Sitobion miscanthi
(indian grain aphid)

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Identity

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Preferred Scientific Name

• Sitobion miscanthi (Takahashi, 1921)

Preferred Common Name

• indian grain aphid

Other Scientific Names

• Macrosiphum eleusines Theobald, 1929
• Macrosiphum miscanthi Takahashi, 1921
• Sitobion eleusines Theobald, 1929

International Common Names

• English: grain aphid

Summary of Invasiveness

Top of page S. miscanthi is an exotic, invasive species in Australia and New Zealand. The alate (winged) forms have the potential for long-distance dispersal. It appears that S. miscanthi first colonized New Zealand as airborne alatae from Australia (Close and Tomlinson, 1975) and population outbreaks have significant plant biosecurity implications (Teulon and Stufkens, 2002).

Once introduced, populations of S. miscanthi have the potential to increase rapidly. Reproduction is by parthenogenesis, so there is no need for females to locate males in order to reproduce. Therefore, when populations are initially low following introduction, the reproduction rate is high. The generation time is short and dispersal via alatae can be rapid. There is also no need for a primary woody host (as occurs for host-alternating aphids) and reproduction can occur all year round on abundant grass species. S. miscanthi colonizes cereal crops from wild grasses, pasture grasses and other cereals.

S. miscanthi populations in Australia and New Zealand have limited gene pools compared to populations in Asia (Turak and Hales, 1994; Wilson et al., 1999). These populations initially arose from a limited number of introduced females. Introduced populations can show founder effects, which lead them to vary in certain characteristics from the species in its native range. The different chromosomal races of S. miscanthi present in Australia have different temperature preferences, growth rates and seasonal abundance (Turak et al., 1998).

Taxonomic Tree

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• Domain: Eukaryota
•     Kingdom: Metazoa
•         Phylum: Arthropoda
•             Subphylum: Uniramia
•                 Class: Insecta
•                     Order: Hemiptera
•                         Suborder: Sternorrhyncha
•                             Unknown: Aphidoidea
•                                 Family: Aphididae
•                                     Genus: Sitobion
•                                         Species: Sitobion miscanthi

Notes on Taxonomy and Nomenclature

Top of page Takahashi (1921) first described S. miscanthi as Macrosiphum miscanthi. It was subsequently reclassified as Macrosiphum (Sitobion) miscanthi (Eastop, 1966) but is now placed within the aphid genus Sitobion Mordvilko, in the tribe Macrosiphini of the subfamily Aphidinae. The main synonym in the literature is Macrosiphum (Sitobion) eleusines (Eastop and Hille Ris Lambers, 1976).

Description

Top of page S. miscanthi is almost exclusively anholocyclic with no sexual forms (oviparae and males) or egg stage. Reproduction is parthenogenetic and viviparous. Populations consist of nymphs, adult apterae (wingless) and adult alatae (winged) females. Four nymphal instars are normal, but it has been suggested that five instars are possible (Sekhar and Singh, 2001b). The nymphs are morphologically similar to the adult apterae, although they are smaller. S. miscanthi is a medium-sized and variably coloured aphid, with relatively long tubular siphunculi, five- or six-segmented antennae that are longer than the body length, and a tapering tongue-shaped cauda (David, 1975; Ghosh, 1975; Blackman and Eastop, 2000).
Apterous Adults

The adult apterae are broadly spindle-shaped. The body is green or reddish-brown to dark-brown, with abdominal tergites variably tanned from pale to very dark. The siphunculi are darker than the body, shiny black or dark-brown, imbricated, with a subapical zone of polygonal reticulation. The cauda is relatively pale. The body length is 1.7-3.0 mm and six to seven times the length of the siphunculi. The siphunculi are 1.4-1.9 times longer than the cauda. The hind tarsus segment II is 1.0-1.3 times longer than the last segment of the rostrum. The processus terminalis is 4.5-6.0 times longer than the base of the last antennal segment (Blackman and Eastop, 2000).

Alate Adults

The alatae are similar in colour to the apterae but with larger and darker dorsal abdominal markings. The alatae body length is 1.9-3.1 mm (Blackman and Eastop, 2000).

In northern India, S. miscanthi is usually smaller than in East Asia, with an adult apterous body length that is typically 1.25-2.50 mm (David, 1975).

Distribution

Top of page S. miscanthi is Asiatic in origin and is widespread in India, China and the Far East and in the Pacific region. It has been introduced into Australia and New Zealand, and possibly into several Pacific islands (Blackman and Eastop, 2000).

The distribution map includes records based on specimens of S. miscanthi from the collection in the Natural History Museum (London, UK): dates of collection are noted in the List of countries (NHM, various dates).

Sitobion akebiae may be a synonym of S. miscanthi, and the taxonomy of the S. miscanthi/akebiae groups requires further clarification. At least some of the records from Poaceae probably apply to S. miscanthi (see CABI/EPPO, 2003).

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.

History of Introduction and Spread

Top of page S. miscanthi was accidentally introduced into Australia and New Zealand. The date of introduction is unknown although Hales et al. (1998) considered that it had probably arrived in Australia from Asia before European settlement. A study of DNA microsatellite and karyotype variation in Australian populations revealed that very few successful colonizations initially occurred, from which all introduced populations are derived. There was no evidence of sexual reproduction, so diversification is by mutation and chromosomal rearrangement alone (Sunnucks et al., 1996).

S. miscanthi subsequently spread from Australia to New Zealand where it was initially found in small numbers on native grasses but not on cereals (Lowe, 1969). From 1957 to 1965, alate (winged) S. miscanthi had only been detected in aphid traps in one of 8 years (Lowe, 1968). The first outbreaks on wheat occurred in Canterbury, New Zealand and nearby coastal regions in October 1967. This closely followed severe outbreaks around Melbourne, Victoria, Australia. The meteorological conditions were favourable on two occasions at this time for the airborne movement of aphids in weather systems from Victoria to New Zealand (Tomlinson, 1973). The evidence strongly suggests that large numbers of alatae were almost simultaneously deposited over a wide area on one (or both) of these occasions. The wheat crop in Melbourne was maturing in October 1967, leading to the production of large numbers of winged migrants, whereas infestations around Canterbury occurred in random patches indicating that the foci of infection were due to settling migrants. These infestations were consistent with the trajectory of air movements over the Tasman Sea (Close and Tomlinson, 1975).

Since 1967, S. miscanthi has become economically significant on cereals in New Zealand. Hybridization may have occurred between introduced populations of S. miscanthi and Sitobion near fragariae, although sexual morphs have not so far been found in the field (Hales et al., 1998; Blackman and Eastop, 2000).

Risk of Introduction

Top of page Around 90% of aphid species present in New Zealand are exotic. They entered via two possible pathways: on infested plant material and produce or by wind dispersal. Although S. miscanthi is capable of being transported on weather systems across water bodies, entry on plant material is the main route for exotic aphids into New Zealand. It is not a post-harvest pest, and leaves and growing inflorescences are not usually traded for cereals. However, this aphid could be present on imported exotic and ornamental grasses. Given that windborne dispersal is likely to predominate, further invasion is difficult to prevent using phytosanitary measures. However, knowledge of weather systems, aerial traps and outbreaks in southern Australia could be used to predict further airborne introductions (Close and Tomlinson, 1975; Teulon and Stufkens, 2002).

Habitat

Top of page S. miscanthi can persist throughout the year in unmanaged grasslands. The aphids can migrate from natural grasslands to infest cereals.

Hosts/Species Affected

Top of page S. miscanthi is polyphagous on many members of Gramineae/Poaceae. It can also sometimes be found on dicotyledons: for example, semi-aquatic species such as Polygonum hydropiper in Australia (Eastop, 1966; Blackman and Eastop, 2000).

Host Plants and Other Plants Affected

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Growth Stages

Top of page Flowering stage

Symptoms

Top of page Cereal aphids can reduce yields of wheat, barley, oats and other crops without producing obvious, visible symptoms. Moreover, S. miscanthi usually occurs within a complex of aphid species on cereals. Therefore, there are no clearly distinctive symptoms attributable to S. miscanthi. However, with heavy infestations, yellowing is often discernible on the leaves and earheads (inflorescences) of cereals and the leaves of pasture grasses.

List of Symptoms/Signs

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Biology and Ecology

Top of page Genetics

The chromosome number for S. miscanthi is 2n=18 in India (Kurl and Chauhan, 1988). However, chromosome number varies from 2n=17 to 2n=21 for introduced populations in Australia and New Zealand (Sunnucks et al., 1996). There are four known chromosomal races of S. miscanthi in Australia. Three of these (2n=17, 2n=18 and 2n=20) are believed to have evolved from a recent common ancestor by mutation and chromosomal rearrangement. Significant differences in relative growth rate, developmental time and adult weight were found for laboratory-reared clones of different chromosomal races (Sunnucks et al., 1998). There is a suggestion that hybridization may have occurred between introduced populations of S. miscanthi and Sitobion near fragariae in New Zealand (Hales et al., 1998).

Physiology and Phenology

In eastern Australia, seasonal differences in abundance occur between the chromosomal races of S. miscanthi and the closely related Sitobion near fragariae. Sunnucks et al. (1998) found significant differences in relative growth rate, developmental time and adult weight in laboratory-reared clones of different chromosomal races.

Reproductive Biology

S. miscanthi is anholocyclic throughout its geographic range. Sexual forms have been described on rare occasions: an oviparae (sexual female) collected on Polygonum chinense in India may be S. miscanthi, whereas a pinkish-brown alate male collected from Poa annua in India has been classified as S. miscanthi (David, 1975). In Japan and Korea, Blackman and Eastop (2000) suggested that Sitobion akebiae may be a synonym, although S. akebiae is holocyclic (lays overwintering eggs) and the taxonomy of the miscanthi/akebiae group requires further clarification. The evidence for holocycly remains inconclusive (Blackman and Eastop, 2000).

Reproduction is by parthenogenesis throughout its range, with both apterous (wingless) and alate (winged) females giving birth to live young. The apterae are adapted to exploit the host plants on which they develop, whereas alatae are adapted for dispersal over long distances and are responsible for the initial colonization of cereals and pasture grasses. Rapid reproduction can occur under favourable conditions, leading to population outbreaks. The alates are produced in response to overcrowding or when the plants have become nutritionally unsuitable.

S. miscanthi is monoecious (non host-alternating), having lost the need for a primary woody host. It lives throughout the year on a wide range of grasses and cereals. The apterae usually live on aerial plant parts.


Environmental Requirements

Seasonal differences between chromosomal races of S. miscanthi corresponded to temperature preferences in laboratory studies. S. miscanthi 2n=18 showed evidence of adaptation to warmer conditions, having the highest population growth around 25°C and being the only race able to reproduce above 28°C. S. miscanthi 2n=17 had an intermediate temperature preference (Turak et al., 1998). Sitobion near fragariae showed adaptations to cooler conditions, with the highest population growth between 12 and 20°C and this is consistent with it being more common earlier in the spring.

S. miscanthi favours warmer conditions than other cereal aphids. It replaces Sitobion avenae in importance on cereals in tropical regions. S. miscanthi is adversely affected by cold winters, an effect compounded by its anholocyclic life cycle, i.e. overwintering is done as adult apterae rather than through a more cold-tolerant egg stage (Dixon, 1987). Aphid populations in northern India decline in response to heavy winter rainfall and hail. Simulated rainfall (1-2 cm) in wheat led to 95% mortality of S. miscanthi. Mortality was greater for nymphs rather than adults, and being greater before earing rather than after. However, sudden high temperatures also cause mortality (Sandhu and Deol, 1975). High temperatures, acting in conjunction with natural enemies and grain maturity, for instance, led to a population decline on wheat in the Punjab (Grewal and Bain, 1975).

Associations

S. miscanthi often occurs within a cereal aphid species complex. In India, it is often found with Rhopalosiphum padi and Rhopalosiphum maidis on wheat and barley (Grewal and Bain, 1975). These three species often have distinctive feeding sites. R. padi infests all above-ground plant parts, R. maidis is confined to leaf whorls and S. miscanthi infests leaves and earheads (Sekhar and Singh, 1999). In Australia, S. miscanthi is associated with R. padi, R. maidis and also S. near fragariae on wheat (Turak et al., 1998). S. near fragariae, in common with S. miscanthi, is an introduced cereal aphid that reproduces parthenogenetically in Australia and has populations that derive from a few initial colonizations (Sunnucks et al., 1996).

Natural enemies

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Notes on Natural Enemies

Top of page Natural enemies contribute to the biological control of S. miscanthi in cereal crops. Ahmad and Singh (1995, 1997) and Das and Chakrabarti (1989) described parasitoids of S. miscanthi in India, with the braconid wasp, Aphidius uzbekistanicus being the most important.

A range of generalist aphid predators also attack S. miscanthi. In northern India, coccinellids, syrphids and Leucopis sp. contribute to population declines in wheat (Grewal and Bain, 1975; Sandhu and Deol, 1975; Sandhu and Kaushal, 1975). Similarly, the coccinellid beetle Hippodamia variegata controls S. miscanthi in barley in Himachal Pradesh, India (Hameed et al., 1975b).

Two entomophagous fungal pathogens, Entomophthora aphidis and Entomophthora planchoniana, were recorded on S. miscanthi from wheat in New Zealand (Lowe and Hall, 1978).

Means of Movement and Dispersal

Top of page S. miscanthi is mainly dispersed through alate (winged) forms being carried in weather systems over long distances. For example, the pest may have been introduced into New Zealand from Australia by weather systems across the Tasman Sea (Close and Tomlinson, 1975). However, apterae may also be dispersed on exotic, ornamental species of grasses. It is unlikely that stages of S. miscanthi would be transported with grain or grain products.

Plant Trade

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Impact Summary

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Impact

Top of page S. miscanthi outbreaks can result in significant economic impacts in cereals. The aphids cause direct feeding damage and indirect damage, through the secretion of honeydew (on which sooty moulds grow) and via the spread of plant pathogenic viruses.

S. miscanthi is a vector of barley yellow dwarf luteovirus (BYDV) and millet red leaf persistent luteovirus (MRLV) (Blackman and Eastop, 2000). It transmits BYDV in a persistent manner. BYDV is most serious when plants are infected early in the season. In Australia, S. miscanthi is a minor vector of BYDV in pasture grasses and cereals. BYDV has been estimated as causing annual production losses in Australian cereals of US$ 40 million, with similar losses in pasture grasses (Hales et al., 1990).

During direct feeding, nutrients, amino acids and carbohydrates are extracted from leaves and earheads, while some plant physiological processes may be disrupted. Honeydew and sooty moulds interfere with light capturing by green tissues and reduce photosynthetic efficiency. Damage is dependent on the number of tillers infested, the number of aphids per tiller and the duration of infestation. The resulting yield loss can be quantified in terms of a reduced number of earheads, reduced number of grains per head or reduced seed weight. Maximum yield losses in cereals are most likely to occur because of attack between ear emergence and flowering. S. miscanthi feeds on leaves, moving to the earheads as they develop.

The crop most affected by S. miscanthi is wheat. S. miscanthi is a major pest of wheat, for instance, in northern India. Its pest status increased during the 1970s in India following the introduction of high-yielding Mexican varieties and the increased use of fertilizers and irrigation. S. miscanthi is most numerous during February and March, with peak populations coinciding with the heading phase of wheat (Grewal and Bain, 1975; Sandhu and Deol, 1975). In a study in the Punjab, India it was estimated that populations of 1-20, 21-40 and 41-60 aphids per earhead reduced the thousand-grain weight by 4.7%, 13.2% and 26.4%, respectively. S. miscanthi was the dominant aphid on wheat recorded in this study, but Rhopalosiphum maidis was also present (Grewal and Bain, 1975).

S. miscanthi can cause significant losses in barley, oats, rye and sorghum (Aggarwal and Hameed, 1972; Hameed et al., 1975a). It is a minor pest of rice (Yano et al., 1983) and a range of other cereal crops.

Detection and Inspection

Top of page The presence of winged migrants (alatae) of S. miscanthi can be detected on a regional scale using suction traps and on a local scale using yellow sticky traps within crops. The detection of migrants in spring can be used to formulate aphid control strategies in cereals.

Within crops, small to moderate colonies of aphids can be found on upper leaves and earheads during routine inspections of cereals.

Similarities to Other Species/Conditions

Top of page A number of aphids cause similar damage to cereals and grasses as S. miscanthi. S. miscanthi can be distinguished from Sitobion avenae by its longer siphunculi (more than 1.4 times longer than the cauda cf. less than 1.4 times longer in S. avenae) and shorter hind tarsi (the hind tarsus segment II is less than 1.3 times longer than the last segment of the rostrum cf. more than 1.3 times longer in S. avenae). S. miscanthi differs from Sitobion fragariae by having shorter siphunculi (1.4-1.9 times longer than the cauda cf. 1.8-2.3 times longer in S. fragariae) and a cauda that is more pointed at the apex. S. miscanthi can be distinguished from Sitobion africanum, on maize, millet, sorghum and tropical pasture grasses, by its variable and ill-defined abdominal pigmentation (cf. well-defined black bars in S. africanum) and longer hairs (over 20 µm) on antennal segment III (Blackman and Eastop, 2000).

Turak and Hales (1994) described an allozyme method for separating the morphologically similar Sitobion species complex on grasses in Australia. S. miscanthi occasionally colonizes bamboo in northern India, where it can be distinguished from Sitobion bambusicola due to its shorter siphunculi (1.4-1.9 times the length of the cauda) compared to S. bambusicola (2.0-2.1 times the length of the cauda) (Blackman and Eastop, 1994).

Prevention and Control

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Host-plant Resistance

The use of resistant varieties of wheat and other cereals can reduce aphid infestations and yield loss. Most of the work on host-plant resistance in cereals has concentrated on other aphid species, such as Schizaphis graminum and Sitobion avenae, rather than S. miscanthi. However, resistant varieties are often effective against cereal aphid species complexes, which may include S. miscanthi. Studies in India have evaluated the impact of S. miscanthi on different wheat varieties (Saikia et al., 1998; Sekhar et al., 2001).

Biological Control

Native parasitoids and predators play a role in controlling outbreaks of S. miscanthi in cereals. The parasitoid Aphidius uzbekistanicus has potential as a biological control agent (Das and Chakrabarti, 1989).

Chemical Control

A range of insecticides have been used against cereal aphids where S. miscanthi is one of the species present, including bifenthrin, cypermethrin, deltamethrin, dimethoate, and pirimicarb. A number of insecticides have been evaluated specifically for use against S. miscanthi on wheat in India and Pakistan (Sekhar and Singh, 2001a; Khan and Maqbool, 2002). Out of 10 insecticides tested in India, fenvalerate was the most effective against S. miscanthi and Rhopalosiphum padi (Sekhar and Singh, 2001a).

Early Warning Systems

The use of suction traps to monitor migrants (alatae) provides a warning of potential outbreaks.

Field Monitoring/Economic Threshold Levels

Monitoring for migrant aphids in suction traps and counting apterous aphids on plants can aid in the forecasting of pest incidence and in determining the necessity and timing of insecticide applications.

Integrated Pest Management

Checking for the presence of aphids on tillers can aid in spraying decisions. The use of aphid-specific insecticides is usually advocated because natural enemies (e.g. parasitoids, ladybirds and syrphid larvae) are helpful in preventing secondary outbreaks of aphids.

References

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Aggarwal VD; Hameed SF, 1972. Occurrence of the Indian grain aphid, Macrosiphum (Sitobion) miscanthi Takahashi on barley crop in Kulu Valley, Himachal Pradesh. Indian Journal of Entomology, 34(1):82

Ahmad ME; Singh R, 1995. Survey of parasitoids of aphids in northeastern Uttar Pradesh for possible use in biological control. Annals of Entomology, 13(2):87-96.

Ahmad ME; Singh R, 1997. Records of aphids and their food plants, parasitoids and hyperparasitoids from North Bihar. Journal of Advanced Zoology, 18(1):54-61.

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CABI/EPPO, 2003. Sitobion miscanthi. Distribution Maps of Plant Pests, No. 651. Wallingford, UK: CAB International.

Calilung VJ, 1969. A host index of Philippine aphids. Philippine Entomologist, 1(3):209-223.

Carver M; Hart PJ; Wellings PW, 1993. Aphids (Hemiptera: Aphididae) and associated biota from The Kingdom of Tonga, with respect to biological control. Pan-Pacific Entomologist, 69(3):250-260.

Close RC; Tomlinson AI, 1975. Dispersal of the grain aphid Macrosiphum miscanthi from Australia to New Zealand. The New Zealand Entomologist, 6(1):62-65.

Das BC; Chakrabarti S, 1989. Aphidiid parasitoids (Hymenoptera: Aphidiidae) of graminaceous aphids in Garhwal, Western Himalaya. Oriental Insects, 23:365-372

David SK, 1975. A taxonomic review of Macrosiphum (Homoptera: Aphididae) in India. Oriental Insects, 9(4):461-493

Dixon AFG, 1987. Cereal aphids as an applied problem. Agricultural Zoology Reviews, 2:1-57

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Grewal TS; Bain SS, 1975. The role of abiotic and biotic factors in the population build-up of wheat aphids and the extent of loss caused by them. Indian Journal of Ecology, 2(2):139-145.

Hales DF; Chapman RL; Lardner RM; Cowen R; Turak E, 1990. Aphids of the genus Sitobion occurring on grasses in southern Australia. Journal of the Australian Entomological Society, 29(1):19-25

Hales DF; Sunnucks P; Wilson ACC, 1998. Sitobion in the South Seas - microsatellite revelations. Aphids in natural and managed ecosystems. Proceedings of the Fifth International Symposium on Aphids, Leon, Spain, 15-19 September, 1997., 69-75; 10 ref.

Hameed SF; Sud VK; Giamzo SP, 1975. New records of aphids from Kulu and Lahaul valleys (Himachal Pradesh). Indian Journal of Entomology, 37(2):203-205

Hameed SF; Sud VK; Kashyap NP, 1975. Adonia variegata (Goeze) (Coccinellidae: Coleoptera), an important predator of the Indian grain aphid, Macrosiphum (Sitobion) miscanthi Tak. in Kulu Valley (Himachal Pradesh). Indian Journal of Entomology, 37(2):209-210

Hsieh JS; Pi CP, 1988. Diallel analysis of resistance to aphid in sorghum. Journal of the Agricultural Association of China, No. 142:67-84

Khan SM; Maqbool R, 2002. Varietal performance of wheat (Triticum aestivum) against wheat aphid (Macrosiphum miscanthi) and its chemical control with different doses of insecticides. Asian Journal of Plant Sciences, 1(2):205-207.

Kurl SP; Chauhan RS, 1987. Chromosome numbers of Indian aphids: their possible evolution and taxonomic significance. Journal of Aphidology, 1(1-2):70-77

Lowe AD, 1968. Alate aphids trapped over eight years at two sites in Canterbury, New Zealand. New Zealand Journal of Agricultural Research, 11:829-848.

Lowe AD, 1969. A preliminary account of Macrosiphum miscanthi Takahashi on grasses and cereals in New Zealand. The New Zealand Entomologist, 4(2):33-35.

Lowe AD; Hall IM, 1978. Additional records of Entomophthora aphidis and E. planchoniana from aphids in New Zealand. New Zealand Journal of Zoology, 5(3):493

Ma Jian; Shen ZuoRui; Jin XiaoHua; Xie AiTing, 2008. Dispersing flight of the winged aphid Macrosiphum (Sitobion) miscanthi within the wheat field. Chinese Bulletin of Entomology, 45(2):214-218.

Naumann-Etienne K; Remaudière G, 1995. A commented preliminary checklist of the aphids (Homoptera: Aphididae) of Pakistan and their host plants. Parasitica, 51(1):61 pp.; 82 ref.

Saharia D, 1981. Biology of Coccinella repanda Thunb. an important predator of mustard aphid Lipaphis erysimi (Kltb.) in Assam. Journal of Research, Assam Agricultural University, 2(1):66-73

Saikia SK; Dutta SK; Saikia DK; Devroy TC, 1998. Reproductive parameters of Indian grain aphid Sitobion miscanthi (Tak.) on wheat varieties. Journal of the Agricultural Science Society of North East India, 11(1):66-69; 10 ref.

Sandhu GS; Deol GS, 1975. New records of pest on wheat. Indian Journal of Entomology, 37(1):85-86

Sandhu GS; Kaushal KK, 1975. Biological studies and host range of Leucopis sp. (Chamaemyiidae: Diptera), a predatory fly on aphids in the Punjab. Indian Journal of Entomology, 37(2):185-187

Sekhar SMV; Singh VS, 1999. Incidence and species composition of aphids infesting wheat. Indian Journal of Entomology, 61(3):291-295; 18 ref.

Sekhar SMV; Singh VS, 2001. Bioefficacy and cost effectiveness of different insecticides against aphids infesting wheat. Indian Journal of Entomology, 63(3):249-254; 14 ref.

Sekhar SMV; Singh VS, 2001. Biology of aphids infesting wheat. Indian Journal of Entomology, 63(3):244-248; 8 ref.

Sekhar SMV; Singh VS; Tomar SMS, 2001. Resistance to foliage feeding aphids in wheat. Indian Journal of Entomology, 63(4):377-380; 5 ref.

Sunnucks P; Chisholm D; Turak E; Hales DF, 1998. Evolution of an ecological trait in parthenogenetic Sitobion aphids. Heredity, 81(6):638-647; 30 ref.

Sunnucks P; England PR; Taylor AC; Hales DF, 1996. Microsatellite and chromosome evolution of parthenogenetic Sitobion aphids in Australia. Genetics, 144(2):747-756; 43 ref.

Takahashi R, 1921. Aphididae of Formosa. Part 1. Taihoku, Taiwan: Government Research Institute, Taiwan Department of Agriculture, pp. 8-9.

Tao CC, 1999. List of Aphidoidea (Hemiptera) of China. Taiwan Agricultural Research Institute, Special Publication No. 77. Taichung, Taiwan: Taiwan Agricultural Research Institute.

Teulon DAJ; Stufkens MAW, 2002. Biosecurity and aphids in New Zealand. New Zealand Plant Protection Volume 55, 2002. Proceedings of a conference, Centra Hotel, Rotorua, New Zealand, 13-15 August 2002, 12-17; 21 ref.

Thackray DJ; Ward LT; Thomas-Carroll ML; Jones RAC, 2005. Role of winter-active aphids spreading Barley yellow dwarf virus in decreasing wheat yields in a Mediterranean-type environment. Australian Journal of Agricultural Research, 56(10):1089-1099. http://www.publish.csiro.au/journals/ajar

Tomlinson AI, 1973. Meteorological aspects of trans-Tasman insect dispersal. New Zealand Entomologist, 5(3/4):253-268

Turak E; Hales DF, 1994. An allozyme method for identifying individual aphids of morphologically similar taxa (Hemiptera: Aphididae). Journal of the Australian Entomological Society, 33(1):57-59

Turak E; Talent R; Sunnucks P; Hales DF, 1998. Different responses to temperature in three closely-related sympatric cereal aphids. Entomologia Experimentalis et Applicata, 86(1):49-58; 21 ref.

van Harten A; Ilharco FA; Remaudière G, 1977. Some Aphidoidea of French Polynesia. [Sur quelques Aphidoidea de la Polynésie française.] Bulletin de la Societe Entomologique de France, 82(5-6):150-155.

Wang YongMo; Shen ZuoRui; Gao LingWang; Zhang GuoLiang, 2008. Seasonal genetic structure in Beijing populations of the grain aphid Sitobion miscanthi (Takahashi): an investigation using microsatellites. Applied Entomology and Zoology, 43(1):97-103. http://odokon.ac.affrc.go.jp/

Wilson ACC; Sunnucks P; Hales DF, 1999. Microevolution, low clonal diversity and genetic affinities of parthenogenetic Sitobion aphids in New Zealand. Molecular Ecology, 8(10):1655-1666; 2 pp. of ref.

Yano K; Miyake T; Eastop VF, 1983. The biology and economic importance of rice aphids (Hemiptera: Aphididae): a review. Bulletin of Entomological Research, 73(4):539-566

Distribution Maps

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