pol', King Edvard, Hajestlc 'pol', Seinge, Patrones, Perviosnic and Up-to-date ta attaclt by Bemisia tabaci. There vas no significant difference betveen varieties in degree of infestation by !!. tabaci. 6 ABDEL-SALAH, A.H.; KAIERY, E. i ABBASY, A.M.; ASSEH, M.A. (1972) Effects of granular insecticides on seme pests of horse bean and peas as vell as on plant grovth and root nodulation. Zei tschrift für Angevandte Entomologie 70, 408-413. Field and pot experiments vere conducted in Egypt to ascertain the effect of 11 granular insecticides on broad bean and pea and on their insect pests. O-ethyl ethylphosphonothioate (C"".) gave good control of Bemisia taDaci. 7 ABDEl.RAIIIIAN, A.A. (1986) The potential of natural enemies of the cotton whitefly in Sudan Ge.ira. Insect Science and its Application 7, 69-73. 8 ABOUZID, A.; JESKE, H. (1986) The purification and characteri.ation of gemini partieles frOID Abutilon mosaic virus infected Halvaceae. Journal of Phvtopathology 115, 344-353. 9 ABU YAHAN, 1.K. (1971) Outbreaks and nev records. FAO Plant Protection Bulletin 19 (6), 140-141 Bemisla tabaci attacks tomato in Saudi Arabia.
------
10 ABUL-NASR, S.; EL-NAHAL, A.K.H. (1969) Seasonal population of Hemiptera-Homoptera infesting cotton plants in Egypt. Bulletin de la Société Entomologique d'tgypte 52 (1968), 371-389. Bemisia tabaci the ':ec tor of cot ton leaf-cur l, 'Jas presen t in
very large numbers. particularly betveen July and mid-August. AnAL, H.; KHAN. H.R. (1978) Life history and feeding behaviour 11 of green laceving, Chryiopa carnea Stephens (Neuroptera, Chrysopidae). Pakistan Journal of Zoo ogy 10 (1), 83-90.
The biology of Chrysopa camea Steph. is described from observations in the laboratory in Pakistan on matenal collected in cotton fields. The precopulation, copulation and preoviposition periods averaged 11.08 h, 33 min and 4.4 days, respectively, and the egg, larval, prepupal and pupal stag~ 4.8, 12.9, 4.2 and 7.8 days. The number of examples of AphIS gossypii Glov. consumed averaged 487.2/Iarva. or of pUl'ae of Bemisia tabaci (Gennadius) averaged 51O.8llarva Aphlds were preferred to whitefly pupae. 12 AGRAI/AL, R.S.; GUPTA. N.K.: PRASAD, V.K.; VISHIIAKARHA, S. (1979) Chemical control of yello'" mosaic of maong. Pesticides 13 (5), 44-47.
Moong yellow mosaic virus is transmitted by. Bemisia tabaci (Gennadius) and causes severe losses of VIgna spp. (moong and urd) in Uttar Pradesh, Indla. Dunng the zald and kharif seasons (March-June and July-September) of 1976 and 1977 field tests of granular and spray formulations containing 9 insecticide compounds were carried out in the field at Hardoi to determine their possible effect on the
disease incidence. Dichlorvos (Nuvan), monocrotophos (Nuvacron) and phosphamidon, as emulsion concell;trates in sprays applied at 500, 375 and 250 ml tOXIcant/ha, respectively, considerably reduced the incidence of the disease and significantly increased the yield in all seasons except March-June 1976. It was conc1uded that yellow mosaic virus can he effectively controlled by 2 applications (a fortnight apart) of dichlorvos, monocrotophos or phosphamidon. 13 AllARONSON, N.; HUSZl Priesner (, Bosny and !!. gossypiperda Kisra • Lamba. 31 AZAS, A.K.; IŒGAIIED, H.H.; EL-HIRSAIII, B.D. (1970b) Effeet of degree of pubescence of host-plant on the nUllber and distribution of dorsal spines in pupa of Bemisia tabaci (Genn.) (Hemiptera-Homoptera: Aleyrodidae). Bulletin de ïiSoc";éte'"Eiltomologique d' %ypte 53 (1969), 353-357. Studies on the IIOrpho1ogy of Bemisia tabac1 in Egypt indicated a relation betveen the shape of the piijiâBii(l the number and distribution of the dorsal spines, on the one band, and the degree of pubescence of the larval food-plant on the other. Pupae taken froc plants such as c:abbage and sveet potato that have glabrous leaves are generally devoid of dorsal spines or possess less tban seven pairs; those fram plants such as dEuPtibia and Lantana that have pubescent leaves have seven pairs of or spines. 32 AZAS, A.K.; 1lEGAIIED, H.H.; EL-HIRSAIII, H.D. (197Oc:) Parasitism of Bemisia tabaci (Genn.) in U.A.R. (Bemiptera-Bomoptera: AleyroER, H.J.; DABAN, R. (1983) Integrated control of the tobacco whitefly Bemisia tabaci in green/lOuse tou.toes in Israel. In 10th InternationaT'COiiO ëSSOf Plant Protection 1983. Volume r. Proc:e
l
0
a con erence
e
a t Bri
ton,
,
Novem
r,
9 . P ant protection or human ve are. Croydon, UK; Bntis Crop Protection CouncÙ. p. 1109. In investigations on the control of Bemisia tabaci (Gennadius) and tomato yellow leaf curl virus (of which the aleyrodid is a vector) on tomatoes in greenhouses in Israel, the use of a yellow mulch, or painting the glasshouse roof
77
Annotated Bibliography yellow, reduced the number of mfected plants. Minimising the contact between the vector and the plants (for example by inducing a positive air pressure~ithin the house or c10sing down the ventilators to the minimum necessary) contributed to a reduction in infection. Covering the plants with polypropylene sheets reduced infection considerably, but could only be used during the first month, when the plants were less than about 60 cm tall. Spraying with pyrethroids when nece~sary 2-4 times a week, especially when integrated with the other methods, prevented virus transmission and also , prevented the establishment of other pests. 54 BERLINGER, Il. J .; DAlIAN, R.; COHDI, s. (1983) Greenhouse tomata pests and their control in Israel. Bulletin SllOP 6 (3), 7-11 The crop is generally severely dâlïâiêd in the early part of the vin ter , vhen temperatures are still high, by Beadsia tabaci vhich is the ll&in pest. --- -- 55 BEIU.IIQ:R, H.J.; DAlIAN, R.; SHEVACII-l/RKIN, E. (1983) Breeding for resistance to vhiteflies in tOllllltoes - in relation ta integrated pest control in greenhouses. Bulletin SllOP 6 (3), 172-176. The authors review the general procedures for pl~nt
breeding for resistance to specifie pests, and then outhne work for this purpose with species of Lycopersicon and Solanum in search of factors conferring resistance to Bemisia tabaci (Gennadius). The accessions di.ffered markedly in their resistance to the whitefly. The reslstance appeared to be polygenic and comprised various mechanisms, such as differences in pH, the content of secondary plant substances and sticky exudations that were probably toxic. It is thought that breeding for resistance ta B. tabaci in tomato is possible but complicated and still needs much research.
Ir is reported from Sudan that 1980-81 vas disastrous for the production of cotton, ...inly because of the eHects of pests, espec:ially ~ tabac!.
61 BIlOlA, O.S.; SIDIIU, A.S.; SDCII, G.; BRAR, R.S. (1973) Control of suclt1ng pestll of cotton br sail application of granular syst....ic insecticides. IncIian Journal of ~icultural Sciences 43, 352-356. In tvo held tests in t PunJab, ÏÏîdia, in 1969-70, five systeaic insecticides wre applied in lTanules ta the soil against Bemisia tabac! and other pests on cotton. AlI treatlDl!llts reduced lnfestation for up to 10 veeks, the be~t results being obtained vi th disulfo ton , phorate and carbofuran at 1-1.5 kg/ha. 62 Bnw;, F.A. (1973) Nouvelle contribution à l'étude de la IIOsaïque du cotOlVlier au Tchad. 1. Syllpta.es, transllission par lIelIisia tabaci Genn. i II. - Observations sur B. tabaci, III. - Autres maladies virales sur cotonniers et plantes voiSIiiëS:' Coton et Fibres Tropicales 28, 365-378. [English edi tion available.l The syaptOllS and properties of cotton IIIOsaic in Olad are described. The causal agent could be transJli tted even by a single individual of Bemisia tabaci on the plant but does not appear to be a stylet-borne virus, since an acquisition period of 24 h proved insuffic!ent. B. tabaci can easily change frOll one of its other food-plants to -Cotton, but not from cotton to another plant. The first individuals naturally infesting cotton appear in June and are the offspring of mgrants frOll vild food-plants. Lists are given of alternative food-plants of B. tabaci; none of these is knovn to harbour cotton IIIOsaic. -- 63 BINK, F.A. (1975) Leafcurl and IIIOsaic diseases of cotton in central Africa. Cotton Groving Reviev 52, 233-241. Investigations on the IDOsalC and leaf-curl diseases of cotton and other plants, the causal agents of vhich are transmi tted by Bemisia tabaci are discussed. 'nie leaf-curl and IIOsaic S)'IIIptoms observed Oii"Kalvaceae in the African cotton belt are cOMected vi th at least four diseases (leaf-curl small-vein thi.cltening, leaf-curl of IIll1vaceous veeds in Olad, leaf-curl œin-vein thickening, and IlOsaic.
56 BERLINGER, Il.J.; HAGAL, z.; BENZIONI, A. (1983) The illportance of pH in food selection by the tobacco whitefly, Bemisia tabaci. Phytoparasi tica 11, 151-160.
Studies wëre carried out in Israel to confirm that the pH of leaf-cell extracts of cotton plants changed with the age of the plants and ta determine the response of adults of Bemisia tabaci (Gennadius), a pest of cotton and other crops there, to different pH values. The whitefly was able to differentiate between pH values at the level of 0.25, and showed a clear preference for a pH of 6-7.25 offered in vitro. in both choice and no-choice situations. They showed a clear preference for a sucrose concentration of 15%. The addition of 10% sucrose to buffers at various pH values did not change the pattern of the pH preference. In in vivo experiments. whiteflies preferred old cotton leaves (120 days) to voung leaves (60 days); the pH of old leaves was 6.8 and tha't of young leaves 5.9. These resu1ts may explain the fact that B. tabaci attacks cotton plants in commercial fields only late in the season. when the pH of the 1eaves exceeded 6.
BIRO, J. (1957) A vhitefly-trlU1Sllli tted IlOsaic of Jatropha 65 gossypifolia. Technical Papers. Agricultural Experiment Station, Puerto Rico 22, 35 pp. Jaaopha gosPa;pifolia is affected br a IIIOsaic disease in Porto Rico, an a simiar disease has been observed on it in Jamaica, Hartinique and Guadeloupe. The virus vas transmitted by 8emisia tabaci. Hinimum acquisi tion and inoculation feeding periods of a:rurTo minutes, respectively, vere required for transmission, and single individuals readily infected healthy plants if fed on immature leaves. 'nie virus vas found to be of the persistent type. Seedling tobacco plants vere infected in the greenhouse by B. tabaci from infected ~. gossypifolia, but not mature plants, and -they developed symptoms elosely resembHng those reported elsevhere for the leaf-cur l virus disease of tobacco. A leaf-curl disease of tobacco vas found te occur in Porto Rico, but attempts to recover the virus by means of B. tabaci failed. The symptoms vere similar to, if not identical vith, those recorded for tobacco leaf-curl in East Africa and IncIia.
57 BHARGAVA, K.S., JOSHI, R.D. (1962) Yellov mosaic, a virus disease of rose in Gorakhpur. Science and Culture 28, 184-185.
66 BIRO, J. (958) Infectious chlorosis of Sida carpinifolia in Puerto Rico. Technical Papers. Agricultural Expedment Sation, Puerto Rico 26, 23 pp.
58 BHATIACIlERJEE. N.S. (1976) Control of the spread of the yellov mosaic virus of 'moong' in soybean. Entomologists' Nevslener 6 (11112), 64-65. A yellov mosaic virus typical1y affecting IlOOng (~ radiatal but also pathogenie to soy bean is transmi tted in IndTaby 8emisia tabaci which infests the plant trom the 2-leaf stage throughout the grovlng period, being heaviest vhile the plants are still tender. B. tabaci infects soy bean during the formation of the sali vary stylët sheath, which becomes contaminated vi th virus partieles from the stylets before actual sap extraction occurs. It vas found that good protection of soy bean against the virus and its vector vas provided by soil treatment at soving vith aldicarb, phorate, disulfoton or carbofuran at 1.5 kg toxicant/ha, fol1oved by foliar sprays containing 0.01': endosulfan or a mixture of 0.025% methyl-demeton and 0.03S% endosulfan. applied 1 month after soving and repeated at intervals of 15-20 days while the plants remained tender.
nour:s
BIRO, J. (1962) A whitefly-transmitted mosaic of RhffchOsia 67 minima and i ts relation to tobacco leaf curl and other virus iseases orpLants in Puerto Rico. (Abstract) - Phytopatholof 52, 286. 'nie leguminous veed Rhynchosia minima is af ected by a mosaic virus in Puerto Rico which vas transmi ttëdlïy Bemisia tabaci. The host range, symptoms and identi ty of the virus are revieved-.- 68
BIRO, J.; IlARAIlOROSOl, K. (eds.) (1975) Tropical diseases of Nev York, USA; Academie Press Inc. 171 pp. The first seven papers deal vith rugaceous diseases; this tem is used for diseases of uncertain etiology, in the spread of which Bemisia tabaci is involved and which are characterised by symptoms such as malformation, leaf curl, and yellov mosaic.
~.
59 BINIJR.A, O.S. (1983) Insect resistance in cotton in Sudan. In Lamberti, F.; \laller, J.H.; van der Graaf. N.A. (eds.) Durable resistance in crops. Nev York, USA; Plenum Press. pp. 227-229. - -
BlRO, J.; KARAHOROSCH, K. (1978) Viruses and virus diseases 69 associated vith vhiteflies. In Lauffer, Il.A.; Bang, F.B.; Karamorosch, K.; Smith, K.II. (eds.) AdvanCes in virus research 22. London; Academic Press Inc. pp. 55-110. This reviev of plant viruses inc1udes a section on Bemisia tabaci. --
The author discusses insect resistance in cotton in Sudan. where the major insect pests are Bemisia tabaci (Gennadius), Heliothis spp., cicadellids and aphids [Aphis gossypii Glov.]. The ways in which resistance to B. tabaci and H. armigera (Hb.) can be developed are discussed. Studies carried out in the country in 1980-81 on varietal resistance indicated that the okra-leaf character might play an important part in resistance to B. tabaci.
70 BIRO, J.; P~Z, J .E.; ALCONERO, R.; VAKIU, N.G.; KElDIDEZ, P.L. (1972) A vhitefly-transmi tted goiden-yellov masaic virus of Phaseolus lunatus in Puerto Rico. Journal of Agriculture of the Univers! ty OIPUërto Rico 56 0), 64-74. A golden-yellov mosaic virus of lima bean (Phaseolus luna tus) vas detected in Puerto Rico. It vas transmi tted in the Iabôratory by adults of race sidae of Bemisia tabaci to tva varieties of P. lunatus, three of ~. vulgaris and one of tOliaCco. Symptoms caused bY the virus in lima bean, which are similar to those caused by golden IDOsaic virus
60 BINDRA. O.S.; IWIMAN, A.A.A. (1983) Cotton integrated-pest-control in the Sudan. In 10th International Congress of Plant Protection 1983. Volume 3. Proceêdin s of a conference held at Bu hton, and, -2 November, 1 . P ant rotection or human ve are. Croydon, UK; British Crop Protection Counci . p.
71 BIRO, J.; SANCllEZ, J. (1971) lIhitefly-transllitted viruses in Puerto Rico. Journal of Agriculture of the Universi ty of Puerto Rico 55, 461-467. 'nie viruses transmitted in Puerto Rico by 8emisia tabaci are
in Brazil and the yelloll moule virus in El Salvador are described.
Bemisia tabaci
78 dlscussed, together vi th the prindpal plants affected. The probable relationships betwen these viruses and simlar ODeS in atller c:ountries are dlscussed. The vlrus c:auslng IIOsaic dlsease ln reRc:hosia II1nl_ ln Puerto Rico appears to be related to tbat c:auslng y av 1IOSaTCTri" Phaseolus lWl4tus (11_ bean) ln Indla and to tbat c:auslng yellav lIOS&ic of !~p
83 BOND, V.E. (1945) In Progress Reports frOII Experiment Stations season 1943-44. London, Ëiapire Cotton Groving Corporation. pp. 159, 161. In Nigeria a native cotton (Go~ilum Sjiruvianum) is resistance te leaf-curl (~ ~ of Bo s, icll is translRi tted by Beaisia sp., but very susceptible to Dysdercus spp., vbic:h attac:k earIy crops IlOSt severely. 84
BONDAR, G. (1923) Aleyrodideos do Brasi!. Bahia Secretaria da icultura, Industria e Obras Publicas: Sec ·0 de Patolo ia V etaI. pp. The nev species described include Semisia tuberculata frOll cassava. 8
85 BONDAR, G. (1928) Aleyrodideos do Brasn (2a contribuic;1o). Boletim do Laborat6rio de pathologia v'lgetal do Estado de Bahia 5, 37 pp. Inc:ludes Semisia bahiana, sp.n. (~. tabaci). 86 BORROR, D.J.; DeLONG, 0.11. (1964) An introduction to the study of insects. Nev York; Bolt, Rinehart & Vinston. 819 pp.
BORTOLI, S.A. de; GIACOIlINI, P.L. (1981) AC;âo de alguns 87 inseticidas granulados sistêmicos sobre Bemisia tabaci (Gennadius, 1889) (BOIIlOptera-Aleyrodidae) e ~ kraemeri Ross" Iloore, 1957 (BOlIIOptera-Cic:adellidae) e seus erertOSna-prodii'thidade do feijoeiro (Phaseolus vulgaris L.). Anais da Sociedade Entomo16gica do Brasn 10, 97-104. (In Portuguese vith EngHsh sum..,,,·y). In tests in Sao Paulo, Brazil, in 197~77 on the control
of Bemisia tabaci (Gennadius) and Empoasca kraemeri Ross & Moore on beans (Phaseolus vulgaris), granular formulations containing aldicarb, thiofanox or bufencarb were applied to the furrow at the time the seeds were p\anted. The aphid populations were samp\ed at weekly intervals. The best control of B. tabaci was afTorded by the aldicarb at 4 kg/ha, followed by aldicarb at 2 or 1 kg/ha and thiofanox at 2 kg! ha. Aldicarb at 4 or 2 kg/ha also gave the best control of E. kraemeri. The only treatment that significantly increased the yield was aldicarb at 4 kg/ha, though aldicarb at 2 or 1 kg! ha and thiofanox at 2 kg/ha increased the yields as compared with no treatment. BOURIOllET, G. (1938) Note concernant les maladies des plantes cultivées à la Réunion. Revue agricole de l'Île Réunion (N.S.) 43, 33-38. . The Arthropods recorded include Semisia manihotis Frappa. Ilosaic disease of c:assava. of vhic:h B. mani~is considered a possible vector in Iladagascar has not beën observe
Tropical Grain Legume Bulletin 19, 26-29.
Seven cultivars of mung bean (Vigna radiata) that had been shown to be promising for resistance to Bemisia tabaci (Genlladius) (the vector of yellow mosaie virus) were further tested in the Punjab. India. in 1976 and 1977. Observations on whitefly populations and the incidence of virus showed that the cultivars MLl. MU. MU and IMl70 were more resistant than a highly susceptible one that was also included in the trials. MU and MU were the best. 128 CIIIlAllRA, K.S.; KOONER, B.S. (1981) Field resistance in black gram. Vigna mungo L. against insect-pests complex and yellov mosaic virus. Indian Journal of Entomology 43. 288-293.
. Field-eage ëxperiments were carried out in the PunJab, which 63 varieties of black gram '- VIgna mungo) were screened for resistance to Bemisia tabaci (Gennadit!s), Empoasca kerri Pruthi, Lampides boeticus (L.), Maruca testulaiIs (Geyer), Heliothis armigera (Rb.) and yeHow mosaic virus, which is transmitted by B. tabaci. The ~n~la. In 1977 and 1978 in
OlRISTIE, R.G.; KO. N.J.; FAll
177 COI/UND, J.II. (1933) Gezira Entomologieal Section, G.A.R.S. Final report on experilllental \l()rk, 1931-32. Report. Gezira Agncul tural Research Service, Anglo Egyptian Sudan 1932, 93-112. An account is given of work on pests of cotton in the Sudan in the year ending 30th April 1932, much of the information on Bemisla gossypiperda. ~Iisra "& Lamba, the vector oi leaf-curl, having been prenously not!ced ,R._~ .E., A, xx, 623';. The exclusion of lubia (Do/ichos /ab/ab) from the rotation ai crops [x.'\:, 624J seems to have given good results, though an unexpected amount of crinkJed ratoon s\lT\ived the dead season and appeared in the fallow fields at the beginning of the rams. Expenments and observations have shawn that ratoon cotton which is probably the chief source of infection, is not readilv attacked by the whitefiies unless in close proximity ta an attractive plant such as !ubJa. The numbe: feeding on the ratoon in fallow and migrating from ~t t? young cotton IS therefore small. Together with a decrease in the mCldence and severity of the disease, there was a reduction in the nwnber of whitefiies, compared with those present in previous years, panly oWlllg ta the complete destruction of Jpomoea cordofana over la~ge areas by larvae of the convoln.ùus hawk-moth [Herse convoit'uli, 1.: when the unmature stages of B. gossypiperda were de\'eloping on the lea\·es. Apart from a few garden shrubs such as Lantana spp. or C:aT'~ gooseberry [Physalis perut'lanaJ, the whiterlies have been found in tne G,:zlra only on herbs, usually on annuals except for WitiUlnia ~omntJera. Expenmellts prond that the\' are able ta transmit leai ;~rl ta seedlings after feeding on diseased ratoon plants of Sakel cotton llJOSSYPl um peruv,anllm X. barbadellse} of differellt ages (2!-4! mont hs), b,,, do not cor.tract mfect!on irom plants that have recovered or from those infected but not showing symptoms. Sorne whiterlies remained mlect!ve after an inten'aI of 4 days on lubia or 8 dan on health\' Sake!. Durmg April-June, the feeding of whitefiies from Sakel resulted in the production ofsyrnptoms of ;n0saic in 4 out of 6 plants of Watts Long Staple (Amencan) cotton [G. htrsutum] lJl 12-20 days, but during December-February, 2 out of 14 de\'eloped slight crinkle vein~nations on 1-2 leaves after Il days. Later le.aves ~howed no syrnptoms, and the subse9uent generatlon of whitefiles failed ta re-infect plants ai elthe.r vanety.. Expenments ta determine whether infection could be obtamed lJl Watts Long Staple by passage of the virus from Hibiscus
esculentus through H. cannabinus, and in one case also through Sakel, gave negative results [cf. xix. 709]. The development of the disease was relatively late and slight in cotton plants sown in October or later.
178 COI1UlNll, J.II. (1934) Gezira EntOlllOlogica1 Section, G.A.R.S. Gezira Final report on experimental york, 1932-33. 20 ~ltural Researeh Service. Anglo Egyptian Sudan 19, 7-12~ On lubia (Dolichos lablab) sown during May and June 1932 in the Sudan, Bemisia gossypiperda, Misra & Lamba, was nUIllerons only during October and November. Apparently, however, it was not controlled by parasites early in the season either on lubia or on cotton, although after the winter its development on cotton was effectively checked by Ereimoce1us diversiciliatus, Silv., ProspaJ.tella sp. and Encarsia spp. A soda-resin spray [R.A .E., A 20 554) used against the nymphs of Bemisia was not very efficient, and the sprayed plants were immedi ately re-infested by adults. Similarly cotton treated with a spray of castor ail and resin emulsitied with ammonia [17 504] was at once re-infested by adults; and the control of the nymphs was offset by the scorching of the leaves. In expenments on leaf-curl, 2-15 whltetlies caged on single leaves of susceptible and resistant varieties of Sakel cotton (Gossypium peruvianurn X barbai/mse) for 4 days failed ta trans mit the infection. When large numbers 01 infected whiteflies were fed for 1-9 days on lubia and uninfected Sakel plants and then transferred to healthy Sakel seedlings, the disease was successfully transmitted ta the latter. Sakel plants only became infective one day before the veins were visibly thickened. Although cotton is susceptible at ail stages,leai-curl is delayed in aider plants, sometimes until the secondary growth has begun. A resistant variety of cotton (XH 1029) maintained its resistance whate\'er the source of infection. The percentages of transmission experiments ta susceptible cotton that were successful were: 100 from susceptible cotton; 43 from the resistant variety; 9 from Hibiscus esculentus, which is itself resistant; and 100 from H. cannabinus. which is very susceptible. Experiments ta fmd out whether certain weeds are symptomless carriers ai the disease gave negative results. Tests showed that new varieties of cotton bred for their resistance ta leaf-curl [cf. preceding paperJ were able ta maimain their resistance at all stages of their development. The percentage of infection for X 1530 so....'Il ear!y (11 th August) was 0·5 in NO\'ember, 1·5 in December, and 6 in :'larch, as against 56,84'5 and 93 for Gezira :Main Crop. Crops sown later (lst September) were slightly more liable ta infection. Arnerican cotton [G. hirsutum} on an expenmental plot showed less susceptibility than Sakel sown at the same time. The percentage of infection was least in the lat est of the three crops sown.
;3 16'
179 COI/UND, J.II. (1935) Gezira Entomological Section, G.A.R.S. Report on experimental vork, 193>34. Report. Gezira agricultural Researeh Service, Anglo Egyptian SüJM 1934, 99-118. Extensive breeding of Bemisia gossypiperda, ~lisra & Lamba, on Jpomoea corda/ana was recorded at the end of July in the. Sennar district of the Sudan. There is evidence that adults that appeared on J. cordofana and other plants maturing later in areas nonh of this district at the beginning of August had migrated tram these breeding grounds. On lubia (Dolichos /ab/ab) sown during :'lay and June, the Aleurodid was not nUmerous until the rains set in [cf. R.A.E., A 22 68IJ. Ta obtain records of infestation on cotton that could be corn pared from year ta year, the nyrnphs found within a central area of 10 sq. cm. on each of 100 leaves of about 3 week.,. growth were counted weekly in 5 localities throughout the season and c1assitied as healthy, parasitised or dead. This way of counting made it possible ta estimate the number of nymphs belonging ta a single generation. An analysis of the data shows that Aleurodids emerged in large numbers from early ta midd.le OctOber in all localities and that two further large genera tions occurred during the midd.le of November and midd.le of December in two localities. C';tton sO....'Il on 7th August was more heavily infested than cotton sown on 5th September, but the infestation fell off sooner. The rapid decrease was due to the increasing number of nymphs of ail stages drying up from causes that appear ta be connected \~ith tem perature and humidity and age of the plant. These deaths began in the midd.le of Octaber, when humidity dropped though temperature remained high. and rapid.ly became more numerous until the end of December. In parasite cages, where batches of 200 leaves collected fortnightly from various localities were kept, large numbers of Eretmo ceTtls diversiciliatus, Silv., emerged in January, but parasites did not exert an effective check. The increase in the number of parasites is much more marked on D. /ab/ab than on cotton. OnIv one of several attempts to transmit leaf-curl by caging one or two infe~tive Aleurodids in small glass tubes on single apical leaves ai cotton seedlings was successful; in this experiment two individuals were fed on fully devel oped leaves after the plant had been cut back. The disease did not seem ta be more virulent when successive fresh lots of infective .'\leuro dids were caused ta feed on the plants than when only one lot fed con tinuously. The results of experiments in leaf-curl transmission by means of grafting among 4 strains of cotton resistant or susceptible ta leaf-curl are sho"'Il. In main crop sakel [Gossyplum perltviallum X barbadenseJ, the percentages ai infected plants were 29 on 2nd ~o\'em ber and 92 on llth December in 1932 whereas in 1933 on two plots they were onlv 0·6 and 0·2 on Il th ~ovember and 9·6 and 8'4, respec tively, on 24th December. A number of new strains and types of cotton continued ta be far more resistant to leaf curl at aH stages of de\'elopment than the Gezira main crop [loc. cil.}. The effect of
Bemisia tabaci
84 manunng vanes with the strains and date of SOwIDg. but in many cases unmanured plants show less infection. 180
195
CROVE, T.J. (1985) Field trOP pests in Burma. An annotated list. 65 pp.
Rangoon, Burma; Office oftliel'AO Repre~enta;ive.
181 CYPRUS (1977) Annual Report for 1976. Report, Cyprus Agricultural Research Institute, 40-47. The results are presented of control lIl!ausres against Betaisia tabaci as a veetor of tDllato yellov luf curl virus on tllllato. 182 CYPRUS (1979) Annual Report for 1978. Report, ~ Agricultural Research Insti tu te 48-50. The investigations revieved in this report include the chemical control of Bemisia tabaci. 183 CYPRUS (1980) Annual report for 1979. Report, Cyprus Agr1cultural Research Insti tute. 33-35, 36-39. 'hM! Investigations revieved include the cheIoical control of 8eIIisia tabaci vi th special reference to i ts status as a veetor of tOll&to yel10v luf curl virus (TYLCV) in tOll&to. Notes on the transaission of TYLCV in tOlllato by this veetor are also lncluded in a section on plant pathology. 184 CYPRUS (1981) Annual report for 1980. Report, Agricul tural Research Insti tute, 34-37. Continues stûdies on ëh...ical control of Betaisia tabaci.
Cyprus
185 d'ARAUJO e SILVA, A.G.; GONÇALVES, C.R.; GALVAO, D.I1.; GONÇALVES, A.J.L.; GOI1ES, J.; SILVA, 11. do N.; SIIIONI, L. de. (1968) Quarto catalOS? dos insetos que viven nu plantas do Brasil seus parasi tas e prédadores. Parte II l' TOIIIO. Rio de Janeiro, Brazil; Hinisterio da Agricultura. pp. 117-118. 186 DA COSTA UHA, A. (1936) Terceiro cata10go dos insectos que vivem nas plantas do Brasil. Rio de Janeiro, BrazÙ; Hinisterio da Agricultura, Escala NacionaI de Agronomia 460 pp. DAHIYA, A.S.; SINGH, R. (1982) Bio-efficacy of sorne systemic 187 insecticides against jassid, thrips and white fly attacking cotton. Pesticides 16 (12), 13-14, 22.
ln Hissar, Haryana, Iodia, vamidothion at 3 diflerent concentrations was compared with monocrotophos, dimethoate, oxydemeton-methyl, formothion and phosphamidon for the control of Sundapteryx biguttuJa bigutruJa (1shida), Bemisia tabaci (Gennadius) and Thrips tabaci Lind. on cotton. Yamidothion at 0.06% was the most effective against the lst species, monocrotophos at 0.03% against the 2nd and monocrotophos and dimethoate at 0.03% and vamidothion at 0.06% against the 3rd. 188 DANZIG, E.H. (1964a) The whiteflies (HOIIIOptera, Aleyroidea) of the caucasus. Éntomologicheskoe obozrenie 43, 633-646. [In Russian; English translation in Entomological Reviev 43, 325-330.)
190 DANZIG, E.H. (1966) The whi teflies (Homoptera, A1eyrodoidea) of the Southern Primar'ye (Soviet Far East). Entomoligeskoe obozrenie 45, 364-386. [In Russian. English trans1a tion in En tomological Reviev. Vashington 45 (2), 197-209·1 191 DATAR, V.V. (1980) Chemical control of chilli leaf curl complex in Haharashtra. Pesticides 14 (9), 19-20.
The incidence of leaf-curl disease of chilli [Capsicum annuum], which is caused by the tobacco leaf-eurl virus and transmitted by Bemisia tabaci (Gennadius), is high in Maharashtra, Iodia, since the chilli variety commonly grown in that state is very susceptible to it. Tests for the control of the insect vector were carried out in the rainy season (kharif) in 1975 and 1976 with 5 insecticides in the fmt year and 8 in the second. Monocrotophos in a spray at 0.03% was found to be highly effective in bath years in reducing the incidence of the disease and in increasing the yield. As a result, 4 applications are recommended of this compound (in the commercial formulation Nuvacron) at intervals of 10 days, beginning 2 weeks after transplanting. 192 DAVID, B.V.; ANAtmlAKRISIlNAN, T.N. (1976) Host correla.ted variation in Trialeurodes rara Singh and Bemisia tabac! (Gennadius) (Aleyrodidae: Homoptera: Insecta). Current SCi"ëilce 43"";"2ï3-225. Examples of intraspecific diversity in Bemisia tabaci on 6 different food-plants, are given in tables shoving the type, presence or size of 12-14 di~ostic characters of the pupal case. 193 DAVID, B.V.; SUBRAHANIAN, T.R. (1976) Studies on some Indian Aleyrodidae. Record of the Zoological Survey of India 70, 133-233. DEANG, R.T. (1969) An annotated list of insect pests 194 vegetables in the Philippines. Philippine EntoDlOlogist l, 313-333.
of
DE5ROT C., E.A.; ORDOSGOIm F., A. (1975) !studios sobre un
IlOsaico _ril10 de la soya en Venezuela. AgronOlli{ Tropical 25, 435-449. ~ yellov IlOsaic disease of say bean characterised by bright yellov IlOttling, vrinlc.ling of the leaves and stunting of the plants bas been observed frequently in the States of Aragua and carabobo, Venezuela. In tests, the causal agent appeared to be tranSllitted by Bemisia tabaci. Field tests shoved that IlOst plants becalIe infected at an early stage of grovth and the reduction in yield vas negatively correlated vi th the age of the plants vhen infeeted. 196 DEBROT C. ,E.; IŒROLO, F.; DAO, F. (1963) Nota preli.inar sobre un "lIOsaico _rElento" del tOllate en Venezuela. Agronomi! tropical 13, 33-41. [Vith English SUlllllary. J Yellov IlOsaic of tDllato is c _ in Venezuela, and an account is given of studies on the disease S)'lIptOll$ and transmission of the caus,al virus by Bemisia tabaci. 197 DELA'ttRE, R. (1947a) Insectes du cotonnier nouveaux ou peu connus en CCte d'Ivoire. Coton et fibres tropicales 2 (1), 28-33. The author records the presence on cotton ln the Ivory Coast of lleoIisia tabaci. 198 DELA'ttRE, R. (1947b) Insectes du cotonnier nouveaux ou peu connus en C5te d'Ivoire (11.) Coton et fibres tropicales 2 (3), 97-100. Leaf-curl on cotton at Bouaké in 1945 and 1946 is revieved. Suggested control _asures include the destruction of Sida carpinifolia, Age'iâtf' con~ides and Acanth0f!à'Cl'1Il\JlII, which are the IlOst important vi oOd-p ts of lleoIisia ta ci (the veetor), and the planting of ils cultivated f~,""'VIilch include cucurbi ts and Dolichos, only at a distance from the cot ton. 199 DELATI'RE, R. (1961) Studies on the infestation and pesticide treatment of cotton in Africa. Outline of the vork of the I.R.C.T. Phytiatrie-phytopharmacie 10 (1), 13-26. [In French.) 200 DENGEL, H.J. (1981) Untersuchungen über das Auftreten der Imagines von Bemisia tabaci (Genn.) auf verschiedenen Hanioksorten. Zeitschrift fürPrraiiz~eitenund Pflanzenschutz 88, 355-366.
Studies were carried out in Togo on the occurrence of Bemisia tabaci (Gennadius), the vector of cassava mosaic virus, on various cassava varieties. The number of adults on the 2 youngest leaves near the vegetation point fluctuated greatly between the varieties investigated. No connection between this phenomenon and the colour of the petioles was observed. No consistent relationship was observed between the numbers of vectors counted on the 3 local cassava varieties; variations occurred according to location and time of planting. In addition, the vector population decreased between the coast and the inland areas of the country. When planting and sowing were carried out at the beginning of and during the rainy season, no extreme fluctuations in the vector population during the vegetation period were observed. However, the population increased when planting took place in the dry season and plants were subsequently watered. The factor responsible for population fluctuations appeared to be the physiological age of the cassava plants and other nearby food-plants of B. tabaci. 201 DtFARTEHENl' oLEAGINEUX ANNUELS DE l'IRHO (1982) Synthesis of studies presented on control of predators and groundnut diseases. Oléagineux 37 (1), 25-28. In Senegal Bemisia spp. vere found on groundnut but did not cause economi c da.ma:ge:- 202 DESSART, P. ; BOURNIER, A. (1971) Thri ps tabaci Lindman (Thysanoptera) hote inattendu d'Aphan09:US fumipennis (Thompson) (Hym. Ceraphronidae). Bulletin et Annales de a Société Royale d'Entomologie de Belgique 107, 116-118. 203 DllINGRA, K.L.; NARIANI, T.K. (1961) Yellov net virus disease of tobacco plant. [Abstract 44.] Indian Journal of Hicrobiology l, 98. DIIURI, A.V.; SINGH, K.H. (1983) Pest complex and succession of 204 insect pests in black gram Vigna mungo (L.) Hepper. Indian Journal of Entomo10~ 45, 396-401 Inêuding~. tabaci at Delhi. India. 205 DUz, R.O.; BELLOTTI, A.C.; SCBOONHOVEN, A. Van. (1978) Insectos y acaros que a tacan al cul ti vo de la yuca en Colombia. Turrialba 28 (1). 43-49. During a survey in Colombia whi teflies (Bemisia sp. and Trialeurodes sp.) vere present in more than 50% oftiiePlantations vlsi ted. 206 DIooON, R.C.; JOHNSON, H. HcD.; LAIRD, E.F. (1954) Leaf crumple, a virus disease of cotton. PhytoratholOgy 44, 479-480. The cot ton gro"" in the Coachel a and Imperial Valleys of south-eastern Ca1ifornia is affected by a virus disease te~ leat crumple. This disease becornes increasingly cotll!lOn during the s.......,r in
sa"" crops, and ratoon cotton ohen shows 100% infection by midsWMler. The virus does not stunt the plants conspicuously or prevent the setting of bolls, and it is not evident whether it reduces yield. It
85
Annotated Bibliography is apparently distinct frOll other cotton vi ruses , including that causing leaf-eurl in Africa. In tests, it VU t~tted by aleurodids. l\elIlisia [tabad J is videspread and SOlMtimes n.-rous. Batches of aduIts coll~fl"Oll severely infected cotton, in vltich !!. tabaci predOllinated, transœi tted the virus to 16.7% of healthy cotton
ji]iii'i'S• 207 Dnç:R, J. (1979) Progress . .de in integrated control of cotton pests vi th special esophasis on its practicabili ty and econoaic interest. In Russ, K.; Berger. a. (eds.) Proc:eedings. International YomSiua of IOBCIVPRS on Integrated control in agriculture ~d orestry. vieJUla, 8th-12th October 1979. vleJUla, Austrla; Intarnational OrganlZatlon for 81010liCâl Control. pp. 211-213.
The development of integrated control methods against pests of cotton in Turkey since 1977 is described. and notes on attitudes to cotton pest control in other countries in the Middle East (especially Pakistan, Iran, Greece, Israel and Syria) are presented. The cost and effectiveness of chemical treatments against various pests, especially Bemisia tabaci (Gennadius), Tetranycbus cinnabarinus (Boisd.), Spodoptera littoralis (Boisd.) and Heliothis armiger (Rb.), in Turkey are discussed. and it is reported that some resistance to acaricides has developed in T. cinnabarinus. A list is presented of 9 predacious insects of potential future value that were found in the field. The importance of monitoring and the acceptance of new methods by growers in the results that have been achieved so far are discussed.
leaves (expressed on a six-point scale), 2 cultivars are listed as highly promising and 17 as promising. 214 OOZIER, a.L. (1937) Descriptions of .iscellaneous chalddoid parasi tes frOll Puerto Rico (Hymenoptera). Journal of Agriculture of the Universi ty of Puerto Rico 21 (2), 121-135. Includes ~ nigricephala frOll Bemisia sp. on Euphorbia hyperid folta. 215 OUBERN, J.; DOUEr, M. (1981) Groundnut crinkle virus, a nev _ber of the carlavirus group. Phytopathologische lei tschrift 101, 337-347. 216 OUFFUS, J.E. (1965) Beet pseudo-yellovs virus, transmitted by the greenhouse vhitefly (Trialeurodes vaporariorum). Phytopathology 55, 450-453. 217 OOFFUS, J.E.; FLOO
Bemisia tabaci
86 through
1
Parafila -.brane 1s described. It cons1sts of tw sections,
an upper refillable diet c:hallher and a lover insect chaaber. Different parts of the device are interc:hangeable. and they can he autoclave
ln connection with the development of a holidic diet for 1Jemisia tabaci (Gennadius), homogenates of adults were analysed by paper chromatography to detennine their amino acid contents. By the use of the two dimensional ascending technique, 17 amino acids (cystine, aspartic acid, glutamic acid, glycine, serine, threonine, alanine, lysine, histidine, arginine, proline, tryosine, valine, methionine, phenylalanine, leucine and isoleucine) were detected in the soluble fractions, and 16 (aIl the foregoing except cystine) in tht: non-soluble fraction. Each of the fractions also contained 2 unidentified amino compounds. 229 EL-HElJILY, M.S.; EL-SlIAZLI. A.Y.; EL-GAYAR, F.H. (1971a)
Morpholorical studies on illlll&ture stages of llenIisia tabac!, Gennadius
(Komoptera, Aleyrodidael.
Jassid, ~ iYbiCa De Berg, and vhitefly, BeIIisia tabaci
(Gennadius~ro on cotton (Hemiptera: Jassidae anrtant disease affecting the crop in the Pacifie coastal plains, has been fOWld to be identical vith the virus disease reporte
. .ltiple-parasit s.~e ajîhelinid 1s an illljlOrtant eontrolling factor
in the Coaehe11a and Imperial valleys, but not in /tiverside and the
c.oastal Brea.
,
278 GERLIl«;, D. (1967) Bionomics of the vilitefly-parasite eomplex
assoeiated vith cotton in southern California (&o.optera: Aleurodidae;
IIymenoptera: Aphelinidae). Annals of the EntOlDOlogieal Society of
America 60, t306-1321.
---Three speeies of vili teflies Alot'rodbû spiraeoides Quaint.,
Beeisia tabaei (Gennadius) and Trialeur es a tHonea (Hald.), infest
ëottoi1 iii"tlii'" Coaehe11a and Imperial Valleys of southern Califomia. FrOll February to April, no green cotton leaves are available. ~. tabad is foWld during that period on Halva parviflora. Six speeies of parasi tes are knovn to attacl< the vilitëII1es. Encarsia forDlOsa Gah., E• •eritoria Cab. and Eretmoeerus haldemani Rov~tise aIl thr.... ipeeies bût the others do not atta~ci. The f_les of a11 the parasi te speeies and the ...les of ErëtiiiOCei'iiS haldemani are soli tary internal parasi tes of the vili tefly luvae, the ïïâles of Encarsia spp. are parasi tes of Enearsia fe...les. B. tabaci predOlliiiatëSTn the Coaehe11a Valley frOiïJUIY onvards. E.- foiiiiOSa""oceurs in lev nUlllbers. E. llleri toria occurs in lov nUlllbers - in~lIperial Valley, but is abundant during the first tvo-thirds of the season in the Coaehella Valley. Eretmocerus haldemani is the major constituent of the parasite fauna in the Coaeherra-VaIley during the late sUlllller and in the Iaperial Valley throughout the season. Treatments vi th both triehlorphon and a eombination of methyl-parathion and endrin greatly reduced the adult populations of vhiteflies and parasites. The reduetion eaused by trlchlorphon vas usually less drastic than that by Ilethyl-parathion and endrin. The vhi teUies and parasites usually regained their earlier population levels tvo veeks after treatment. 279 GERLIl«;, D. (1972) Notes on thr.... spedes of Eretmocerus Haidetlllll oceurring in Israel "ith a description of a nev spedes. EntOlllOlogisehe Berichten 32, 156-161. ln Israel Eretmocerus mundus Mere., parasi tises Beeisia tabaei. 280 GDU.I~, D. (1983) Factors affecting the establishment' of Bemisia tabaci on cotton in Israel. In tOth International Congress of PIailtPiotewon 1983. Volwne 3. Proceedinllants. Iktwf,en 21st ~Iarch and tith Apnl. iil4 adults of H. nil;mtnsis colIeckd from infected p1:lnts in the !Jdd w~re Introduced in batehes of :!.'!.-H7 lntn on
Annotated Bibliography near Baghdad did not appear severe, but in country districts where cotton was generally unthinned and over-irrigated, great losses were experienced. The plants are probably weakened and sa become more susceptible to adverse c1imate or poor cultivation. The honey-dew covering the leaves encourages the growth of fungus in damp situations where the aeration is not sufficient ; the lint is alsa damaged. The premature opening of the bolls in August was probably due to this insect, which began to diminish in numbers in September and had almost entirely:disappeared by the end of October. 311 GUPTA, P.C. (1972) External DOrphology of BMisia JOSSYJi~rda (Il. & L.) a vector of plant virus diseases (8olIoptera: eur i e). Z00logische !leitrage 18, 1-23. 312 GUPTA, P.C.; CBAUlllIRY, R.S. (1972) New record of lIeIIIitarsoneIIIUS latus Banks (Tarson....idae) as a parasi te of Ilemisia gossypiperda H. & L. Indian Journal of EntOllOlogy 33 (1971), 4 ~ Dûring a survey of the food-plAn ts of 1leIIisia tabaci in the Gorakhpur Region of Uttar Pradesh, India, nevly eIlergedaa.:iIts of the aleyrodid vere observed to be a t taclted by .Hes. The species ci ted by the authors is pol~tarsonemus latus (Banks) [vllich is c~nly regarded as being exc usively phytophagous). 313
GUPTA, P.K.; SImH, J. (1982) Important insect pests of covpea (~ W!gUiculata L.) in agroeCOsystClll of eastern Uttar Pradesh. IiIiflaii Jourriâl of Zootoay 22 (2), 91-95. Inclûding iêÎÎisia tabaci. 314 GUPl'A, P.K.; SINGB, J. (1983) Effect of systemic granular insecticides on viii tefly population and yellov-llOsaic infection in greengru. Indian Journal of Agricultural Sciences 53, 737-742.
When applied at 2 kg a.i.jha in summer as weil as in the rainy season. the insecticides controlled Bemisia tabaci and concomitantly [mung bean] yellow mosaic virus on Vigna radiata. 315 HABER, s.; IlŒGAHI, H.; BAJET, N.B.; GOODMAN, R.Il. (1981) Evidence for a divided genome in bean golden DOsaic virus, a geminivirus. ~ , London 289, 324-326. HABIB, A.; FARAG, F.A. (1971) Studies on nine cOllllllOn aleurodids 316 of Egypt. Bulletin de la Société Entomologique d'Égypte 54 (1970), 1-41. Keys are provided for the separation of the tvo species of Remisia. HABIB, R.; HOHYUOOIN, A.1. (1981) Possibili ties of biocontrol of 317 some pests of cotton in Pakistan. Bio\ogia 27 (1), 107-113. The possibili ties of biologica control of llemisia tabac! in Pakistan are discussed. --- -- 318 HABIBI, J. (1975) The cotton vhite fiy Remisia tabaci Gen. Bioecology and methods of control. Entomologie---etPhytoPatllologie Appliquées 38, 13-36. [In Persian, English summary pp. 3-4.] Since 1955 Remisia tabaci has been observed in aIl the cotton-grololing areasOf"Southern and central Iran. Large populations significantly reduce the moisture and nitrogen content of the plants and exode large quantities of honeydelol; as a result, most of the leaves, flololers and immature bolls fall prematurely and the number of buds and the quantity and quality of cotton yield are reduced. The late-instar 1arvae and pupae overvinter under the leaves of IoIeedS that remain green in the vinter, such as Convolvulus and Althaea sPf' The adults emerge in spring vhen the mean temperature reaChes 12 C and feed on the leaves of IoIeedS and cucurbi ts, but during the hot vinds of summer the preferred food-plant is melon, vhich bas thick juicy leaves; at the end of August vhen the leaves of melon vilt, B. tabaci migrates to cotton fields, and the decreasing temperatüre~ increasing humidi ty favour the rapid build-up of large populations in a short time. In the Garmsar district, vhere the bionomics vere studied, the mild vinters usually enabled populations to carry over from one year to the next. In years of heavy infestation, integrated control is necessary and should indude early soving of cotton to provide a harvest vhile B. tabaci is still on other food-plants; cultural measures such as 1ertiHsing, irrigation and veed control to promete vigorous and early grolo"th; the si ting of cotton and melon fields as far apart as possible; the destruction of infested melon leaves alter the harves ting of the frui t; and chemical sprays of 35% endosulfan (Thiodan) at 3 l i tres/ha, or of mixtures of 49% dimethoate at 2 litres vith 25% DDT at 4 litres/ha or 49% dimethoate at 1 kg vith 85% carbaryl (Sevin) at 3 kg/ha. 319 HAFEZ, H.; TA1lFIK, M.F.S.; AIIADALLAH, K.T.; SARIWI, A.A. (1983a) Natural enemies of the cot ton vhi tefly, !lemisia tabaci (Genn.), in the IoIorld and in Egypt. Bulletin de la sociTtTEn"tOmoIOi"ïque d'fgypte 62 (1978/1979), 9-13.
Bem1sJa tabaci (Gennadius) is an important pest of especially of cotton in Egypt, sinee as weil as damage it makes the cotton lint stieky and ~eult to spin with its honeydew. Numbers have increased ID re:eent y~, possibly owing to the killing of natura! ~enues by Wldespread application of pesticides. A list is glven of 18 species of parasites (mostly Encarsia and v~getables ~d ~ect feeding
Ere~ocerus spp.) and 2 predators (Chrysopa spp.) of B. tabac! t.hat are round in the. worId. During a field survey in Egypt ID 1973-75, E. medius Mereet and an unidentified species of Enearsia were the only parasites found on B.
91 tabaci, but ChrysoperJa carnes (Steph.) (Chrysopa cames), Drius spp. and Scymnus syriscus Mars. were all found preying on eggs, larvae and pupae of the aleyrodid. 320 IIAFEZ, Il.; TAIIFIK, H.F.S.; AIIADALLAII, K.T.; SARRAN, A.A. (1983b) Studies on EretDOcerus mundus Hercet, a parasite of the cotton vIIitefly, Semisia tabaci (Genn.) in Egy t (iIylIIenoptera: Aphelinidae). Bulletin deIiiSOciTtTEntODlOl i ue d' te 62 (1978/1979), 15-22.
. A method is described of rearing Eretmocerus mundus Mere., a parasite of the cotton· and vegetable pest Bemisis tabaci (Gennadius), in the laboratory in Egypt. Adults of B. tabaci were reared from pupae collected from cotton, oUa. cabbage and Lantana camara in the field, and were released into oviposition cages containing a cut stem of sweet potato in a jar of water. A generation was reared in about 20 days under these conditions. Aduit parasites emerging from field collected hosts on the food·plants mentioned were introduced intoglass lamp-chimneys· containing sweet potato stems in water. New aduit parasites emerged iD 13-14 days at 27-34·C and 65-75% RH. Although E. mundus was previously recorded from Spain, Illinois (USA), Greece and Sudan, it was found 10 be widespread in Egypt though less common in the south. Its only hosts in Egypt appeared to be B. rabaci and Aleyrodes proJetelJa (L.), a pest of cauliflower. Notes are given on the morphology of the immature stages of E. mundus. The eggs were laid externally under the hast larva, but the parasite larva on hatching immediately entered the body of the host and developed internal1y. Pupation took place within the dried body of the hast larva. 321 HAFEZ, H.; TAIIFIK, M.F.S.; AIIADAlJjJj, K.T.; SARRAN, A.A. (1983c) Impact of the parasite, Eretmocerus mundus Mercet on population of the cotton viii tefly , Bemisia tabaci (Genn.) , in Egypt. Bulletin de la §acUité Entomelogi~gy!ite6""2(1978/1979), 23-32.
In 197+'75, the rates of parasitism by Eretmocerus mundus Mercet on Bemisia tabsci (Gennadius) on sprayed cotton and cabbage in both Upper and Lower Egypt were compared with the_rate on populations on unsprayed plants of Lantana camara at Giza. In 1974, parasitism on cotton averaged 46% in Lower Egypt and 31 % in Upper Egypt, and the corresponding rates for 1975 were 42 and 71 %. On cabbage, parasitism averaged 31 and 36% in the 2 localities in 1974 and 47 and 52% in 1975. On L. camara at Giza, the rates of parasitism were 75% in 1974 and 69% in 1975.' The highest rates of parasitism generally occurred in October and the lowest ones in August. E. mundus thus appea.rs to be a major factor in regulating populations of B.· tabaci in Egypt, but its activity is hindered by insecticide treatments; its raIe in pest control must therefore be considered before any programmes of integrated control on cotton or vegetables are planned. 322 IlAMILTON, Il.0.0.; BI SARO , D.H.; Bua
