Doctoral seminar- II

(ENT- 692)

INSECT BIOTYPES

Submitted to: Course Instructor: Dr. A. L. Narangalkar Head, Dept. of Agril. Entomology

Submitted by: H. R. Sawai Ph.D. Scholar (Regd. No. 126)

DEPARTMENT OF AGRICULTURAL ENTOMOLOGY COLLEGE OF AGRICULTURE, DAPOLI DR. BALASAHEB SAWANT KONKAN KRISHI VIDYAPEETH, DAPOLI - 415712, DIST - RATNAGIRI, (MS)

DEPARTMENT OF AGRICULTURAL ENTOMOLOGY COLLEGE OF AGRICULTURE, DR. BSKKV, DAPOLI

Name Degree course No. Course instructor : : : : H. R. Sawai Ph. D. ENT-692 Dr. A. L. Narangalkar Regd No. Discipline Course title Date : : : : 126 Agril.Entomology Doctoral Seminar - II 15-05-2012

Abstact - Insect Biotypes

The concept of insect biotypes has been around for 100 years, having been first touched upon by Walsh (1864). First biotype was recognized by Painter 1930 who referred to them as biological strain. The existence of biotype discovered by Painter 1930 on Hessian fly, Mayetiola destructor on resistance varieties of winter wheat. Biotype is known to occur in 36 arthropod species belonging to 17 families of five orders. Aphids constitute about 50 % of the spp. with known biotype (Dhaliwal and Arora 2006). The continuous growing of insect resistant variety may lead to certain physiological and behavioral changes in insect pest. So that, they are capable of feeding and developing on resistant varieties (kogan, 1998). The biotypes are more prone in rice, wheat, soybean, vegetables crops. The main causes for occurrence of biotypes viz., genetic changes in insect pests, continuous growing of insect resistant varieties, single gene resistance and mutation. Biotype development is, one of the major constraints encountered in breeding programmers for varietal resistance (Dhaliwal and Arora 2006). Q biotype whitefly reported from Queensland Australia and it is severe on vegetable crops but differs from B biotypes as Q biotype does not induce the physiological changes normally as B biotype, also it has the ability to develop resistance quickly to some insecticide groups, particularly if they are used repeatedly (Fancelli, and Vendramim, 2002). In 2010 the soybean aphid biotypes 3 have been identified by university of Illinois which can multiply on aphid resistant soybean varieties consisting Rag1 and Rag-2 resistant gene (Hill et al. 2010). Presently in rice five biotypes of BPH, six in rice gall midge in India, seven in Russian wheat aphid have occurred. For prevention of the insect biotype development systematic surveillance programme, adaptation of IPM, injudicious use of insecticides, sequential cultivar release, maintenance of refugia, and use of tolerant varieties and phytosanitary measures are to be utilized. References Dhaliwal, G.S. and Arora, R. (2006). Integrated Pest Management: Concepts and Approaches. Ludhiana, Kalyani Publishers. Pp135-137. Fancelli, M., and Vendramim, J. D. (2002). Development of Bemisia tabaci Gennadius biotype B on Lycopersicon Spp. Scientia Agricola, 59: Pp.665-669. Hill, C.B., Herman, T. K., Voegtlin, D. J. and Hartman, G. L. (2010). A New Soybean Aphid (Hemiptera: Kogan, M. (1998). Integrated pest management: Historical perspectives and contemporary developments. A. Rev. Ent, 43: 243-270. Aphididae) Biotype Identified. J. Econ. Entomol., 103(2): 509- 515

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www.planthealthaustrelia.com (Q biotypes whitefly)

History of insect biotype

The concept of insect biotypes has been around for 100 year, having been first touched upon by Walsh (1864).The existence of biotype discovered by painter 1930 on Hessian fly Mayetiola destructor on resistance varieties of winter wheat. First biotypes were recognized by Painter 1930 who referred to them as biological strain. Occurrences of gall midge biotype in India were suspected by Khan and Murthy (1955) even when no resistance varieties were developed (Benture et al 2003). Biotype development in Corn leaf aphid, Rhapalosiphum maidis was first detected by Cartier and Painter 1956 was observation in population. Carties (1959) demonstrated the existance of three Acyrthisiphon pisium biotype on peas in field and green house.First biotype alfalfa aphid Theroaphis maculata was identified by Pesho et al. 1960 Arizona. Eastop (1973) reviewed the concept of biotype with particular reference to aphid and suggested that the term was synonymous with clone. First reports of a newly evolved biotype of B. tabaci, the B biotype, appeared in the mid-1980s (Brown et al., 1995). Commonly referred to as the silverleaf whitefly or Poinsettia strain. Diehl and Bush ( 1984) have presented an evolutionary and applied perspectives of insect biotype. Saxena and Barrion (1987) have listed over 35 spp. of insect pest Agriculture importance for which biotype has been reported. Willam and Shambaaugh (1988) used biological studies to demonstrate that two Phylloxera biotype exited the clinton biotype unable to feed on concord and six other grape spp. and concord biotype unable to feed on clinton and nine other spp. of grape. A detailed analysis on gall midge biotypes in India has been made by Bentur et al in 2003.March 2005, University of California and Arizona researchers identify Biotype Q on poinsettias. University of Illinois researchers (May 2010) recently identified a new soybean aphid biotype that can multiply on aphid-resistant soybean varieties.

Introduction
The continuous growing of insect resistant variety may lead to certain physiological and behavioral changes in insect pest. So that they are capable of feeding and developing on resistant varieties.The term biotype is generally used to describe a population of insects capable of damaging and surviving on plants previously known to be resistant to other populations of the same species (Kogan, 1994). Biotype refers to the populations within a species which can survive Page17

on and destroy varieties that have gene for resistance (Heinrichs et al., 1985). Bohssini et al. (2001) insect biotype studies genotypic interaction between resistance in plant and virulence gene in insect. The challenge for the host-plant resistance strategy is the constant development of new biotypes that can overcome the resistance of deployed genes (Ratcliffe et al. 2000).In these cases, an insect biotype is considered to be a population of insects that is able to survive, reproduce on, and/or cause injury to a cultivated plant that is resistant to other populations of the same species. Biotypes are morphologically similar with normal insect types but they are physiologically differing from them.Biotypes are convenient and useful designations for applied problems in agricultural pest management, especially involving host plant resistance. First, biotypes are intraspecic categories. Second, biotypes are usually morphologically indistinguishable. Third, bio- types differ in expressed biological attributes (Claridge and et al 1983). Biotype development is, one of the major constraints encountered in breeding programmes for varietal resistance. (Dhaliwal and Arora, 2005). Biotypes are developed more on varieties having more biochemical defense than the varieties offering physical defense. The development of insect biotype has based has posed a serious threat to the success of plant resistance for the management of insect pests. Aphids constitute about 50 % of the spp. with known biotype (Dhaliwal and Arora. 2005) because of parthenogenesis. Although the occurrence of biotype among insect is comparatively less frequent than in plant pathogen (Dhaliwal and Arora, 2005).

Causes of insect biotype

1. Many factors are associated with the ability of an insect to overcome plant resistance. 2. Biotype are known to develops on varieties where antibiosis (biochemical defense) is the major component of resistance and rarely develop on varieties where antixenosis or tolerance is mechanism of resistance 3. The continuous growing of insect-resistant varieties may lead to development of biotype. 4. Biotypes of the insects are evolving as a result of selection pressure exerted by large scale growing of resistant cultivars (Kindler, 1999; Naber, 2000). Due to a high degree of host specificity, the insect may evolve into a resistance breaking biotype. Page17 The widespread use of one resistance gene is decreasing the genetic diversity of a host. As a result some insect species will break the resistance gene (Heinrichs, 2001). Use of multiple resistance genes conferring different causes of resistance.

Puterka and Burton (1990) reasoned that biotype developed from 1. Mutation or a Pre-existing variability of virulence: Even a single mutant aphid capable of feeding on a resistant genotype can build up into a new biotype (Pathak, 1970). Insect biotypes are strains of the pest insect that mutate to express virulence genes that overcome resistance. 2. Biotypes of the insects are evolving as a result of selection pressure exerted by large scale growing of resistant cultivars. Resistance due to antibiosis will put high pressure on biotype development. Biotype are known to develops on varieties where antibiosis (biochemical defense)is the major component of resistance. 3. Resistance due to Antixenosis will put little pressure on biotype development. Biotypes are known to rarely develop on varieties where antixenosis is mechanism of resistance. 4. Complete tolerance will put no pressure on biotype development. Biotype not develops on varieties where tolerance is mechanism of resistance

Example of insect biotype

Biotype are known to be occurs in 36 arthropod species belonging to 17 families of six order (Saxena and Barrion, 1987). Aphids constitute about 50 % of the spp. with known biotype. Ten of 18 are aphid species in which parthenogenic reproduction contribute greatly successfully development of biotype (Smith, 2004). 1. Coleoptera Species Callosobruchus maculatus Phaedonia inclusa Sitophilius oryzae Common name Cowpea weevil Soyabean weevil Rice weevil Family Bruchidae Crop Cowpea Biotype recent reference 1 Shade et al. (1996) 1 2

Cuculionidae Cuculionidae

Soyabeen Corn wheat

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2. Diptera

Species Agromyza oryzae Mayetiola destructor Orseolia oryzae Chlorops oryzae Rhagolitis cerasi 3. Lepidoptera Species Ostrinia nubilalis Laspeyresia pomonella Yponomeuta padells 4. Thysanoptera Species Thrips tabaci

Common name Rice leaf miner Hessian fly Asian rice gall midge Rice stem maggot Cherry Fruit fly

Family Agromyzidae Cecidomyiidae Cecidomyiidae Chloropidae Tephritidae

Crop Rice Wheat, Barley Rice Rice Cherry

Biotype no. Recent reference 2 14 Bohissini et al. 2001 Naber, 2000 Indian -6 Chinese-4 Katitar et al. 2000 2 2

Common name European corn borer Walnut Coddling Moth Small ermine moth

Family Pyralidae Tortricidae Yponomeutidae

Crop Corn Apple, Walnut Apple, Walnut

Biotype no. Recent reference 4 3 1

Common name Onion thrips

Family Thripidae

Crop Tobbaco

Biotype no. Recent reference 2

5. Homoptera

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Species Bemisia tabacci Acyrthosiphon pisum Aphis kondi Amphoraphora rubi A. idaei Aphis craccivora Aphis fabae Aphis gossypii Aphis nasturtii Aulacorthum solani Brevicoryne brassicae Chaetosiphon fragaefoli Dysaphis directa D. plantaginea Eriosoma lanigerum Macrisiphum euphorbiae Myzus persicae Rhopalosiphum maidis Rhopalosiphum padi Schizaphis graminum Therioaphis maculata Diuraphis

Common name Whitefly Pea aphid Aphid Raspbery Aphid Cowpea aphid Bean aphid Cotton aphid Buckhorn aphid Green spotted potato aphid Cabbage Aphid Strawberry aphid Apple aphid Wooly apple aphid Potato aphid Green peach aphid Corn leaf aphid Oat-bird cherry aphid Green bug Spotted alfalfa aphid Russian wheat

Family Aleyrodide Aphididae Aphididae Aphididae Aphididae Aphididae Aphididae Aphidida Aphididae Aphididae Aphididae Aphididae Aphididae Aphididae Aphididae Aphididae

Crop

Biotype no. Recent reference

Cotton, 9 Okra,Cassava, Cervera et al. 2000 9 Alfalfa Zarrabi, 1995 1 Zarrabi et al 1995 6 Beeries Panda and Khush 1995, Jones, 2000 Groundnut 2 Bush sitao 5 Broad bean 2 Pepper 2 Potato Potato Cabbage Strawberry Apple Apple Potato Potato Barley, Corn, Sorghum Cherry Barley, oat, sorghum, wheat Alfalfa Wheat 2 1 2 2 3 Morris, 1998 3 Young, 1982 2 3 5 5, 2 Panda and Khush, 1995 2 11 Porter, 2000 6 Panda and Khush, 1995 Zsuzsa, 2001

Aphididae Aphididae Aphididae Aphididae

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noxia Mueleerianella farmairei Nephotettix virescens Saisstia oleae Nilaparvata lugens Lepidosahes ulmi Daktulosphaira Vitrifoliae

aphid Leaf hopper Green leaf hopper Black scale Brown plant hopper Oystershell scale Grapr phylloxera

Cicadellidae Cicadellidae Coccidae Delphacidae Diaspididae Phylloxeridae

Rice Melon Rice Apple Grapevine

1 3(lab) Panda and khush, 1995 1 5 Heinrichs, 2001; Huang, 2001 2 2

Insects biotypes known in different crop pest systems

Rice In brown plant hoppers five different biotypes have been reported from Indian. Most of the biotypes having morphological variations among biotypes such as rostrum, legs and antennae got modified both in males and females

BPH Resistance Varieties Released The host resistance of rice against BPH was first reported for the variety Mudgo in 1969. Following gene inserted in different variety of rice for development of resistance of brown plant hopper.

Biotypes Biotypes 1 & 2 Biotype 3 Biotype 4

Location Southeast Asia Philippines Indian subcontinent Page17

BPH Resistance Varieties Released

BPH Resistance Varieties IR26, IR 1561-228-3 , Mudgo, TKM6 IR36, IR38 ,ASD 7, IR1154-243 IR50, IR60 , Rathu Heenati Babawee ARC10550 Swarnalatha T12 Chin Saba Kharamana, Balamwee and Pokkali Introgression line of O. australiensis

BPH Resistance Gene Bph 1 gene Bph 1 gene Bph 3 gene Bph4 bph5 (Khush et al., 1985) Bph6 bph7 bph8 Bph9 Bph10 (Kabir and Khush, 1988) (Kabir and Khush, 1988) (Nemoto et al., 1989) (Ikeda, 1985) (Jena and Khush, 1990).

Resistance and Susceptible to Insect Biotype The following table shows that the different resistance varieties of rice develop biotypes. Gene Bph3,bph4 Bph5, Bph6, Bph7 Bph8, Bph9,Bph 10 Resistance and susceptible to insect biotype Resistant to all four biotype Susceptible to biotypes 1,2,3, Resistant to biotype 4 Resistant to biotypes1, 2,3

Scheme for selection of new Biotypes of Nilaparvata lugens Insect biotype in rice gall midge (GM) Biotype Biotype 1 Biotype 2 Biotype 3 Biotype 4 Response of Cultivar Resistance IR26,ASD7,Rathu Heenati, Babawee Rathu Heenati, Babawee, ASD7 IR26, Rathu Heenati, Babawe Babawee(Coimbatore) RathuHeenati,Babawee (Bangladesh and hyderabad) Susceptible TN1 IR26,TN1 ASD7,TN1 IR26,ASD7,Rathu Heenati (Coimbatore) IR26,TN1, ASD7 (Bangladesh and Hyderabad Page17

The losses of rice due to rice gall midge are about 477 thousand tons of grain or 0.8% of total production (Rs 330 Crores) in India. The occurrence of biotype in India suspected by Khan and Murthy in 1955.The Biotype developed in rice gall midge in Indian-6 and Chinese-4 (Katiyar

et al 2000).Genetic studies have identified, so far, 10 major genes conferring resistance (Kumar et al., 2005). Most of the 60 plus gall midge resistant rice varieties developed to date contain one of the three major genes viz., Gm1, Gm2 and an unidentified gene(s) in Ptb21 conferring immune level of resistance. Six distinct gall midge biotypes have been characterized so far the reaction of biotype among different group of cultivars of rice against rice gall midge is follows. Biotype in rice gall midge Biotype 1 2 3 4 5 6 Location Andhra Pradesh, Tamil Nadu, Chhattisgarh, Madhya Pradesh Orissa, Maharashtra, Karnataka Bihar, Manipur Andhra Pradesh, Maharashtra (Sakoli) Kerala Manipur

Important biotype in whitefly Bemisia tabaci Common names of whitefly are Tobacco whitefly, Sweet potato whitefly, Cotton whitefly.The host of whitefly are Cassava, Cotton, Sweet potatoes, Tobacco, Tomatoes ,Capsicum, Cucurbita pepo, Cucumbers , Hibiscus, Gerbera, lettuces, poinsettia, soybeans, green beans, cabbage, broccoli, cauliflower. Bemisia tabaci (Gennadius), is one of the most devastating pest insects of agriculture and horticulture in the world (Denholm et al. 1996) but it is also an efficient vector of a great many plant viruses (Brown 2007). The pest status of this insect is further complicated by the recognition of more than 20 distinct strains or biotypes worldwide(Brown et al. 1995; Perring 2001).Of these, the B(silverleaf whitefly or poinsettia strain) and Q biotypes represent the greatest threat to growers. First reports of a newly evolved biotype of B. tabaci, the B biotype, appeared in the mid1980s (Brown et al., 1995) B biotype has been shown to be highly polyphagous. The B biotype is able to cause phytotoxic disorders in certain plant species, e.g. silverleaf in squashes (Cucurbita sp.) Although indistinguishable in appearance from silverleaf whitefly (biotype B), these insects are much less susceptible to insect growth regulators and many neonicotinoid insecticides. The Q biotype was detected in the United States in 2004 on greenhouse poinsettia plants (Dennehy et al. Page17

2005). Q-biotype populations are generally less susceptible to many insecticides currently used in this country, including pyriproxifen (Distance), buprofezin (Talus, Applaud), imidacloprid (Marathon), thiamethoxam (Flagship), and acetamiprid (Tristar). Important biotype in soyabean aphid Rhamnus cathartica L.(buckthorn) is the primary or overwintering host ofA.glycines on which sexual reproduction occurs; how-ever, eggs can be laid on Rhamnus alnifolia L. (Voegtlin et al. 2004). Soybean aphids are the No. 1 insect threat to soybean production in the North Central region of the United States.Soybean is the most important secondary or summer host of A. glycines (Hill et al. 2004). Soybean aphid has the ability to transmit plant viruses to soybean such as Alfalfa mosaic virus, Soybean dwarf virus, and Soybean mosaic virus (Hartman et al. 2001, Hill et al. 2001, Clark and Perry 2002, Wang and Ghabrial 2002, Domier et al. 2003). Earlier study identifying a soybean aphid biotype that could colonize plants with the Rag1 resistance geneThis gives the pest a high potential to adapt to and reduce the effective life of resistance genes deployed in production." When farmers plant aphid-resistant soybean varieties, they provide protection against Biotype 1. However, recent research indicates that Biotype 2, which was first discovered in 2006, can overcome some aphid-resistant varieties University of Illinois researchers recently identified a new soybean aphid biotype that can multiply on aphidresistant soybean varieties The most recently identified soybean aphid, Biotype 3, was discovered in Springfield Fen, Ind., on an overwintering host, glossy buckthorn They found it was capable of feeding and multiplying on varieties carrying the resistance genes Rag1 and Rag2. Biotype 3 was capable of feeding and multiplying on varieties carrying the resistance genes Rag1 and Rag2 in soybean.Other resistance genes against soybean aphid Rag 3 Rag 4, are currently on the breeding pipelines for incorporation into elite varieties to suppress biotype 3 Prevention of Insect Biotype Development Systemic surveillance programme should be designed for monitoring the insect biotype development. (Ratcliffe, 2001). Adaptation of integrated pest management techniques likes inters- cropping mixed Page17 cropping.

Reduction of the alate phase will slow down the development of the insect towards a resistance breaking biotype. Use of insect growth regulator: Example-Buprofezin inhibits acetylcholinesterase activity in B-biotype Bemisia tabaci (Cottage and Gunning, 2006). Use good non-chemical controls: Remove older leaves, keep areas free from weeds, segregate infested from non-infested plants, discard heavily infested material. E.g. Q biotype of white flies (Sanderson et al 2005). Insecticides for rotation include Azatin, Insecticidal soap, Horticultural oil, Botanigard etc.

No single control treatment can be used on a long-term basis against this pest.Example: There have also been good results with applications of Orthene followed several days later by a pyrethroid. Give good result to control Q and B biotype of white fly. (Sanderson et al 2005).

Maintenance of Refugia: Provide harborage for susceptible moth production to reduce the chance of resistant (R).To support avirulent individuals to mate with virulent individuals in the insect population.

Use of susceptible cultivars for at least 20% of the growing area to offer

refugees to the main

avirulent biotype in the insect population. (Sloderbeck, 1997 and Kerlin, 2002). 1. To preserve and promote the activity of parasites and predators.Avoid the use of broad-spectrum insecticides such as pyrethroids, organophosphates, Oraganochlorine. 2. Increasing the number of miner gene in a variety may enhance the level of horizontal resistance and its stability. 1. Miner gene may be combined with major gene. Page17

2. Gene rotating: Use cultivars with one gene in one season, then a different resistance gene the next (gene rotation).To minimize selection pressure on given resistance varieties on insect population may be reduced. 3. DNA marker technology is currently used to map and identify specific gene structures conferring resistance traits in plants. Understanding the genetics of resistance in plants will provide the knowledge to improve resistance deployment strategies. 4. Use of tolerant cultivars that tolerant varieties would not only check yield loss caused by insects, but this character combined with major genes. 5. Crop multiline: Different resistance genes in different plants of the same crop within a single field of area 6. Sequential cultivar release: Use until failure, switch to next gene or when varieties with a major gene become susceptible due to selection for new biotype another a variety with a new major gene for resistance is released. 7. Gene pyramiding combines multiple resistance genes (against one pest) in the hybrid or cultivar OR two or more major gene for vertical resistance is incorporated into a variety to impart resistance to more biotype. Since monogenic resistance is generally more vulnerable to biotype development than polygenic resistance. 8. Phytosanitary risk Very few countries remain free from B. tabaci, illustrating the difficulty of preventing its movement in international trade. Emergence of the B biotype of B. tabaci, with its ability to feed on many different host plants has given whitefly-transmitted viruses the potential to infect new plant species.(Tobacco leaf curl, transmissible tomato yellow leaf curl bean golden mosaic, squash leaf curl. Particular attention is needed from countries where certain B. tabaci isted viruses. Conclusion Insect biotypes are main limitation of host plant resistance. An average period of 5-8 years is required for development of variety; however, it has taken less than 10 years for breakdown of resistance in the field. Insect biotypes are more developed in Homopteran insect (50% in aphids). Developed more on varieties having more biochemical defense than the varieties offering physical defense. Monogenic resistance is generally more vulnerable to biotype development than polygenic resistance. Page17

References Cervera, M, Pina J. A, Jurez, J, Navarro L, Pea, L. (2000). A broad exploration of a transgenic population of citrus: stability of gene expression and phenotype. Theor. Appl. Genet. 100: 670-677. Claridge, K. L. ( 1983). The biotype concept and its application to insect pests of agriculture. Crop Protection. 2: 85-95. Dhaliwal, G.S. and Ram Singh, (2004). Host plant resistance to insect: Concepts and Applications. New Delhi Panima Publishing Corporation. Pp.141-187. Dhaliwal, G.S. and Ramesh Arora (2006). Integrated Pest Management: Concepts and Approaches. Ludhiana, Kalyani Publishers. Pp.135-137.
Fancelli, M. and Vendramim, J. D. (2002). Development of Bemisia tabaci Gennadius biotype B on Lycopersicon Spp. Scientia Agricola, 59: Pp.665-669.

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(Hemiptera: Aphididae) Biotype Identified. J. Econ. Entomol., 103(2): 509- 515

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Jones, A. T., Mcgavin, W. J. and Birch, A. N. E., 2000, Effectiveness of resistance genes to the large raspberry aphid, Amphorophora idaei Brner, in different raspberry (Rubus idaeus L.) genotypes and under different environmental conditions. Annal of Appl. Biol. 136: 107-113. Kabir, M.A., and G.S. Khush. 1988. Genetic analysis of resistance to brown planthopper in rice (Oryza sativa L). Plant Breed. 100:5458. Katiyar, S.K. Tan, Y. Huang, B. Chandel, G. Zhang, Y. and Bennett, J. (2001). Molecular mapping of gene Gm-6(t) which confers resistance against four biotypes of Asian rice gall midge in China. Theor Appl Genet, 103: 953961 Khush, G. S. (1985). Rice varietal improvement. Int.Rice Commn. Newsl. 34 (2):111-125.
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and Caddel. J. L. (1995). New biotype of Acyrthosiphon

kondoi (Homoptera: Aphididae) on alfalfa in Oklahoma. J. Econ. Entomol ., 88:14611465. Zsuzsa B., Keith. Hopper, R, Jordan, J, Saayman, T, (2001). Biotype differences in Russian wheat aphid (Diuraphisnoxia) between South African and Hungarian agro-ecosystems. Agric. Ecosystems & Environment, 83: 121128

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