Hymenopteran parasitoids and dipteran predators of the invasive aphid Diuraphis noxia after enemy introductions: Temporal variation and implication for future aphid invasions

Shifts in prevalence and abundance of hymenopteran parasitoids and dipteran predators, Diuraphis noxia, and other aphids were measured in the west-central Great Plains of North America, April–September, in 2001 and 2002, corresponding to over a decade after first detection of D. noxia and first release of D. noxia enemies. Significant temporal shifts in enemy species prevalence and diversity were detected in this study and more broadly during an 11 year time span. At any given time, some species were relatively common. One parasitoid had been predominant throughout (Aphelinus albipodus), two had shifted in dominance (Lysiphlebus testaceipes and Diaeretiella rapae), three parasitoids had been detected infrequently (Aphidius avenaphis, Aphidius matricariae, and Aphelinus asychis), one parasitoid was detected in the 1990s but not during 2001 and 2002 (Aphelinus varipes), two predatory flies occurred at occasional significant levels (Leucopis gaimarii and Eupeodes volucris), and two parasitoids may have been minor members of the fauna (Aphidius ervi and Praon yakimanum). Aphid populations detected were usually very low or not detected, precluding estimation of percent parasitism. The best evidence of suppression was observations of parasitoids in the rare case of D. noxia exceeding economic thresholds, which complemented past studies using high aphid densities. The D. noxia enemies detected were primarily endemic or long-time residents derived from previous introductions. This enemy community may provide flexibility in responding to a future aphid invasion, allowing more strategic use of biological control and other pest management approaches.     

Effect of wheat resistance, the parasitoid Aphidius rhopalosiphi, and the entomopathogenic fungus Pandora neoaphidis, on population dynamics of the cereal aphid Sitobion avenae

The influence of wheat (Triticum aestivumL.) resistance, the parasitoid Aphidius rhopalosiphiDe Stephani-Perez (Hymenoptera: Braconidae) and the entomopathogenic fungus Pandora neoaphidis(Remaudière et Hennebert) Humber (Zygomycetes: Entomophthorales) on the density and population growth rate of the cereal aphid Sitobion avenae(F.) (Hemiptera: Aphididae) was studied under laboratory conditions. Partial wheat resistance was based on hydroxamic acids, a family of secondary metabolites characteristic of several cultivated cereals. The partial resistance of wheat cultivar Naofén, the action of the parasitoid and the joint action of the parasitoid and fungus, reduced aphid density. The lowest aphid densities were obtained with the combination of the parasitoid and the fungus, but wheat resistance under these circumstances did not improve aphid control. Significant reductions of population growth rate (PGR) of aphids were obtained with the joint action of wheat resistance and natural enemies. In particular, the combined effects of parasitoids and fungi showed significantly lower PGR than the control without natural enemies in both wheat cultivars. Our results support the hypothesis that wheat resistance and the utilization of biological control agents could be complementary strategies in an integrated pest management program against cereal aphids.     

Interactions of Russian wheat aphid, a hymenopterous parasitoid and resistant and susceptible slender wheatgrasses

Interactions among three trophic levels of resistant and susceptible slenderwheat grasses, Elymus trachycaulum (Link) Goule ex Shinners ex. H.F. Lewis, Russian wheat aphid, Diuraphis noxia (Mordvilko), and a hymenopterous parasitoid were studied in the laboratory and greenhouse. These relationships were compared with a commercial susceptible wheat Triticum aestivum L. variety. Aphids reared on the resistant entries showed significantly lower weights and numbers. Significant reduction of parasitoid mummy weight and adult size was positively correlated with the effects on the aphids. Resistant entries also induced a longer prereproductive period for both the aphids and parasitoids. Numbers of aphids and aphid damage were significantly modified by the addition of parasitoids. Parasitism was higher on plants that did not have leaf rolling. These findings may indicate that antibiosis resistance studied here is not the most desirable because it decreases natural enemy vitality.     

Aphelinus albipodus(Hymenoptera: Aphelinidae) andDiaeretiella rapae(Hymenoptera: Braconidae) Parasitism onDiuraphis noxia(Homoptera: Aphididae) Infesting Barley Plants Differing in Plant Resistance to Aphids

Russian wheat aphid,Diuraphis noxia(Mordvilko), as a pest of small grains, has prompted research into biological control and host plant resistance. In the presence of Russian wheat aphid, leaves of a susceptible barley (Morex) are curled and chlorotic and sustain large densities of this aphid, while leaves of a resistant barley (STARS-9301B) remain flat and green and sustain fewer aphids. Might parasitism of Russian wheat aphid byAphelinus albipodusHayat & Fatima andDiaeretiella rapaeMcIntosh be affected differently by these plant types? When presented the plants separately and based on parasitism rate relative to aphid density, the largerD. rapaewas more effective in parasitizing relatively high densities of aphids within curled leaves of Morex than relatively low densities of aphids on uncurled leaves of STARS-9301B. Parasitism byA. albipodusdid not significantly differ among the plants. When given a choice of plants, approximately equal rates of parasitism occurred on the two plant lines for both parasitoid species, and parasitism byD. rapaewas greater thanA. albipodus.These data indicate that using parasitoid size as an indicator of success in a physically restricted environment may be misleading, when considered in a plant environment responsive in several manners to aphids (chlorosis, curling, and ability to sustain Russian wheat aphid). We expect that use of resistant barley will result in decreased parasitoid abundance as aphid densities decrease. However, parasitism rates are expected to be approximately equal on resistant and susceptible barley. In this system, plant resistance and biocontrol are compatible management strategies.     

Hymenopteran parasitoids and dipteran predators of <Emphasis Type="Italic">Diuraphis noxia</Emphasis> in the west-central Great Plains of North America: Species records and geographic range

Parasitoids and predatory flies were sampled in the wheat production region of the west-central Great Plains (southeastern Wyoming, western Nebraska, and north-central Colorado) of North America using plant material infested with the Russian wheat aphid, Diuraphis noxia (Mordvilko) (Hemiptera: Aphididae). Samples were taken April through October in 2001 and 2002, which was 15–16 years after first detection of D. noxia and 5–6 years after the last release of natural enemies for its control in this region. The natural enemies detected were (in order of high to low detection frequencies across three states and 2 years): Aphelinus albipodus Hayat and Fatima (Hymenoptera: Aphelinidae), Eupeodes volucris Osten Sacken (Diptera: Syrphidae), Lysiphlebus testaceipes (Cresson) (Hymenoptera: Braconidae, Aphidiinae), Leucopis gaimarii Tanasijtshuk (Diptera: Chamaemyiidae), Aphidius avenaphis (Fitch), Aphidius matricariae Haliday, Diaeretiella rapae (MIntosh), Aphidius ervi Haliday, Praon yakimanum Pike and Starý (Hymenoptera: Braconidae, Aphidiinae), and Aphelinus asychis Walker (Hymenoptera: Aphelinidae). The results confirmed establishment of one of the 10 exotic parasitoid species released for D. noxia control (A. albipodus) in the west-central Great Plains. It is unknown whether detection of A. asychis, A. matricariae, and D. rapae can be attributed to exotic introductions or preexisting populations. Other species detected in this study have been previously documented from the western US, although the recognized distributions have expanded for A. avenaphis, L. gaimarii, and P. yakimanum compared to the first few years after initial detection of D. noxia. Thus, there is definitive establishment of one exotic introduced for D. noxia and considerable range expansion of preexisting species that prey upon D. noxia.     

Comparison of categories of resistance in wheat and barley genotypes against biotype 2 of the Russian wheat aphid, Diuraphis noxia (Kurdjumov)

The Russian wheat aphid Diuraphis noxia (Kurdjumov) (Homoptera: Aphididae) is a global pest of wheat and barley. This arthropod is difficult to manage with pesticides or biological control agents due to the aphid’s ability to seek shelter in rolled leaves and also to develop virulent biotypes. During the past 20 years, the use of aphid-resistant cereal cultivars has proven to be an economically and ecologically beneficial method of protecting crops from D. noxia damage. Our research reports the results of experiments to determine the categories of D. noxia biotype 2 resistance present in Cereal Introduction Triticeae (CItr) 2401, and a barley genotype (IBRWAGP4-7), compared to control resistant and susceptible wheat and barley genotypes. CItr2401 and IBRWAGP4-7 exhibit no antixenosis, but both genotypes demonstrated antibiosis to D. noxia in the form of reduced aphid populations. Reduced leaf dry weight change, a measure of plant tolerance of D. noxia feeding, was significantly less in CItr2401 and IBRWAGP4-7 plants than in plants of susceptible control varieties. However, tolerance was negated when a tolerance index was calculated to correct for differences in aphid populations. Barley IBRWAGP4-7 is a new source of D. noxia biotype 2 resistance. D. noxia foliar leaf damage and population growth were significantly less on IBRWAGP4-7 plants than on plants of the susceptible barley variety Morex. IBRWAGP4-7 plants were equal in resistance to plants of the resistant barley STARS 9301 and wheat genotype CItr2401. Handling editor: Heikki Hokkanen     

Survey of the parasitoids of the tuliptree aphid,Illinoia liriodendri (Hom: Aphididae), in northern California

A survey of the parasitoids ofIllinoia liriodendri (Monell) in northern California conducted from 1988–1990 revealed the presence of 12 primary and 14 hyper-parasitoid species. The most common primary parasitoid wasAphidius polygonaphis (Fitch), which was imported from the eastern United States in the 1970's and is now established throughout the area. New host records were noted forA. ervi Haliday,A. avenaphis (Fitch), Praon occidentale Baker,P. unicum Smith,Diaeretiella rapae M'Intosh,Lysiphlebus testaceipes (Cresson), andMonoctonus nervosus (Haliday) (all Hymenoptera: Braconidae: Aphidiinae), andAphelinus sp. nr.asychis Walker (Hymenoptera: Aphelinidae). The most common hyperparasitoid species werePachyneuron aphidis (Bouché) andAsaphes californicus Girault (both Hymenoptera: Pteromalidae). New hyperparasitoid host records were noted forPachyneuron californicum Girault on Aphidiine and Aphelinidae spp. andCoruna clavata Walker (Hymenoptera: Pteromalidae) onAphelinus sp.     

Ecology of the aphid pests of protected pepper crops and their parasitoids

Myzus persicae, Macrosiphum euphorbiae, Aphis gossypii and Aulacorthum solani (Homoptera: Aphididae) are principal pests of protected pepper crops in southeastern Spain. Our goal was to determine the incidence of aphids on pepper crops and the role of vegetation surrounding greenhouses as a source of aphids and their parasitoids. The population dynamics were followed in six commercial greenhouses during 3 years. Another 82 greenhouses and their surrounding vegetation were surveyed occasionally. Myzus persicae had the highest incidence in pepper greenhouses followed by M. euphorbiae and A. solani. Parasitism of all aphid species in greenhouses was low, Aphidius matricariae and Aphidius colemani being the most abundant parasitoids. Myzus persicae and Macrosiphum euphorbiae were the most abundant and polyphagous aphids, being present on 77 and 55% of the plants sampled outside greenhouses, respectively; species of Brassicaceae were the main hosts for both aphids. Aulacorthum solani was only present on Malva parviflora and at low numbers. Outside greenhouses, A. matricariae was the most common parasitoid of M. persicae, followed by Diaeretiella rapae and A. colemani. Aphidius matricariae was the most polyphagous, being present in 10 out of 22 aphid species. Macrosiphum euphorbiae and A. solani were both parasitised by A. ervi and Praon volucre. Aphelinus asychis was found on A. solani. Parasitoids were found in other aphids not attacking pepper. The role of natural vegetation as a reservoir of aphid pests of pepper and of parasitoids is discussed.     

Electrophoresis as a tool for estimating levels of hymenopterous parasitism in field populations of the cereal aphid, Sitobion avenae

Polyacrylamide gel electrophoresis was used to monitor levels of parasitism in field populations of the cereal aphid, Sitobion avenae (F.) (Hemiptera: Aphididae). Electrophoretic keys previously prepared from laboratory-reared parasitoids were used to identify parasitoids within field-collected aphids. Using this approach, the primary parasitoids Aphidius rhopalosiphi De Stefani Perez, Aphidius ervi (Haliday), Aphidius picipes (Nees), Praon volucre Haliday, Ephedrus plagiator Nees and Aphelinus abdominalis Dalman were detected. Some unidentified banding patterns were also obtained, most of which were probably attributable to the presence of hyperparasitoids. Electrophoresis was compared with the more conventional sampling techniques of live-rearing and mummy counts as a means of estimating percentage parasitism. Electrophoresis was found to be a much quicker method than live-rearing and gave similar results for both levels of parasitism and parasitoid species composition, whereas mummy counts tended to give lower estimates than the other two methods.

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Résumé L'électrophorèse sur gel de polyacrylamide a servi à déterminer l'importance du parasitisme de populations naturelles de S. avenae F. Des clés électrophorétiques, préparées à partir de parasitoïdes élevés au laboratoire, ont été utilisées pour identifier les parasitoïdes provenant des pucerons récoltés dans la nature. Ainsi ont été décelés les parasites primaires: Aphidius rhopalosiphi De Stefani Perez, A. ervi Haliday, A. picipes Nees, Praon volucre Haliday, Ephedrus plagiator Nees et Aphelinus abdominalis Dalman. Quelques bandes non-identifiées devraient être attribuées à des hyperparasitoïdes. L'électrophorèse a été comparée aux techniques d'échantillonnage plus conventionnelles de décompte des momies et d'élevage des individus vivants utilisées pour estimer le taux de parasitisme. L'électrophorèse et les méthodes d'élevage donnent généralement des résultats semblables pour les 2 niveaux de parasitisme et la composition en espèces de parasitoïdes, tandis que le décompte des momies tend à donner des résultats inférieurs.

     

The temperature-dependent duration of development and parasitism of three cereal aphid parasitoids, Aphidius ervi, A. rhopalosiphi, and Praon volucre

Temperature dependencies were established for the egg-to-mummy and mummy-to-adult phases, for mummy mortality, and for parasitism of Aphidius ervi Haliday, Aphidius rhopalosiphi De Stefani-Perez, and Praon volucre (Haliday) (Hymenoptera, Aphidiidae), three parasitoids of Sitobion avenae (Fabricius) (Homoptera, Aphididae), at 8°C, 12°C, 16°C, 20°C, and 25°C on winter wheat (cv. Haven). A physiological model described temperature-dependent development over the full temperature range, whereas a linear model was fitted for data above 8°C and used to estimate the lower temperature thresholds and day-degrees (° D) required for development. The thresholds for A. ervi were 2.2°C for egg-mummy development and 6.6°C for mummy-adult development, those for A. rhopalosiphi were 4.5°C and 7.2°C, and those for P. volucre were 3.8°C and 5.5°C. The time to develop into mummies and adults differed significantly between the three species: A. ervi development into mummies required an average of 159 ° D, while development into adults took an average of 73 ° D. The corresponding average times required for A. rhopalosiphi and P. volucre to develop mummies were 124° D and 126° D, while their development into adults required an average of 70° D and 150° D, respectively. Mummy mortality was 25–35% at 8°C and less at the higher temperatures tested, but began to increase again at 25°C, showing a quadratic relationship between mortality and temperature. Parasitization was very low or, in the case of P. volucre, absent up to 12°C and thereafter increased with increasing temperature. The relationship between parasitization, recorded as percent aphids mummified, and temperature was linear at the temperatures tested and depended on species. A. ervisuperparasitized 11.1% aphids at 20°C and 16.6% aphids at 25°C, whereas superparasitism was low in A. rhopalosiphi and absent in P. volucre. From 16°C to 25°C the P. volucre sex ratio increased. For A. ervi and A. rhopalosiphi there was no trend with temperature, but at 20°C and 25°C it was close to even. Field data for 1996 and 1997 allowed for a comparison of actual and expected emergence of overwintering mummies. In both years, parasitoids were predicted to have emerged from overwintering mummies well in advance of the onset of aphid infestation, and more than a month earlier than the first parasitized aphids were found in winter wheat. Observations from trap plants in other crops supported the predictions of the models. Other factors that can affect biological control by cereal aphid parasitoids are discussed.     

Prevalences of fungal pathogens and othernatural enemies of cereal aphids (Homoptera:Aphididae) in wheat under dryland and irrigatedconditions in South Africa

Surveys were conducted in the summer andwinter rainfall wheat producing regions of SouthAfrica in a first attempt to investigate theidentity and impact of entomopathogenic fungi withinthe cereal aphid complex. Wheat produced underdryland and irrigated conditions was surveyed duringthe 1996 and 1997 seasons. Six cereal aphid specieswere recorded of which the Russian wheat aphid, Diuraphis noxia, was the most abundantunder dryland conditions in the summer rainfallregion as opposed to the oat aphid, Rhopalosiphum padi, in the winter rainfallregion. Rose grain aphid, Metopolophiumdirhodum, was most prevalent underirrigated conditions in the summer rainfall region.Five species of entomopathogenic fungi were recordedincluding four entomophthorales and the hyphomycete,Beauveria bassiana. TheEntomophthorales included Pandora neoaphidis, Conidiobolus obscurus, C.thromboides, and Entomophthoraplanchoniana. Pandora neoaphidis wasthe most important etiological agent recorded fromD. noxia, with up to 50% mycosis recordedunder dryland conditions in the Bethlehem summerrainfall region. Similarly, P. neoaphidis wasthe most prevalent species within populations ofM. dirhodum. under irrigated conditions in theBergville/Winterton summer rainfall region (up to77% mycosis). However, mycoses of R. padi didnot exceed 1.7% in samples from these areas,suggesting that R. padi may be lesssusceptible to P. neoaphidis than M.dirhodum. Epizootics in populations of D.noxia under dryland conditions in both the winterand summer rainfall regions indicated a high levelof susceptibility to P. neoaphidis.Occurrences of hymenopterous parasitoids andpredators in populations of D. noxia were low,although a parasitism level of 25% was recorded inone small sample of R. padi collected from anirrigated field in the summer rainfall region.     

A survey of aphids and aphid parasitoids in cereal fields in Denmark, and the parasitoids' role in biological control

In 1996 and 1997 a field survey of the abundance and species composition of cereal aphid primary and secondary parasitoids in spring barley, winter wheat and durum wheat was conducted in Zealand, Denmark. The purpose was to create a better understanding of the mechanisms underlying aphid–parasitoid dynamics in the field. Such an understanding can be used when developing biological control methods in cereals. In both years aphid attacks in cereals began in late June and never exceeded the economic threshold. In 1996 the first aphids were found in wheat on 26 June; in 1997 the first aphids were found on 24 June on both crops. The highest densities reached in 1996 were an average of six aphids per shoot in winter wheat and one aphid per shoot in spring barley. In 1997 the highest densities reached were 11 aphids per shoot in winter wheat and four aphids per shoot in spring barley. The aphid population collapsed by the end of July to early August in 1996, but it collapsed by mid-July in 1997. The onset and peak of parasitization were delayed in comparison to aphid infestation. Parasitism was 20–60% by the end of the cropping season in spring barley, and 30–80% in winter wheat and durum wheat in 1996. In 1997 parasitism did not exceed 3–11% in barley and was less than 2% in one winter wheat field but more than 40% in the other winter wheat field sampled. In both years most parasitism was due to Aphidiidae (Hymenoptera). The two dominant species were Aphidius ervi Haliday and Aphidius rhopalosiphi De Stefani-Perez. Hyperparasitism began after primary parasitism and increased progressively during the cropping season. The two years were similar in many respects, including for species composition of aphids and parasitoids. The late start of the aphid infestation may have contributed to the high level of parasitization found in 1996, but in 1997 the aphid infestation period was so short that a parasitoid population did not have time to build up.     

Influence of wheat and oat cultivars on the development of the cereal aphid parasitoid Aphidius rhopalosiphi and the generalist aphid parasitoid Ephedrus plagiator

The effects of three wheat cultivars and two oat cultivars on the development of the cereal aphid parasitoid Aphidius rhopalosiphi De Steph. and the generalist aphid parasitoid Ephedrus plagiator (Nees) (Hymenoptera: Braconidae) were evaluated in the laboratory. The level of hydroxamic acids, a family of secondary metabolites that can affect the mean relative growth rate of cereal aphids in cereals, were measured in the different cultivars. The parasitoids were reared in Sitobion avenae (F.) (Homoptera: Aphididae), using plants grown under greenhouse conditions. A. rhopalosiphi showed a longer developmental time on wheat relative to oat cultivars. This effect was accounted for by a significant increase in the time from oviposition to pupation (mummy formation), while the duration of the pupal stage remained constant between treatments. No further effects were observed in other variables evaluating A. rhopalosiphi performance, such as adult longevity, adult body weight and secondary sex ratio. The generalist E. plagiator did not show significant differences in any of the variables analysed, both between cultivars and cereal species. Hydroxamic acids levels correlated negatively with mean relative growth rates of S. avenae, but positively with the observed developmental time of A. rhopalosiphi. The results are discussed in terms of tritrophic effects and the development of breeding programmes trying to improve plant resistance to aphids.     

Possible roles of esterase, glutathione S-transferase, and superoxide dismutase activities in understanding aphid–cereal interactions

Activities of the detoxification enzymes esterase, glutathione S‐transferase, and of superoxide dismutase in aphids and aphid‐infested cereal leaves were assayed using polyacrylamide gel electrophoresis and a spectrophotometer to elucidate the enzymatic mechanisms of aphid resistance in cereal plants. A chlorosis‐eliciting Russian wheat aphid, Diuraphis noxia (Mordvilko), and non‐chlorosis‐eliciting bird cherry‐oat aphid, Rhopalosiphum padi (L.), and four cereals were used in this study. The four cereal genotypes were ‘Arapahoe’ (susceptible) and ‘Halt’ (resistant) wheat (Triticum aestivum L.), ‘Morex’ (susceptible) barley (Hordeum vulgare L.), and ‘Border’ (resistant) oat (Avena sativa L.). Esterase isozymes differed between the two aphid species, although glutathione S‐transferase and superoxide dismutase did not. Esterase, glutathione S‐transferase, and superoxide dismutase activities in either aphid species were not affected by the level of resistance of a cereal to D. noxia. The assays of cereal leaf samples showed that D. noxia feeding elicited an increase in esterase activity in all four cereal genotypes, although R. padi feeding did not. The increase of esterase activity in cereals, however, was not correlated to aphid resistance in the cereals. The time‐series assays of aphid‐infested cereal leaves showed that D. noxia‐infested Morex barley had a significant increase in esterase activity on all sampling dates (3, 6, and 9 days) in comparison with either uninfested or R. padi‐infested barley. No difference in glutathione S‐transferase activity was detected among either aphid infestations or sampling dates. The electrophoretic assays, however, revealed that aphid feeding elicited a significant increase in superoxide dismutase activity, which served as the control of glutathione S‐transferase activity assays. The increase in esterase and superoxide dismutase activities suggested that D. noxia feeding imposes not only toxic, but also oxidative stresses on the cereals. The ramification of using these enzyme activity data to understand the etiology of D. noxia‐elicited chlorosis is discussed.     

Supercooling capacity of Eurasian and North American populations of parasitoids of the Russian wheat aphid, >Diuraphis noxia

Supercooling points were estimated for seven populations of >Aphelinus albipodus, five populations of >Aphelinus asychis, and four populations of >Diaeretiella rapae to assess whether their supercooling points were sufficiently low to provide the potential for overwintering survival in colder temperate climatic areas. Test individuals from all 16 of the parasitoid populations were collected originally from mummies of the Russian wheat aphid, >Diuraphis noxia. Mummies containing parasitoid pupae were maintained for 1 wk under three different temperature conditions (treatments): at room temperature (24.8 ± 0.2 °C), 1 wk at 0 °C, and 1 wk –5 °C, and the supercooling points across treatments, and within and among species were compared. Statistical differences in supercooling points were found among populations of >A. albipodus for each treatment, and for >A. asychis when maintained for 1 wk at room temperature. No differences in supercooling points were found among populations of >D. rapae mummies maintained under the three temperature treatments. The lowest supercooling points obtained for the three parasitoid species maintained at room temperature were the >A. albipodus population from Montana (–31.68 °C), the >A. asychis population from Greece (–32.04 °C), and the >D. rapaepopulation from the Caucasus (–33.12 °C). Preconditioning the parasitoid mummies to cold had no effect on the supercooling points for >A. albipodus, and in some cases unexpectedly increased the supercooling points for >A. asychisand >D. rapae. In comparing the overall mean supercooling points of the three parasitoid species, no differences were found within species (among temperature treatments), nor among species (within temperature treatments). It was concluded that observed differences in supercooling points of only a few degrees Centigrade among parasitoid populations and species would not be expected to cause differences in their overwintering success, especially given the expected variability in temperatures within and among overwintering sites.     

Effect of honeydew and hosts on plant colonization by the aphid parasitoidEphedrus cerasicola

Females ofEphedrus cerasicola Stary were released into small glasshouses (13.6 m³) or cages (0.125 m³) which contained paprika plants in 2 of the following categories: fresh plants (without honeydew and aphids), honeydew-contaminated plants (aphids removed) and aphid-infested plants (with honeydew). Two hundred females were released into each glasshouse with 20 plants, 10 of each category, while 10 females were released into the cages with 2 plants, one of each category. Control experiments with only fresh plants were conducted in both the glasshouses and the cages. The female parasitoids on each plant were counted at regular intervals for 24 or 48 h in the glasshouse and for 4 h in the cage experiments; this number was used as an indicator of plant preferences. The following preferences were established (preferred plant > less preferred plant): honeydew-plant > fresh plant, aphid-plant > fresh plant, aphid-plant > honeydew-plant. The number of females on the aphid-plants increased throughout the experimental period, whereas the number on the honeydew-plants tended to be more stable.      

Effects of honeydew and insecticide residues on the distribution of foraging aphid parasitoids under glasshouse and field conditions

The opposing effects of attraction to host-derived kairomones and repellency from the pyrethroid insecticide deltamethrin were investigated with aphid parasitoids from the genus Aphidius (Hymenoptera: Aphidiinae). The spatial distribution of female parasitoids was recorded in a series of experiments conducted in a small glasshouse containing wheat plants either infested with cereal aphids, Sitobion avenae (F.) (Homoptera: Aphididae), uninfested or treated with the recommended field concentration of deltamethrin. The number of parasitoids per plant were counted at 0.5 h, 1 h and then at one hourly intervals up to 8 h after release. Parasitoids showed a strong aggregation response to aphid-infested plants compared to adjacent uninfested plants. With the introduction of insecticidetreated plants around the aphid-infested plants, parasitoids showed a greater tendency to disperse away, resulting in fewer parasitoids on plants and significantly lower rates of aphid parasitism. The degree of aphid fall-off from plants was a good indicator of parasitoid foraging activity. In field studies, using sticky traps to measure the activity of parasitoids in plots sprayed with water, deltamethrin and/or an artificial honeydew solution, repellent properties were evident for up to 2 days after application. The attraction/arrestment stimuli associated with the honeydew solution were sufficient for parasitoids to continue searching insecticide-treated areas. The implications of these findings for parasitoids searching crops contaminated with aphid-derived kairomones and insecticides are discussed.     

Release and recovery in South Africa of the exotic aphid parasitoid <Emphasis Type="Italic">Aphelinus hordei</Emphasis> verified by the polymerase chain reaction

The exotic aphid parasitoid Aphelinushordei Kurdjumov (Hymenoptera: Aphelinidae) was released at five Russian wheat aphid [Diuraphis noxia (Mordvilko)] infested wheat fields in the eastern parts of the Free State Province of South Africa during the 1998 and 1999 growing seasons. Except for differences in the setae on the ventralside of the forewings, this species is verysimilar in colour and structure to A.varipes (Foerster) and A. albipodusHayat & Fatima, which also parasitise D.noxia. It is therefore difficult todistinguish between them, and to determinetheir establishment. Therefore a polymerasechain reaction, using specific primers for bothITS2 and mitochondrial 16s DNA sequencesof the three parasitoids, followed by HinfIrestriction endonuclease digestion and agarosegel electrophoresis, was tested to distinguishbetween them. Parasitoid recoveries were madeat the release sites within weeks after theywere released. Putative individuals of A.hordei were also collected during 1999 and2000 on D. noxia in Lesotho, whichis situated on the eastern border of the FreeState Province. The procedure separated the three Aphelinus spp.reliably. Between 94 and 100% of theindividuals recovered during the two fieldseasons were identified as being A.hordei, thus verifying recovery of this exoticaphid parasitoid in South Africa.     

Evidence for regional diversity and host adaptation in Iranian populations of the Russian wheat aphid

Effective pest management is greatly facilitated by knowledge of the genetic structure and host adaptation of the pest species in question. The Russian wheat aphid (RWA), Diuraphis noxia (Mordvilko) (Homoptera: Aphididae: Macrosiphini), is an important economic pest in many cereal‐growing areas of the world, and in this study we investigated these aspects of its populations, using microsatellite markers and host plant response assays. Diuraphis noxia was sampled from 38 locations in Iran and genotyped at four polymorphic microsatellite loci that had been isolated from various Sitobion species. We identified 50 multilocus genotypes in 376 individuals. The overall observed heterozygosity was 0.134. F‐statistics showed a regional partitioning in D. noxia populations with overall F_ST = 0.231. In addition, there was a significant correlation between genetic and geographic distances. In order to test for the ecological consequences of genetic variability in D. noxia, biotypic variation amongst the isolates collected from wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.) was evaluated on a number of resistant and susceptible wheat varieties. The plant variables we measured were damage rating (based on leaf chlorosis, leaf rolling, wilting, and death of the host plant), host plant dry weight, and root length. Damage rating was the best criterion for detecting biotypic variation in D. noxia. Discriminant analysis correctly classified the isolates in respective groups in 80–91.8% of the cases. The barley isolate showed no differences in performance on resistant and susceptible wheat, indicating a lack of gene‐by‐gene relationship with wheat plants. In contrast, wheat isolates differentially damaged the resistant and susceptible plants and showed moderate to severe virulence.     

Oviposition behaviour and patch-time allocation in two aphid parasitoids exposed to deltamethrin residues

Neurotoxic insecticides are widely used for crop protection. One consequence is that changes in behaviour can be expected in surviving beneficial insects because of an impairment of host perception and motor abilities. Under laboratory conditions, we studied the impact of deltamethrin, a pyrethroid, on the oviposition behaviour of two hymenopterous parasitoids of aphids, Aphidius matricariae (Haliday) and Diaeretiella rapae (McIntosh) (Hymenoptera: Braconidae). They both parasitize Myzus persicae (Sulzer) (Homoptera: Aphididae), which is the preferred host of A. matricariae, regardless of the host plant, whereas D. rapae is a major parasitoid of aphids on Cruciferae crops, including M. persicae. After exposure to deltamethrin, the different items of oviposition behaviour and the total time spent on the patch were recorded. The results showed that the patch time allocation by both parasitoid species was not significantly affected by deltamethrin treatment, when compared with the controls. Nor were the frequencies and sequences of behavioural items modified (e.g., frequency of sting). It therefore appeared that the patch use of A. matricariae and D. rapae on new colonies of M. persicae was not disturbed by deltamethrin at the three doses tested. The possibility that parasitoid strains are partially tolerant to deltamethrin is discussed.