Bemisia tabaci (MED) (silverleaf whitefly)

Identity

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Preferred Scientific Name

Bemisia tabaci (MED)

Preferred Common Name

silverleaf whitefly

Other Scientific Names

Bemisia tabaci biotype Q
Bemisia tabaci Q
Mediterranean (MED) species (Bemisia tabaci) (Gennadius, 1889)

International Common Names

English: poinsettia whitefly; Q biotype; tobacco whitefly

English acronym

MED

Summary of Invasiveness

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The exact origin of the MED species of Bemisia tabaci, and the reasons why it became such an important pest are still not fully known. MED species has been identified as a distinct member within the B. tabaci species complex (Tay et al., 2012). MED species is also an effective vector of many different plant viruses which, in conjunction with its high level of polyphagy, make it extremely problematic within agricultural regions where crops may be susceptible to viruses acquired from indigenous plants. Despite Bemisia in general being a tropical/sub-tropical whitefly species, MED species can easily be transported on plant species to temperate regions of the world (Cuthbertson and Vänninen, 2015). Within these cooler regions, MED species can survive within a protected environment and could feasibly spread virus diseases to new locations. It is for this reason that B. tabaci and members of its species complex, including MED species are on EPPO A2 Alert list.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Arthropoda
Subphylum: Uniramia
Class: Insecta
Order: Hemiptera
Suborder: Sternorrhyncha
Unknown: Aleyrodoidea
Family: Aleyrodidae
Genus: Bemisia
Species: Bemisia tabaci (MED)

Notes on Taxonomy and Nomenclature

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The pest status of B. tabaci insects is complicated and through the comparison of mitochondrial cytochrome oxidase 1 (mtCO1) gene it is generally accepted that, rather than one complex species, B. tabaci is a complex of 11 genetic groups. These genetic groups are composed of at least 34 morphologically indistinguishable species, which are merely separated by a minimum of 3.5% mtCOI nucleotide divergence (Dinsdale et al., 2010; De Barro et al., 2011; Boykin and De Barro, 2014). The MED species, which was originally described in Greece as Aleyrodes tabaci in 1889 by Gennadius, was recently confirmed using molecular markers as the Mediterranean species (Tay et al., 2012). Within the B. tabaci complex, the Middle East-Asia Minor 1 (MEAM1) cryptic species, formerly referred to as 'B biotype', and Mediterranean (MED) cryptic species, formerly referred to as 'Q biotype' are the two most widely distributed, and as a result, best known of the species. These two species present the greatest threat to glasshouse crops worldwide (Bethke et al., 2009). The damaging MEAM1 is described as an aggressive coloniser and is an effective vector of many viruses, whereas the MED species characteristically shows strong resistance to novel insecticides (Jones et al., 2008; McKenzie et al., 2009).

Description

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Eggs

Eggs are pear shaped with a pedicel spike at the base, approximately 0.2 mm long.

Puparium

A flat, irregular oval shape, about 0.7 mm long, with an elongate, triangular vasiform orifice. On a smooth leaf the puparium lacks enlarged dorsal setae, but if the leaf is hairy, 2-8 long, dorsal setae are present.

Adult

Adults are approximately 1 mm long, the male slightly smaller than the female. The body and both pairs of wings are covered with a powdery, waxy secretion, white to slightly yellowish in colour.

Distribution

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MED species continues to be detected around the globe (Chu Dong et al., 2006).

Certain areas within Europe are still MED species free, e.g. Finland, Sweden, Republic of Ireland and the UK (Cuthbertson and Vänninen, 2015).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 25 Feb 2021

Continent/Country/Region	Distribution	First Reported	Invasive	Reference	Notes
Africa
Algeria	Present			Ahmed et al. (2009) Tahiri et al. (2006) Shatters et al. (2009)
Burkina Faso	Present			Gnankine et al. (2013)
Egypt	Present			Ahmed et al. (2012) Simón et al. (2007) Ahmed et al. (2009)
Ghana	Present			Shatters et al. (2009)
Morocco	Present			Ahmed et al. (2009) Monci et al. (2000) Tahiri et al. (2006) Simón et al. (2007) Shatters et al. (2009)
Senegal	Present			Hélène et al. (2015)
South Africa	Present	2013		Esterhuizen et al. (2013)
Sudan	Present			Shatters et al. (2009)
Tanzania	Present			Tajebe et al. (2015) Tajebe et al. (2015a)
Togo	Present			Gnankine et al. (2013)
Tunisia	Present			Ben Abdelkrim et al. (2017) Saleh et al. (2012)
Uganda	Present			Shatters et al. (2009)
Asia
China	Present			CABI (Undated) Zhang et al. (2005) Chu Dong et al. (2006) Chu et al. (2007) Hsieh ChiaHung et al. (2007) Ma DeYing et al. (2007) Ahmed et al. (2009) Rao et al. (2011) Chu Dong et al. (2012) Li RuMei et al. (2013) Jahan et al. (2015)	Present based on regional distribution.
-Anhui	Present			Rao et al. (2011)
-Beijing	Present			Li RuMei et al. (2013) Zhang et al. (2005) Chu Dong et al. (2006) Hsieh ChiaHung et al. (2007) Ahmed et al. (2009)
-Gansu	Present			Teng Xi et al. (2010)
-Guangdong	Present			Sun XiuXin et al. (2013)
-Guangxi	Present			Teng Xi et al. (2010)
-Guizhou	Present			Teng Xi et al. (2010)
-Hainan	Present			Teng Xi et al. (2010)
-Henan	Present			Hsieh ChiaHung et al. (2007) Teng Xi et al. (2010)
-Hubei	Present			Ahmed et al. (2009) Teng Xi et al. (2010) Rao et al. (2011)
-Hunan	Present			Teng Xi et al. (2010)
-Jiangsu	Present		Invasive	Li et al. (2015) Ahmed et al. (2009) Teng Xi et al. (2010)
-Jiangxi	Present			Rao et al. (2011)
-Shaanxi	Present			Teng Xi et al. (2010)
-Shandong	Present			Chu Dong et al. (2010) Chu et al. (2007) Ahmed et al. (2009) Teng Xi et al. (2010) Jahan et al. (2015)
-Shanxi	Present			Teng Xi et al. (2010)
-Tianjin	Present			Teng Xi et al. (2010)
-Tibet	Present			Lu ShaoHua et al. (2016)
-Yunnan	Present			Ahmed et al. (2009) Zhang et al. (2005) Chu Dong et al. (2006) Hsieh ChiaHung et al. (2007) Chu Dong et al. (2012)
-Zhejiang	Present			Ahmed et al. (2009) Hsieh ChiaHung et al. (2007) Teng Xi et al. (2010)
Israel	Present			Horoxitz et al. (2008) Horowitz et al. (2003) Ahmed et al. (2009) Kontsedalov et al. (2012)
Japan	Present			CABI (Undated) Hsieh ChiaHung et al. (2007) Ahmed et al. (2009)	Present based on regional distribution.
-Ryukyu Islands	Present			Kijima et al. (2013)
Malaysia	Present			Shadmany et al. (2019) Mohammad Shadmany et al. (2013)
-Peninsular Malaysia	Present			Mohammad Shadmany et al. (2013)
-Sabah	Present			Shadmany et al. (2019) Mohammad Shadmany et al. (2013)
South Korea	Present			Kim EunSung and Kim YongGyun (2014) Hsieh ChiaHung et al. (2007) Ahmed et al. (2009) Jahan et al. (2015)
Syria	Present			Ahmed et al. (2012)
Taiwan	Present			Hsieh ChiaHung et al. (2011) Hsieh ChiaHung et al. (2007)
Turkey	Present			Karut et al. (2014) Ahmed et al. (2009) Shatters et al. (2009) Fidan et al. (2019)
Vietnam	Present			Götz and Winter (2016)
Europe
Cyprus	Present			Ahmed et al. (2009) Papayiannis et al. (2009)
Finland	Absent, Eradicated	2014		Lemmetty and Vänninen (2014)
France	Present			Ahmed et al. (2009) Tahiri et al. (2006) Dalmon et al. (2008) Shatters et al. (2009)
Germany	Present			Nauen et al. (2002)
Greece	Present			CABI (Undated) Roditakis et al. (2009)	Present based on regional distribution.
-Crete	Present			Roditakis et al. (2009) Ahmed et al. (2009)
Italy	Present			Cavalieri et al. (2014) Nauen et al. (2002) Simón et al. (2003) Bosco et al. (2004)
-Sardinia	Present			Demichelis et al. (2005) Simón et al. (2003)
-Sicily	Present			Simón et al. (2007) Simón et al. (2003) Bosco et al. (2004)
Netherlands	Present			Ahmed et al. (2009)
Portugal	Present			Ahmed et al. (2009)
Serbia	Present			Konjević et al. (2018)
Spain	Present			Ahmed et al. (2009) Monci et al. (2000) Nauen et al. (2002) Stansly et al. (2005) Simón et al. (2007) Shatters et al. (2009)
-Canary Islands	Present			Simón et al. (2007) Monci et al. (2000)
Sweden	Absent, Eradicated	2014		Lemmetty and Vänninen (2014)
United Kingdom	Absent, Eradicated	2012		Powell et al. (2012) Cuthbertson et al. (2012)
North America
Bermuda	Present			McKenzie et al. (2012)
Canada	Present			CABI (Undated) McKenzie et al. (2012) Frewin et al. (2014)	Present based on regional distribution.
-British Columbia	Present			McKenzie et al. (2012)
-Ontario	Present			McKenzie et al. (2012) Frewin et al. (2014)
Costa Rica	Present			Guevara-Coto et al. (2011)
Guatemala	Present	2009		Bethke et al. (2009)
Mexico	Present	2007		Martinez-Carrillo and Brown (2007) McKenzie et al. (2012)
United States	Present		Invasive	McKenzie et al. (2012) Ahmed et al. (2009) McKenzie et al. (2009) Shatters et al. (2009)
-Alabama	Present			McKenzie et al. (2012)
-Arizona	Present			McKenzie et al. (2012)
-California	Present			McKenzie et al. (2012) Ahmed et al. (2009)
-Connecticut	Present			McKenzie et al. (2012)
-Florida	Present			McKenzie et al. (2012) Schuster et al. (2007) McKenzie et al. (2009)
-Georgia	Present			McKenzie et al. (2012) Ahmed et al. (2009)
-Hawaii	Present			McKenzie et al. (2012)
-Illinois	Present			McKenzie et al. (2012)
-Indiana	Present			McKenzie et al. (2012)
-Kentucky	Present			McKenzie et al. (2012)
-Louisiana	Present			McKenzie et al. (2012)
-Maine	Present			McKenzie et al. (2012)
-Maryland	Present			McKenzie et al. (2012)
-Massachusetts	Present			McKenzie et al. (2012)
-Michigan	Present			McKenzie et al. (2012)
-New Hampshire	Present			McKenzie et al. (2012)
-New Jersey	Present			McKenzie et al. (2012)
-New York	Present			McKenzie et al. (2012)
-North Carolina	Present			McKenzie et al. (2012)
-Ohio	Present			McKenzie et al. (2012)
-Oregon	Present			McKenzie et al. (2012)
-Pennsylvania	Present			McKenzie et al. (2012)
-South Carolina	Present			McKenzie et al. (2012)
-Texas	Present			McKenzie et al. (2012)
-Vermont	Present			McKenzie et al. (2012)
-Washington	Present			McKenzie et al. (2012)
Oceania
New Zealand	Present			Scott et al. (2007)
South America
Argentina	Present	2011		Grille et al. (2011)
Brazil	Present	2015		Barbosa et al. (2015) Bello et al. (2020)
-Rio Grande do Sul	Present			Barbosa et al. (2015)
-Sao Paulo	Present			Bello et al. (2020)
Uruguay	Present	2011		Grille et al. (2011) Arruabarrena et al. (2014) Arruabarrena et al. (2015)

History of Introduction and Spread

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The B. tabaci Mediterranean species is currently increasing its global presence. It is continually being distinguished from Middle East-Asia Minor 1 species of the B. tabaci complex around the world (Powell et al., 2012; Cavalieri et al., 2014).

Risk of Introduction

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The risk to the EPPO region is primarily to the glasshouse industry in northern countries, and mainly concerns MED species and also MEAM1 species. Present legislation relates to B. tabaci, so reference to B. argentifolii should be avoided to prevent confusion in these areas. Since the recent introduction of this whitefly to several of these countries, the pest has proved particularly difficult to combat because of its polyphagy, its resistance to many insecticides and its disruption of biological control programmes (Prabhaker et al., 1985; Della Giustina et al., 1989; Mushtaq Ahmad et al., 2002).

Countries that already contain many begomoviruses that infect indigenous plants and weeds, and are associated with localized populations of B. tabaci that exhibit narrow host ranges may be particularly at risk from MED species. Should MED species become established in these countries its polyphagous feeding activities could enable viruses to move into new susceptible host plants, causing new crop protection problems.

Hosts/Species Affected

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MED species is documented as being polyphagous, having a host range of around 600 different plant species. This includes many glasshouse and field crops, as well as weeds. However, a study by de Courcy Williams et al. (1996) indicated that only a small number of individuals within a population can readily change host plants. It is the progeny of these particular individuals that lead to the species as a whole being highly polyphagous.

Host Plants and Other Plants Affected

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Plant name	Family	Context	References
Abelmoschus esculentus (okra)	Malvaceae	Other	Hélène et al. (2015)
Acer (maples)	Aceraceae	Other
Amaranthus (amaranth)	Amaranthaceae	Other
Anacardiaceae	Anacardiaceae	Wild host
Annona	Annonaceae	Wild host
Araceae	Araceae	Wild host
Aristolochia (dutchman's pipe)	Aristolochiaceae	Wild host
Asclepias (Silkweed)	Asclepiadaceae	Wild host
Asteraceae (Plants of the daisy family)	Asteraceae	Wild host
Balsaminaceae	Balsaminaceae	Wild host
Begonia	Begoniaceae	Unknown	Shatters et al. (2009)
Bignonia	Bignoniaceae	Wild host
Bixa	Bixaceae	Wild host
Bombacaceae	Bombacaceae	Other
Boraginaceae	Boraginaceae	Wild host
Brassica	Brassicaceae	Main
Brassica oleracea var. botrytis (cauliflower)	Brassicaceae	Main
Brassica rapa subsp. pekinensis	Brassicaceae	Other
Brassicaceae (cruciferous crops)	Brassicaceae	Other
Cannabidaceae	Cannabidaceae	Wild host
Caprifoliaceae	Caprifoliaceae	Wild host
Capsicum annuum (bell pepper)	Solanaceae	Main	Ahmed et al. (2009); Hélène et al. (2015); Muñiz and Nombela (2001); Stansly et al. (2005); Stansly et al. (2005)
Capsicum frutescens (chilli)	Solanaceae	Unknown	Hélène et al. (2015); Tahiri et al. (2006)
Carica papaya (pawpaw)	Caricaceae	Main
Caryophyllaceae	Caryophyllaceae	Wild host
Catharanthus roseus (Madagascar periwinkle)	Apocynaceae	Other
Chenopodiaceae	Chenopodiaceae	Wild host
Cistacaea		Wild host
Clusiaceae	Clusiaceae	Wild host
Commelinaceae	Commelinaceae	Wild host
Convolvulaceae (Plants of the bindweed family)	Convolvulaceae	Wild host
Cucumis (melons, cucuimbers, gerkins)	Cucurbitaceae	Unknown	Shatters et al. (2009)
Cucumis melo (melon)	Cucurbitaceae	Unknown	Ben et al. (2017); Tahiri et al. (2006)
Cucumis sativus (cucumber)	Cucurbitaceae	Unknown	Ahmed et al. (2009); Hsieh et al. (2007)
Cucurbita (pumpkin)	Cucurbitaceae	Main	Ahmed et al. (2009)
Cucurbita maxima (giant pumpkin)	Cucurbitaceae	Unknown	Hélène et al. (2015); Hsieh et al. (2007)
Cucurbita moschata (pumpkin)	Cucurbitaceae	Unknown	Ahmed et al. (2012)
Cucurbita pepo (marrow)	Cucurbitaceae	Unknown	Ben et al. (2017); Tahiri et al. (2006)
Dittrichia viscosa	Asteraceae	Unknown	Tahiri et al. (2006)
Ericaceae	Ericaceae	Wild host
Euphorbia (spurges)	Euphorbiaceae	Unknown	Bethke et al. (2009)
Euphorbia pulcherrima (poinsettia)	Euphorbiaceae	Main	Ahmed et al. (2009); Chu et al. (2006); Frewin et al. (2014); Hsieh et al. (2007)
Euphorbiaceae	Euphorbiaceae	Wild host
Fabaceae (leguminous plants)	Fabaceae	Main
Fagaceae	Fagaceae	Other
Ficus	Moraceae	Unknown	Ahmed et al. (2012); Ahmed et al. (2009)
Flacourtiaceae	Flacourtiaceae	Wild host
Geraniaceae	Geraniaceae	Other
Gerbera (Barbeton daisy)	Asteraceae	Unknown	Shatters et al. (2009)
Gerbera jamesonii (African daisy)	Asteraceae	Unknown	Ahmed et al. (2009)
Gossypium (cotton)	Malvaceae	Unknown	Shatters et al. (2009)
Gossypium herbaceum (short staple cotton)	Malvaceae	Unknown	Ahmed et al. (2009)
Gossypium hirsutum (Bourbon cotton)	Malvaceae	Main	Chu et al. (2006)
Grossulariaceae	Grossulariaceae	Wild host
Helianthus annuus (sunflower)	Asteraceae	Unknown	Chu et al. (2006)
Hibiscus (rosemallows)	Malvaceae	Unknown	Ahmed et al. (2009)
Hibiscus rosa-sinensis (China-rose)	Malvaceae	Unknown	Shatters et al. (2009)
Ipomoea nil (white edge morning-glory)	Convolvulaceae	Unknown	Ahmed et al. (2009); Hsieh et al. (2007)
Lactuca (lettuce)	Asteraceae	Unknown	Bethke et al. (2009)
Lactuca sativa (lettuce)	Asteraceae	Main
Lamiaceae	Lamiaceae	Wild host
Lantana camara (lantana)	Verbenaceae	Unknown	Ben et al. (2017); Ahmed et al. (2012); Ahmed et al. (2009); Tahiri et al. (2006)
Lauraceae	Lauraceae	Other
Malva parviflora (pink cheeseweed)	Malvaceae	Unknown	Papayiannis et al. (2009)
Malvaceae	Malvaceae	Wild host
Manihot	Euphorbiaceae	Unknown	Shatters et al. (2009)
Manihot esculenta (cassava)	Euphorbiaceae	Other
Menispermum (moonseed)	Menispermaceae	Wild host
Mentha (mints)	Lamiaceae	Other
Myrtaceae		Main
Nicotiana tabacum (tobacco)	Solanaceae	Main	Shatters et al. (2009)
Nyctaginaceae	Nyctaginaceae	Wild host
Ocimum	Lamiaceae	Unknown	Ahmed et al. (2009)
Oleaceae	Oleaceae	Other
Origanum majorana (sweet marjoram)	Lamiaceae	Main
Oxalis (wood sorrels)	Oxalidaceae	Wild host
Parthenocissus tricuspidata (Boston ivy)	Vitaceae	Unknown	Ahmed et al. (2009)
Passifloraceae	Passifloraceae	Other
Pedaliaceae	Pedaliaceae	Wild host
Phaseolus vulgaris (common bean)	Fabaceae	Unknown	Ahmed et al. (2009); Papayiannis et al. (2009)
Plantaginaceae	Plantaginaceae	Wild host
Poinsettia		Unknown	Lemmetty and Vänninen (2014)
Polygonaceae	Polygonaceae	Wild host
Portulacaceae	Portulacaceae	Wild host
Proteaceae	Proteaceae	Wild host
Punicaceae	Punicaceae	Wild host
Ranunculaceae	Ranunculaceae	Wild host
Rhamnaceae	Rhamnaceae	Wild host
Rosaceae	Rosaceae	Other
Rubiaceae	Rubiaceae	Other
Rutaceae	Rutaceae	Wild host
Salicaceae	Salicaceae	Wild host
Scrophulariales		Wild host
Solanaceae	Solanaceae	Other
Solanum aethiopicum (african scarlet eggplant)	Solanaceae	Unknown	Hélène et al. (2015)
Solanum americanum	Solanaceae	Unknown	Kijima et al. (2011); Ben et al. (2017)
Solanum lycopersicum (tomato)	Solanaceae	Main	Ahmed et al. (2009); Hélène et al. (2015); Hsieh et al. (2007); Stansly et al. (2005); Tahiri et al. (2006); Ben et al. (2017); Chu et al. (2006); Götz and Winter (2016); Shatters et al. (2009)
Solanum melongena (aubergine)	Solanaceae	Main	Ahmed et al. (2009); Chu et al. (2006); Hélène et al. (2015); Hsieh et al. (2007); Tahiri et al. (2006)
Solanum tuberosum (potato)	Solanaceae	Unknown	Ahmed et al. (2012); Ahmed et al. (2009)
Sonchus oleraceus (common sowthistle)	Asteraceae	Unknown	Tahiri et al. (2006)
Sphenocleaceae	Sphenocleaceae	Wild host
Thymelaeaceae	Thymelaeaceae	Wild host
Tiliaceae	Tiliaceae	Wild host
Umbelliferae (Plants of the parsley family)	Umbelliferae	Wild host
Verbena (vervain)	Verbenaceae	Wild host
Violaceae	Violaceae	Wild host

Growth Stages

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Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

Symptoms

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Early indication of infestation may consist of chlorotic spots caused by larval feeding, which may also be disfigured by honeydew and associated sooty moulds. Leaf curling, yellowing, mosaics or yellow-veining may also indicate the presence of whitefly-transmitted viruses. These symptoms are also observed in B. tabaci infestations, however phytotoxic responses such as a severe silvering of courgette and melon leaves, mis-ripening of tomato fruits, stem whitening of brassicas and yellow veining of some solanaceous plants may also be seen (Costa et al., 1993; Secker et al., 1998).

The feeding of adults and nymphs causes chlorotic spots to appear on the surface of the leaves. Depending on the level of infestation, these spots may coalesce until the whole of the leaf is yellow, apart from the area immediately around the veins. Such leaves are later shed. The honeydew produced by the feeding of the nymphs covers the underside of leaves and can cause a reduction in photosynthetic potential when colonized by moulds. Honeydew can also disfigure flowers and, in cotton, can cause problems in lint processing. Following heavy infestations, plant height, the number of internodes, and yield quality and quantity can be affected, for example, in cotton.

Phytotoxic responses in many plant and crop species caused by larval feeding include severe silvering of courgette leaves, white stems in pumpkin, white streaking in leafy Brassica crops, uneven ripening of tomato fruits, reduced growth, yellowing and stem blanching in lettuce and kai choy (Brassica campestris) and yellow veining in carrots and honeysuckle (Lonicera) (Bedford et al., 1994a,b).

A close observation of leaf undersides will show tiny, yellow to white larval scales. In severe infestations, when the plant is shaken, numerous small and white adult whiteflies will emerge in a cloud and quickly resettle. These symptoms do not appreciably differ from those of Trialeurodes vaporariorum, the glasshouse whitefly, which is common throughout Europe.

List of Symptoms/Signs

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Sign	Life Stages	Type
Leaves / abnormal colours
Leaves / abnormal leaf fall
Leaves / abnormal patterns
Leaves / fungal growth
Leaves / honeydew or sooty mould
Leaves / necrotic areas
Leaves / wilting
Leaves / yellowed or dead
Stems / honeydew or sooty mould
Whole plant / early senescence

Biology and Ecology

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Eggs are usually laid in circular groups on the underside of leaves, with the broad end touching the surface and the long axis perpendicular to the leaf. They are anchored by a pedicel which is inserted into a fine slit made by the female in the plant tissue, and not into stomata as is the case with many other members of the Aleyrodidae. Eggs are whitish in colour when first laid, but gradually turn brown. Hatching occurs after 5-9 days at 30°C but this depends very much on host species, temperature and humidity.

On hatching, the first instar, or 'crawler', is flat, oval and scale-like in shape. The first instar is the only larval stage of this whitefly which is mobile. It moves from the egg site to a suitable feeding location on the lower surface of the leaf, after which its legs are lost in the next moult and the larva becomes sessile. It does not move again throughout the remaining nymphal stages. The first three nymphal stages each last 2-4 days, according to temperature. The fourth nymphal stage is termed the puparium, and is approximately 0.7 mm long. True pupation within the whitefly life-cycle does not occur, although the last (fourth) nymphal instar is typically referred to as a pupa after apolysis has been completed. Metamorphosis to adult occurs over about 6 days.

The adult emerges through a 'T'-shaped rupture in the skin of the puparium and spreads its wings for several minutes before beginning to powder itself with a waxy secretion from glands on the abdomen. Copulation begins 12-20 hours after emergence and takes place several times throughout the life of the adult. The lifespan of the female can extend to 60 days. The life of the male is generally much shorter, being between 9 and 17 days. Each female can oviposit over 300 eggs during her lifetime, these are often arranged in an arc around the female as she rotates on her stylet. Some 11 to 15 generations can occur within 1 year.

Virus Transmission

MED species is also responsible for vectoring many plant viruses in the genera Geminivirus, Closterovirus, Nepovirus, Carlavirus and Potyvirus. Whitefly-transmitted geminiviruses, now designated begomoviruses (Mayo and Pringle, 1998), are the most important of these agriculturally, causing yield losses to crops of between 20 and 100% (Brown and Bird, 1992; Cathrin and Ghanim, 2014). Begomoviruses cause a range of different symptoms which include yellow mosaics, yellow veining, leaf curling, stunting and vein thickening. Mansoor et al. (1993) reported that 1 million hectares of cotton were decimated in Pakistan by Cotton leaf curl virus (CLCuV) and important African subsistence crops such as cassava are affected by begomoviruses such as African cassava mosaic virus (ACMV) (see datasheet on African cassava mosaic disease). Tomato crops throughout the world are particularly susceptible to many different begomoviruses, and in most cases exhibit yellow leaf curl symptoms. This has caused their initial characterization as Tomato yellow leaf curl virus (TYLCV). A number of different species of TYCLV have now been recorded from within both the New World and Old World where MED occur (Jones, 2003). MED species has also been associated with transmission of several other begomoviruses such as Tomato mottle virus, Tobacco leaf curl virus (TLCV), Sida golden mosaic virus (SiGMV), Squash leaf curl virus (SLCV), Cotton leaf crumple virus (CLCV), Bean golden mosaic virus (BGMV) and Cotton leaf curl virus (Simon et al., 2003), some of which cause heavy yield losses in their respective hosts. Dual infections have also been shown to occur (Bedford et al., 1994c).

Natural enemies

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Natural enemy	Type	Life stages	References	Biological control in	Biological control on
Amblyseius limonicus	Predator	Larvae	Cuthbertson (2014)	UK	poinsettia plants
Amblyseius swirskii	Predator		Cuthbertson (2014)	UK	poinsettia
Amitus bennetti	Parasite
Aschersonia aleyrodes	Pathogen	Nymphs
Beauveria bassiana	Pathogen
Ceraeochrysa cubana	Predator
Chrysoperla carnea	Predator
Chrysoperla rufilabris	Predator			Texas
Delphastus pusillus	Predator
Encarsia formosa	Parasite
Encarsia nigricephala	Parasite
Encarsia pergandiella	Parasite
Encarsia strenua	Parasite
Encarsia transvena	Parasite
Eretmocerus eremicus	Parasite
Eretmocerus mundus	Parasite
Eretmocerus orientalis	Parasite
Isaria poprawskii	Pathogen		Cabanillas et al. (2013)
Nephaspis oculata	Predator
Paecilomyces fumosoroseus	Pathogen
Transeius montdorensis	Predator	Larvae	Cuthbertson (2014)	UK	poinsettia plants

Notes on Natural Enemies

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Natural enemies of whiteflies will, in most cases, have co-evolved with their prey and may, in some regions, be more efficient at controlling their native prey than an introduced one.

Various species of predatory mites have also been shown to be effective in feeding upon Mediterranean species of B. tabaci populations, including; Amblyseius limonicus, A. swirskii and Transeius montdorensis (Cuthbertson, 2014). No doubt they will also feed on other Bemisia species. A large range of natural enemies of B. tabaci have been recorded in China (Li et al., 2011). Their specificity to individual members of the B. tabaci species complex is unknown.

Entomopathogenic nematodes and fungi have been shown to offer much potential in controlling what has now been determined as MEAM1 species Bemisia populations (Cuthbertson et al., 2003a, 2005a, 2007a,b; Cuthbertson and Walters, 2005). The impact of these agents upon MED species has yet to be determined.

Means of Movement and Dispersal

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Adults do not fly very efficiently, but once airborne, can be transported long distances by convection or by wind. The greater dispersive capacity of MED species has been instrumental in its greater economic importance. All stages of this whitefly are likely to be transported within the international trade of ornamental plants and cut flowers. The international trade in poinsettia and gerbera has played a significant role in the dispersal of Bemisia (Cuthbertson, 2013) to all continents.

Natural dispersal

As with other species of whitefly and biotypes of B. tabaci, MED species can easily disperse over short distances. This dispersal can occur in vast numbers when host plants become heavily infested and begin to senesce. Large 'clouds' comprising many millions of individuals have been recorded leaving a dying host crop. Dispersal is almost certainly assisted by wind.

Agricultural practices

Physical movement of infested plants, whether it be through plant care, harvesting or spraying, can result in adult MED species dispersal from an infested plant.

Movement in trade

Any susceptible plant or crop, where leafy material is produced for distribution and export, can act as a means of dispersing MED species. Seasonal plants such as poinsettia, bedding plants, grafted crop plants and cut flowers are all potential means for MED species distribution. However, this usually involves dispersal of whitefly larvae and pupae rather than adults.

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne externally	Visibility of pest or symptoms
Flowers/Inflorescences/Cones/Calyx	adults; pupae	Yes	Pest or symptoms usually visible to the naked eye
Leaves	adults; pupae	Yes	Pest or symptoms usually visible to the naked eye
Seedlings/Micropropagated plants	eggs; larvae	Yes	Pest or symptoms not visible to the naked eye but usually visible under light microscope
Stems (above ground)/Shoots/Trunks/Branches	adults; pupae	Yes	Pest or symptoms usually visible to the naked eye

Plant parts not known to carry the pest in trade/transport
Bark
Bulbs/Tubers/Corms/Rhizomes
Fruits (inc. pods)
Growing medium accompanying plants
Roots
True seeds (inc. grain)
Wood

Impact Summary

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Category	Impact
Animal/plant collections	None
Animal/plant products	None
Biodiversity (generally)	None
Crop production	None
Cultural/amenity	None
Economic/livelihood	None
Environment (generally)	None
Fisheries / aquaculture	None
Forestry production	None
Human health	None
Livestock production	None
Native fauna	None
Native flora	None
Other	None
Rare/protected species	None
Tourism	None
Trade/international relations	None
Transport/travel	None

Impact

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Currently difficult to determine specific economic damage caused for MED species.

Environmental Impact

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The appearance of MED species within new areas is in most cases, the result of movement of infested plant material. The movement and establishment of MED species populations through this route brings along the possibility of insecticide resistance genes. This invariably leads to an increase in the use of insecticides as whitefly control becomes increasingly more difficult. This in turn can produce an ever increasing spiral in the levels of insecticide resistance and insecticide use, having a direct impact on the environment.

Detection and Inspection

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Numerous chlorotic spots develop on the leaves of affected plants, which may also be disfigured by honeydew and associated sooty moulds. Leaf curling, yellowing, mosaics or yellow veining could indicate the presence of whitefly-transmitted viruses, and phytotoxic responses such as a severe silvering of courgette and melon leaves indicate the presence of MED species, the immature stages being mainly responsible for this symptom (Costa et al., 1993). Other phytotoxic responses to themed species include mis-ripening of tomato fruits (Maynard and Cantliffe, 1989), white streaking of Brassica leaves (Brown et al., 1992) and yellow veining of some solanaceous plants (Bedford et al., 1998).

Close observation of the undersides of the leaves will show the tiny, yellow/white larval scales and in severe infestations, when the plant is shaken, numerous small, white adult whiteflies will flutter out and quickly resettle.

Similarities to Other Species/Conditions

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B. tabaci is now widely regarded to be a multi-species complex. Consisting of as many as 34 species that are morphologically indistinguishable from each other. They can however, be distinguished molecularly (De Barro et al., 2011; Tay et al., 2012; Boykin and De Barro, 2014).

Differentiation of MED species from other whitefly species on the basis of adult morphology is often difficult, although close observation of adult eye morphology may often show differences in ommatidial arrangements between some species. At rest, MED species has wings more closely pressed to the body than Trialeurodes vaporariorum, which is a larger whitefly and more triangular in appearance.

The fourth instar or puparium can also be used to distinguish MED species from T. vaporariorum as a glasshouse pest. T. vaporariorum is 'pork-pie shaped', regularly ovoid, has straight sides (viewed laterally) resulting from the vertical wax pallisade surrounding each puparium, and in most instances, 12 large setae. MED species has an irregular, 'pancake-like' oval shape, oblique sides and shorter, finer setae. Although the number of enlarged setae in the MED species and wax rods in T. vaporariorum can vary according to host plant morphology, the two caudal setae are always stout and nearly always as long as the vasiform orifice in the B biotype. The length of caudal setae can be used to distinguish some Bemisia species.

For more information on the identification of B. tabaci from slide-mounted pupae, see Martin (1987).

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Cultural Control

Intercropping practices using non-hosts have been used in many countries aiming to reduce numbers of whiteflies on specific crops. However, intercropping with susceptible crops can promote whitefly populations, by offering a greater leaf area for feeding.

Weed species can play an important role in harbouring whiteflies between crop plantings and attention should be paid to removing these in advance of planting susceptible crops. Weeds also often harbour whitefly-transmitted viruses (Bedford et al., 1998) and may be a major source of crop virus epidemics, especially where MED species is present, due to its polyphagous nature.

Cultural control is generally much more effective where whiteflies are physical pests rather than virus vectors.

Host-Plant Resistance

The development of transgenic resistant plant and crop species through genetic engineering must be considered and accepted as a future method of control where whitefly-transmitted viruses are already endemic and causing severe crop losses (Wilson, 1993; Raman and Altman, 1994). Traditional sources of resistance have been used successfully for the control of other whitefly species.

Chemical Control

The following active ingredients have been reported as effective in controlling MED species worldwide: bifenthrin, buprofezin, imidacloprid, fenpropathrin, amitraz, fenoxycarb, deltamethrin, azadirachtin, pymetrozine. Chemical resistance is an ever increasing concern within the B. tabaci species complex. The MED species is widely considered to evolve stable resistance to the neonicotinoid insecticides commonly used for Bemisia control more rapidly than MEAM1 (Nauen, 2005). Hence, MED species neonicotinoid resistance is becoming increasingly widespread and problematic, with numerous cases being reported worldwide (Horowitz et al., 2005; Fernandez et al., 2009; Dennehy et al., 2010; Luo et al., 2010; Wang et al., 2010). However sequential chemical treatments have been shown under laboratory conditions to offer excellent potential in eradicating MED species on poinsettia plants (Cuthbertson et al., 2012).

A rotation of insecticides that offer no cross-resistance must therefore be used to control infestations.

A new group of environmentally safer insecticides that effectively kill whitefly by a physical mode of action are appearing on the market in many countries (Cuthbertson and Collins, 2015). These products do not have a specific active ingredient, but appear to utilise surfactant-like properties to overcome the protective waxes on whitefly larvae and adults.

IPM

The increased fecundity and polyphagous habit of MED species has exacerbated many control problems in field and glasshouse crops worldwide, compounded by insecticide resistance. It appears that no single control treatment can be used on a long-term basis against this pest, and that approaches should be integrated to achieve an effective level of control.

IPM appears to offer the best option for controlling MED species infestations without causing contamination of the environment. Beneficial insects are used alongside chemicals that offer a high level of selectivity, such as insect growth regulators. However, if whitefly-transmitted viruses are present, it is unlikely that the threshold of whitefly vectors would ever be reduced to a level where virus transmission would cease by using these methods, because MEAM1 is such an efficient viral vector. Plant and crop species that exhibit a high level of resistance to both vector and virus must also be considered when designing an IPM system.

Entomopathogenic nematodes and fungi have been shown to be successfully tank-mixed with several chemical products for use in eradication programmes against what has now been deemed as MEAM1 species in the UK (Cuthbertson et al., 2003b, 2005b, 2007b). Their impact upon MED species populations has not been fully determined. Cuthbertson et al. (2012) devised a sequential treatment programme that successfully eradicated MED species on poinsettia plants under laboratory conditions using both chemicals and entomopathogenic fungi.

Phytosanitary Measures

In countries where MED species is not already present, the enforcement of strict phytosanitary regulations as required for B. tabaci in general, may help to reduce the risk of this whitefly becoming established (Cuthbertson and Vänninen, 2015).

Because of the difficulty of detecting low levels of infestation in consignments, it is best to ensure that the place of production is free from the pest (OEPP/EPPO, 1990). Particular attention is needed for consignments from countries where certain B. tabaci-listed viruses, now on the EPPO A1 or A2 quarantine lists, are present. These viruses can also be transmitted by MED species.

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