The value of three cereal aphid species as food for a generalist predator

The value of the cereal aphid species Metopolophium dirhodum (Wlk.), Sitobion avenae (F.) and Rhopalosiphum padi (L.) as prey for the linyphiid spider Erigone atra (Bl.) was assessed. Fecundity of females was determined for spiders fed on eight experimental diets: three single‐species aphid diets, a mixed diet of all three aphid species, three mixed diets with each aphid species in combination with fruit flies Drosophila melanogaster (Meig.), and pure D. melanogaster as a high quality comparison diet. The development and survival of first‐instar juveniles fed on three diets of single aphid species, and on a diet of Collembola were compared with those subjected to starvation. Prey value for adult females was assessed by egg production, hatching success and offspring size. In pure diets all three aphid species were of low value to the spiders, causing a rapid decline in egg production and supporting no growth of significance of first‐instar juveniles. No difference in value of aphid species of single‐species aphid diets was found in the fecundity experiment, while a ranking of aphid species of M. dirhodum > R. padi > S. avenae was revealed in the survivorship experiment. A mixed‐aphid diet was not found to be advantageous compared with single‐species aphid diets, and no advantage of including aphids in mixed diets with fruit flies was found. Metopolophium dirhodum and R. padi were neutral in mixed diets, while a diet of S. avenae and fruit flies caused reduced egg production compared with the pure diet of fruit flies, revealing a toxic effect of S. avenae on the spider. The value‐ranking of aphid species in mixed diets was similar to that of single‐species diets. A similar ranking of aphid species was found for different fitness parameters (fecundity of adult females and development of juveniles). A ranking of aphids by offspring size of mothers on aphid‐only diets was S. avenae > M. dirhodum > R. padi. All aphid‐fruit fly diets resulted in larger offspring than a diet of only D. melanogaster, with the overall largest offspring being produced on the diet of M. dirhodum and fruit flies.     

Microcosm studies on intraguild predation between female erigonid spiders and lacewing larvae and influence of single versus multiple predators on cereal aphids

Abstract: Intraguild predation between female erigonid spiders [Erigone atra (Blackwall) and Oedothorax apicatus (Blackwall), Araneae, Erigonidae] and lacewing larvae (second instar larvae of Chrysoperla carnea (Stephens), Neuropt., Chrysopidae) and interaction effects of predator combinations on cereal aphids were investigated in a microcosm system under laboratory conditions. The microcosm experiments were run for 7 days and consisted of 15wheat seedlings, 15 Sitobion avenae (F) (Hom., Aphididae) as start population, plus a female spider or a lacewing larva or a combination of a spider plus a lacewing larva. The mortality rate of lacewing larvae was significantly increased by 44 and 31% due to intraguild predation by female spiders of E. atra and O. apicatus in comparison with lacewing larvae that were kept alone. The final aphid numbers in the microcosms were significantly reduced by all single predator treatments (spiders, lacewing larvae) and the predator combinations in comparison with controls without predators. The predation effect on aphid populations due to both spider species was similar and not statistically different. An additive effect of the predator combinations ‘spider plus surviving lacewing larva’ was found for both spider species resulting in reduced aphid numbers compared with the single predator treatments. When the lacewing larva was killed by an E. atra female the effects on aphids were non‐additive, but aphid numbers were not statistically increased compared with the lacewing larva treatment. When the lacewing larva was killed by an O. apicatus female, the effects of spider and C. carnea larva were additive on aphid numbers. In the presence of additional prey (fruit flies and Collembola) intraguild predation was not found and E. atra females had no significant effect on the survival of lacewing larvae. In addition, E. atra females had no significant effect on aphid numbers in the presence of fruit flies and Collembola, but in combination with a lacewing larva that survived, a significantly greater reduction of the aphid population was observed compared with the lacewing larva treatment. The body mass of lacewing larvae at the end of the experiment was not statistically influenced by the presence or absence of an E. atra female.     

Fly disturbance suppresses aphid population growth

1. The presence of predators is known to suppress prey populations not only by consumptive but also by non-consumptive effects, as it stresses the prey inducing costly changes to behaviour and physiology. However, there is recent evidence that disturbance from non-predacious, non-competing commensals can also negatively affect herbivore performance. 2. Populations of cherry-oat aphids (Rhopalosiphum padi L.) were initiated with adult aphids in mesocosms containing seedling wheat grass. Following aphid establishment, fruit flies (Drosophila melanogaster Meigen) were added at a relatively high density to half of the mesocosms and the aphids were left for another 5 days to reproduce. The experiment was performed over two blocks at 24 ± 4 °C. It was hypothesized that a relatively high density of commensals would stress the aphids and reduce their fitness, causing lower population growth and resulting in lower population sizes. 3. Aphid numbers were significantly lower in mesocosms with commensal flies after 5 days of fly presence across the two experimental blocks, documenting that fly disturbance suppresses aphid fitness and population growth. 4. The negative effect of fruit flies on aphid population growth must have come from the disturbance that flies imposed on the aphids in their search for food, indicating that the flies stressed the aphids. Thus, our study indicates that commensals may stress herbivores that do not distinguish between enemies and other active species in their environment, adding in the overall herbivore top-down control through fitness costs.     

Food quality of Ephestia eggs,the aphid Rhopalosiphum padi and mixed diet for Orius majusculus

We studied the food quality of the aphid Rhopalosiphum padi to the pirate bug Orius majusculus using Ephestia eggs as high-quality comparison prey. Several performance parameters were tested on individuals that had been reared and maintained on each of the two single-prey diets or on a mixed diet. All fitness parameters were lower in individuals fed aphids only, indicating poor food quality of this prey. Compared with the pure Ephestia egg diet, the mixed diet enhanced teneral mass, while adult survival and female starvation tolerance were negatively affected and all other traits were unaffected. Body protein proportions were constant across diets, whereas lipid proportion was low in the aphid treatment. Preference for aphids was lower following a monotypic aphid diet than when reared on Ephestia eggs or a mixed diet. The results confirm that R. padi is low-quality food for O. majusculus as it is for other generalist predators, even though O. majusculus may contribute significantly to population suppression of the aphid.     

Predation of Rhopalosiphum padi (Homoptera: Aphididae) by polyphagous predatory arthropods during the aphids' pre-peak period in spring barley

Polyphagous predators (e.g. Araneae, Carabidae and Staphylinidae), collected from spring barley fields during 1981-85, were examined by either gut dissection or a R. padi-specific antiserum in enzyme linked immunosorbent assay (ELISA) in order to detect predation of Rhopalosiphum padi during the aphids' establishment and exponential growth phases. Overall 18% of c. 3000 carabids dissected were shown to feed on R. padi during the aphids' pre-peak period. No overall relationship was found between percentage carabids with R. padi in the diet and peak R. padi densities. Relatively high proportions of Bembidion spp. (particularly B. lampros) and Pterostichus cupreus fed on R. padi during the aphids' establishment phase, and proportions of those predator taxa containing R. padi increased with increasing R. padi densities in both high and low aphid density years. P. melanarius and Harpalus rufipes mainly fed on R. padi during the aphids' exponential growth phase. Overall 11% of c. 1350 predators examined in ELISA gave positive reactions to the R. padi antiserum. Relatively high proportions of linyphiid and lycosid spiders were positive throughout the aphid pre-peak period. Several Acari, Opiliones, Trechus spp. (Carabidae), Philonthus spp. (Staphylinidae), Cantharidae and Chilopoda were positive mainly during the aphids' exponential growth phase. Sample sizes were small, however. Very few of the Bembidion spp. tested in ELISA were positive compared with those examined by gut dissection. The maximum period of R. padi protein (antigen) detection in B. lampros was related to temperature, i.e. 8.5 h at 30°C, 20.5 h at 20°C and 34.5 h at 10°C, respectively. It is suggested that the rate of R. padi protein digestion in B. lampros is faster at higher temperatures than the rate of elimination of prey solids from the guts. Several key predators (in this case B. lampros, P. cupreus and linyphiid spiders) which are abundant in spring cereal fields at a time when R. padi emigrants arrive in the crop and which feed on R. padi during the aphids' establishment phase, are identified. The results are compared with those from similar investigations elsewhere with predators of Sitobion avenae and Metopolophium dirhodum.     

Searching behaviour and cereal aphid consumption by Bembidion lampros and Pterostichus cupreus, in relation to temperature and prey density

Maximum consumption rates were determined for two carabids, Bembidion lampros Herbst. and Pterostichus cupreus L., feeding on the cereal aphid Rhopalosiphum padi L. at different temperatures in the laboratory. Mean daily consumption increased with increasing temperature for both species, B. lampros consuming a maximum of 16 1–3 instar nymphs and 9 apterous adult aphids at 25°C. P. cupreus was particularly voracious and consumed 125 apterous adult R. padi per day at 20°C. The behaviour of both species was analysed by video filming starved beetles, maintained at different constant temperatures, in arenas sown with spring barley. The behavioural components (1) still; (2) run/walk; (3) search and (4) confrontation were identified and were common to both species. P. cupreus was more active over the temperature range tested; B. lampros was inactive under 10°C. The proportion of time spent searching, number of plants searched, and velocity increased with increasing temperature for both species. When observed in similar arenas seeded with R. padi colonies, individuals of P. cupreus significantly increased their time spent searching in arenas with increasing aphid density. Following discovery of an aphid colony, individuals climbed and searched the host plant and its nearest neighbours. Plants in aphid free arenas were rarely climbed. B. lampros was not observed climbing in either aphid free arenas or in arenas with increasing aphid densities, and did not significantly increase its time spent searching in response to increased prey density. The few B. lampros that found aphids caught them walking on the soil surface. The relative efficiences of these two carabids as predators of R. padi are discussed, and the results are compared with similar studies elsewhere with predators of Sitobion avenae on winter wheat.

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Résumé Les taux maximum de consommation de R. padi L. à différentes températures ont été déterminés au laboratoire chez deux carabes, B. lampros Herbstet P. cupreus L. La consommation moyenne a augmenté avec la température chez les deux espèces, B. lampros consommant un maximum de 15,9 larves des stades 1 à 3 et 9,1 pucerons adultes aptères, à 25°C. P. cupreus a été particulièrement vorace et a consommé 125,3 adultes aptères par jour à 20°C. Le comportement des deux espèces a été observé en filmant en vidéo des carabes à jeun, maintenus à différentes températures constantes, dans des enceintes semées en orge de printemps. Des éléments du comportement, communs aux deux espèces, ont été définis: 1) immabilité, 2) marche et course, 3) recherche, 4) affrontement. P. cupreus a été plus actif à toutes les températures, B. lamprosa été inactif au-dessous de 10°C. La part de temps consacrée à la recherche, le nombre de plantes prospectées, et la vitesse ont augmenté avec la température chez les deux espèces. Dans des enceintes similaires colonisées par R. padi, P. cupreus a significativement augmenté le temps consacré à la recherche dans les enceintes, parallèlement à l'augmentation de la densité des pucerons. Après la découverte d'une colonie de pucerons, P. cupreus escalade et prospecte la plante et ses voisines immédiates; tandis que les plantes des enceintes sans pucerons sont rarement escaladées. B. lampros n'a pas été observé escaladant des plantes d'enceintes avec ou sans pucerons, et il n'a pas accru son temps de prospection en fonction de la densité de pucerons. Les quelques B. lampros qui ont capturé des pucerons l'ont fait lorsque ceux-ci marchaient sur la surface du sol. La discussion a porté sur l'efficacité relative des deux carabes comme prédateurs de R. padi, et les résultats ont été comparés à ceux d'études du même type, menées ailleurs, avec des prédateurs de Sitobion avenae sur blé d'hiver.

     

Consumption by carabid beetles of three cereal aphid species relative to other prey types

The cereal aphidRhopalosiphum padi has previously been found to be a low quality prey for a range of generalist arthropod predators. The aim of this study was to reveal, using food consumption experiments whether this applies to other cereal aphids. The question of whether predator feeding capacity increased when several aphid species were offered relative to a single aphid species was also addressed by measuring food consumption on a mixed aphid diet relative to single aphid diets. Food consumption by five carabid beetles of the three cereal aphid speciesRhopalosiphum padi, Sitobion avenae andMetopolophium dirhodum was determined relative to fruit fliesDrosophila melanogaster and the collembolanIsotoma anglicana. Feeding rate was measured as food consumption over 24 hour both for previously satiated and beetles starved for 7 days. Generally the largest aphid consumption was ofM. dirhodum and the lowest ofR. padi, withS. avenae in between. The mixed aphid consumption experiments did not reveal a higher feeding rate on mixed aphid diets relative to single aphid diets. The results indicate low preference forR. padi andS. avenae.     

Evaluation of lycosid,micryphantid and linyphiid spiders as predators ofRhopalosiphum padi (Hom.: Aphididae) and their functional response to prey density-laboratory experiments

Laboratory experiments were performed to determine the potential of dominant spider species in winter wheat in Germany,Erigone atra (Blackwall),Lepthyphantes tenuis (Blackwall) andPardosa agrestis (Westring) adults and youngs, in suppressing the population ofRhopalosiphum padi (L.) on wheat plants and their functional response to different aphid densities. The presence of spiders significantly caused between 34 and 58% reduction in aphid population development on wheat plants compared to the aphid population in the absence of spiders. The functional response curves for these spiders as predators ofR. padi seem to descrive a typical type II functional response with the prey consumed increasing to a plateau as aphid densities increased. Prey killed without eating was linear on prey density.     

Consumption of aphids by spiders and the effect of additional prey: evidence from microcosm experiments

Spiders are common generalist predators in agroecosystems and have been suggested to lower herbivore abundance in crops. It is not clear, however, if spiders can effectively suppress pest populations, and if so, by what mechanisms. In a microcosm experiment, we examined the consumption of the bird cherry-oat aphid, Rhopalosiphum padi L. (Homoptera: Aphididae), a pest species in wheat fields, by three spider species that differ in their hunting methods. We then tested the effect of additional prey type on the ability of erigonid spiders to reduce aphids. In a 48-h experiment Mermessus denticulatus (Banks) (Araneae: Linyphiidae; Erigoninae) consumed more aphids than did Enoplognatha gemina Bosmans and Van Keer (Araneae: Theridiidae) and Bathyphantes cf. extricatus (O·P.-Cambridge) (Araneae: Linyphiidae; Linyphiinae). This difference may be due to the ability of erigonids to forage actively on the vegetation in addition to using their webs to catch prey. In a 7-week experiment, we provided springtails (Collembola) in high and low densities as additional prey to mated erigonids, prior to aphid introduction. Spiders in the low-density springtail treatment built more webs on the vegetation, and caused a 50% reduction in aphid populations. There were significantly fewer aphids in the low-density springtail treatment, but not in the high-density treatment, in comparison to the control (high-density springtails without spiders). The results suggest that additional prey density affects predatory interactions between M. denticulatus and R. padi and that erigonids, which occur in high densities in wheat fields in the Negev desert, may be involved in aphid suppression in these agroecosystems.

Prey preference and egg production of the carabid beetleAgonum dorsale

In order to elucidate whether the aphidRhopalosiphum padi is low quality food for the carabid beetleAgonum dorsale, as it has previously been found to be for cereal spiders, we performed a series of experiments using fruit fliesDrosophila melanogaster as a standard alternative prey (‘control prey’): 1) Prey preference, 2) Aphid consumption for satiated and starved beetles, 3) Egg production on different diets and 4) Preference experiments with fruit flies coated with a taste of different prey types. Other alternative prey types used were earthworms and sciarid midges, all potential prey in the field.A. dorsale has a low preference for aphids compared to fruit flies. Apparently there is an upper limit to the consumption of aphids independent of hunger and much below the beetles' total food demand. Egg production on a pure diet of aphids is lower than on a pure diet of fruit flies; it is extremely low on a pure diet of earthworms; the highest fecundity is found on a mixed diet. As fruit flies coated with a taste of aphids or sciarid midges are less prefered than flies covered with a taste of fruit flies there may be a chemical factor, acting through taste, involved in determining prey preference.     

Predator density manipulation and its effects on populations of Rhopalosiphumpadi (Horn.: Aphididae) in spring barley

Experiments were carried out in 1981-83 to assess the impact of polyphagous predators (e.g. Carabidae, Staphylinidae and Araneae) on populations of cereal aphids in spring barley in Sweden. Barriers were erected around mid-field and field-edge plots at different times during the aphids' establishment phase in order to manipulate the predation pressure during the aphid pre-peak period; predators were emigrating from field boundaries. Rhopalosiphum padi was the dominant species of cereal aphid in 1981 and 1982, and was abundant in both years. Reduction of the predation pressure resulted in R. padi populations that were two to six times larger compared with those in unenclosed control plots. Aphid populations were much larger when the predation pressure was reduced earlier in June than when this was done later, in both years, since late erection of barriers excludes fewer predators because most had already entered the enclosed areas. In 1983, R. padi and Sitobion avenae were equally common but occurred in low numbers, and the effects of reducing the predation pressure were less obvious. Bembidion spp. (B. lampros and B. quadrimaculatum) and linyphiid spiders were the most abundant predators during the early establishment phase of aphid populations in each year, and significant inverse correlations were found between peak numbers of R. padi per shoot and numbers of predators from these taxa in 1981 and 1982. Similar relationships were found between total aphids and numbers of Staphylinidae, B. quadrimaculatum and Coccinella septempunctata in 1983. It was concluded that differences between the manipulated numbers of polyphagous predators caused the observed differences in peak population levels of R.padiin 1981,1982 and to a lesser extent, 1983. Apparent levels of parasitism and fungal disease were low during the pre-peak period. Aphid specific predators, abundant after the peak, were considered partly responsible for the decline of aphid populations each year. The results are compared with those from similar experiments elsewhere with S. avenae and Metopolophium dirhodum in winter wheat. Reasons why the effects of polyphagous predators are greater, and occur over a much shorter period of time in Sweden are discussed.     

Can polyphagous predators control the bird cherry-oat aphid (Rhopalosiphum padi) in spring cereals?: A simulation study

Predation by two carabid species,Bembidion lampros (Herbst.) andPterostichus cupreus L., on the cereal aphid,Rhopalsiphum padi L., was described using a discrete event simulation model. The results were incorporated into an aphid population growth model. The aphid model was run using actual temperatures,R. padi emigration rates, field densities and distributions from three years with distinctly different aphid population levels. Predation was incorporated at different time periods. Predation during the arrival of emigrants and establishment of aphid populations had approximately the same effect as predation over the entire aphid period, whereas predation during the 20 days prior to peak had very little effect on resulting peak populations. We conclude that a) yearly levels ofR. padi infestation are mainly determined by the volume and intensity of emigrants; b) peak levels are sensitive to changes in predator density; and c) in some cases an outbreak ofR. padi may be prevented by the action of spring active polyphagous predators alone.     

Foliar and ground-foraging predators of aphids associated with alfalfa crops in Chile: Are they good or bad partners?

Carabids and coccinellids are the most abundant aphid predators in alfalfa. Depending on the amount of prey consumed, the impact of their combined effect can be additive, antagonistic, or synergistic. Laboratory trials demonstrated that a higher proportion of Aphis craccivora Koch dropped from the alfalfa plants in the presence of coccinellids, and that the interaction between these predators was additive or synergistic, but never antagonistic, suggesting that they might be good partners in controlling aphids.     

The aggregative numerical response of polyphagous predators to aphids in cereal fields: attraction to what?

We tested in a field experiment two hypotheses for why polyphagous predators aggregate at concentrations of aphids: 1) because they are attracted directly to aphids as prey, or 2) because they are attracted to alternative prey that aggregate around the honeydew produced by aphids. Small plots were established in the field with two experimental treatments, aphid addition and honey spraying, and a watersprayed control, each replicated 10 times. Arthropods were recorded by pitfall traps and sticky traps in each plot. Diptera were significantly more abundant in the honey plots. Of the predators, Agonum dorsale, “All carabids” and Philonthus sp. were most abundant in the honey plots; Tachyporus spp. and carabid and staphylinid larvae were most abundant in the aphid addition plots. It is suggested that these results reflect differences among the predators in their ability to tolerate and utilise aphids as food.     

The influence of time and temperature on molecular gut content analysis： Adalia bipunctata fed with Rhopalosiphum padi

Gut content analysis is a useful tool when studying arthropod predator-prey interactions. We used polymerase chain reaction （PCR） technique to examine how detection of prey DNA in the gut content of predators was influenced by digestion time and temperature. Such knowledge is critical before applying PCR-based gut content analysis to field collected predators. Larvae of the two-spotted ladybeetle （Adalia bipunctata L.） were fed with the bird cherry-oat aphid （Rhopalosiphum padi L.） at either 21℃ or 14℃. After consuming one aphid, the predators were allowed to digest the prey for a range of time periods up to 24 hours. The influence of temperature on A. bipunctata feeding behavior was also recorded. From the fed larvae, total DNA was extracted and PCR reactions with R. padi specific primers were run. The number ofA. bipunctata that tested positive for R. padi DNA was negatively related to the length of digestion time. Temperature influenced larval feeding behavior but did not have a significant effect on R. padi DNA detection. After pooling the data from both temperature treatments we estimated the time point when R. padi DNA could be amplified from 50% of the fed A. bipunctata by PCR to be 4.87 hours. With such a rapid decrease in prey DNA detection success, positive PCR reactions will most likely be the result of predation events occurring shortly before capture. If a defined digestion temperature range has proven not to influence prey detection, PCR data obtained from predators collected within that particular range can be interpreted in the same way.     

Life history characteristics of Orius insidiosus (Say) fed diets of soybean aphid, Aphis glycines Matsumura and soybean thrips, Neohydatothrips variabilis (Beach)

Field studies in soybeans have demonstrated that the endemic predator, Orius insidiosus (Say), is an important natural enemy of the soybean aphid, Aphis glycines Matsumura. Soybean thrips, Neohydatothrips variabilis (Beach), serve as an important prey resource for O. insidiosus in soybeans and may be important in sustaining O. insidiosus populations before the arrival of soybean aphid. Because soybean aphid is new to the US soybean system, the effects of a mixed diet of soybean aphid and soybean thrips on O. insidiosus life history is not known. We measured the survival, development, and reproduction of O. insidiosus when fed soybean thrips, and a mixed prey diet of soybean aphids and soybean thrips, and compared these results to a previous study of O. insidiosus life history fed soybean aphid alone. Nymphal development to adulthood (15.9 days) and fecundity (68.8 eggs per female) was improved for O. insidiosus fed ad libitum soybean thrips daily compared to O. insidiosus fed ad libitum soybean aphids daily. The contribution of alternative prey to O. insidiosus life history characteristics can be complex depending on the amount and quality of a particular prey item. At low levels of prey, the addition of prey appears to enhance O. insidiosus survival, development, and fecundity. However, as predators are fed more often, the predator’s response depends on the type of prey that predominates in the mixed prey diet. We discuss soybean thrips impact on O. insidiosus population ecology and soybean aphid dynamics.     

Tracking aphid predation by lycosid spiders in spring-sown cereals using PCR-based gut-content analysis

Detection of prey DNA-remains in arthropod predators by polymerase chain reaction (PCR) is useful when investigating food webs. In this study we estimated how long after a feeding event it was possible to detect mitochondrial COII DNA (331 bp) from an important aphid pest, Rhopalosiphum padi (Homoptera: Aphididae), in spiders of the genus Pardosa (Araneae: Lycosidae). Following laboratory evaluations we tested spiders collected in spring-sown cereals for aphid predation during two seasons. Aphids were digested rapidly in laboratory-fed predators and the time point when prey DNA could be amplified from 50% of the spiders was estimated to be 3.7 h. A total of 372 field- collected predators were analyzed by PCR and despite low aphid densities many spiders (26% in 2004 and 19% in 2005) tested positive for R. padi, indicating consumption of at least one aphid within a few hours before capture. The percentage of spiders that tested positive for R. padi DNA varied considerably between fields and logistic regression analysis revealed that the probability of detecting aphid DNA was significantly influenced by location and year. We conclude that Pardosa spiders, under certain conditions, frequently feed on R. padi and deserve special attention in conservation biological control.     

Influence of drought on interactions between Rhopalosiphum padi and ground dwelling predators – A mesocosm study

Climate change will lead to extreme droughts, but it is difficult to predict how this will affect crop pests. In particular, it is unclear how interactions between natural enemies and pests will be influenced. In the field, bird cherry-oat aphids (Rhopalosiphum padi (L.)) have been observed to reside close to, or below the ground surface during dry conditions. We hypothesized that this will increase the niche overlap between R. padi and ground-dwelling predators such as carabid beetles and wolf spiders and that aphid numbers will therefore decline during dry conditions. A fully factorial mesocosm experiment was conducted testing the combined effects of drought and predator presence on aphid position and abundance on barley (Hordeum vulgare) plants. In support of our hypothesis, we found that (a) aphids moved below ground during dry conditions, (b) predators reduced aphid numbers, but only during dry conditions, and (c) predators reduced the proportion of aphids below ground in dry conditions. This increased predation effect during dry conditions was, however, compensated for by a corresponding increase in aphid performance on the plants and so the net effect of drought on aphid numbers ended up being neutral. Thus, pests can be affected by drought in complex ways via a combination top-down and bottom-up mechanisms. Predicting how pest populations will be affected by droughts in the future is thus a formidable research challenge.     

Foraging responses of Coccinella septempunctata,Hippodamia variegata and Chrysoperla carnea to changing in density of two aphid species

The successful use of predators in classical biocontrol programmes needs several background laboratory investigations, one of which is the evaluation of predator behavioural responses to changes in the density of their prey. The impact effect of the density of two prey species [Myzus persicae Sulzer and Aphis craccivora Koch (Hemiptera: Aphididae)] on the predation rates of third-instar Chrysoperla carnea Stephens (Chrysopidae: Neuroptera) and fourth-instar Coccinella septempunctata L. and Hippodamia variegata Goeze (Coccinellidae: Coleoptera) larvae was studied. Although prey species, predator species, prey density, and their interactions all had significant effects on the numbers of aphids consumed, the type of functional response did not vary, remaining a type II response in all treatments. However, the type II parameters differed among predator species on the same prey species, and for each predator species on the two prey species. Chrysoperla. carnea on M. persicae and H. variegata on A. craccivora were more voracious than other predators. In the context of functional response and biological control, the release of these predators, that show inverse density-dependent mortality, has to be started in early season to build up their population on low aphid densities and attack later high aphid populations.     

Weak responses to dietary enrichment in a specialized aphid predator

Some ladybeetles are specialist predators of aphids, coccids or other prey, although they often eat a variety of species from their focal prey taxon. In addition, the diet is often supplemented with alternative prey. How larvae of the aphidophagous Coccinella septempunctata L. utilize a non‐aphid alternative prey (fruit‐fly larvae Drosophila melonogaster Meigen) is compared with adequate (i.e. high‐quality) aphid prey provided alone (monotypic diet) or in mixed diets. The alternative prey are presented either nutrient‐enriched (i.e. raised on dog food supplemented medium) or not (raised on pure medium). Ladybird performance (survival, growth and development) is poor on the pure fly larvae diets, and also reduced when given mixed diets compared with the pure aphid diet. Nutrient enrichment of the fly larvae has no positive effects. The physiological background for the differences in food value, as indicated by performance in life‐history parameters, is a strong pre‐ingestive effect (i.e. reduced consumption of fly larvae compared with aphids) and a post‐ingestive effect (i.e. reduced utilization of assimilated larval fly tissue), whereas the assimilation efficiency of the consumed fly larvae is as high as that of aphids. The results show a physiological trade‐off resulting from prey specialization that reduces the possibility of utilizing alternative prey when the availability of aphids is scarce. Connected with this is a high robustness against variation in prey nutrient diversity and composition; the ladybird shows little positive response to dietary mixing (i.e. neither mixing of adequate aphids, nor of aphids and alternative prey) or to nutrient enrichment of prey. This contrasts with the results from generalist predators (spiders), where similar treatments lead to strong effects on life‐history parameters.