Factitious prey and artificial diets for predatory lady beetles: current situation,obstacles, and approaches for improvement: a review

Predatory lady beetles (Coleoptera: Coccinellidae) are important natural enemies of many pests in crop ecosystems throughout the world. Although several species are currently mass-reared and sold by biocontrol companies, there is an urgent need to reduce rearing costs. Cost-effective mass rearing of lady beetles involves the use of factitious prey/foods or artificial diets in lieu of natural prey. This review summarises the literature on the various kinds of factitious prey/foods and artificial diets as well as the feeding efficiency of predatory lady beetles. Factitious prey/foods generally satisfy the nutritional requirements for development of lady beetles, and lepidopteran immatures are the most widely used species. However, artificial diets, based on vertebrate protein and devoid of arthropod constituents, often yield disappointing results. We also highlight several successful examples and novel approaches for improving feeding efficiency. Moreover, available approaches and perspectives for speeding up the development and screening of cost-effective diets for mass rearing are stated in this review.     

Artificial diet and rearing methods for Thanasimus dubius (Coleoptera: Cleridae), a predator of bark beetles (Coleoptera: Scolytidae)

Clerid beetles are common natural enemies of bark beetles, and could potentially be used as biological control agents if they could be reared in sufficient numbers. We developed an artificial diet devoid of insect components for rearing Thanasimus dubius (Fabricius), a clerid that attacks several economically important bark beetles in eastern North America. We reared larvae of this predator using the artificial diet, and then used either natural or factitious prey to feed the adults so produced. Two different methods of presenting the diet were also examined. We then compared the performance of T. dubius reared on the artificial diet with newly-emerged wild individuals collected from the field. Our results suggest that adult predators reared on the diet are near in quality to wild ones, and high R₀ values can be obtained. No difference in prey preference was found between wild and diet-reared individuals after five generations in the laboratory. Sufficient numbers of predators could be generated using these techniques to permit limited field trials of augmentative biological control.     

Life table parameters, reproductive rate, intrinsic rate of increase, and estimated cost of rearing Podisus maculiventris (Heteroptera: Pentatomidae) on an artificial diet.

The impact of an insect-free artificial diet provided at nymphal and/or adult stage upon the developmental rate, life table parameters, and fertility table parameters was examined for Podisus maculiventris (Say). This study showed that when fed an insect-free artificial diet during both the nymphal and adult stage, developmental time was prolonged, preoviposition period was extended, and reproductive rate (R0) and intrinsic rate of increase (r) were significantly lower than when fed larval insect prey at both nymphal and adult stages. Additionally, feeding larval prey to adults reared as nymphs on an artificial diet significantly increased the proportion of fertile females, the number of eggs laid by mated females, the reproductive rate and intrinsic rate of increase, but the mean generation time was not significantly different. Likewise, feeding artificial diet to adults reared on larval prey resulted in a significant reduction in reproductive rate and intrinsic rate of increase. The "realized" cost to rear P. maculiventris on the artificial diet was calculated (as the cost to double the population size) using raw material cost, fertility table parameters and doubling time values. Raw material cost for rearing P. maculiventris colony on Trichoplusia ni (Hübner) (Lepidoptera: Noctuidae) was only 1.4 times higher than the cost of artificial diet raw materials required to rear the same size colony. However, the realized cost of rearing was 3.5 times higher when rearing on artificial diet because of the prolonged developmental time and reduced reproductive output. The cost efficiency of rearing a beneficial insect on an artificial diet that decreases the intrinsic rate of increase of a colony is discussed, as well as the advantages and disadvantages of supplementing adult diets with natural prey at the reproductive stage.     

Quality assessment of the predatory bugs Podisus maculiventris (Say) and Podisus sagitta (Fab.) (Heteroptera: Pentatomidae) after prolonged rearing on a meat—based artificial diet

The predatory stinkbugs Podisus maculiventris and Podisus sagitta were reared for more than 15 consecutive generations on a meat—based artificial diet. When they were returned to a diet of live prey (larvae of the pyralid Galleria mellonella) after different generations on the artificial diet, developmental and reproductive traits were similar to those of bugs continuously reared on live prey. Under laboratory conditions, predation rates for nymphs and adults of both pentatomids on larvae of the noctuid Spodoptera exigua were found to be unaffected by the previous diet. The results suggest that long—term rearing of P. maculiventris and P. sagitta on the meat—based artificial diet did not substantially affect the quality of the predators.     

Development and reproduction of Adalia bipunctata on factitious and artificial foods

The native coccinellid Adalia bipunctata (L.) (Coleoptera: Coccinellidae) may be an alternative to exotic species like Harmonia axyridis (Pallas) (Coleoptera: Coccinellidae) for the biological control of aphid pests in Europe. The availability of adequate factitious or artificial foods may help optimize its mass production. This study examines the nutritional value of Ephestia kuehniella Zeller (Lepidoptera: Phycitidae) eggs plus bee pollen, pea aphids Acyrthosiphum pisum (Harris) (Hemiptera: Aphididae) and mixtures of bee pollen and cysts of Artemia franciscana Kellogg (Branchiopoda: Artemiidae) and/or a lyophilized artificial diet based on bovine meat and liver. Reproducing adults of A. bipunctata were obtained on all diets tested, but immature survival, adult weights, development times and reproductive rates differed among diets. Over 84% of first instars fed on E. kuehniella eggs plus pollen or aphids survived to adulthood. Feeding predator larvae on pollen combined only with A. franciscana cysts or artificial diet yielded 40–55% immature survival, but survival increased to 74% when all of these components were mixed. Adult weights of A. bipunctata on the mixtures of pollen, A. franciscana cysts and/or artificial diet were 55–75% of those on aphids or on E. kuehniella eggs plus pollen. Lifetime fecundity was superior on E. kuehniella eggs plus pollen (1,864 eggs) to that on the other diets (264–889 eggs). The use of mixtures of plant and animal foods for A. bipunctata and other predators may contribute to increasing the cost-effectiveness of commercial mass production by reducing inputs of natural prey like aphids, or of nutritious but expensive factitious foods like lepidopteran eggs.     

Fitness and predation potential of Macrolophus pygmaeus reared under artificial conditions

Abstract The biological parameters of Macrolophus pygmaeus Rambur after prolonged rearing in the absence of plant materials were compared with those of conventionally plant‐reared predators. When eggs of Ephestia kuehniella Zeller were provided as food, developmental and reproductive fitness of M. pygmaeus reared for over 30 consecutive generations using artificial living and oviposition substrates was similar to that of predators kept on tobacco leaves. Plantless‐reared fifth instars of the predator also had similar predation rates on second instars of the tobacco aphid, Myzus persicae nicotianae Blackman, as their peers maintained on plant materials. In a further experiment, predation on aphid prey by fifth instar M. pygmaeus fed one of two egg yolk‐based artificial diets was compared with that of nymphs fed E. kuehniella eggs. Despite their lower body weights, predators produced on either artificial diet killed similar numbers of prey as their counterparts reared on lepidopteran eggs. Our study indicates that artificial rearing systems may be useful to further rationalize the production of this economically important biological control agent.     

Performance of Lymantria xylina (Lepidoptera: Lymantriidae) on artificial and host plant diets

Lymantria xylina Swinhoe (Lepidoptera: Lymantriidae) is a serious defoliator of hardwood and fruit trees in Taiwan. The larvae of L. xylina feed on >63 species of host plants, belonging to 29 families. Because a large number of larvae are needed for the production of nucleopolyhedrosis virus (NPV) or other related studies, the development of a suitable artificial diet is very important for the mass rearing of this moth in the laboratory. In this study, eight artificial diets, modified from different formulas, and one host plant, Liquidambar formosana Hance, were used to feed L. xylina caterpillars. Through various bioassays (first instar survival trial and long- and short-term feeding trials), the most suitable diet for the L. xylina was selected by performance comparisons with L. formosana. After the first instar survival trial, two of the diets were discarded, because no larva survived on these diets. The results of the long-term feeding trial indicated that the larvae grew successfully on only three kinds of artificial diet. Finally, results of the short-term feeding trial revealed that a diet (diet A), modified from the gypsy moth, Lymantria dispar (L.), formula diet, was the most appropriate for the L. xylina. Larvae fed on diet A had better survival rate, pupal weight, adult size, efficiency of conversion, and relative growth rate than larvae fed on other diets; they did not grow as well as those fed on L. formosana, however, except for pupal and adult weight, and approximate digestibility. In summary, diet A was found to be the best of the artificial diets for the L. xylina and is suitable for mass rearing of this moth in the laboratory.     

Life history and cost analysis for continuous rearing of Podisus maculiventris (Say) (Heteroptera: Pentatomidae) on a zoophytophagous artificial diet

The impact of a zoophytophagous, insect-free artificial diet upon the developmental rate, life table parameters, and fertility table parameters was examined over 11 consecutive generations for domesticated and wild colonies of Podisus maculiventris (Say) (Heteroptera: Pentatomidae). This study showed that the developmental time, preoviposition period, fecundity, and nymphal survival improved in the domestic colony when fed an insect-free artificial diet for 11 consecutive generations, but remained relatively constant for the wild colony, as did reproductive rate and intrinsic rate of increase. Although, after 11 generations of adaptation to an artificial diet feeding regime, all reproductive and fertility table parameters were still significantly lower than when fed on T. ni larvae as the natural prey, the realized cost of rearing either colony on the artificial diet approached 1.2 times the cost of rearing these insects on a natural prey. This is a significant achievement in the effort to develop cost-effective artificial diets for the mass-rearing of beneficial pentatomids, and has positive implications for the use of one artificial diet to efficiently rear several beneficial insects.     

Isotopic marking of natural enemies fed on C4 honey for habitat management studies in agroecosystems

1. To improve biological control and habitat management, how pest predators spread from natural habitats to crops must be understood. We studied whether intrinsic differences in stable-isotopic ratios of C and N from an artificial C₄ (sugar-cane) or C₃ (muti-flower honey) diet could help mark and track predators that feed on them.
2. Two aphid predators solely feeding on sugar resources as adults were used: the green lacewing Chrysoperla carnea s.l., and the parasitic wasp Aphidius colemani. δ¹³C and δ1⁵N values from wild individuals helped distinguish them from marked ones and determine the habitat resources they used.
3. Green lacewings fed on C₄ showed significantly higher δ¹³C values than those on C3 and wild individuals. However, parasitoid values were unaffected, with no mark acquired. Logistic regression was fitted to assess the probability of green lacewings having either diet with a probability of 0.93. Marks were acquired after 5 days and were detectable 20 days after switching diets with a probability of 0.67. Similar δ¹⁵N values for both wild populations indicated both predators had similar prey, but different vegetal resources (different δ¹³C values).
4. Sugar-cane honey is a natural and reliable marker for tracking lacewing populations in the field, but not for parasitoids.

     

Potential cotton aphid,Aphis gossypii,population suppression by arthropod predators in upland cotton

The cotton aphid, Aphis gossypii Glover, predation rate of convergent lady beetle, Hippodamia convergens Guerin‐Meneville, was determined by assigning a single predator randomly to each of four prey density treatments in the laboratory. Prey densities included 25, 50, 100, and 200 aphids per Petri dish arena. Predation response was recorded at 1, 4, 8, 16, 24, and 48 h after assigning predators to their prey treatments. Rate of consumption increased through time, with all 25 aphids consumed during the first 4 h of the experiment. At the highest density, adult lady beetle consumed on average 49, 99, 131, 163, 183, and 200 aphids within 1, 4, 8, 16, 24 and 48 h, respectively. Predators showed a curvilinear feeding response in relation to total available time, indicating that convergent lady beetles have the potential to suppress larger populations of aphids through continuous feeding by regulating their predation efficiency during feeding. The analysis of age‐specific mortality in absence of prey revealed that lady beetles could survive for an extended period of time (more than 2 weeks) without prey. The ability of a predator to survive without prey delays or prevents the rebound of pest populations that is a significant factor in natural biological control. A two‐year field sampling of 10 cotton arthropod predator species showed that spiders (27%) were the most dominant foliage dwelling predators in the Texas High Plains cotton followed by convergent lady beetles (23.5%), hooded beetles (13.5%), minute pirate bugs (11%), green lacewings (9.5%), bigeyed bugs (7.5%), scymnus beetles (3%), soft‐winged flower beetles (2%), damsel bugs (1.5%), and assassin bugs (1.5%). A field cage study showed that one H. convergens adult per plant released at prey density of one aphid per leaf kept the aphid population below economic threshold for the entire growing season.     

Developmental and reproductive fitness of Orius laevigatus (Hemiptera: Anthocoridae) reared on factitious and artificial diets

The developmental and reproductive fitness of the polyphagous predator Orius laevigatus (Fieber) (Hemiptera: Anthocoridae) was compared on two factitious foods and four artificial diets. Adults fed factitious foods (Ephestia kuehniella Zeller eggs and Artemia franciscana Kellogg cysts) performed better than those fed artificial diets. Among the artificial diets, a diet composed of liver and ground beef scored better than meridic diets based on egg yolk. Within the egg yolk-based artificial diets, the developmental fitness varied proportionally with the amount of egg yolk present in the diet. A food switching experiment, in which nymphs and adults of the predator were fed either E. kuehniella eggs or an egg yolk-based artificial diet, showed that the impact of adult food on reproductive capacity was greater than that of nymphal food. An optimal adult food was able to wholly compensate for deficiencies incurred by an inferior artificial diet in the nymphal stage. A strong correlation was found between oocyte counts, lifetime oviposition, and the number of eggs laid after 8 d. A rapid dissection assay may thus be effective to reliably and economically assess the fitness of O. laevigatus as a function of the diet. This method also may prove useful as part of quality assurance procedure for commercially produced predators.     

Eggs of Ephestia kuehniella and Ceratitis capitata,and motile stages of the astigmatid mites Tyrophagus putrescentiae and Carpoglyphus lactis as factitious foods for Orius spp.

Several factitious foods were assessed for rearing the anthocorid predators Orius thripoborus (Hesse) and Orius naivashae (Poppius) (Hemiptera: Anthocoridae) in the laboratory. Developmental and reproductive traits of both Orius species were examined when offered frozen eggs of the Mediterranean flour moth, Ephestia kuehniella Zeller, frozen processed eggs of the medfly, Ceratitis capitata Wiedemann, or mixed motile stages of the astigmatid mites Tyrophagus putrescentiae (Schrank) or Carpoglyphus lactis (L). Whereas C. lactis and T. putresecentiae proved to be an inferior food for rearing O. thripoborus and O. naivashae, eggs of C. capitata fully supported development and reproduction of both predators. Results on medfly eggs were similar or slightly inferior to those on E. kuehniella eggs, which is the standard food for culturing these anthocorid bugs. O. thripoborus could be maintained for 4 consecutive generations on C. capitata eggs indicating that processed medfly eggs can be a suitable and cheaper alternative to E. kuehniella eggs for prolonged rearing of these Orius spp.     

Optimization and evaluation of microencapsulated artificial diet for mass rearing the predatory ladybird Propylea japonica (Coleoptera: Coccinellidae)

Artificial diet optimization is a key aspect in mass rearing of natural enemies since it influences the quality and feeding effectiveness, and thus the success of the biological control program. Here, we introduced the microencapsulation method to package liquid artificial diet for feeding of the ladybird Propylea japonica. An orthogonal test of the quality of microencapsulated artificial diets (ADMs) was performed on key variables in production; Ca‐alginate concentration, chitosan concentration and weight ratio of wall material to inner diet. We compared the development and reproduction of P. japonica fed on the ADMs under different cold‐stored periods with those fed on fresh aphids and liquid artificial diets, in addition to a comparison of respiration, locomotion and predation. Our results indicated that chitosan concentration and ratio of shell to core significantly influence the quality of ADMs. The optimal recipe is 1.0% Ca‐alginate, 1.6% chitosan and shell : core = 1 : 2. Insects reared on fresh optimized ADMs were similar to those fed on fresh prey in all developmental and reproductive characteristics except for survival ratio and female fertility. ADMs appeared more beneficial than using a liquid artificial diet, although this may decrease with the prolonging of the cold‐storage period. P. japonica fed either on fresh ADMs or fresh prey showed improved respiration and predation abilities compared to where liquid artificial diet was used. Our study indicates advantages of microencapsulation in the production of artificial diet for predatory ladybird rearing. A microencapsulated diet can directly increase the efficiency and stability of mass rearing.     

Performance of the Wheat Armyworm, Pseudaletia sequax Franclemont, on Natural and Artificial Diets

Armyworms in the genus Pseudaletia Franclemont are important pests of cereal crops worldwide. The development of pest management strategies is largely dependent on studies involving the relationship among the pest, its host plants and natural enemies, as well as its susceptibility to chemical and biological control strategies. Part of these studies requires rearing the species of interest under laboratory conditions, which can be greatly facilitated by the use of artificial diets. In the present study, the effects of three artificial diets on development, survival, reproduction and demographic parameters of P. sequax were compared to a natural host plant, Pennisetum clandestinum. A diet based on pinto beans, wheat germ and torula yeast provided the best results in comparison to those obtained with the host plant. The most important life history parameters used in the evaluation of the nutritional quality of the diets were fecundity, egg viability and pupae deformities. The intrinsic rate of increase and the net reproductive rate were higher when individuals were fed on the artificial diet if compared to the host plant, indicating the suitability of the tested diet for the rearing of P. sequax. This artificial diet has been used successfully for the rearing of P. sequax without any fitness effect for over 18 consecutive generations.     

Two Meridic Diets for Perillus bioculatus (Heteroptera: Pentatomidae), a Predator of Leptinotarsa decemlineata (Coleoptera: Chrysomelidae)

Two artificial diets, devoid of insect components, were developed for the predator Perillus bioculatus (Fabricius). The diets based on chicken liver and tuna fish were designed to approach the nutritional characteristics of the primary insect prey, Leptinotarsa decemlineata (Say) eggs. P. bioculatus was reared on diet 2 for 11 consecutive generations without cumulative detrimental effects on its biology. Developmental time and preovipositional period were significantly longer, and egg viability, survival from egg to adult, and fecundity were significantly lower in P. bioculatus individuals reared on artificial diets than in those reared on L. decemlineata eggs. A significant reduction in developmental time, increase in survival from egg to adult, and reduction in length of the preovipositional period were observed after 11 generations of in vitro reared P. bioculatus on diet 2. These changes may indicate that the predators experience some degree of adaptation to the diet after several consecutive generations of in vitro rearing. No changes in egg viability were observed after 11 generations of in vitro rearing. Adult female weight was unaffected when reared on artificial diet, but a significant increase was observed after 11 generations of in vitro rearing. Females under all treatments weighed an average of 11.6 mg more than the males.     

Development and reproduction of the predatory mite Amblyseius swirskii on artificial diets

Development, survival and reproduction of the predatory mite Amblyseius swirskii (Athias-Henriot) (Acari: Phytoseiidae) were assessed when fed on cattail pollen (Typha latifolia L.), dried fruit mite (Carpoglyphus lactis L.), or on two artificial diets. The basic artificial diet (AD1) was composed of honey, sucrose, tryptone, yeast extract, and egg yolk. This diet was enriched (AD2) by adding hemolymph from oak silkworm pupae (Antheraea pernyi (Guérin-Méneville)). Mites fed on C. lactis and AD2 had shorter immature and preoviposition periods than those fed on the other diets. The total number of deposited eggs was significantly higher for females fed on AD2 than for those maintained on the other diets. The intrinsic rate of increase (r_m) of A. swirskii was highest on AD2 and C. lactis, followed by T. latifolia pollen, and AD1. In conclusion, the artificial diet AD2 supported development and reproduction of A. swirskii to the same extent as a factitious prey which is routinely used in the mass rearing of the phytoseiid. Our findings indicate the potential of artificial diets for the mass production of this economically important predatory mite.     

Deliberately added and "cryptic" antioxidants in three artificial diets for insects

Characteristics of both deliberately added and "cryptic" antioxidants were assayed from hydrophilic and lipophilic extracts from artificial diets for plant bugs, lepidopteran larvae, and green lacewings. Cryptic antioxidants are defined as substances naturally existing in diet ingredients but not deliberately added because of their antioxidant potential. Diets were tested after 1) being freshly produced, 2) stored for 48 h at 4 degrees C, or 3) held for 48 h under rearing room conditions at 27 degrees C. Tests included 1) a general assay of antioxidant capacity known as the ferric-reducing antioxidant power (FRAP) assay. 2) a cation radical-scavenging assay, 3) an ascorbic acid assay, and 4) an assay of inhibition of lipid peroxidation. In all assays, the lepidopteran diet had the highest values for protection against reactive oxygen species (ROS). The lepidopteran diet (with 0.17-0.23-mg equivalents of gallic acid equals total phenolic compounds per gram of diet) had three- to four-fold higher concentrations of phenolic compounds than did either the plant bug diet or the lacewing diet. Unexpectedly, the plant bug and the lacewing diets caused more lipid peroxidation than did the positive controls. This was attributed to the high concentrations of iron in these diets (mainly from chicken eggs), causing an ascorbate-ferric ion-induced lipid peroxidation. Diet storage, measured after 2 d at 27 or 4-6 degrees C, caused no significant declines in overall antioxidant potential. However, storage did lead to decline in ascorbic acid. The FRAP assay offered the best potential as a general, routine test of the potential of various insect diets to resist the destructive effects of ROS. The importance of addressing issues of protection against ROS in insect diets is discussed.     

Combining lacewings and parasitoids for biological control of foxglove aphids in sweet pepper

The role of natural enemy diversity in biological pest control has been debated in many studies, and understanding how interactions amongst predators and parasitoids affect herbivore populations is crucial for pest management. In this study, we assessed the individual and combined use of two species of natural enemies, the parasitoid Aphidius ervi Haliday, and the predatory brown lacewing Micromus variegatus (Fabricius), on their shared prey, the foxglove aphid, Aulacorthum solani (Kaltenbach), on sweet pepper. We hypothesized that the presence of intraguild predation (IGP) and predator facilitation (through induced aphid dropping behaviour) might have both negative and positive effects on aphid control, respectively. Our greenhouse trial showed that overall, the greatest suppression of aphids occurred in the treatment with both the parasitoid and the lacewing. While the combination of lacewings and parasitoids significantly increased aphid control compared to the use of parasitoids alone, the effect was not significantly different to the treatment with only predators, although there was a clear trend of enhanced suppression. Thus, the combined effects of both species of natural enemies were between additive and non‐additive, suggesting that the combination is neither positive nor negative for aphid control. High levels of IGP, as proven in the laboratory, were probably compensated for by the strong aphid suppression provided by the lacewings, whether or not supplemented with some level of predator facilitation. For aphid management over a longer time scale, it might still be useful to combine lacewings and parasitoids to ensure stable and resilient aphid control.     

人工饲料与天然饲料饲养条件下烟青虫的生长发育和繁殖力比较

报道饲养烟青虫HelicoverpaassultaGue澖ee的一种人工饲料,比较了烟青虫取食该饲料和天然饲料的主要生物学指标,测定了6龄幼虫对2种食料的转化和利用率。该人工饲料的主要成分是麦胚、黄豆粉、酵母粉和烟叶粉,天然饲料为烟草叶片。结果表明:取食人工饲料的烟青虫幼虫生长发育快,化的蛹较大,成虫寿命长,产卵量大,所产卵孵化率高;取食人工饲料的烟青虫与取食烟草蒴果的相比,前者相对取食量和近似消化率较高,相对生长率无显著性差异。说明烟青虫对该人工饲料有较好的适应性,可以作为繁殖烟青虫的实用饲料。     

Comparison of artificial diets for rearing Corcyra cephalonica (Stainton, 1865) (Lep., Pyralidae) for Trichogramma mass production

The objective of this research was, based on biological studies, to determine and adequate diet for rearing Corcyra cephalonica (Stainton, 1865) in the laboratory so as to permit the rearing of this factitious host for Trichogramma mass production. The research was conducted at 25 +/- 1 degrees C, RH 60 +/- 10% and photophase of 14 hours. Six artificial diets were compared: a) whole wheat flour (48.5%), ground rice (48.5%) and sugar (3%); b) ground rice (97%) and sugar (3%); c) whole wheat flour (48.5%), rice flour (48.5%) an sugar (3%); d) whole wheat flour (97%) and yeast (3%); e) wheat germ (97%) and yeast (3%); f) rice bran (94%), sugar (3%) and yeast (3%); f) rice bran (94%), sugar (3%) and yeast (3%). All of the diets studied permitted the development of C. cephalonica although the diets with wheat germ and yeast and that consisting of rice bran, sugar and yeast proved to be the most adequate for rearing the moth. These diets reduced the total (egg-adult) cycle, shortened the egg laying period, and produced heavier adults. Studies on the fertility life tables showed that higher net reproduction rates (Ro) and finite ratio of increase (lambda) were obtained from adults reared on these diets.