Yellow mutant of the Neotropical green lacewing Chrysoperla externa: trait inheritance and predator performance

The Neotropical green lacewing Chrysoperla externa (Hagen) (Neuroptera: Chrysopidae) is a key predator of various small soft‐bodied pest species. Chrysopidae species are known as ‘green lacewings’ due to their overall green body coloration. However, yellow mutant individuals were observed emerging from our lacewing rearing colony. Thus, the mode of inheritance of the yellow trait was studied and the hypothesis of an autosomal recessive allele for yellow color was tested using hybridization and backcrossing techniques. Furthermore, the possible implications of this color variation on specific life‐history characteristics of C. externa and the predation rates of each morph were evaluated. In both yellow and green morphs, basic life‐history characteristics were monitored, including time to hatching and viability of eggs, duration, and viability of larval and pupal stages, emergence rate and survivorship of adults, and fecundity and longevity of females. The yellow and green morphs were indistinguishable with respect to all life‐history traits evaluated and the predation rate of their larvae. Crossing experiments revealed the yellow color to be caused by a homozygous recessive allele, without sex‐linked expression. We conclude that the allele for yellow color is occurring at high frequency in the laboratory colony, supporting the existence of a genetic polymorphism for body ground color.     

Impact of Cheiracanthium pelasgicum (Araneae: Miturgidae) and Chrysoperla carnea (Neuroptera: Chrysopidae) intraguild predation on the potential control of cotton pest Helicoverpa armigera (Lepidoptera: Noctuidae)

Antagonist interactions such as intraguild predation (IGP) or cannibalism among predatory arthropods can reduce the impact of these invertebrates on pest limitation in agroecosystems. Here, the effects of IGP between two major natural enemies of cotton pests, the cursorial spider Cheiracanthium pelasgicum (C.L. Koch) and the common green lacewing Chrysoperla carnea (Stephens), were studied under laboratory conditions. First, a feeding preference test was carried out to determine the degree of C. pelasgicum preference for lacewing larvae, using second-instar Helicoverpa armigera larvae as alternative prey. In a second bioassay, the effects of predator interactions on potential predation of H. armigera larvae were analysed using three treatment combinations (plus a control with no predator): (1) spider alone, (2) lacewing larvae alone, (3) spider + lacewing larvae. Potential predation by C. pelasgicum on lacewing eggs was also studied. C. pelasgicum showed no significant preference for either of the two species, indicating that this spider may impact negatively on the green lacewing population. Findings revealed no additive effects and an antagonist interaction between C. pelasgicum and green lacewing larvae, which adversely affected H. armigera suppression; both predators displayed lower predation rates when kept together than either predator alone. However, presence of lacewing larvae and subsequent unidirectional IGP did not affect the predation capacity of C. pelasgicum. Finally, predation rates of C. pelasgicum on lacewing eggs were very low (mean 2.35 ± 0.71 eggs, 24 h after offering) indicating that the impact of C. pelasgicum on lacewing populations may be limited.     

Implications of alternative prey on biocontrol of pests by arthropod predators in strawberry

A range of naturally occurring predator species or commercially produced predators can be used in biocontrol strategies for pests. However, multiple potential prey species or other alternative food sources are often present for predatory insects at any one time. The availability of this ‘alternative’ prey may affect specific pest control by predators and thus influence the release rates of predators required for economic pest control. Strawberry aphid (Chaetosiphon fragaefolii), western flower thrips (Frankliniella occidentalis) and European tarnished plant bug (Lygus rugulipennis) are important and damaging pests in strawberry. In this study, laboratory, glasshouse and field experiments were undertaken to assess the effects of the availability of multiple prey species on biocontrol of specific pests. Results indicated that two of the predators tested showed preferences for prey species such that biocontrol of a particular pest was often less effective when a combination of pest species was present than would have been expected from results of experiments with single prey species alone. The experiments indicated that Orius laevigatus preferred C. fragaefolii to F. occidentalis or to L. rugulipennis, and preferred L. rugulipennis to F. occidentalis. Chrysoperla carnea was shown to prefer C. fragaefolii to L. rugulipennis, and C. fragaefolii over F. occidentalis. Therefore, it is important to consider the effects of alternative prey on suppression of pest species when deciding on management strategies and release rates of predators.     

Effect of host plant,prey species and intergenerational changes on the prey preferences of the predatory mirid <Emphasis Type="Italic">Macrolophus caliginosus</Emphasis>

The outdoor establishment of non-native biocontrol agents released for inundative control of glasshouse pests is determined primarily by two factors: ecophysiological compatibility with local climate, particularly winter cold tolerance, and ability to locate and utilise wild prey. Observations on the number and diversity of acceptable wild prey as part of an assessment of establishment potential therefore overlap with more focused studies to determine host range. This study investigated two aspects of the interactions between biocontrol agents and non-target prey that are rarely considered in tests for establishment or host range: the role of different host plant–prey associations in modifying the development and reproduction of biocontrol agents, and the longer term sustainability of such relationships beyond the single generation observed in most laboratory studies. Using the glasshouse whitefly (Trialeurodes vaporariorum) predator Macrolophus caliginosus Wagner (Hemiptera: Miridae) as a case study, the mirid was able to sustain viable populations over three generations on the related cabbage whitefly Aleyrodes proletella (Linnaeus) (Hemiptera: Aleyrodidae) and the aphid Myzus persicae (Sulzer) (Hemiptera: Aphididae), including when these prey were feeding on different host plants (Chinese cabbage, cabbage and Brussels sprout). However, the rate of development, fecundity and mortality of the predator varied between the different prey and host plant combinations, and in all cases differed than when feeding on its glasshouse prey T. vaporariorum reared on tobacco (Nicotiana tabacum). The results are discussed in the light of the current debate on methods for conducting host range testing as part of an environmental risk assessment. Handling Editor: Dirk Babendreier     

An immunological approach to distinguish arthropod viviphagy from necrophagy

Scavenging activity of predators inhabiting agroecosystems has not been thoroughly investigated. Understanding the prevalence of necrophagy in predators is paramount to determining the effectiveness of biological control agents. A molecular predator gut content assay is described that can differentiate necrophagy from viviphagy. Cadaver sweetpotato whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) and green lacewing, Chrysoperla rufilabris Burmeister (Neuroptera: Chrysopidae) serving as targeted prey items were marked with rabbit immunoglobulin G (IgG) protein and live prey items were marked with chicken IgG, respectively. The marked prey items were fed to convergent lady beetles, Hippodamia convergens Guérin-Méneville (Coleoptera: Coccinellidae) and soft-winged flower beetles, Collops vittatus (Say) (Coleoptera: Melyridae). The frequency of detection of the protein-marked prey items in the gut of the predaceous beetles was assessed at 0, 3, 6, 12, 24 and 48 h after feeding using a rabbit-IgG-specific or chicken-IgG-specific enzyme-linked immunosorbent assay (ELISA). Each IgG-specific ELISA detected the presence of the marker proteins in the gut of 90 % of the predators up to 12 h after prey consumption. A laboratory feeding study was also conducted to determine the propensity that each predator species engages in viviphagy and necrophagy. The laboratory feeding observations revealed that C. vittatus prefer carrion prey items. Finally, the laboratory observations of necrophagy were confirmed in a field study where C. vittatus was observed, directly and indirectly, feeding on H. convergens carcasses. The methodologies described here are useful for future studies on various aspects of insect predation.     

Predation and reproduction of the generalist predator Nabis pseudoferus preying on Tuta absoluta

The damsel bug, Nabis pseudoferus Remane (Hemiptera: Namidae), is a generalist predator of small arthropods, including key insect pests of vegetable crops. In this study, we characterized the predation and development of various N. pseudoferus life stages when preying on the eggs and first- and fourth-instar nymphs of the invasive South American tomato pinworm, Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae). These findings were compared to those of the bug’s common prey, cotton aphid, Aphis gossypii Glover (Hemiptera: Aphididae). Of the life stages tested, females showed the highest predation rate on all tested prey, due to their high longevity, large body, and great energy requirements for reproduction. The predator’s oviposition rate was clearly influenced by prey type, and was highest when feeding on fourth instars of T. absoluta. Considering the zoophytophagy of most of the life stages of other mirid species of tomato, and the lower propensity of N. pseudoferus to feed on plants, these results show that this species can be considered as a key indigenous natural enemy for sustainable pest control packages against T. absoluta in newly invaded areas.     

Reciprocal intraguild predation between two mirid predators,Cyrtorhinus lividipennis and Tytthus chinensis (Hemiptera: Miridae)

Interactions between predator species shape ecological communities and are crucial in the relationship between biodiversity and ecosystem functioning. They contribute to improving the natural biological control. Cyrtorhinus lividipennis Reuter and Tytthus chinensis Stål (Hemiptera: Miridae) are important natural enemies of brown planthopper, Nilaparvata lugens Stål (Hemiptera: Delphacidae), the most destructive rice pest in Asia. Comparative functional responses of both sexes of the two mirids at different development stages were estimated and the direction, symmetry, as well as intensity of intraguild predation (IGP) were studied in the laboratory. Prey preferences of the two mirid species and the influence of alternative prey on functional responses were also evaluated. The mirids exhibited reciprocal IGP in addition to consuming eggs of BPH. All predator–prey combinations exhibited a type II functional response. Females and fourth instar nymphs of C. lividipennis showed greater voracity for eggs of BPH than of T. chinensis. The two mirids were more voracious to intraguild prey than to extraguild prey. Alternative prey released the functional responses of C. lividipennis to both intra- and extraguild prey. In contrast, alternative prey had no effect on the functional response of T. chinensis. RIGP was symmetrical in the absence of extraguild prey. And in the presence of extraguild prey that was common in the field, resulted in different responses by the mirids.     

Ontogenetic change of prey preference in the generalist predator Zelus renardii and its influence on predator–predator interactions

1. Two field experiments were conducted to test the hypothesis that the intensity of predation by a generalist predator on two species of prey changes with the developmental stage of the predator. The generalist predator studied was Zelus renardii Kolenati (Hemiptera: Reduviidae) and the prey were the lacewing larva, Chrysoperla carnea Stephens, and the cotton aphid, Aphis gossypii Glover.
2. Zelus renardii and lacewings feed on aphids, thereby acting as potential competitors. In addition, Z. renardii feeds on lacewings. Thus, Z. renardii is an intraguild predator of lacewings.
3. Zelus renardii exhibited changes in prey preferences across developmental stages. The older stages of Z. renardii exerted greater mortality on lacewings and fed on larger lacewing larvae than did the younger stages.
4. Lacewings suppressed aphid population growth strongly. In contrast, none of the stages of Z. renardii was an effective control agent of the cotton aphid.
5. The addition of Z. renardii frequently disrupted the effective control of aphids generated by lacewings. In one of the two replicates of the experiment, the disruption increased with the developmental stage of Z. renardii , paralleling the increase in lacewing mortality.
6. Although the developmental stage of Z. renardii can influence the prevalence of intraguild predation and the intensity of the disruption of the aphid biological control, these experiments have demonstrated that even the youngest instars of Z. renardii can cause substantial lacewing mortality and release aphid populations from regulation.     

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.     

An energy budget approach for evaluating the biocontrol potential of cotton aphid (Aphis gossypii) by the ladybeetle Propylaea japonica

Biological control of economically important crop pests is an important component of integrated pest management (IPM) strategies. Predator–prey energy relationships are critical to the success of biocontrol strategies; however, these relationships are often ignored in many IPM programs. In this study, the biocontrol potential of cotton aphid, Aphis gossypii (Glover) (Hemiptera: Aphididae), by the ladybeetle Propylaea japonica (Thunberg) (Coleoptera: Coccinellidae) was estimated in terms of energy budgets calculated at 27 ± 1 °C. The energy equivalent of prey subjects (aphids) consumed was estimated from bomb calorimetry and partitioned into the energy associated with ingestion, assimilation, respiration, reproduction, and waste for each developmental stage of the lady beetle. The average assimilation efficiencies for larval and adult ladybeetles were 88.2 and 91.1%, respectively, whereas net ecological efficiencies were 17.6% for larvae and 2.6% for adults. Similarly, assimilation efficiencies of cotton aphids were 71.5 and 74.4% for nymphs and adults, respectively. Based on energy budget calculations, approximately 520, 3‐day‐old aphids and 5 356, 3‐day‐old aphids were estimated to be consumed by the ladybeetle larval stage and the female adult stage, respectively. These estimates were similar to the actual number of aphids consumed by the ladybeetles, based on actual counts. The current data demonstrate that P. japonica is an important natural enemy of the cotton aphid, and that predator–prey energy relationships can play a critical role in biocontrol strategies and IPM programs.     

Suitability of Aphis craccivora (Hemiptera: Aphididae) and Bemisia tabaci (Hemiptera: Aleyrodidae) biotype-B as prey for Chrysopa pallens (Neuroptera: Chrysopidae)

The suitability of Aphis craccivora Koch (Hemiptera: Aphididae) and Bemisia tabaci Gennadius (Hemiptera: Aleyrodidae) biotype-B eggs and nymphs as prey for pre-imaginal development and survival, adult longevity, and fecundity of the lacewing, Chrysopa pallens (Rambur) (Neuroptera: Chrysopidae) was evaluated under laboratory conditions at 25±1°C, 50±10% RH and a photoperiod of 16 h L:8 h D. Survival of C. pallens from first instar to adult eclosion was significantly different between the larvae that fed on the two prey species. C. pallens fed on A. craccivora completed development from egg to adult emergence, but those fed on eggs and nymphs of B. tabaci could not complete development, resulting in abnormal pupae and no normal adults emerged. The net reproductive rate (R ₀), intrinsic rate of natural population increase (r _m ), finite rate of increase (λ), mean generation time (T), index of population trend (I), doubling time (DT), and gross reproductive of rate (GRR) of C. pallens that fed on A. craccivora were 201.9 eggs per female, 0.13/d, 1.1/d, 40.1 d, 68.5, 5.2 d, 203.1 eggs per female, respectively. These results could be useful for mass-rearing of C. pallens and for understanding its population dynamics in the field in relation to the availability of different prey species.     

Inundative field releases and evaluation of three predators for Bemisia tabaci (Hemiptera: Aleyrodidae) management in three vegetable crops

Abstract The sweetpotato whitefly, Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae), is a global pest on numerous crops, including vegetables. Weekly inundative releases of a coccinellid predator (Coccinella undecimpunctata L. [Coleoptera: Coccinellidae]), a common green lacewing predator (Chrysoperla carnea Stephen [Neuroptera: Chrysopidae]), and a mirid predator (Macrolophus caliginosus[Wagner][Hemiptera: Miridae]) were independently made in three vegetable crops (cabbage [Brassica oleracea var. capitata L.], cucumber [Cucumis sativus L.], and squash [Cucurbita pepo L.]) for the management of the sweetpotato whitefly. Approximately 1 million to 2.5 million larvae or nymphs of each predator were released in the vegetable crops during 20 weeks. Whitefly populations were reduced by ≈ 25%–45% during most of the season in each crop where each predator was released. The effect of each predator was similar on whitefly population reduction. Late in the season (October) when whitefly populations were low, generally no benefit was obtained from releasing the predators. Numbers of predators recovered during sampling in all crops were greatest for C. carnea, but this corresponded with the fact that more individuals of this predator were released than any other predator in the experiment. These results help define the utility of these natural enemies for managing B. tabaci in vegetable crops.     

Predation of cotton bollworm by green lacewings in the presence of cotton aphid as alternative prey on transgenic Bt and conventional cotton

Experiments were conducted in small arenas and on whole plants to explore the effect of cotton aphids, Aphis gossypii Glover (Hemiptera: Aphididae), as alternative prey on the predation of Helicoverpa armigera Hübner (Lepidoptera: Noctuidae) larvae by green lacewing larvae, Mallada signatus Schneider (Neuroptera: Chrysopidae). Transgenic Bt (Bollgard II^®) and conventional cotton plants were included to explore potential differences in the predator's performance on these cotton types. In small arenas, the presence of 20 aphids reduced predation on H. armigera larvae by 22% (from 5.5 to 3.3 of 10) by a single lacewing larva over a 24‐h period. The presence of H. armigera reduced predation on aphids by ca. 29% (from 16.8 to 11.0 of 20) over 24 h. On whole plants, the presence of alternative prey had no effect on the number of H. armigera larvae or aphids remaining after 3 days. The presence of H. armigera larvae alone, without the predator, caused a 24% reduction in the numbers of aphids on conventional, but not on Bt cotton plants. The combination of Bt cotton and lacewing larvae caused a 96.6% removal of early‐stage H. armigera larvae, a statistically significant increase over the addition of the proportions (91.6%) removed by each factor measured separately, providing evidence of synergism. These studies suggest that the presence of aphids as alternative prey would not necessarily disrupt the predation by green lacewing on larvae of H. armigera, especially on Bt cotton.     

Impacts of biological control and invasive species on a non-target native Hawaiian insect

The potential for classical biological control to cause unintended harm to native species was evaluated in the case of the endemic Hawaiian koa bug, Coleotichus blackburniae White (Hemiptera: Scutelleridae), and parasitoids introduced to Hawaii for control of an agricultural pest, the southern green stink bug, Nezara viridula (L.) (Hemiptera: Pentatomidae). Parasitism of C. blackburniae eggs, nymphs and adults by biocontrol agents was quantified across a wide range of habitats and compared to other sources of mortality. Egg mortality due to the biocontrol agent Trissolcus basalis Wollaston (Hymenoptera: Scelionidae) was low (maximum 26%) and confined to elevations below 500 m on a single host plant. Predation, mainly by alien spiders and ants, was the greatest source of egg mortality (maximum 87%). Parasitism of adult C. blackburniae by the biocontrol agent Trichopoda pilipes (F.) (Diptera: Tachinidae) was near zero at 21 of 24 sites surveyed. Three sites with high bug density had higher levels of T. pilipes parasitism, reaching maxima of 70% among adult female bugs, 100% among males and 50% among fifth instars. Male-biased parasitism indicated that T. pilipes is adapted to using male aggregation pheromone for finding C. blackburniae hosts. The relative impacts of biocontrol agents and other sources of mortality were compared using life tables. Invasive species, particularly generalist egg predators, had the greatest impacts on C. blackburniae populations. Effects of intentionally introduced parasitoids were relatively minor, although the tachinid T. pilipes showed potential for large impacts at individual sites. In retrospect, non-target attacks by biological control agents on C. blackburniae were predictable, but the environmental range and magnitude of impacts would have been difficult to foresee.     

Bt Crops Producing Cry1Ac,Cry2Ab and Cry1F Do Not Harm the Green Lacewing,Chrysoperla rufilabris

The biological control function provided by natural enemies is regarded as a protection goal that should not be harmed by the application of any new pest management tool. Plants producing Cry proteins from the bacterium, Bacillus thuringiensis (Bt), have become a major tactic for controlling pest Lepidoptera on cotton and maize and risk assessment studies are needed to ensure they do not harm important natural enemies. However, using Cry protein susceptible hosts as prey often compromises such studies. To avoid this problem we utilized pest Lepidoptera, cabbage looper (Trichoplusia ni) and fall armyworm (Spodoptera frugiperda), that were resistant to Cry1Ac produced in Bt broccoli (T. ni), Cry1Ac/Cry2Ab produced in Bt cotton (T. ni), and Cry1F produced in Bt maize (S. frugiperda). Larvae of these species were fed Bt plants or non-Bt plants and then exposed to predaceous larvae of the green lacewing Chrysoperla rufilabris. Fitness parameters (larval survival, development time, fecundity and egg hatch) of C. rufilabris were assessed over two generations. There were no differences in any of the fitness parameters regardless if C. rufilabris consumed prey (T. ni or S. frugiperda) that had consumed Bt or non-Bt plants. Additional studies confirmed that the prey contained bioactive Cry proteins when they were consumed by the predator. These studies confirm that Cry1Ac, Cry2Ab and Cry1F do not pose a hazard to the important predator C. rufilabris. This study also demonstrates the power of using resistant hosts when assessing the risk of genetically modified plants on non-target organisms.     

Biological Activity of Cry1Ab Toxin Expressed by Bt Maize Following Ingestion by Herbivorous Arthropods and Exposure of the Predator Chrysoperla carnea

A major concern regarding the deployment of insect resistant transgenic plants is their potential impact on non-target organisms, in particular on beneficial arthropods such as predators. To assess the risks that transgenic plants pose to predators, various experimental testing systems can be used. When using tritrophic studies, it is important to verify the actual exposure of the predator, i.e., the presence of biologically active toxin in the herbivorous arthropod (prey). We therefore investigated the uptake of Cry1Ab toxin by larvae of the green lacewing (Chrysoperla carnea (Stephens); Neuroptera: Chrysopidae) after consuming two Bt maize-fed herbivores (Tetranychus urticae Koch; Acarina: Tetranychidae and Spodoptera littoralis (Boisduval); Lepidoptera: Noctuidae) by means of an immunological test (ELISA) and the activity of the Cry1Ab toxin following ingestion by the herbivores. Moreover, we compared the activity of Cry1Ab toxin produced by Bt maize to that of purified toxin obtained from transformed Escherichia coli, which is recommended to be used in toxicity studies. The activity of the toxin was assessed by performing feeding bioassays with larvae of the European corn borer (Ostrinia nubilalis (Hübner); Lepidoptera: Crambidae), the target pest of Cry1Ab expressing maize. ELISA confirmed the ingestion of Bt toxin by C. carnea larvae when fed with either of the two prey species and feeding bioassays using the target pest showed that the biological activity of the Cry1Ab toxin is maintained after ingestion by both herbivore species. These findings are discussed in the context of previous risk assessment studies with C. carnea. The purified Cry1Ab protein was more toxic to O. nubilalis compared to the plant-derived Cry1Ab toxin when applied at equal concentrations according to ELISA measurements. Possible reasons for these findings are discussed.     

Effect of plant nutrition on aphid size,prey consumption,and life history characteristics of green lacewing

Plant quality can directly and indirectly affect the third trophic level. The predation by all the instars of green lacewing, Chrysoperla carnea （S.） （Neuroptera： Chrysopidae） on the cereal aphids, Rhopalosiphum padi （L.）, and Sitobion avenae （F.） at varying nitrogen fertilizer levels was calculated under laboratory conditions. Wheat plants were grown on four nitrogen fertilizer levels and aphids were fed on these plants and subsequently offered as food to the C. carnea. Aphid densities of 10, 30, and 90 were offered to first, second, and third instar larvae of green lacewing. Increased nitrogen application improved nitrogen contents of the plants and also the body weight of cereal aphids feeding on them. Aphid consumption by green lacewings was reduced with the increase in nitrogen content in the host plants of aphids. Predation of both aphid species by first, second, and third instars larvae of C. carnea was highest on aphids reared on plants with the lowest rate of fertilization, suggesting a compensatory consumption to overcome reduced biomass （lower aphid size）. Total biomass devoured by C. carnea on all nitrogen fertilizer treatments was not statistically different. Additionally, the heavier host prey influenced by the plant nutrition had an effect on the life history characteristics of green lacewings. The larval duration, pupal weight, pupal duration, fecundity, and male and female longevity were significantly affected by the level of nitrogen fertilization to the aphid＇s host plants, except for pupal duration when fed on S. avenae. This study showed that quantity of prey supplied to the larvae affects the prey consumption and thereafter the life history characteristics of green lacewings.     

Infestation of Glycaspis brimblecombei (Hemiptera: Psyllidae) on three Eucalyptus species in selected ecological zones in Malawi

The study determined variations in incidence, severity and damage of Glycaspis brimblecombei among Eucalyptus species and ecological zones, and the interaction between Eucalyptus species and ecological zones. Additionally, the study determined variation in pest abundance between the upper, middle and lower parts of the tree crown. The study was conducted in six districts located in four ecological zones in Malawi. Eucalyptus stands established in 2014–2016 or coppices from trees cut in that period were sampled for G. brimblecombei infestation. A significant interaction was found between Eucalyptus species and ecological zones on G. brimblecombei infestation. Significant differences were observed between ecological zones in G. brimblecombei infestation with lowest incidence, severity and damage being found in the cooler ecological zone M. Eucalyptus camaldulensis and Eucalyptus tereticornis showed high susceptibility to G. brimblecombei compared to Eucalyptus grandis. There were no significant differences in abundance between the upper, middle and lower tree crown parts of each Eucalyptus species. Control efforts for this psyllid should focus on breeding and planting of resistant Eucalyptus varieties in specifically recommended sites. The uniform distribution of the pest on different tree crown parts implies that use of contact insecticides on a large scale would be tedious and expensive.     

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.     

Essential versus alternative foods of insect predators: benefits of a mixed diet

Although many predatory insects appear to be opportunistic generalists in their selection of prey, only a subset of prey species may in fact serve as “essential foods” capable of supporting immature growth and adult reproduction. It has been suggested that other, “alternative foods” serve only to maintain the predator when essential foods are not available, but little research has evaluated the significance of a mixed diet of essential and alternative foods for predator growth or reproduction. Here we test the general hypothesis that although alternative prey may be inadequate to support reproduction when consumed alone by adult predators, consumption of such prey may enhance the predator's reproductive output when the predator also has access to essential prey. We compared egg production by two aphidophagous lady beetles, Coccinella septempunctata and C. transversoguttata, provided with diets of aphids (essential prey) and weevils (alternative prey). As predicted, female predators produced greater numbers of eggs when a diet of pea aphids in limited number was supplemented by alfalfa weevil larvae. The predators laid no eggs when provided only with weevils or only with sugar. But once aphids were added to the diet, females of C. transversoguttata (but not C. septempunctata) laid eggs in greater numbers when they had fed previously on weevils than on sugar. Females of both species also produced eggs in modest numbers when provided with both weevils in excess and sugar, but this diet supported a lower rate of egg production than did a diet of weevils in excess plus a limited number of aphids. Although C. septempunctata has a longer history of association with the alfalfa weevil than does C. transversoguttata, the former species was not more effective in exploiting this alternative prey in support of reproduction. The tendency of generalist predators such as adult lady beetles to consume alternative as well as essential prey probably enhances considerably their ability to capitalize on short-lived and scattered opportunities as they seek out suitable sites in which to reproduce. Received: 11 May 1998 / Accepted: 1 May 1999