Diuraphis noxia (Russian wheat aphid)
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- Risk of Introduction
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Species Vectored
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Impact Summary
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Diuraphis noxia (Kurdjumov, 1913)
Preferred Common Name
- Russian wheat aphid
Other Scientific Names
- Brachycolus noxia Mordvilko
- Brachycolus noxius Mordvilko
- Brachysiphoniella noxius (Mordvilko ex Kurdjumov)
- Cavahyalopterus graminearium Mimeur
- Cavahyalopterus noxius (Mordvilko)
- Cuernavaca noxia (Mordvilko)
- Cuernavaca noxius
- Diuraphis muehlei (Borner)
- Diuraphis noxius (Mordvilko ex Kurdjumov)
- Holcaphis noxius (Mordvilko ex Kurdjumov)
- Quernavaea noxia
International Common Names
- English: barley aphid; Russian grain aphid
Local Common Names
- Germany: Russische Getreide-Blattlaus; Russische Weizen-Blattlaus
- BRAYNO (Diuraphis noxia)
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Hemiptera
- Suborder: Sternorrhyncha
- Unknown: Aphidoidea
- Family: Aphididae
- Genus: Diuraphis
- Species: Diuraphis noxia
Notes on Taxonomy and NomenclatureTop of page
D. noxia was first recognized as a separate species by Mordvilko (Kurdjumov, 1913), which he named Brachycolus noxius. Since then it has been described under various names by Aizenberg (1935), Mimeur (1942), Bodenheimer and Swirsky (1957) and Anon. (1963). Eastop and Hille Ris Lambers (1976), Blackman and Eastop (1984) and Stoetzel (1987) all place it in the genus Diuraphis Aizenburg.
DescriptionTop of page
D. noxia is a yellow-green to grey-green spindle-shaped aphid, 1.4-2.6 mm long as an adult. Although its integument is slick just after moult, it soon becomes covered with a waxy white exudate. The eyes and distal third of the antennae are dark; the head and thorax of winged adults are also dark. It has six-segmented antennae about half the length of the body. The rostrum reaches to about the middle coxae. The cornicles are pale and very truncated (50-60 µm long), being about as wide as long, so that they are inconspicuous. The cauda is elongate, and above the cauda is a supracaudal process on the 8th abdominal tergite, which gives it the appearance of having two caudae and distinguishes it from all other cereal aphid species. The supracaudal process is almost as long as the cauda on apterous adults, but it is smaller and less conspicuous on alates and nymphs.
Blackman and Eastop (2000), Stoetzel (1987), Pike et al. (1990) and Olsen et al. (1993) give descriptions and keys to D. noxia and other cereal aphids. Aalbersberg et al. (1987) and Olsen et al. (1993) provide keys to the immature stages.
DistributionTop of page
Although found in Xinjiang-Uigur Autonomous Region of the Peoples Republic of China for about 70 years, D. noxia has not spread to the major wheat-growing area of central China, but there is concern that it may do so in the future (Zhang, 1991). D. noxia has recently been found in Chile and Argentina but has not yet become a pest (Reed and Kindler, 1994). D. noxia has been found in Kenya where it is considered a pest, and biological control is being investigated.
D. noxia spread to 17 western states in the USA and three western provinces in Canada within 3 years of its introduction, it has not spread eastward in the USA or Canada (Webster and Amosson, 1994). D. noxia has not spread much into central or northern Europe.
D. noxia has been detected in South Australia for the first time where it has been found in cereal crops in the Mid-North (Government of South Australia, 2016). There is also a preliminary report of the pest in Victoria (IPPC, 2016).
A record of D. noxia in the UK (EPPO, 2006) published in previous versions of the Compendium is now considered erroneous and has been removed.
Distribution TableTop of page
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.
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|Afghanistan||Present||CABI, 1991; EPPO, 2014|
|Iran||Present||CABI, 1991; EPPO, 2014|
|Iraq||Present||Ali et al., 1985; EPPO, 2014|
|Israel||Present||CABI, 1991; Hopper et al., 1997; EPPO, 2014|
|Jordan||Present||Tanigoshi et al., 1995; EPPO, 2014|
|Kazakhstan||Present||Gruber et al., 1991; Hopper et al., 1997; EPPO, 2014|
|Kyrgyzstan||Widespread||Gruber et al., 1991; Hopper et al., 1997|
|Saudi Arabia||Present||Alhag et al., 1996|
|Turkey||Present||CABI, 1991; Gruber et al., 1991; EPPO, 2014|
|Uzbekistan||Widespread||Hopper et al., 1997|
|Yemen||Present||Stary and Erdelen, 1982; EPPO, 2014|
|Algeria||Present||CABI, 1991; Miller et al., 1993; EPPO, 2014|
|Egypt||Present||CABI, 1991; EPPO, 2014|
|Ethiopia||Present||Haile and Megenasa, 1987; EPPO, 2014|
|Kenya||Present||Introduced||1995||Invasive||Macharia et al., 1999; IPPC-Secretariat, 2005|
|Libya||Present||CABI, 1991; EPPO, 2014|
|Morocco||Present||Tanigoshi et al., 1995; Hopper et al., 1997; EPPO, 2014|
|South Africa||Present||Introduced||1978||Walters et al., 1980; EPPO, 2014|
|Canada||Restricted distribution||EPPO, 2014|
|-Alberta||Present||Jones et al., 1989; Butts, 1992; EPPO, 2014|
|-British Columbia||Present||Jones et al., 1989; EPPO, 2014|
|-Saskatchewan||Present||Jones et al., 1989; Butts, 1992; EPPO, 2014|
|Mexico||Present||Gilchrist et al., 1984; EPPO, 2014|
|USA||Restricted distribution||EPPO, 2014|
|-Arizona||Widespread||Webster and Amosson, 1994|
|-California||Widespread||Webster and Amosson, 1994|
|-Colorado||Present||Webster and Amosson, 1994; EPPO, 2014|
|-Idaho||Widespread||Webster and Amosson, 1994|
|-Kansas||Present||Webster and Amosson, 1994; EPPO, 2014|
|-Montana||Present||Webster and Amosson, 1994; EPPO, 2014|
|-Nebraska||Present||Webster and Amosson, 1994; EPPO, 2014|
|-Nevada||Widespread||Webster and Amosson, 1994|
|-New Mexico||Present||Webster and Amosson, 1994; EPPO, 2014|
|-North Dakota||Restricted distribution||Webster and Amosson, 1994|
|-Oklahoma||Present||Webster and Amosson, 1994; EPPO, 2014|
|-Oregon||Widespread||Webster and Amosson, 1994|
|-South Dakota||Restricted distribution||Webster and Amosson, 1994|
|-Texas||Present||Introduced||1986||Webster and Amosson, 1994; EPPO, 2014|
|-Utah||Widespread||Webster and Amosson, 1994|
|-Washington||Present||Webster and Amosson, 1994; EPPO, 2014|
|-Wyoming||Present||Webster and Amosson, 1994; EPPO, 2014|
|Argentina||Present||Introduced||1991||Reed and Kindler, 1994; EPPO, 2014|
|Chile||Present||Introduced||1987||Reed and Kindler, 1994; EPPO, 2014|
|Bulgaria||Present||Gruber et al., 1991|
|Croatia||Present||Culjak and Barcic, 2002|
|Czech Republic||Present||Stary, 1996|
|France||Present||Gruber et al., 1991; Hopper et al., 1997; EPPO, 2014|
|Germany||Present||Thieme et al., 2001|
|Greece||Widespread||Hopper et al., 1997; EPPO, 2014|
|Hungary||Present||Basky, 1993; EPPO, 2014|
|Italy||Present||Hopper et al., 1997; EPPO, 2014|
|Macedonia||Widespread||Hopper et al., 1997|
|Moldova||Widespread||Gruber et al., 1991|
|-Central Russia||Widespread||Grossheim, 1914|
|-Russia (Europe)||Widespread||Grossheim, 1914; Kovalev et al., 1991|
|Slovakia||Present||Barta and Cagan, 2005; Barta and Cagan, 2007; Barta and Cagan, 2007|
|UK||Absent, invalid record||EPPO, 2010; EPPO, 2014|
|Ukraine||Present||Gruber et al., 1991; EPPO, 2014|
|Yugoslavia (Serbia and Montenegro)||Present||Hopper et al., 1997|
|-South Australia||Present||Introduced||Invasive||Government of South Australia, 2016; IPPC, 2016||In the Mid-North, south of Tarlee|
|-Victoria||Restricted distribution||IPPC, 2016||Preliminary report.|
Risk of IntroductionTop of page
D. noxia was found in Chile in 1987 and in Argentina in 1991, and it is spreading rapidly (Reed and Kindler, 1994). Although it has not yet become a pest in Argentina and Chile, its rapid spread suggests that it may soon become one. D. noxia has not yet been found in several other major wheat-growing areas, such as Australia and central China. However, Hughes and Maywald (1990), using a climate-matching model, predicted potentially severe infestation of Australian cereals if D. noxia were introduced.
Although nymphs and adults are very unlikely to be transported alive on harvested grains or fodder, overwintering eggs could be transported in this way. However, D. noxia is anholocyclic in North America so far, despite the discovery of a few oviparae (Kiriac et al., 1990). Although it is possible that the sexual phase was lost after colonization, it appears more likely that the founders were also anholocyclic.
Habitat ListTop of page
Hosts/Species AffectedTop of page
Kindler and Springer (1989) found that D. noxia could develop on 47 out of 48 cool-season grass species and 18 out of 32 warm-season grass species to which it was exposed in the greenhouse. They found that it did not develop on any of 27 legume species or 17 forb species to which it was exposed. Pike and Allison (1991) compiled a list of host plants from the literature, comprising 140 grass species with varying degrees of suitability for D. noxia reproduction. Zea mays does not appear suitable for D. noxia reproduction, and Sorghum bicolor is only marginally suitable (Webster et al., 1987). Thus it appears that D. noxia is restricted to grasses. Furthermore, it reproduces best and does most damage on cool-season grasses.
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage
SymptomsTop of page
Although D. noxia feeds on leaves and flowers/seedheads of grasses, it appears to inject a polypeptide toxin that affects the entire plant (Hewitt et al., 1984). Colonies are found most frequently on the youngest leaves or on newly emerged flowers/seedheads. Even very small colonies of D. noxia cause characteristic white to purple streaking and leaf-rolling on wheat and barley leaves. The streaks usually extend most of the long axis of the leaf and are irregularly distributed across the short axis of the leaf. Rolling extends in severity from simple folding of the leaf along the mid-vein, to one side of the leaf rolled in upon itself, to the whole leaf being tightly rolled around the aphid colony. Large colonies can roll the flag leaf to the point where the tip of the inflorescence becomes trapped, giving it a fish-hook shape. Young plants are often stunted and even killed. Plants attacked after flowering show few to no obvious symptoms. Duration of infestation may have more impact than aphid density on yield loss (Burd and Burton, 1992; Kieckhefer and Gellner, 1992).
List of Symptoms/SignsTop of page
|Inflorescence / discoloration panicle|
|Leaves / abnormal colours|
|Leaves / leaves rolled or folded|
|Leaves / necrotic areas|
Biology and EcologyTop of page
Hughes (1988, 1996) and Araya et al. (1990) review the literature on D. noxia, and Poprawski et al. (1992) give a complete bibliography from 1886 to 1992.
D. noxia is multivoltine with a development time from birth to reproduction of 11 days at 20°C. With a daily fecundity of 1-3 nymphs per day and adult longevity of about 80 days in the laboratory, D. noxia has a high potential capacity for increase with rm = 0.06/day for a temperature regime of 5-15°C (Michels and Behle, 1989). Aalbersberg et al. (1987) and Kieckhefer and Elliott (1989) found somewhat similar values for development rate, survival, and reproduction, although differences in techniques make comparisons problematic.
Plant growth stage, temperature, and their interaction affect D. noxia life span and reproduction (Girma et al., 1990). Reproduction is greatest at 18-21°C on wheat in growth stages from jointing to heading. D. noxia develops faster, lives longer, produces more progeny, and thus has a higher rate of increase on wheat than on rye (Behle and Michels, 1990). D. noxia from wheat have higher levels of reproduction when shifted to less suitable host plants, than those grown for several generations on the less suitable plants (Schotzko and Smith, 1991; Worrall and Scott, 1991; Robinson, 1993).
In the northern part of the USA and in Canada, overwintering mortality can be 100% if temperatures are low enough for long enough (Butts, 1992; Armstrong et al., 1992). For D. noxia, supercooling point is weakly if at all related to ability to survive cold winters; most winter mortality occurs at temperatures well above the supercooling point (Butts, 1992). Winter survival can depend on the details of aspect and snow cover so that within a field, survival can be high on the south-facing sides of furrows receiving greater solar insolation (Hammon and Peairs, 1992).
The geographical range of D. noxia (particularly the areas where it is a pest) is restricted to regions of low rainfall. Furthermore, even in areas with low rainfall, D. noxia is rarely a problem in irrigated cereals, and populations decline after heavy rainfall. These observations suggest that precipitation and/or humidity may directly or indirectly reduce survival or reproduction of D. noxia.
Alates are formed when plants are water stressed, not when aphids are crowded (Baugh and Phillips, 1991). Messina (1993) found only 15-35% of variation in alate production was explained by crowding, and prenatal experience has a strong effect on alate formation. D. noxia density has little effect on per capita growth rate (Messina, 1993). The rapid spread of D. noxia across the western USA and its ability rapidly to infest newly germinated wheat suggest a high dispersal rate. However, quantitative measurements are lacking, and it seems likely that it was present in many areas at low numbers before being detected, so that its rate of spread may have been less than it would appear.
Wheat and barley can provide suitable habitat for 9-11 months of the year, but wild grasses are very important for persistence and growth of D. noxia populations (Kriel et al., 1986; Aalbersberg et al., 1988a; Armstrong et al., 1991; Montandon et al., 1993). D. noxia can oversummer on a variety of wild grass species (Kindler and Springer, 1989; Clement et al., 1990; Messina et al., 1993b), which thus provide a bridge for infestation of autumn-planted cereals.
In Eurasia, some populations have both sexual and asexual generations (i.e. are holocyclic) with sexual forms and egg laying in the fall so that the aphids overwinter as eggs (Grossheim, 1914; Basky, 1993). However, in other European populations (e.g. in southern France) and in all populations in the USA only asexual reproduction has been found (i.e. populations are anholocyclic), although a few ovipariae have been observed in USA populations (Kiriac et al., 1990). Besides differences in life-cycle among populations, D. noxia populations vary in the damage done to host plants (Bush et al., 1989; Puterka et al., 1992, 1993), in fecundity (Webster et al., 1993a), and in randomly amplified polymorphic DNA markers (Black et al., 1992; Puterka et al., 1993).
Despite initial indications that D. noxia could vector diseases such as Barley yellow dwarf virus (Rybicki and von Wechmar, 1982; von Wechmar and Rybicki, 1984), subsequent experiments showed little if any persistent transmission of viruses (Kriel et al., 1986; Fouche et al., 1984; Hewitt et al., 1984; Halbert et al., 1992; Damsteegt et al., 1992).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Adalia bipunctata||Predator||Adults/Larvae/Nymphs||South Africa||wheat|
|aphid lethal paralysis virus||Pathogen||Adults/Nymphs|
|Coccinella septempunctata||Predator||Adults/Nymphs||North America; USSR||wheat|
|Coccinella transversoguttata biinterupta||Predator||Adults/Eggs/Larvae/Nymphs/Pupae|
|Diaeretiella rapae||Parasite||Adults/Nymphs||USSR; Washington||wheat|
|Ephedrus plagiator||Parasite||Adults/Nymphs||USSR; Washington||wheat|
|Hippodamia convergens||Predator||Adults/Nymphs||South Africa||wheat|
|Hippodamia undecimnotata||Predator||Adults/Nymphs||North America|
|Hippodamia variegata||Predator||Adults/Nymphs||North America|
|Oenopia conglobata||Predator||Adults/Nymphs||North America|
|Orius niger compressicornis||Predator||Adults/Eggs/Larvae/Nymphs/Pupae||USSR||wheat|
|Praon sp. nr callaphis||Parasite||Adults/Eggs/Larvae/Nymphs/Pupae|
|Propylea quatuordecimpunctata||Predator||Adults/Nymphs||North America; USSR||wheat|
|Scymnus frontalis||Predator||Adults/Nymphs||North America|
|Sphaerophoria sp. nr annulipes||Predator||Adults/Eggs/Larvae/Nymphs/Pupae|
Notes on Natural EnemiesTop of page
Coccinellids were found associated with D. noxia throughout its Eurasian range (Hopper et al., 1997). Coccinella septempunctata was the species most frequently found in association with D. noxia, Hippodamia variegata the next most frequent, and Propylea quattuordecimpunctata the next most frequent (Hopper et al., 1997). Although some of the other species, for example Scymnus spp., are smaller and thus may be more able to attack D. noxia in rolled leaves (Kauffman and LaRoche, 1994), they were much rarer during exploration for introductions into USA. Yu and Liang (1998) described 17 species of coccinellid attacking D. noxia in Xinjiang, China. Both larvae and adults of coccinellids can prey on aphids; however, they attack other prey as well: for example, eggs of Lepidoptera. Furthermore, these predators do not in general show strong prey preferences or great differences in development when exposed to various aphid species (Michels and Flanders, 1992; Formusoh and Wilde, 1993).
Syrphids were associated with D. noxia throughout its Eurasian range (Hopper et al., 1997). The most common species were Episyrphis balteatus, Eupeodes corollae and Sphaerophoria scripta (Hopper et al., 1997). Syrphid adults eat nectar, honeydew and pollen, but their larvae eat aphids. The number of eggs laid by adult female E. balteatus and E. corollae increases with aphid abundance (Chambers, 1991). E. balteatus and E. corollae larvae can eat 413±5 and 311±6 D. noxia, respectively, over their development times of 9-10 and 7-8 days (Rojo et al., 1996). Furthermore, third instars can crawl 2 m/h and live 3-6 days without aphids (Rojo et al., 1996).
Leucopis spp. were found in association with D. noxia throughout its Eurasian range, however they were not as abundant as syrphids. Most have been identified as L. ninae. Like syrphids, adult Leucopis spp. eat nectar, honeydew, and pollen, but their larvae eat aphids, coccids, and psyllids (Tanasijtshuk, 1986). Although they occur in a variety of habitats, their small size has led some to argue that they should excel at attacking D. noxia in the rolled leaves which this aphid often causes. Adult female L. ninae can lay 20±4 eggs per day and 78±6 eggs over a 9-day period (Dabire, 1995). Females oviposit preferentially in aphid colonies and lay more eggs where aphid density is higher (Dabire, 1995). Although females tend to lay more eggs in leaf sheaths, neither they nor their larvae prefer D. noxia over other cereal aphids (Dabire, 1995). Larvae can consume 66±10 D. noxia over their development time of 9±1 days in the laboratory at 20°C. The number of aphids consumed has a great impact on larval development and survival and on adult size and fecundity (Dabire, 1995). Only 4% of second instars find aphid colonies 10 cm away (Dabire, 1995).
Aphelinus spp. were collected from D. noxia throughout most of its Eurasian range, although they were not found in several trips to the Middle East (Hopper et al., 1997). They are solitary endoparasitoids of nymphs and adults of many aphid species: A. asychis attacks at least 39 aphid species (Wilbert, 1964; Kalina and Stary, 1976). A. asychis females search for their hosts by walking and occasionally flitting from leaf to leaf. Females are only weakly attracted by volatiles from the plant-host complex (Farias, 1995). A. asychis females can parasitize 17±5 D. noxia per day and 230±30 D. noxia over their lifetime of 40±5 days in the laboratory at 20°C (K Hopper, Beneficial Insects Introduction Research Laboratory, USDA, USA, unpublished data). Adult females of Aphelinus spp. kill a few aphids per day by feeding on them to obtain nutrients for egg production (Cate et al., 1973, Bai and Mackauer 1990). At 20°C, Aphelinus spp. take about 3 weeks to develop from egg to adult emergence in D. noxia (Lajeunesse and Johnson, 1992). Wasp larvae kill their hosts and cause hardening of the aphid exoskeleton to form black, ellipsoidal mummies 7-10 days after oviposition at 20°C.
Aphelinus spp. are relatively rare on cereal aphids other than D. noxia (Rabasse and Dedryver, 1983). However, in the Montpellier region (of France), levels of parasitism of D. noxia by Aphelinus spp. can reach 55% in spring in wheat (Farias, 1995). The taxonomy of the genus Aphelinus needs revision. A. albipodus was only distinguished from A. varipes late in the project to introduce natural enemies for control of D. noxia in the USA; and it appears from reciprocal crosses among populations from different regions that what has been identified as A. asychis is also a species complex (Kazmer et al., 1996). A. varipes and A. albipodus abundances appear to vary inversely in space and time with abundance of the A. asychis complex (Farias, 1995). Although parasitism of D. noxia by Aphelinus spp. can reach high levels (Farias, 1995), there are often few mummies per colony (Chen and Hopper, 1997).
Parasitoid species in the braconid subfamily Aphidiinae were collected from D. noxia throughout its Eurasian range (Hopper et al., 1997). Diaeretiella rapae was the most frequently collected from D. noxia, and Aphidius spp. the next most frequently collected; other species were much more rare (Hopper et al., 1997). Aphidiines are solitary endoparasitoids of nymphs and adults of a wide variety of aphid species: D. rapae and Aphidius colemani collected from D. noxia successfully parasitized 7 and 9 species, respectively, of 14 aphid species to which they were exposed (Elliot et al., 1994a, b). Unlike aphelinids, they do not host feed. A. colemani develops in 12 days at 20°C and produces 316 offspring per female on D. noxia (Prinsloo et al., 1993). Aphidius matricariae develops on D. noxia in 273.1 ± 5.9 degree-days above a threshold of 4.5 ± 0.4°C (Miller and Gerth, 1994). D. rapae and Aphidius spp. can parasitize 40-100 aphids per day and 212-532 aphids over their lifetimes of 7-21 days in the laboratory at 20°C; D. rapae produces a maximum of 60 mummies per day at 21°C (Hågvar and Hofsvang, 1991). They are attracted to volatiles from the plant-host complex in general and from D. noxia on wheat in particular (Farias, 1995; Reed et al., 1995). Parasitism of D. noxia by aphidiines was often lower than that by aphelinids when both were present (Farias, 1995; Chen and Hopper, 1997).
Fungal pathogens were collected from D. noxia in fewer areas in Eurasia than predators and parasitoids (Hopper et al., 1997). This may be in part because some collectors lacked the training to collect and culture entomopathogenic fungi. Nonetheless, epizootics of fungal pathogens in D. noxia populations appear to be rare, perhaps because of the relatively arid environments in which this aphid is usually found (Wraight et al., 1993). However, D. noxia populations were found in the Jezreel Valley and upper Galilee Valley in Israel in 1994 that appeared to have been destroyed by fungal epizootics (Hopper et al., 1997). Prevalence of fungal pathogen infections has also been low in the US (Feng et al., 1991; Wraight et al., 1993). At least two viruses, aphid lethal paralysis virus and Rhopalosiphum padi virus, are known to attack D. noxia (Laubscher and Von Wechmar, 1991, 1992).
Impact SummaryTop of page
|Fisheries / aquaculture||Negative|
ImpactTop of page
D. noxia has a great economic impact on cereal crops (Brooks et al., 1994). It is a phloem feeder like other aphids and the symptoms evident on plants are a result of this feeding mechanism. By feeding on the phloem, the aphid damages the plants through nutrient drainage (Dixon, 1985) which results in chlorosis, necrosis, wilting, stunting, and curling of the leaves, misshapen or nonappearance of new growth, and localised cell death at the site of aphid feeding. D. noxia further elicits an increase in essential amino acids in the phloem sap by triggering the breakdown of proteins in infested wheat leaves (Burd and Burton, 1992; du Toit, 1986; Ma et al., 1998; Miller et al., 2001). The damage to the foliar tissue is thought to play a role in the pest’s ability to increase nutritional quality of the host plant (Botha et al., 2006).
Barley and wheat are the most important cultivated hosts of D. noxia and considerable yield losses can occur in these crops. It is particularly injurious to late-sown barley in continental climates. Spring wheat suffers greatest yield loss when attacked during tillering to boot stage; winter wheat suffers greatest loss after vernalization (Gray et al., 1990). Early, heavy infestations on barley can cause total crop loss (Adisu et al., 2003) and significantly affect grain quality (Bregitzer et al., 2003). Aphid feeding on susceptible genotypes causes chlorosis and longitudinal streaking of leaves, and emerging leaves remain tightly rolled, which traps spikes and prevents their normal development (Mornhinweg, 2011). Infestation of wheat seedlings by D. noxia in the autumn can reduce the ability of seedlings to survive the winter (Thomas and Butts, 1990; Storlie et al., 1993). In general, damage is greatest when crop ripening coincides with peak aphid numbers.
D. noxia has been a pest of small grains in Russia since 1912, when the first serious outbreaks on barley were recorded. In the Crimea, D. noxia has reduced harvests by up to 75% in some years. In Eurasia generally, where it is thought to have originated, it is only occasionally a serious pest; with short-lived outbreaks being reported (Grossheim, 1914; Tuatay and Remaudière, 1964; Fernández et al., 1992).
Since its appearance in Texas in 1986 (Stoetzel, 1987), D. noxia has become a major pest of wheat and barley in the USA, causing over US$850 million in direct and indirect losses from 1987 to 1992 (Brooks et al., 1994). During the 1992/93 cropping season, over 7 million acres (20%) of dryland winter wheat and 1 million acres (33%) of barley were infested throughout the western USA (Webster and Amosson, 1994). In Canada, yield losses ranging from 25 to 37% without insecticide treatments in field trials (Butts et al., 1997).
Since its introduction in 1978, it has also become the major pest of wheat in South Africa (Walters et al., 1980), where it can cause over 80% yield loss if not controlled in the summer rainfall region (Tolmay and Prinsloo, 1994). In Ethiopia, it has been a major pest of barley for over two decades, with 20-30% yield loss in some areas (Haile and Megenasa, 1987). D. noxia is also a pest in Yemen (Erdelen, 1981).
Macedo et al. (2009) showed that D. noxia affected net photosynthesis of wheat. Wheat plants tolerant to D. noxia often exhibit increased photosynthetic rates, increased growth rates, increased stored root carbon and/or an enhanced ability to shunt stored carbon from roots to shoots (Burd and Elliott, 1996; Haile et al., 1999; Reidel and Blackmer, 1999; Smith and Chuang, 2013). In addition to direct feeding damage, major indirect losses in wheat and barley can be caused as a result of D. noxia transmitting Barley yellow dwarf virus (BYDV-PAV). However, D. noxia is a less effective vector of BYDV than Rhopalosiphum padi, R. maidis, Schizaphis graminum or Sitobion avenae (El-Yamani and Bencharki, 1997).
There have been conflicting reports on the effect of D. noxia on yield and yield components of host plants. For instance, in Canada, Ba-Angood and Stewart (1980) found that aphid feeding significantly reduced the number of grains per ear, whereas Milne and Delves (1999) in Australia found that aphid feeding did not affect grain numbers per ear. However, aphid feeding usually significantly reduces 1000-seed weight and grain yield per ha (e.g., Ba-Angood and Stewart, 1980; Milne and Delves, 1999; Damate, 2015). The percentage yield loss from D. noxia on irrigated wheat in Ethiopia ranged from 15 to 93%, depending on variety and season (Damate, 2015).
Damage ratings based on leaf chlorosis have frequently been used to evaluate resistance to D. noxia. However, some workers (e.g., Frank et al., 1989; Smith et al., 1991; Assad et al., 2004) suggest that leaf chlorosis may not be effective in separating resistant entries. To increase the accuracy of evaluation, leaf damage other than chlorosis, i.e. leaf rolling and leaf trapping, should also be considered (Nematollahy et al., 1998).
Detection and InspectionTop of page
D. noxia is most easily detected in cereal crops by visual examination of plants. This aphid causes characteristic symptoms in cereals: leaves of infested plants are streaked and rolled. The streaks are white to purple, extend most of the long axis of the leaf, and are irregularly distributed across the short axis of the leaf. Purple, as opposed to white, streaks appear more frequently when the plant has been exposed to low temperatures. Rolling extends in severity from folding along the mid-vein, to one edge of the leaf rolled, to the leaf being rolled completely upon itself forming a straw-like tube. Heavily infested plants are stunted. Seed heads can be trapped by rolled flag leaves, giving the seed head a fish-hook shape. Symptoms are less severe on some wild grasses.
Like other aphids, alate D. noxia can be captured in suction traps, yellow pan traps, and sticky traps (Labonne et al., 1983; Aalbersberg et al., 1984; Melia et al., 1990; Armstrong et al., 1991; Halbert et al., 1992b; Basky, 1993). Nymphs and apterous adults are rarely captured in suction traps and are caught much less than alate adults in yellow pan traps and sticky traps (Labonne et al., 1983). D. noxia can also be sampled effectively by collecting plant material and placing it directly in Berlese funnels.
Similarities to Other Species/ConditionsTop of page
No other cereal aphids have both short, inconspicuous cornicles and a supracaudal process. Furthermore, no other pest causes the symptoms typical of D. noxia attack on cereals (see Symptoms, and Detection and Inspection Methods).
Prevention and ControlTop of page
Walker and Peairs (1994) reviewed cultural control methods for D. noxia. These include planting date, planting density, furrow orientation, irrigation, fertilization, grazing, manipulation of crop residue, and alternative host plants.
Delaying planting beyond the period at the beginning of a season when alate D. noxia are abundant can reduce initial infestations. However, other agronomic constraints prevent large shifts in planting date. Although suction trap catches indicated sufficient autumn dispersal in Idaho (USA) to warrant delayed planting to avoid D. noxia infestation (Halbert et al., 1990), in more southerly regions, where early spring densities of alate D. noxia are higher, late-planted wheat may suffer more from spring infestations. D. noxia density is sometimes higher where plant density is lower (Walker and Peairs, 1994), so that increasing plant density could reduce D. noxia infestation.
North-South-oriented irrigation furrows in some cases give lower winter survival and slower development of D. noxia than East-West-oriented furrows because of reduced solar insolation (Hammon and Peairs, 1992). Thus, running irrigation furrows from north to south where possible in regions with marginal winter survival for D. noxia could reduce spring infestations and damage.
Nitrogen fertilization alone did not affect D. noxia numbers in several studies (Walker et al., 1991; Archer et al., 1995), although there was a shift to earlier reproduction with high-nitrogen fertilized wheat (Moon et al., 1995). On the other hand, Larson et al. (1990) found that nitrogen and sulphur together increased D. noxia infestation and Walker and Peairs (1994) found the opposite. It appears that individual nutrients can have varied effects depending on other nutrients, irrigation, and perhaps other variables. Irrigation in some cases reduced D. noxia population growth (Archer et al., 1995; Farias et al., 1995), but in others did not (Farias et al., 1995). A phenotypic study showed delayed development and less severity of symptoms with decreased numbers of aphids feeding on wheat plants treated with potassium phosphate in comparison with untreated plants for both resistant (Tugela DN) and susceptible (Scheepers) cultivars (Venter et al., 2014).
Although spring grazing can increase subsequent D. noxia population growth on the regrown plants (Messina et al., 1993a), it can also delay D. noxia growth and thus insecticide applications (Walker et al., 1990). Because D. noxia populations must survive the period between the harvest of one year's crop and the planting of the next, management of crop residues and alternative host plants appear promising. The current practice in much of the western USA is to plough under all crop residues and strictly control non-crop plants with herbicides. Although this does reduce reservoirs for D. noxia and other pests, it also reduces sources of natural enemies of these pests. Replacement of wild grasses currently used for forage and for conservation reserve programmes with grasses less suitable for D. noxia growth appears promising. Grass species vary in suitability for D. noxia development (Kindler et al., 1991a, 1992; Mowry et al., 1995). Furthermore, Kindler et al. (1991c, 1992) have found resistance to D. noxia within species of forage grasses. Fungal endophytes may provide some of this resistance (Kindler et al., 1991b; Clement et al., 1992).
Imidacloprid is effective as a seed treatment for dryland wheat (Pike et al., 1993; van der Westhuizen et al., 1994), and Pike et al. (1993) consider it an environmentally safer approach than aerial or in-furrow applications of broader spectrum insecticides. When compared with several other foliar applications chlorpyrifos caused the greatest reduction in D. noxia density (Hill et al., 1993). Chlorpyrifos was effective at half the dose currently used in Canada, but residual impact on D. noxia survival was low (Hill et al., 1993).
Variation in D. noxia susceptibility to insecticides suggests the possibility that D. noxia may develop resistance to them (Brewer and Kaltenbach, 1995). However, Bayoun et al. (1995) found some insecticides not in current use against D. noxia that were effective for its control and relatively less toxic to natural enemies.
The limited evidence available suggests that indigenous natural enemies in South Africa and in the USA have had little impact to date on D. noxia population dynamics (Aalsbersberg et al., 1988b; Feng et al., 1992; Wraight et al., 1993). However, there are several reasons to expect that introducing Eurasian natural enemies would reduce D. noxia abundances in these countries. Introduced natural enemies have substantially controlled several introduced aphid pests, e.g. Therioaphis trifolii (spotted alfalfa aphid) and Chromaphis juglandicola (walnut aphid) (Laing and Hamai, 1976). Although these successes were in orchards or perennial crops, introduced natural enemies have also substantially reduced the abundance of introduced cereal pests, e.g. the cereal leaf beetle, Oulema melanopus in the USA (Haynes et al., 1974), and greenbug, Schizaphis graminum in Chile (Zuniga, 1990). Furthermore, a wide variety of predators and parasitoids attack cereal aphids, and many of these have been reported in association with D. noxia (Pike et al., 1991). Lastly, exclosure experiments suggest natural enemies (including predatory coccinellid, predatory syrphids and parasitoid wasps) play a major role in limiting D. noxia populations in southern France (Hopper et al., 1995).
Studies conducted from 1993 to 1994 in southeastern Wyoming (Brewer et al., 1998), after establishment of Aphelinus asychis and A. albipodus, documented high levels of parasitism of D. noxia by these parasitoids, especially A. albipodus. Lee et al. (2005) showed there may be regional differences in the importance of parasitoids in biological control of D. noxia.
Currently, D. rapae is being released for biological control of D. noxia. Studies of the impact of aphidophagous parasitoids and predators on D. noxia populations soon after invasion by the pest indicated that native natural enemies were ineffective at controlling D. noxia (Rice and Wilde, 1991; Wraight et al., 1993). In contrast, Hopper et al. (1995) observed that natural enemies played a significant role in suppressing D. noxia populations in Europe, where the Russian wheat aphid is presumably native (Lee et al., 2005). So far, four species of introduced parasitoids are known to have established in the USA: Aphelinus asychis (in Texas, Michels and Whitaker-Deerberg, 1993), Aphelinus albipodus, Aphidius colemani and A. uzbekistanicus (not from D. noxia; Halbert et al., 1996). No predators are known to have established.
Researchers in biological control at universities, state departments of agriculture, the US Department of Agriculture, and the International Institute for Biological Control carried out an extensive programme of foreign exploration and importation of predators, parasitoids and entomopathogens into the USA (e.g., Gonzàlez, 1989; Hopper et al., 1990; Gruber et al., 1991; Hopper, 1992; Gonzàlez et al., 1993; Hopper et al., 1993; Hopper et al., 1993; McKinnon et al., 1993; Gonzàlez et al., 1994; Hopper et al., 1994a, b[N1] ; Pike et al., 1994; Reed and Kindler, 1994; Tanigoshi et al., 1995).
Syrphids are promising candidates for D. noxia biological control because of their frequent co-occurrence with D. noxia and because of the great consumption rate and vigour of their larvae. Leucopis spp. are less promising candidates than syrphids because of the lower number of aphids their larvae consume and the lower capacity of their larvae to find aphid colonies. However, because of their relatively high fecundity, the dependence of oviposition on aphid density, and their potential to attack aphids in rolled leaves, they should not be neglected. Aphelinus spp. are promising candidates for D. noxia biological control because they are active during the seasons when D. noxia is active, they occur throughout most of the geographic range of D. noxia, and they live long and can kill a large number of aphids. Aphidiines appear to be less promising than aphelinids for biological control of D. noxia because they parasitize relatively few of these aphids. However, they have been successful in the control of other aphid pests and thus were included among the introductions into the USA.
A more limited programme of introductions has been conducted in South Africa (Tolmay and Prinsloo, 1994). This included introductions of Aphelinus hordei from the Ukraine (via CSIRO, Australia) and Leucopis ninae from Pakistan, Iran, and China (via USDA, USA). Aalbersberg et al. (1989) found that planting a refugium around wheat, to provide prey/hosts for D. noxia natural enemies, increased natural enemy abundance in wheat in South Africa. However, the natural enemies were still not able to keep D. noxia in check, partly because of poor timing of prey/host dynamics in the refugium. Hughes et al. (1994) released A. varipes into Australia as a pre-emptive measure against D. noxia which may eventually be introduced. However, there is no evidence that A. varipes established in Australia.
Feng and Johnson (1991) and Wang and Knudsen (1993) explored the use of the fungal pathogen Beauveria bassiana for D. noxia control. Wraight et al. (1993) suggested that use of pathogens to control D. noxia in the semi-arid western USA will require increased humidity, e.g. from irrigation.
Resistance has been identified in barley and different wheat genotypes, including bread and non-bread wheat (e.g., Du Toit, 1989; Calhoun et al. 1991b; Quick et al., 1991; Smith et al., 1991; Souza et al., 1991; Robinson et al., 1991; Robinson, 1992a, 1993; Sarafrazi and Ahmadi, 1993; Formusoh et al., 1992, 1994; Nematollahy et al., 1998; Kazemi et al., 2001; Najafi-Mirak et al., 2003; Bregitzer et al., 2005, 2008; Hamedanian et al., 2010; Pourhaji and Ahmadi, 1999; Moharramipour et al., 2002; Mornhinweg et al., 2006, 2007a, 2007b, 2008, 2009, 2011; Pourhaji et al., 2011). Webster et al. (1991) published a uniform rating system for classifying foliar damage, but other rating systems have also been used (Du Toit, 1988; Nkongolo et al., 1990; Souza et al., 1991; Robinson et al., 1992a; Burd et al., 1993; Porter et al., 1993). Resistance to foliar damage symptoms correlated well with resistance to yield loss (Calhoun et al., 1991b). In vitro tests (Zemetra et al., 1993; Dong et al. 1994) and behavioural tests (Girma et al., 1992) for screening accessions for resistance to D. noxia have been developed.
Resistance in some wheat lines appears to be mostly from antibiosis with some tolerance (Du Toit, 1989; Quisenberry and Schotzko, 1994). One resistant wheat line showed moderate tolerance, high antibiosis, and little antixenosis (Baker et al., 1992). Other lines are tolerant with low levels of antibiosis (Smith et al., 1992). Nematollahy et al. (1999) showed that out of 15 bread and non-bread wheat genotypes, ‘5172’, a T. monococcum genotype, and ‘4898’ a T. aestivum line, had the highest level of tolerance and antibiosis resistance. Barley resistance is in part from antibiosis (D. noxia feed and grow less on resistant lines (Robinson et al., 1991, 1992b; Webster et al., 1993b) but also from antixenosis (Robinson, 1992b).
Resistance varies with environment, e.g. susceptible seedling survival in harsh winters is compromised by D. noxia infestation (Thomas and Butts, 1990; Storlie et al., 1993) so it is best to screen for resistance in the target environment (Calhoun et al., 1991b). Resistance may be related to concentration of hydroxamic acid (DIBOA) in barley (Gianoli and Niemeyer, 1998).
The genetic mechanisms of resistance in various accessions has been investigated (Du Toit, 1989; Nkongolo et al., 1991a, b, 1992; Robinson et al., 1992b; Elsidaig and Zwer, 1993; Marais et al., 1994; Mornhinweg et al., 1993; Schroeder-Teeter et al., 1993-4; Nieto-Lopez and Blake, 1994; Assad et al., 1999; Najafi-Mirak et al., 2003; Fazel-Najafabadi et al., 2009, 2015). Six major genes have been identified (Quick, 1994). More resistance genes may be available in perennial Triticeae (Kindler et al., 1993), wild Hordeum spp. (Kindler and Springer, 1991), and triticale (Webster, 1990; Scott et al., 1991). Effects of resistance on other characters have been explored (Zwer et al., 1994). The physiological/biochemical basis of resistance have also been studied (Miller et al., 1994).
There is a lot of research on screening resistance to D. noxia in the laboratory and greenhouse at the seedling stage, but reports of field screening are scarce (e.g., Calhoun et al., 1991a, b; Nematollahi, 2010).Six resistant accessions have been released for commercial production in the USA (Quick, 1994). Two resistant cultivars have been released for commercial production in South Africa (Tolmay and Prinsloo, 1994). Screening of the more than 23,000 accessions in the USDA-ARS National Small Grains Collection (NSGC) has identified 116 accessions with some level of resistance to the United States biotype USA1 (Mornhinweg, 2011). From 40 of these accessions, 4 cultivars and 58 adapted germplasm lines have been developed and released (Mornhinweg et al., 2006, 2007a, b, 2008, 2009, 2011; Bregitzer et al., 2005, 2008). Several studies have examined the genetics of RWA resistance in barley. Two studies indicated that single dominant or semi-dominant genes provided RWA resistance in lines from CIMMYT tested with a Mexican RWA population (Robinson et al., 1992b) and from Iran tested with RWA from Iran (Assad et al., 1999).
Resistance and biotypes problem
Because virulence of D. noxia appears to vary within and between geographic regions (Bush et al., 1989; Puterka et al., 1992, 1993), new biotypes may arise that can overcome the defences of the new cultivars being released. This is especially true for antibiotic forms of resistance because these select for aphids that are not susceptible to antibiosis and may reduce the efficacy of natural enemies (Reed et al., 1992a, b). Resistant cultivars carrying Dn-resistance genes were classically used to control outbreaks of D. noxia (Botha and Venter, 2000). However, this control is increasingly more difficult, with the development of new D. noxia resistance-breaking biotypes putting pressure on farmers to revert to using pesticide applications to increase yields.
Since the occurrence of the first biotype (RWA2) in 2003, biotypes of D. noxia have become a serious threat to the development and deployment of plant resistance in cereals (Haley et al., 2004). Previous to the appearance of RWA2, resistance genes for wheat designated Dn1 to Dn9, Dnx and Dny were found to be effective against the original Russian wheat aphid (RWA1) that invaded the USA in 1986 (Haley et al., 2004). Since the discovery of RWA2, six additional biotypes (RWA3- RWA8) have been described (Burd et al., 2006; Weiland et al., 2008; Randolph et al., 2009).
Biotypic diversity within D. noxia populations has been identified in wheat based on differential reactions associated with the 11 identified genes and QTLs for resistance (Dn1-9, x, y). At least eight biotypes are present in the USA (USA1–8; Burd et al., 2006; Puterka et al., 2006; Weiland et al., 2008). USA1 and USA2 are prevalent in the seven western states most affected by D. noxia (Puterka et al., 2007). In a recent study, Puterka et al. (2014) found that biotypes RWA1, RWA2, RWA6 and RWA8 differed in virulence, while biotypes RWA3, RWA4, RWA5 and RWA7 produced similar virulence profiles. Thus, they consolidated these biotypes as RWA3/7, and indicated that in USA the five main biotypes RWA1, RWA2, RWA3/7, RWA6 and RWA8 can be identified using only four wheat genotypes containing Dn3, Dn4, Dn6 and Dn9.
Jankielsohn (2014) had been provided guidelines for the sampling, identification and designation of D. noxia biotypes in South Africa. There are currently four D. noxia biotypes recorded in South Africa. RWASA1 is the first biotype that was recorded in 1978. RWASA2, virulent against the Dn1 resistant gene in wheat, was recorded in 2005 in wheat-producing areas, especially in the Eastern Free State (Tolmay et al., 2007). RWASA3, virulent against the Dn4 resistant gene in wheat, was recorded in 2009, also predominantly in the Eastern Free State (Jankielsohn, 2014). During 2011 RWASA4, virulent against the Dn5 resistant gene, was recorded near Bethlehem in the Eastern Free State (Jankielsohn, 2014). Additional biotypes have been identified in Hungary, Chile, Ethiopia, Argentina and the Czech Republic that differentially affect resistance in wheat (Basky, 2003; Smith et al., 2004; Ricci et al., 2012).
The biotypic diversity in D. noxia has limited useful resistant germplasm to 94M370 (Dn7 gene), CI2401 and STARS 2414-11. In addition, two sources of resistance in barley, STARS 9301B and STARS 9577B, still exhibit strong resistance to all eight biotypes (Puterka et al., 2006). In a recent study, 23 lines resistant to all biotypes, included 10 unadapted germplasm accessions and 13 improved germplasm lines, providing useful germplasm for developing new barley cultivars with resistance to multiple D. noxia biotypes (Daheen et al., 2014).
Resistance genes in wheat appear to function in a biotype-specific manner with single genes providing resistance to one or more biotypes (Puterka et al., 2007; Weiland et al., 2008). In contrast, the two most widely-used barley germplasm lines for breeding resistant cultivars, STARS9301B and STARS9577B, show resistance to all eight United States biotypes, as defined by their differential reactions on wheat (Puterka et al., 2006; Weiland et al., 2008). So it seems that the apparent lack of biotype diversity with respect to resistance in barley is probably an artefact of insufficient study (Daheen et al., 2014).
Although new biotypes develop in various parts of the world it appears that there is very limited genetic variation between the different biotypes and also between their endosymbionts (Weiland et al., 2008; Swanevelder et al., 2010).
Integrated Pest Management
Integrated pest management (IPM) based on sampling pest density before deciding whether or not to treat requires cheap and rapid sampling plans for determining pest densities and accurate assessment of economic impact of these densities. Various plans have been developed for sampling D. noxia density (Schaalje et al., 1991; Feng and Nowierski, 1992; Feng et al., 1993, 1994; Nowierski et al., 1994), and for assessing whether D. noxia density exceeds an action threshold (Bechinski and Hohman, 1991; Legg et al., 1991, 1994; Butts and Schaalje, 1994). Legg et al. (1993) developed software for supporting decisions concerning chemical treatments.
Various relationships between D. noxia density and yield loss have been published. Du Toit (1986) reported an economic injury level of 14% infested plants when the ear had emerged from the flag leaf, and 4-7% infested plants when the first node had formed for winter wheat in South Africa. Girma et al. (1993) reported an economic injury level of 1-2 aphids per 7 plants for spring infestations, and 2-4 aphids per 7 plants for autumn infestations for Kansas wheat. Archer and Bynum (1992) found 4.6% yield loss for each 10% increase in damaged tillers and 4.8% yield loss for each 10% increase in infested tillers in dryland winter wheat in Colorado.
It may be possible to improve the level of aphid control by combining plant resistance and biological control (van Emden and Wratten, 1991). Control remains relatively limited when each method is applied separately. The compatibility between these management tactics was studied by Farid et al. (1998) and Brewer et al. (1999). Brewer et al. (1999) showed that the use of resistant barley in areas where the parasitoid, Diaretiella rapae, is established is justified. In contrast, Reed et al. (1992) found that the development time of D. rapae was prolonged when it developed on D. noxia reared on resistant wheat cultivars with antibiotic factors compared to aphids reared on susceptible cultivars. It seems that plants can have a positive or negative effect on parasitoids, depending on the level and category of resistance (van Emden, 1991; Hare, 1992) and understanding the interactions that may arise from plant-aphid-parasitoid systems can therefore be important in developing a pest management system for these insects.
ReferencesTop of page
Aalbersberg YK, Toit F du, Westhuizen MC van der, Hewitt PH, 1987. Development rate, fecundity and lifespan of apterp of the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae), under controlled conditions. Bulletin of Entomological Research, 77(4):629-635
Aalbersberg YK, Walters MC, Van Rensburg NJ, 1984. The status and potential of biological control studies on Diuraphis noxia (Aphididae). Tech. Commun. South Afr. Dep. Agric., 191:44-66.
Aalbersberg YK, Westhuizen MC van der, Hewitt PH, 1987. A simple key for the diagnosis of the instars of the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae). Bulletin of Entomological Research, 77(4):637-640
Aalbersberg YK, Westhuizen MC van der, Hewitt PH, 1988. Natural enemies and their impact on Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae) populations. Bulletin of Entomological Research, 78(1):111-120
Aalbersberg YK, Westhuizen MC van der, Hewitt PH, 1989. Japanese radish as a reservoir for the natural enemies of the Russian wheat aphid Diuraphis noxia (Hemiptera: Aphididae). Phytophylactica, 21(3):241-245
Adisu B, Freier B, Büttner C, 2003. Effectiveness of predators and parasitoids for the natural control of Diuraphis noxia (Homoptera: Aphididae) on barley in Central Ethiopia. Communications in Agricultural and Applied Biological Sciences [Proceedings of the 55th International Symposium on Crop Protection, Part 1, Gent, Belgium, 6 May 2003.], 68(4a):179-188. http://fltbwww.rug.ac.be/
Aizenberg E, 1935. New genera and two new species of Aphididae. Bulletins de la station biologique a Bolchevo, 7-8:158-60.
Alhag EA, Al-Rokaibah AA, Zaitoon AA, 1996. Natural enemies of cereal aphids in sprinkler-irrigated wheat in central Saudi Arabia. Bulletin of Faculty of Agriculture, University of Cairo, 47(4):649-663; 19 ref.
Anonymous, 1963. Insects not known to occur in the United States. Barley Aphid [Cuernavaca noxius (Mordvilko)]. Coop. Econ. Insect Rep. U. S., 13:1357-8.
Araya JE, Quiroz C, Wellso SG, 1990. Pest status and control of the Russian Wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae). A review. Station Bulletin, 588.
Armstrong JS, Peairs FB, Pilcher SD, Russell CC, 1993. The effect of planting time insecticides and liquid fertilizer on the Russian wheat aphid (Homoptera: Aphididae) and the lesion nematode (Pratylenchus thornei) on winter wheat. Journal of the Kansas Entomological Society, 66(1):69-74
Armstrong S, Peairs F, Nielsen D, Roberts E, Holtzer T, Stushnoff C, 1992. The overwintering biology of Diuraphis noxia on the Northeastern Plains of Colorado. Great Plains Agric Council Publ., 142:211-212.
Assad MT, Mehrabi AM, Pakniyat H, Nematollahy MR, 2004. The effect of resistance components on reducing yield and its related characters in wheat as infected by Diuraphis noxia (Hemiptera: Aphididae). Cereal Research Communications, 32(1):69-73.
Barta M, Cagan L, 2007. Natural control of Diuraphis noxia and Rhopalosiphum maidis (Aphidoidea) by parasitic entomophthorales (Zygomycota) in Slovakia. Cereal Research Communications, 35(1):89-97. http://www.akademiai.com/content/40787021m3kj1422/fulltext.pdf
Basky Z, 2003. Biotypic and pest status differences between Hungarian and South African populations of Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae). Pest Management Science, 59(10):1152-1158.
Bayoun IM, Plapp FW Jr, Gilstrap FE, Michels GJ Jr, 1995. Toxicity of selected insecticides to Diuraphis noxia (Homoptera: Aphididae) and its natural enemies. Journal of Economic Entomology, 88(5):1177-1185
Bechinski E, Homan H, 1991. Sequential decision cards: a tool for Russian wheat aphid control. Bull. Idaho Agric. Exp. Stn., 884.
Black WC IV, DuTeau NM, Puterka GJ, Nechols JR, Pettorini JM, 1992. Use of the random amplified polymorphic DNA polymerase chain reaction (RAPD-PCR) to detect DNA polymorphisms in aphids (Homoptera: Aphididae). Bulletin of Entomological Research, 82(2):151-159
Bodenheimer FS, Swirsky E, 1957. The Aphidoidea of the middle east. Jerusalem, Israel: Weizmann Science Press of Israel.
Botha AM, Lacock L, Niekerk Cvan, Matsioloko MT, Preez FBdu, Loots S, Venter E, Kunert KJ, Cullis CA, 2006. Is photosynthetic transcriptional regulation in Triticum aestivum L. cv. 'TugelaDN' a contributing factor for tolerance to Diuraphis noxia (Homoptera: Aphididae)? Plant Cell Reports, 25(1):41-54.
Bregitzer P, Mornhinweg DW, Jones BL, 2003. Resistance to Russian wheat aphid damage derived from STARS 9301B protects agronomic performance and malting quality when transferred to adapted barley germplasm. Crop Science, 43(6):2050-2057.
Brewer MJ, Mornhinweg DW, Huzurbazar S, 1998. Compatibility of insect management strategies: Diuraphis noxia abundance on susceptible and resistant barley in the presence of parasitoids. BioControl, 43(4):479-491.
Brewer MJ, Mornhinweg WD, Huzurbazar S, 1999. Compatibility of insect management strategies: Diuraphis noxia abundance on susceptible and resistant barley in the presence of parasitoids. BioControl, 43:479-491.
Brooks LS, Amosson G, Hein G, Johnson D, Legg B, Massey B, Morrison P, McBride D, Peairs F, 1994. Economic impact of the Russian wheat aphid in the western United States: 1990-1992. Great Plains Agricultural Council Publication, GPAC-143., USA: Great Plains Agricultural Council.
Burd JD, Burton RL, Webster JA, 1993. Evaluation of Russian wheat aphid (Homoptera: Aphididae) damage on resistant and susceptible hosts with comparisons of damage ratings to quantitative plant measurements. Journal of Economic Entomology, 86(3):974-980
Burd JD, Elliott NC, 1996. Changes in chlorophyll a fluorescence induction kinetics in cereals infested with Russian wheat aphid (Homoptera: Aphididae). Journal of Economic Entomology, 89(5):1332-1337.
Burd JD, Porter DR, Puterka GJ, Haley SD, Peairs FB, 2006. Biotypic variation among North American Russian wheat aphid (Homoptera: Aphididae) populations. Journal of Economic Entomology, 99(5):1862-1866. http://www.bioone.org/doi/full/10.1603/0022-0493-99.5.1862
Bush L, Slosser JE, Worrall WD, 1989. Variations in damage to wheat caused by wheat aphid (Homoptera: Aphididae) in Texas. J. Econ. Entomol., 82:466-71.
Butts RA, 1992. Russian wheat aphid summary Alberta, Canada 1991. Great Plains Agric Council Publ.,142:22.
Butts RA, Schaalje GB, 1994. Spatial distribution of fall populations of Russian wheat aphid (Homoptera: Aphididae) in winter wheat. J. Econ. Entomol., 87:1232-1236.
Butts RA, Thomas JB, Lukow O, Hill BD, 1997. Effect of fall infestations of Russian wheat aphid (Homoptera: Aphididae) on winter wheat yield and quality on the Canadian prairies. Journal of Economic Entomology, 90(4):1005-1009; 12 ref.
Cate RH, Archer TL, Eikenbary RD, Starks KJ, Morrison RD, 1973. Parasitization of the greenbug by Aphelinus asychis and effect of feeding by the parasitoid on aphid mortality. Environ. Entomol., 2:549-553.
Chambers RJ, 1991. Oviposition by aphidophagous hoverflies (Diptera: Syrphidae) in relation to aphid density and distribution in winter wheat. In: Dixon AFG, Hodek I, eds. Behaviour and Impact of Aphidophaga. Hague, The Netherlands: Academic.
Chen K, Hopper KR, 1997. Diuraphis noxia (Homoptera: Aphididae) population dynamics and impact of natural enemies in the Montpellier region of southern France. Environ. Entomol., in press.
Clement SL, Johnson RC, Pike KS, 1990. Field populations of the Russian wheat aphid (Homoptera: Aphididae) and other cereal aphids on cool-season perennial grass accessions. Journal of Economic Entomology, 83(3):846-849
Clua A, Castro AM, Ramos S, Gimenez DO, Vasicek A, Chidichimo HO, Dixon AFG, 2004. The biological characteristics and distribution of the greenbug, Schizaphis graminum, and Russian wheat aphid, Diuraphis noxia (Hemiptera: Aphididae), in Argentina and Chile. European Journal of Entomology, 101(1):193-198.
Culjak TG, Barcic JI, 2002. Diuraphis noxia (Kurdjumov) - a new pest of cereals in Croatia. (Diuraphis noxia(Kurdjumov) - nova prijetnja strnim zitaricama u Hrvatskoj.) Agriculturae Conspectus Scientificus (Poljoprivredna Znanstvena Smotra), 67(1):47-55.
Dabire R, 1995. Impact of Leucopis ninae Tanasijtschuk (Diptera: Chamaemyiidae) on Diuraphis noxia (Mordvilko) (Homoptera: Aphididae). Ph.D. diss. Montpellier, France: Ecole Nationale Superieure Agronomique de Montpellier.
Dahleen LS, Mornhinweg D, Bregitzer P, Vitou J, Cakir M, 2014. Biotype differences for resistance to Russian wheat aphid in barley. Crop Science, 54(4):1505-1513. https://www.crops.org/publications/cs/abstracts/54/4/1505
Damate T, 2015. Occurrence of, and yield response to, Russian wheat aphid, Diuraphis noxia (Hemiptera: Aphididae) in irrigated wheat in Ethiopia. International Journal of Tropical Insect Science, 35(1):3-10.
Damsteegt VD, Gildow FE, Hewings AD, Carroll TW, 1992. A clone of the Russian wheat aphid (Diuraphis noxia) as a vector of barley yellow dwarf, barley stripe mosaic, and brome mosaic viruses. Plant Disease, 76(11):1155-1160
De Gouveia MA, 1984. Contribuicao para o estudo dos afideos dos cereais em Portugal. Bol. Soc. Port. Entomol., 2:289-320.
Du Toit F, 1989a. Inheritance of resistance in two Triticum aestivum lines to Russian wheat aphid (Homoptera: Aphididae). J. Econ. Entomol., 82:1251-1253.
Du Toit F, 1989b. Components of resistance in three bread wheat lines to Russian wheat aphid (Homoptera: Aphididae) in South Africa. J. Econ. Entomol., 82:1779-1781.
Elliott NC, French BW, Burd JD, Kindler SD, Reed DK, 1994. Parasitism, adult emergence, sex ratio, and size of Aphidius colemani (Hymenoptera: Aphidiidae) on several aphid species. Great Lakes Entomologist, 27(3):137-142
Elliott NC, French BW, Reed DK, Burd JD, Kindler SD, 1994. Host species effects on parasitization by a Syrian population of Diaeretiella rapae M'Intosh (Hymenoptera: Aphidiidae). Canadian Entomologist, 126(6):1515-1517
El-Yamani M, Bencharki B, 1997. Transmission of Moroccan isolates of barley yellow dwarf virus by the Russian wheat aphid (Diuraphis noxia) and other aphid species. Phytopathologia Mediterranea, 36(3):129-134; 21 ref.
Emden HFvan, Wratten SD, 1991. Tritrophic interactions involving plants in the biological control of aphids. Aphid-plant interactions: population to molecules [ed. by Peters, D. C. \Webster, J. A. \Chlouber, C. S.]., USA: USDA/Agricultural Research Service and Oklahoma State University, 29-43.
EPPO, 2010. Diuraphis noxia does not occur in the United Kingdom. EPPO Reporting Service 2010/087. Paris, France: European and Mediterranean Plant Protection Organization. http://archives.eppo.org/EPPOReporting/2010/Rse-1004.pdf
EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
Erdelen C, 1981. Biologie und Bekampfung der Getreideblattlaus Diuraphis noxia Mordv. in der arabischen Republik Jemen. Med. Fac. Landbouww. Rijksuniv. Gent., 46:613-21.
Farias A, 1995. Parasitoids and hyperparasitoids associated with Diuraphis noxia (Mordvilko) (Homoptera: Aphididae) in the Monpellier region. Ph.D. diss. Montpellier, France: Ecole Nationale Superieure Agronomique de Montpellier.
Farias AMI de, Hopper KR, Leclant F, 1995. Damage symptoms and abundance of Diuraphis noxia (Homoptera: Aphididae) for four wheat cultivars at three irrigation levels. Journal of Economic Entomology, 88(1):169-174
Farid A, Johnson JB, Shafii B, Quisenberry SS, 1998. Tritrophic studies of Russian wheat aphid, a parasitoid, and resistant and susceptible wheat over three parasitoid generations. Biological Control, 12(1):1-6.
Fazel-Najafabadi M, Peng JunHua, Peairs FB, Simkova H, Kilian A, Lapitan NLV, 2015. Genetic mapping of resistance to Diuraphis noxia (Kurdjumov) biotype 2 in wheat (Triticum aestivum L.) accession CI2401. Euphytica, 203(3):607-614. http://rd.springer.com/journal/10681
Feng MG, Johnson JB, 1991. Bioassay of four entomophthoralean fungi (Entomophthorales) against Diuraphis noxia and Metopolophium dirhodum (Homoptera: Aphididae). Environmental Entomology, 20(1):338-345
Feng MG, Johnson JB, Halbert SE, 1991. Natural control of cereal aphids (Homoptera: Aphididae) by entomopathogenic fungi (Zygomycetes: Entomophthorales) and parasitoids (Hymenoptera: Braconidae and Encyrtidae) on irrigated spring wheat in southwestern Idaho. Environmental Entomology, 20(6):1699-1710
Feng MG, Johnson JB, Halbert SE, 1992. Parasitoids (Hymenoptera: Aphidiidae and Aphelinidae) and their effect on aphid (Homoptera: Aphididae) populations in irrigated grain in southwestern Idaho. Environmental Entomology, 21(6):1433-1440
Feng MG, Nowierski RM, 1992. Spatial distribution and sampling plans for four species of cereal aphids (Homoptera: Aphididae) infesting spring wheat in southwestern Idaho. Journal of Economic Entomology, 85(3):830-837
Feng MG, Nowierski RM, Zeng Z, 1994. Binomial sequential classification sampling plans for Russian wheat aphid (Homoptera: Aphididae) management: robustness varying with tally thresholds of aphids in sample units. Journal of Economic Entomology, 87(5):1237-1250
Formusoh ES, Wilde GE, 1993. Preference and development of two species of predatory coccinellids on the Russian wheat aphid and greenbug biotype E (Homoptera: Aphididae). Journal of Agricultural Entomology, 10(1):65-70; 12 ref.
Fouche A, Verhoeven RL, Hewitt PH, Walters MC, Friel CF, De JaGer J, 1984. Russian aphid (Diuraphis noxia) feeding damage on wheat, related cereals and a Bromus grass species. Pretoria, South Africa: Tech Commun South Afr Dep Agric., 191:22-33.
Geng MG, Nowierski RM, Zeng Z, Scharen AL, 1993. Estimation of population density of the Russian wheat aphid (Homoptera: Aphididae) from the proportion of grain tillers with different tally thresholds of aphids. Journal of Economic Entomology, 86(2):427-435
Gilchrist LI, Rodriguez R, 1984. The extent of Freestate streak and Diuraphis noxia in Mexico. Barley yellow dwarf, a proceedings of the workshop December 6-8, 1983 CIMMYT Mexico, sponsored by the United Nations Development Programme and CIMMYT [edited by Burnett, P.A.; Cuellar, E.] Mexico, D.F.; Mexico; Centro Internacional de Mejoramiento de Maiz y Trigo, 157-163
Girma M, Wilde G, Reese JC, 1990. Influence of temperature and plant growth stage on development, reproduction, life span, and intrinsic rate of increase of the Russian wheat aphid (Homoptera: Aphididae). Environmental Entomology, 19(5):1438-1442
Gonzalez D, Gilstrap F, Zhang W, McKinnon L, Stary P, Waggoner M, Zhang G, 1994. Foreign exploration for natural enemies of Russian wheat aphid in southern Xinjiang, People's Republic of China. In: Proceedings Sixth Russian Wheat Aphid Workshop, 23 January 1994, 208-222.
Government of South Australia, 2016. Detection of Russian Wheat Aphid., Australia: Primary Industries and Regions SA. http://pir.sa.gov.au/alerts_news_events/news/biosecurity/detection_of_russian_wheat_aphid
Gray ME, Hein GL, Walgenbach DD, 1990. Effects of russian wheat aphid (Homoptera: Aphididae) on winter and spring wheat infested during different plant growth stages under greenhouse conditions. J. Econ. Entomol., 83:2434-42.
Grossheim NA, 1914. The barley aphid, Brachycolus noxius Mordwilko. Mem. Nat. Hist. Mus. Zemstwo Province Tavria, 3:35-78.
Gruber F, Poprawski TJ, Rey E, 1991. Survey for natural enemies of Diuraphis noxia (Mordvilko) in Eurasia. IOBC/WPRS Bull//Bull. OILB / SROP., XIV:102-9.
Halbert S, Connelly J, Sandvol L, 1990. Suction trapping of aphids in western North America (emphasis on Idaho). Acta Phytopathol. Entomol. Hung., 25:411-422.
Halbert S, Elberson L, Johnson J, 1992. Suction trapping of Russian wheat aphid: what do the numbers mean. Great Plains Agric Council Publ., 142:282-297.
Halbert SE, Connelly BJ, Bishop GW, Blackmer JL, 1992. Transmission of barley yellow dwarf virus by field collected aphids (Homoptera: Aphididae) and their relative importance in barley yellow dwarf epidemiology in southwestern Idaho. Annals of Applied Biology, 121(1):105-121
Hamedanian P, Hosseinzadeh A, Mirak TN, Nourbakhsh AH, 2010. Evaluation of resistance of bread wheat cultivars to Russian wheat aphid at seedling and stem elongation growth stages. Seed and Plant Improvement Journal, 26-1(1):Pe27-Pe42, en3. http://spij.spii.ir
Hammon RW, Peairs FB, 1992. Distribution of overwintering Russian wheat aphid (Homoptera: Aphididae) in furrow-irrigated small grains in western Colorado. Journal of Economic Entomology, 85(6):2452-2458
Hare JD, 1992. Effects of plant variation on herbivore- natural enemy interactions. In: Plant resistance to herbivores and pathogens: ecology, evolution, and genetics [ed. by Fritz, R. S.\Simms, E. L.]. Chicago, USA: University of Chicago Press, 278-298.
Heimpel GE, Ragsdale DW, Venette R, Hopper KR, O'Neil RJ, Rutledge CE, Wu ZS, 2004. Prospects for importation biological control of the soybean aphid: anticipating potential costs and benefits. Annals of the Entomological Society of America, 97(2):249-258.
Hewitt PH, Niekerk GJJ van, Walters MC, Kriel CF, Fouche A, 1984. Aspects of the ecology of the Russian wheat aphid, Diuraphis noxia, in the Bloemfontein district. I. The colonization and infestation of sown wheat, identification of summer hosts and cause of infestation symptoms. Technical Communication, Department of Agriculture, South Africa, No. 191:3-13
Hill BD, Butts RA, Schaalje GB, 1993. Reduced rates of foliar insecticides for control of Russian wheat aphid (Homoptera: Aphididae) in western Canada. J. Econ. Entomol., 86:1259-1265.
Hopper K, Randolph T, Boylan J, Cepaitis A, Fauvergue X, Gould J, Prokrym D, 1994b. Natural enemy impact on Diuraphis noxia (Mordvilko) (Homoptera: Aphididae) in northeastern Colorado compared to southern France. In: Proceedings Sixth Russian Wheat Aphid Workshop, 23 January 1994, 223-228.
Hopper KR, 1992. Report on European Biological Control Laboratory program on Diuraphis noxia. Agricultural Research Service Progress Report: the Russian Wheat Aphid Fourth Annual Report, 25-28.
Hopper KR, Aidara S, Agret S, Cabal J, Coutinot D, Dabire R, Lesieux C, Kirk G, Reichert S, Tronchetti F, Vidal J, 1995. Natural enemy impact on the abundance of Diuraphis noxia (Homoptera: Aphididae) in wheat in southern France. Environmental Entomology, 24(2):402-408
Hopper KR, Coutinot D, Chen K, Halbert SE, Kazmer DJ, Mercadier G, Miller RH, Pike KS, Tanigoshi LK, 1997. Exploration for natural enemies to control Diuraphis noxia in the United States. Thomas Say Publications, Entomological Society of America, in press.
Hopper KR, Lacey LA, Kazmer DJ, 1993. Report on European Biological Control Laboratory program on Diuraphis noxia. Agricultural Research Service Progress Report: the Russian Wheat Aphid Fifth Annual Report, 34-39.
Hopper KR, Lacey LA, Kazmer DJ, 1994a. Report on European Biological Control Laboratory program on Diuraphis noxia. Agricultural Research Service Progress Report: the Russian Wheat Aphid Sixth Annual Report, 23-25.
Hopper KR, Poprawski TJ, Gruber F, 1990. Report on European Biological Control Laboratory program on Diuraphis noxia. Agricultural Research Service Progress Report: the Russian Wheat Aphid Third Annual Report, 19-20.
Hughes RD, 1988. A synopsis of information on the Russian wheat aphid, Diuraphis noxia (Morvilko). Australia: CSIRO, Division of Entomology Technical Paper No. 28.
Hughes RD, 1996. A synopsis of information on the Russian wheat aphid, Diuraphis noxia (Mordwilko). (Revised edition). CSIRO Australia Division of Entomology Technical Paper, No. 34:97 pp.; 45 pp. of ref.
Hughes RD, Maywald GF, 1990. Forecasting the favourableness of the Australian environment for the Russian wheat aphid, Diuraphis noxia (Homoptera: Aphididae), and its potential impact on Australian wheat yields. Bulletin of Entomological Research, 80(2):165-175
IPPC, 2016. Detection of Russian wheat aphid (Diuraphis noxia) in South Australia and Victoria. IPPC Official Pest Report, No. AUS-74/2. Rome, Italy: FAO. https://www.ippc.int/
IPPC-Secretariat, 2005. Identification of risks and management of invasive alien species using the IPPC framework. Proceedings of the workshop on invasive alien species and the International Plant Protection Convention, 22-26 September 2003. xii + 301 pp.
Jankielsohn A, 2011. Distribution and diversity of Russian wheat aphid (Hemiptera: Aphididae) biotypes in South Africa and Lesotho. Journal of Economic Entomology, 104(5):1736-1741. http://esa.publisher.ingentaconnect.com/content/esa/jee/2011/00000104/00000005/art00035
Jankielsohn A, 2014. Guidelines for the sampling, identification and designation of Russian wheat aphid (Diuraphis noxia) biotypes in South Africa. Journal of Agricultural Research, 1:36-43.
Kalina V, Stary P, 1976. A review of the aphidophagus aphelinidae (Hym., Chalcidoidea), their distribution and host range in Europe. Studia Entomologica Forestalia, 2:143-170.
Kazemi MH, Talebi-Chaichi P, Shakiba MR, Mashhadi-Jafarlou M, 2001. Evaluation of susceptibility to Russian wheat aphid, Diuraphis noxia (Homoptera: Aphididae) in several wheat cultivars at stem elongation growth stage. Iranian Journal of Agricultural Sciences, 11(2):103-111.
Kazmer DJ, Maiden K, Ramualde N, Coutinot D, Hopper KR, 1996. Reproductive compatibility, mating behavior, and random amplified polymorphic DNA variability in some Aphelinus asychis (Hymenoptera: Aphelinidae) derived from the Old World. Annals of the Entomological Society of America, 89(2):212-220; 28 ref.
Kieckhefer RW, Elliott NC, 1989. Effect of fluctuating temperatures on development of immature Russian wheat aphid (Homoptera: Aphididae) and demographic statistics. J. of Economic Entomolgy, 82:119-22.
Kindler SD, Breen JP, Springer TL, 1991. Reproduction and damage by Russian wheat aphid (Homoptera: Aphididae) as influenced by fungal endophytes and cool-season turfgrasses. Journal of Economic Entomology, 84(2):685-692
Kindler SD, Breen JP, Springer TL, 1991b. Endophytes in fescue species and associated enhancement of Russian wheat aphid resistance. Misc. Publ. Agric. Exp. Stn. Okla. State Univ., 132:258.
Kindler SD, Greer LG, Springer TL, 1992. Feeding behaviour of the Russian wheat aphid (Homoptera: Aphididae) on wheat and resistant and susceptible slender wheatgrass. Journal of Economic Entomology, 85(5):2012-2016; 18 ref.
Kindler SD, Springer TL, 1989. Alternate Hosts of russian wheat aphid (Homoptera: Aphididae). J. Econ. Entomol., 82:1358-62.
Kiriac I, Gruber F, Poprawski T, Halbert S, Elberson L, 1990. Occurrence of sexual morphs of Russian wheat aphid, Diuraphis noxia (Homoptera: Aphididae), in several locations in the Soviet Union and the northwestern United States. Proceedings of the Entomological Society of Washington, 92(3):544-547
Kovalev OV, Poprawski TJ, Stekolshchikov AV, Vereshchagina AB, Gandrabur SA, 1991. Diuraphis Aizenberg (Hom., Aphididae): key to apterous viviparous females, and review of Russian language literature on the natural history of Diuraphis noxia (Kurdjumov, 1913). Journal of Applied Entomology, 112(5):425-436
Kriel CF, Hewitt PH, Westhuizen MC van der, 1986. The Russian wheat aphid Diuraphis noxia (Mordvilko): population dynamics and effect on grain yield in the western Orange Free State. Journal of the Entomological Society of Southern Africa, 49(2):317-335
Kurdjumov NB, 1913. Notes on European species of the genus Aphelinus Dalm (Hymenoptera, Chalcidodea), parasitic upon the plant-lice. Revue d'Entomologie russe, 13:266-70.
Laing JE, Hamai J, 1976. Biological control of insect pests and weeds by imported parasites, predators, and pathogens. Theory and Practice of Biological Control; ed. by C.B. Huffaker and P.S. Messenger. Academic Press. New York USA, 685-743
Laubscher JM, Von WMB, 1991. Detection of aphid lethal paralysis virus by immunofluorescence. J. Invertebr. Pathol., 58:52-56.
Lee JH, Elliott NC, Kindler SD, French BW, Walker CB, Eikenbary RD, 2005. Natural enemy impact on the Russian wheat aphid in Southeastern Colorado. Environmental Entomology, 34(1):115-123. HTTP://www.esa.catchword.org
Legg DE, Nowierski RM, Feng MG, Peairs FB, Hein GL, Elberson LR, Johnson JB, 1994. Binomial sequential sampling plans and decision support algorithms for managing the Russian wheat aphid (Homoptera: Aphididae) in small grains. Journal of Economic Entomology, 87(6):1513-1533
Macedo TB, Peterson RKD, Weaver DK, Ni XinZhi, 2009. Impact of Diuraphis noxia and Rhopalosiphum padi (Hemiptera: Aphididae) on primary physiology of four near-isogenic wheat lines. Journal of Economic Entomology, 102(1):412-421. http://esa.publisher.ingentaconnect.com/content/esa/jee/2009/00000102/00000001/art00055
Macharia M, Muthangya PM, Wanjama JK, 1999. Response to seed-dressing aphicides in commercial varieties for preventing Russian wheat aphid (Diuraphis noxia) damage in Kenya. Proceedings of the Tenth Regional Wheat Workshop for Eastern, Central and Southern Africa, University of Stellenbosch, South Africa, 14-18 September, 1998., 418-425; 5 ref.
McKinnon LK, Gilstrap FE, Gonzalez D, Woolley JB, Stary P, Wharton RA, 1992. Importations of natural enemies for biological control of Russian wheat aphid, 1988-1991. Great Plains Agric Council Publ., 142:136-145.
Melia A, Seco MV, Duenas ME, Nunez E, Nieto JM, 1990. Alate aphids (Hom. Aphidoidea) captured with suction traps in Castellon, Leon and Salamanca during 1989. Boletin de Sanidad Vegetal, Plagas, 16(3):635-643
Messina FJ, 1993. Effect of initial colony size on the per capita growth rate and alate production of the Russian wheat aphid (Homoptera: Aphididae). Journal of the Kansas Entomological Society, 66(3):365-371
Messina FJ, Jones TA, Nielson DC, 1993. Performance of the Russian wheat aphid (Homoptera: Aphididae) on perennial range grasses: effects of previous defoliation. Environmental Entomology, 22(6):1349-1354
Michels GJ Jr, Behle RW, 1989. Influence of temperature on reproduction, development, and intrinsic rate of increase of Russian wheat aphid, greenbug, and bird cherry-oat aphid (Homoptera: Aphididae). Journal of Economic Entomology, 82(2):439-444
Michels GJ, Flanders RV, 1992. Larval development, aphid consumption and oviposition for five imported coccinellids at constant temperature on Russian wheat aphids and greenbugs. Southwestern Entomologist, 17(3):233-243
Miller GL, Stoetzel MB, Kane EC, 2005. A systematic reappraisal of the genus Diuraphis Aizenberg (Hemiptera: Aphididae). Proceedings of the Entomological Society of Washington, 107(3):700-728. http://apt.allenpress.com/aptonline/?request=get-archive&issn=0013-8797
Miller H, Porter DR, Burd JD, Mornhinweg DW, Burton RL, 1994. Physiological effects of Russian wheat aphid (Homoptera: Aphididae) on resistant and susceptible barley. Journal of Economic Entomology, 87(2):493-499
Miller RH, Pike KS, Tanigoshi LK, Buschman LL, Kornosor S, 1993. Distribution and ecology of the Russian wheat aphid, Diuraphis noxia Mordvilko (Homoptera: Aphididae) in western Asia and northern Africa. Arab Journal of Plant Protection, 11(1):45-52
Mimeur J, 1942. Aphididae Nord-Americain. Bull. de la societe des sciences naturelles du Maroc., 21:67-70.
Mirak TN, Hossainzadeh A, Zali A, Zeinali H, Saidi A, Rassoulian G, 2004. Inheritance of resistance to Russian wheat aphid, Diuraphis noxia (Mordvilko), based on leaf rolling in wheat. Seed and Plant, 20(2):Pe245-Pe257, en20.
Moharramipour S, Movahedi S, Saidi A, Talebi AA, Fathipour Y, 2002. Evaluation of resistance to the Russian wheat aphid, Diuraphis noxia (Mordvilko), in some advanced wheat lines. Seed and Plant, 18(2):Pe215-Pe228, en17.
Montandon R, Slosser JE, Frank WA, 1993. Factors reducing the pest status of the Russian wheat aphid (Homoptera: Aphididae) on wheat in the rolling plains of Texas. Journal of Economic Entomology, 86(3):899-905
Moon CE, Lewis BE, Murray L, Sanderson SM, 1995. Russian wheat aphid (Homoptera: Aphididae) development, reproduction, and longevity on hydroponically grown wheat with varying nitrogen levels. Environmental Entomology, 24(2):367-371
Mornhinweg DW, 2011. Biotic stress in barley: Insect problems and solutions. In: Barley: Production, improvement, and uses [ed. by Ullrich]., UK: Blackwell Publication, 355-390.
Mornhinweg DW, Bregitzer PP, Porter DR, Peairs FB, Baltensperger DD, Hein GL, Randolph TA, Koch M, Walker T, 2009. Registration of 'Sidney' spring feed barley resistant to Russian wheat aphid. Journal of Plant Registrations, 3(3):214-218.
Mornhinweg DW, Bregitzer PP, Porter DR, Peairs FB, Baltensperger DD, Hein, GL, 2011. Registration of 'Stoneham' spring feed barley resistant to Russian wheat aphid. Journal of Plant Registration, 6:1-5.
Mornhinweg DW, Porter DR, Webster JA, 1993. Inheritance of Russian wheat aphid resistance in spring barley germplasm line, STARS-9301B. Barley Genetics Newsletter, 23:40; 2 ref.
Najafi-Mirak T, Zali, AA, Hosseizadeh AH, Zeinali H, Rasoulian GR, Saeidi A, 2003. Evaluation of resistance to Russian wheat aphid, Diuraphis noxia (Mordwilko) in several durum and bread wheat genotypes. Journal of Science and Technology of Agriculture and Natural Resources, 4:115-126.
NAPIS, 2000. Diuraphis noxia. National Agricultural Pest Information Service, USA: http://www.ceris.purdue.edu/napis/.
Nematollahi MR, 2010. Evaluation of Russian wheat aphid resistance in cultivars and advanced lines of wheat under field conditions in Isfahan province, Iran. Journal of Entomological Society of Iran, 30(1):Pe1-Pe13.
Nematollahy MR, Ahmadi AA, 1998. Characterization of resistance to Russian wheat aphid, Diuraphis noxia (Mordvilko), in several wheat (Triticum spp.) genotypes. Iran Agricultural Research, 18(2):91-106.
Nematollahy MR, Ahmadi AA, 1999. Characterization of resistance components to Russian wheat aphid, Diuraphis noxia (Mordvilko), in several wheat (Triticum spp.) genotypes. Iran Agricultural Research, 18(2):91-105.
Nkongolo KK, Quick JS, Limin AE, Fowler DB, 1991. Sources and inheritance of resistance to Russian wheat aphid in Triticum species amphiploids and Triticum tauschii. Canadian Journal of Plant Science, 71(3):703-708
Olsen CE, Pike KS, Boydston L, Allison D, 1991. Identification of immature and adult apterous forms of the major aphids on small grains. Stillwater, USA: Misc Publ Agric Exp Stn Okla State Univ., 132:323.
Olsen CE, Pike KS, Boydston P, Allison D, 1993. Keys for identification of apterous viviparae and immatures of six small grain aphids (Homoptera: Aphididae). Journal of Economic Entomology, 86(1):137-148
Pike K, Tanigoshi L, Miller R, Kornosor S, 1994. Summary report: biological control agents of Russian wheat aphid in Syria and Turkey. In: Proceedings Sixth Russian Wheat Aphid Workshop, 23 January 1994, 232-237.
Pike KS, Boydston L, Allison D, 1990. Alate aphid viviparp associated with small grains in North America: a key and morphometric characterization. Journal of the Kansas Entomological Society, 63(4):559-602
Pike KS, Reed GL, Graf GT, Allison D, 1993. Compatibility of imidacloprid with fungicides as a seed treatment control of Russian wheat aphid (Homoptera: Aphididae) and effect on germination, growth, and yield of wheat and barley. J. Econ. Entomol., 86:586-592.
Poprawski TJ, Underwood NL, Mercadier G, Gruber F, 1992. Diuraphis noxia (Kurdjumov) - a bibliography on the Russian wheat aphid 1886-1992. Ithaca, USA; USDA Plant Soil & Nutrition Laboratory, 153 pp.
Pourhaji A, 2011. Mass screening of 76 barley genotypes for resistance to Russian wheat aphid Diuraphis noxia (Horn.: Aphididae), in greenhouse condition. Applied Entomology and Phytopathology, 79(1):Pe1-Pe16. http://www.sid.ir/en/JournalList.asp?ID=306&Name=APPLIED+ENTOMOLOGY+AND+PHYTOPATHOLOGY
Pourhaji A, Ahmadi AA, 1999. Greenhouse comparison of resistance of 23 barley genotypes to the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Hom.: Aphididae). Journal of Entomological Society of Iran, 19:57-78.
Pourhaji AR, 2010. Mass screening of 76 barley genotypes for resistance to Russian wheat aphid Diuraphis noxia (Hom.: Aphididae), in greenhouse condition. Applied Entomology and Phytopathology, 79(1):1-16.
Prinsloo GJ, Hewitt PH, Westhuizen MC van der, 1993. The effect of temperature on oviposition behaviour and success of two parasitoids of the Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Hemiptera: Aphididae). African Entomology, 1(2):189-193
Puterka GJ, Black WC IV, Steiner WM, Burton RL, 1993. Genetic variation and phylogenetic relationships among worldwide collections of the Russian wheat aphid, Diuraphis noxia (Mordvilko), inferred from allozyme and RAPD-PCR markers. Heredity, 70(6):604-618
Puterka GJ, Burd JD, Mornhinweg DW, Haley SD, Peairs FB, 2006. Response of resistant and susceptible barley to infestations of five Diuraphis noxia (Homoptera: Aphididae) biotypes. Journal of Economic Entomology, 99(6):2151-2155. http://www.bioone.org/doi/full/10.1603/0022-0493-99.6.2151
Puterka GJ, Burd JD, Porter D, Shufran K, Baker C, Bowling B, Patrick C, 2007. Distribution and diversity of Russian wheat aphid (Hemiptera: Aphididae) biotypes in North America. Journal of Economic Entomology, 100(5):1679-1684. http://www.bioone.org/doi/abs/10.1603/0022-0493%282007%29100%5B1679%3ADADORW%5D2.0.CO%3B2
Puterka GJ, Nicholson SJ, Brown MJ, Cooper WR, Peairs FB, Randolph TL, 2014. Characterization of eight Russian wheat aphid (Hemiptera: Aphididae) biotypes using two-category resistant-susceptible plant responses. Journal of Economic Entomology, 107(3):1274-1283. http://esa.publisher.ingentaconnect.com/content/esa/jee/2014/00000107/00000003/art00050
Quick JS, 1994. Development of cultivars resistant to the Russian wheat aphid. In: Proceedings Sixth Russian Wheat Aphid Workshop, 23 January 1994, 37-41.
Quisenberry SS, Schotzko DJ, 1994. Russian wheat aphid (Homoptera: Aphididae) population development and plant damage on resistant and susceptible wheat. Journal of Economic Entomology, 87(6):1761-1768
Rabasse JM, Dedryver CA, 1983. Biology of cereal aphids in western France. III. Effect of hymenopteran parasites on populations of Sitobion avenae F., Metopolophium dirhodum Wlk. and Rhopalosiphum padi L. Agronomie, 3(8):779-790
Randolph TL, Peairs F, Weiland A, Rudolph JB, Puterka GJ, 2009. Plant responses to seven Russian wheat aphid (Hemiptera: Aphididae) biotypes found in the United States. Journal of Economic Entomology, 102(5):1954-1959. http://esa.publisher.ingentaconnect.com/content/esa/jee/2009/00000102/00000005/art00028
Reed D, Kindler S, 1994. Report of trip to Argentina and Chile. In: Proceedings 6th Russian Wheat Aphid Workshop, 23 January 1994, 163-168.
Reed DK, Kindler SD, Springer TL, 1992. Interactions of Russian wheat aphid, a hymenopterous parasitoid and resistant and susceptible slender wheatgrasses. Entomologia Experimentalis et Applicata, 64(3):239-246
Reed HC, Tan SH, Haapanen K, Killmon M, Reed DK, Elliott NC, 1995. Olfactory responses of the parasitoid Diaeretiella rapae (Hymenoptera: Aphidiidae) to odor of plants, aphids, and plant-aphid complexes. Journal of Chemical Ecology, 21(4):407-418
Ricci M, Cakir M, Castro M, 2012. Diuraphis noxia (Hemiptera: Aphididae): Identification of biotypes present in Argentine populations. Revista de la Sociedad Entomológica Argentina, 71:105-113.
Rice ME, Wilde GE, 1991. Aphid predators associated with conventional and conservational tillage winter wheat. Journal of Kansas Entomological Society, 64:245-250.
Rojo S, Hopper KR, Marcos-Garcfa M-A, 1996. Fitness of the hover flies Episyrphus balteatus and Eupeodes corollae faced with limited prey. Entomol. Exp. Appl., 81:53-59.
Rybycki EP, Wechmar VMB, 1982. Characterisation of an Aphid-transmitted virus disease of small grains. Phytopath., 103.
Sarafrazi AM, Ahmadi AA, 1993. Components of resistance to Russian wheat aphid, Diuraphis noxia (Kurdjimov) (Homoptera: Aphididae) in small grains in Fars province. Iran Agricultural Research, 12:79-97.
Schaalje GB, Butts RA, Lysyk TJ, 1991. Simulation studies of binomial sampling: a new variance estimator and density predictor, with special reference to the Russian wheat aphid (Homoptera: Aphididae). Journal of Economic Entomology, 84(1):140-147
Schroeder-Teeter S, Zemetra RS, Schotzko DJ, Smith CM, Rafi M, 1994. Monosomic analysis of Russian wheat aphid (Diuraphis noxia) resistance in Triticum aestivum line PI137739. Euphytica, 74(1/2):117-120
Seco Fernandez MV, Nunez Perez E, Duenas Santero E, Nieto Nafria JM, 1992. Situation in the north of Spain of the Russian cereal aphid Diuraphis noxia (Mordvilko) (Hom. Aphididae). Georgica, No. 1:9-24
Smith CM, Belay T, Stauffer C, Stary P, Kubeckova I, Starkey S, 2004. Identification of Russian wheat aphid (Homoptera: Aphididae) populations virulent to the Dn4 resistance gene. Journal of Economic Entomology, 97(3):1112-1117. http://www.esa.catchword.org
Smith CM, Chuang WP, 2013. Plant resistance to aphid feeding: behavioral, physiological, genetic and molecular cues regulate aphid host selection and feeding. Pest Management Science, 70:528-540.
Smith CM, Schotzko DJ, Zemetra RS, Souza EJ, 1992. Categories of resistance in plant introductions of wheat resistant to the Russian wheat aphid (Homoptera: Aphididae). Journal of Economic Entomology, 85(4):1480-1484
Stary P, 1996. The expansive Russian wheat aphid, Diuraphis noxia (Mordw.) detected in the Czech Republic. Anzeiger fu^umlaut~r Scha^umlaut~dlingskunde, Pflanzenschutz, Umweltschutz, 69(1):19-20; 8 ref.
Stoetzel BM, 1987. Information on and identification of Diuraphis noxia (Homoptera: Aphididae) and other aphid species colonizing leaves of wheat and barley in the United States. J. Econ. Entomol., 80:696-704.
Swanevelder ZH, Surridge AKJ, Venter E, Botha AM, 2010. Limited endosymbiont variation in Diuraphis noxia (Hemiptera: Aphididae) biotypes from the United States and South Africa. Journal of Economic Entomology, 103(3):887-897. http://esa.publisher.ingentaconnect.com/content/esa/jee/2010/00000103/00000003/art00039
Tanigoshi LK, Pike KS, Miller RH, Miller TD, Allison D, 1995. Search for, and release of, parasitoids for the biological control of Russian wheat aphid in Washington State (USA). Agriculture, Ecosystems & Environment, 52(1):25-30
Thieme T, Heimbach U, Schliephake E, 2001. Detection of the 'Russian wheat aphid', Diuraphis noxia (Kurdjumov), in Germany. (Nachweis der 'Russischen Weizenlaus', Diuraphis noxia (Kurdjumov), in Deutschland.) Nachrichtenblatt des Deutschen Pflanzenschutzdienstes, 53(2):35-40.
Thomas JB, Butts RA, 1990. Effect of Russian wheat aphid on cold hardiness and winterkill of overwintering winter wheat. Can. J. Plant Sci., 70:1033-41.
Tolmay V, Pinsloo G, 1994. Russian wheat aphid (Diuraphis noxia) in South Africa. In: Proceedings of the sixth Russian Wheat Aphid Workshop, 23 January 1994, 182-184.
Tolmay VL, Lindeque RC, Prinsloo GJ, 2007. Preliminary evidence of a resistance-breaking biotype of the Russian wheat aphid, Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae), in South Africa. African Entomology, 15(1):228-230. http://journals.sabinet.co.za/essa
Tuatay N, Remaudiere G, 1964. Premiere contribution au catalogue des Aphididae (Hom.) de la Turquie. Rev. de Path. Veg. et Ent. Agr. de Fr., 43:243-78.
van der Westhuizen M, de Jager J, Manson M, Deall M, 1994. Cost benefit analysis of Russian wheat aphid (Diuraphis noxia) control in South Africa using a seed treatment. Seed treatment: progress and prospects: Proceedings of a Symposium held at the University of Kent, 5 January 1994. Farnham, UK: British Crop Protection Council Registered Office, 109-116.
Venter E, Mansoor CV, Sibisi P, Botha AM, 2014. Potassium phosphate induces tolerance against the Russian wheat aphid (Diuraphis noxia, Homoptera: Aphididae) in wheat. Crop Protection, 61:43-50. http://www.sciencedirect.com/science/journal/02612194
von Wechmar M, Rybicki EP, 1981. Aphid transmission of three viruses causes Freestate Streak disease. S. African Journal of Science, 77:488-92.
Walker CB, Peairs FB, 1994. Cultural control of Russian wheat aphid. Great Plains Agric Council Publ., 42-52.
Walker CB, Peairs FB, Echols J, 1990. Applications of nitrogen in spring and its effect on Russian wheat aphid infestations in winter wheat. In: Proceedings Fourth Russian Wheat Aphid Workshop, 10 October 1990. Bozeman, Montana: Montana State University,
Walker CB, Peairs FB, Echols J, 1991. Applications of nitrogen in spring and its effect on Russian wheat aphid in winter whaet. In: Proceedings 4th Russain Wheat Aphid Workshop, 10-12 October 1990. Bozeman, Montana, USA: Montana State University, 81-88.
Walters MC, Penn F, Du Toit F, Botha TC, Aalbersberg K, Hewitt PH, Broodryk SW, 1980. The Russian wheat aphid. Farming in South Africa, Leafl. Ser., Wheat G.3:1-6.
Webster JA, 1990. Resistance in Triticale to the Russian wheat aphid (Homoptera: Aphididae). Econ. Entomol., 83:1091-5.
Webster JA, Amosson S, 1994. Economic impact of the Russian wheat aphid in the western United States: 1992-1993. Great Plains Agr. Council Publ. No. 152.
Webster JA, Porter DR, Baker CA, Mornhinweg DW, 1993. Resistance to Russian wheat aphid (Homoptera: Aphididae) in barley: effects on aphid feeding. Journal of Economic Entomology, 86(5):1603-1608; 20 ref.
Webster JA, Starks KJ, Burton LR, 1987. Plant resistance studies with Diuraphis noxia (Homoptera: Aphididae), a new United States wheat pest. J. Econ. Entomol., 80:944-9.
Webster JA, Toit F du, Popham TW, 1993. Fecundity comparisons of the Russian wheat aphid (Homoptera: Aphididae) in Bethlehem, South Africa, and in Stillwater, Oklahoma. Journal of Economic Entomology, 86(2):544-548
Weiland AA, Peairs FB, Randolph TL, Rudolph JB, Haley SD, Puterka GJ, 2008. Biotype diversity in Colorado Russian wheat aphid (Hemiptera: Aphididae) populations. Journal of Economic Entomology, 101:569-574.
Wilbert H, 1964. Das Ausleserverhalten von Aphelinus semiflavus Howard und die Abwehreaktionen seiner Wirte (Hymenoptera: Aphelinidae). Beitr. Entomol., 14:159-221.
Wraight SP, Poprawski WL, Meyer WL, Peairs FB, 1993. Natural enemies of Russian wheat aphid (Homoptera: Aphididae) and associated cereal aphid species in spring-planted wheat and barley in Colorado. Environmental Entomology, 22(6):1383-1391
Yu GY, Liang HB, 1998. Coccinellidae collected in the wheat fields of Xinjiang, China with description of a new species (Coleoptera). Entomotaxonomia, 20(2):127-132.
Zhang G, 1991. Russian wheat aphid (RWA) in China. Stillwater, USA: Misc Publ Agric Exp Stn Okla State Univ., 132:327-328.
Zuniga SE, 1990. Biological control of cereal aphids in the southern zone of South America. In: World Perspectives on Barley Yellow Dwarf. Mexico, DF: CIMMYT, 362-367.
ContributorsTop of page
07/11/15 Impact and Prevention and Control sections updated by:
Mohammad Reza Nematollahi, Assistant Professor of Entomology, Department of Plant Protection, Isfahan Research Center for Agriculture and Natural Resources, Isfahan, Iran.
Distribution MapsTop of page
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