Importance of carbohydrate sources to the reproductive output of the wheat armyworm Pseudaletia sequax

• 1 The importance of nutrients consumed by adults differs among lepidopteran species. Knowledge of adult nutritional requirements is particularly useful when adopting biological control strategies that involve the use of flowering plants to improve the effectiveness of natural enemies. This is because nectar may also serve as a carbohydrate source for the pest.
• 2 We evaluated the effect of different sources and concentrations of carbohydrate consumed by adults on the performance and fitness of the wheat armyworm Pseudaletia sequax Franclemont, which is an important pest of cereal crops.
• 3 The consumption of carbohydrates by the adults significantly increased the length of the oviposition period, fecundity, fertility and longevity of P. sequax. Adult nutrition also affected the daily oviposition pattern of the females.
• 4 The demographic parameters estimated for each diet regime (i.e. intrinsic rate of increase, finite rate of increase and net reproductive rate) were influenced by adult nutrition. These parameters also varied according to the source and concentration of carbohydrates.
• 5 The results obtained in the present study suggest that the use of perennial flowering plants to provide a food source for natural enemies can also favour the pest.

     

Does natural vegetation from olive groves benefit the olive moth,Prays oleae?

Conservation biological control aims to enhance the efficacy of biological control agents, such as predators and parasitoids, by providing them access to key requisites, for example floral or other alternative food resources. However, the occurrence of floral resources in agricultural systems may have positive effects not only on natural enemies but also on the pests themselves. For this reason, we have studied in laboratory common Mediterranean plants as potential food resources for an important lepidopteran olive pest, the olive moth, Prays oleae. The effects of 15 wild flowering plants from olive grove ecosystems, and four sugars present in floral nectars, on key biological parameters of P. oleae (i.e. longevity, pre-oviposition and oviposition periods, fertility and fecundity) were evaluated. The three pest generations were analysed. Only few of the plants tested increased some reproductive parameters of the insect individuals compared to the water-fed controls, while sugars generally improved them. This could be due to a higher concentration of sugars provided by individual sugars than by flower nectar. These results indicate that, in general, P. oleae will not benefit from the availability of the floral resources tested.     

SEXUAL SIZE DIMORPHISM AND SELECTION IN THE WILD IN THE WATERSTRIDER AQUARIUS REMIGIS: LIFETIME FECUNDITY SELECTION ON FEMALE TOTAL LENGTH AND ITS COMPONENTS

Darwin's fecundity advantage model is often cited as the cause of female biased size dimorphism, however, the empirical studies of lifetime selection on male and female body size that would be required to demonstrate this are few. As a component of a study relating sexual size dimorphism to lifetime selection in natural populations of the female size-biased waterstrider Aquarius remigis (Hemiptera: Gerridae), we estimated coefficients for daily fecundity selection, longevity selection, and lifetime fecundity selection acting on female body size and components of body size for two consecutive generations. Daily fecundity was estimated using females confined in field enclosures and reproductive survival was estimated by twice-weekly recaptures. We found that daily fecundity selection favored females with longer total length through direct selection acting on abdomen length. Longevity selection favored females with smaller total length. When daily fecundity and reproductive longevity were combined to estimate lifetime fecundity we found significant balancing selection acting on total length in both years. The relationship between daily fecundity and reproductive longevity also reveals a significant cost of reproduction in one of two years. We relate these selection estimates to previous estimates of sexual selection on male body size and consider the relationship between contemporary selection and sexual size dimorphism.     

Evaluation of the impact of natural enemies on Plutella xylostella L. (Lepidoptera: Yponomeutidae) populations on commercial Brassica farms

Abstract 1 Accurate assessment of the impact of natural enemies on pest populations is fundamental to the design of robust integrated pest management programmes. In most situations, diseases, predators and parasitoids act contemporaneously on insect pest populations and the impact of individual natural enemies, or specific groups of natural enemies, is difficult to interpret. These problems are exacerbated in agro‐ecosystems that are frequently disrupted by the application of insecticides. 2 A combination of life‐table and natural enemy exclusion techniques was utilized to develop a method for the assessment of the impact of endemic natural enemies on Plutella xylostella populations on commercial Brassica farms. 3 At two of the experimental sites, natural enemies had no impact on P. xylostella survival, at two other sites, natural enemy impact was low but, at a fifth site, natural enemies drastically reduced the P. xylostella population. 4 The calculation of marginal death rates and associated k‐values allowed the comparison of mortality factors between experimental sites, and indicated that larval disappearance was consistently the most important mortality factor, followed by egg disappearance, larval parasitism and pupal parasitism. The appropriateness of the methods and assumptions made to calculate the marginal death rates are discussed. 5 The technique represents a robust and easily repeatable method for the analysis of the activity of natural enemies of P. xylostella, which could be adapted for the study of other phytophagous pests.     

Dietary sugars and amino acids increase longevity and enhance reproductive parameters of Bracon cephi and B. lissogaster,two parasitoids that specialise on wheat stem sawfly

The wheat stem sawfly, Cephus cinctus Norton (Hymenoptera: Cephidae), is a major pest of wheat grown on the Northern Great Plains and Canadian Prairies of North America, causing economic losses of as much as $350 million annually. Two species that parasitize wheat stem sawfly larvae in wheat stems are Bracon cephi (Gahan) and B. lissogaster Muesebeck (Hymenoptera: Braconidae). Carbohydrate-rich diets increase adult parasitoid longevity and reproductive parameters, enhancing their success as natural enemies. In previous studies, these species had increased longevity, egg load, and volume when fed sucrose solutions, encouraging further research on their nutritional needs. Therefore, we conducted experiments with artificial diets using adult females fed sucrose, glucose, and fructose solutions. Females were also fed these sugars in combination with a mixture of amino acids. We assessed individuals daily for longevity. Both species benefited from diets containing carbohydrates, with an increase in longevity from an average of 9 to 51 days for B. cephi, and from 6 to 34 days for B. lissogaster. Additional experiments assessed egg load and volume after 2, 5, and 10 days of cumulative feeding in B. cephi. These females produced 1.4-fold more eggs when fed amino acids, 2.5-fold more with sugar, and 2.7-fold more when fed sugar with amino acids. They had a 1.3-fold increase in egg volume when fed amino acids, 1.9-fold with sugar, and 2.1-fold when fed sugar with amino acids. Our study reveals the nutritional requirements of these braconid parasitoids and the benefits of nutritional sources when implementing conservation biological control strategies.     

Influence of adult diet on fitness and reproductive traits of the egg parasitoid Anagrus virlai (Hymenoptera: Mymaridae), a potential biocontrol agent against the corn leafhopper

Dalbulus maidis is the most important leafhopper pest of maize in the Americas. Anagrus virlai is an egg parasitoid commonly associated with the corn leafhopper. We evaluated whether the performance of A. virlai is dependent on different diets provided during 24 hr or throughout adult female lifetime. Additionally, functional response of A. virlai on D. maidis eggs using maize leaves containing honeydew plus honey was described. A. virlai is a mostly pro-ovigenic autogenic species whose females are able to parasitize eggs immediately after emergence. We found that wasps oviposit the majority of their eggs in the first day of their adult life. Realized lifetime fecundity and lifetime potential fecundity did not differ significantly among treatments, but longevity and egg production increased when honey was added to diet. Data were consistent with an intermediate functional response between type II and III, but closer to type II, indicating a high parasitism rate at low host densities and a decrease in the oviposition rate at high host densities, due to a possible egg limitation. Our results suggest that carbohydrate food sources (honeydew and honey) might not be the factor limiting reproductive success during the first 24 hr. Food supply, however, might influence egg maturation and survivorship of wasps, thus potentially enhancing biological pest control when hosts are scarce in the course of the first few days of adulthood.     

Effects of food variability on growth and reproduction of Aedes aegypti

Despite a large body of knowledge about the evolution of life histories, we know little about how variable food availability during an individual's development affects its life history. We measured the effects of manipulating food levels during early and late larval development of the mosquito Aedes aegypti on its growth rate, life history and reproductive success. Switching from low to high food led to compensatory growth: individuals grew more rapidly during late larval development and emerged at a size close to that of mosquitoes consistently reared at high food. However, switching to high food had very little effect on longevity, and fecundity and reproductive success were considerably lower than in consistently well‐fed mosquitoes. Changing from high to low food led to adults with similar size as in consistently badly nourished mosquitoes, but even lower fecundity and reproductive success. A rapid response of growth to changing resources can thus have unexpected effects in later life and in lifetime reproductive success. More generally, our study emphasizes the importance of varying developmental conditions for the evolutionary pressures underlying life‐history evolution.     

A host shift from wild blue cohosh to cultivated potato by the phytophagous ladybird beetle, Epilachna yasutomii (Coleoptera, Coccinellidae)

Life history traits of the phytophagous ladybird beetle Epilachna yasutomii were compared between a nonpest population feeding on wild blue cohosh and a pest population feeding on cultivated solanaceous crops, mainly potato. Newly emerged adults of the nonpest population entered diapause early in midsummer when blue cohosh withered, while adults of the pest population were found in tomato and eggplant fields until late autumn. The pest population had larger females, a higher population growth rate, a shorter larval developmental period, and reduced longevity of overwintered females, compared with the nonpest population. ANOVA indicated that all these life history traits were influenced by the food plant, and that the number of eggs laid per female and the longevity of overwintered females were also affected by the population type. These findings suggest that the life history pattern of E. yasutomii changed to high fecundity with a short life span from low fecundity with a long life span as a result of the host shift from wild blue cohosh to cultivated solanaceous crops. Received: May 22, 1998 / Accepted: January 13, 1999     

The effect of starvation and subsequent feeding on the reproductive potential of the grain aphid,Sitobion avenae

The short‐term starvation tolerance of alate and apterous morphs and the effect of periods of starvation on the longevity and fecundity of alate adults were evaluated in the grain aphid, Sitobion avenae (Fabricius) (Hemiptera: Aphididae). Alate adults exhibited a proportionally larger length of survival compared with apterous adults under continuous starvation conditions. Newly molted pre‐reproductive adults were starved for 0–96 h and their survival rate on the 1st day after recovering with food was not significantly different from that of control aphids. Starvation reduced lifetime fecundity, but increased the reproductive rate immediately after nutrition being improved. Fecundity and longevity after 24, 48, 72, and 96 h of starvation were significantly higher than after 120 or 144 h of starvation. However, no significant differences were observed for alate adults after 24, 48, 72, or 96 h of starvation. This study suggests that the ability of alatae to adapt to brief periods of starvation could be one of the important factors affecting the reproductive success of aphids during delays in locating host plants.     

Uniformity of petroleum‐derived spray oils: lethal and sublethal effects on a herbivore pest and its parasitoid

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Lifetime gains of host-feeding in a synovigenic parasitic wasp

Abstract. Understanding behavioural decisions relative to host use for feeding or reproduction by foraging parasitoids requires not only the study of metabolic pathways followed by nutrients, but also the quantification of lifetime fitness gains of each alternative behaviour. By using a combination of observational and manipulative approaches, the lifetime host‐feeding gains are measured both in terms of fecundity and longevity in the parasitoid Eupelmus vuilletti. Host‐feeding increases both egg production and longevity. The increase in fecundity is mainly determined by the amount of lipids obtained whereas the lifespan extension is mainly determined by carbohydrates. Proteins obtained through host‐feeding have been implicated previously in egg production by parasitoids but protein intake has no effect on fecundity and longevity in E. vuilletti. The amount of nutrients gained through host‐feeding and invested in eggs is variable and changes over the lifetime of the animal. Therefore, lifetime feeding gains are best understood through the construction of dynamical budgets running over the entire lifespan of an insect.     

Tritrophic Choice Experiments with Bt Plants,the Diamondback Moth (Plutella xylostella) and the parasitoid Cotesia plutellae

Parasitoids are important natural enemies of many pest species and are used extensively in biological and integrated control programmes. Crop plants transformed to express toxin genes derived from Bacillus thuringiensis (Bt) provide high levels of resistance to certain pest species, which is likely to have consequent effects on parasitoids specialising on such pests. A better understanding of the interaction between transgenic plants, pests and parasitoids is important to limit disruption of biological control and to provide background knowledge essential for implementing measures for the conservation of parasitoid populations. It is also essential for investigations into the potential role of parasitoids in delaying the build-up of Bt-resistant pest populations. The diamondback moth (Plutella xylostella), a major pest of brassica crops, is normally highly susceptible to a range of Bt toxins. However, extensive use of microbial Bt sprays has led to the selection of resistance to Bt toxins in P. xylostella. Cotesia plutellae is an important endoparasitoid of P. xylostella larvae. Although unable to survive in Bt-susceptible P. xylostella larvae on highly resistant Bt oilseed rape plants due to premature host mortality, C. plutellae is able to complete its larval development in Bt-resistant P. xylostella larvae. Experiments of parasitoid flight and foraging behaviour presented in this paper showed that adult C. plutellae females do not distinguish between Bt and wildtype oilseed rape plants, and are more attracted to Bt plants damaged by Bt-resistant hosts than by susceptible hosts. This stronger attraction to Bt plants damaged by resistant hosts was due to more extensive feeding damage. Population scale experiments with mixtures of Bt and wildtype plants demonstrated that the parasitoid is as effective in controlling Bt-resistant P. xylostella larvae on Bt plants as on wildtype plants. In these experiments equal or higher numbers of parasitoid adults emerged per transgenic as per wildtype plant. The implications for integrated pest management and the evolution of resistance to Bt in P. xylostella are discussed.     

Larval survival,host plant preferences and developmental responses of the diamondback moth Plutella xylostella (Lepidoptera: Plutellidae) on wild brassicaceous species

The diamondback moth Plutella xylostella (L.) (Lepidoptera: Plutellidae) is an important pest of cultivated brassicaceous crops worldwide. The host plant preferences, developmental biology and survival and longevity of P. xylostella are relatively well understood on commercial crop species; however, its relationship with brassicaceous weeds is poorly known. Sinapis arvensis L., Erysimum cheiranthoides L. and Capsella bursa‐pastoris (L.) Medicus are among the most common brassicaceous weeds worldwide and can serve as important bridge hosts of P. xylostella. In this study, preference and performance of P. xylostella were compared on these weed species. In free‐choice situations, females deposited 5.5 and 18.8 times more eggs on S. arvensis than on E. cheiranthoides and C. bursa‐pastoris, respectively. Survival from neonate to pupa and from pupa to adult was highest on S. arvensis and E. cheiranthoides and lowest on C. bursa‐pastoris. Development was fastest, foliage consumption was greatest, pupae and silk cocoons were heaviest, adult body masses and longevities were highest and forewings were largest for both females and males when reared as larvae on S. arvensis. Realized fecundity of new generation adults was highest for individuals reared on S. arvensis compared to those reared on E. cheiranthoides or C. bursa‐pastoris. Relative growth rates of pupae and adults were highest on S. arvensis, suggesting that this plant species is a high‐quality host for P. xylostella compared with other species tested. Potential impacts of these wild brassicaceous species on P. xylostella populations are discussed.     

The effect of mating frequency and mating pattern on female reproductive fitness in cabbage beetle,Colaphellus bowringi

It is generally thought that females can receive more of the material benefits from males by increasing mating frequency and polyandry can lead to greater reproductive success. The cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), is a highly promiscuous species, in which females or males can readily mate repeatedly with a given partner or multiple partners at a very high frequency. In the present study, the effect of mating frequency (number of matings) and mating pattern (polyandry vs. monogamy) on female reproductive fitness was investigated by measuring fecundity, fertility, and female longevity. The results indicated that increased female mating frequency with the same male did not result in variation in lifetime fecundity, but significantly increased fertility and decreased female longevity. Moreover, five copulations were sufficient to acquire maximal reproductive potential. Female lifetime fecundity also did not differ between polyandrous and monogamous treatments. However, monogamous females exhibited a significant increase in fertility and significant prolongation of longevity compared with polyandrous females, further demonstrating that monogamy is superior to polyandry in this beetle.     

Dietary glycerol and adult access to water: effects on fecundity and longevity in the almond moth

The quality of food eaten by larval insects will affect traits such as gamete production, fat reserves, muscle bulk and body size in the adult. Moreover, larvae also depend on high moisture content in the diet for survival. The almond moth (Ephestia cautella) (W.) (Lepidoptera; Pyralidae) does not feed as an adult although it continues to drink water. We tested the idea that an almond moth could compensate for a low-water diet as a larva by increasing its water intake as an adult. We reared larvae on two different food sources with different moisture regimes; standard laboratory diet with glycerol (relatively wet) and standard diet without glycerol (relatively dry). Half the adult moths from each treatment were given water to drink before their first and only mating. Our results show that wet larval diets (i.e. containing glycerol) significantly decreased fecundity (total number of eggs laid and the proportion of hatched larvae), whilst it significantly increased male and female longevity. The interaction effect of water access for adult males and females was significant, independent of the glycerol in the larval diet. Longevity in females that were not presented with water as adults was slightly higher if mated with a male that had had access to water, suggesting a mating donation of water. However, females that received water as adults showed a decreased longevity if mated with a male who had also had access to water as an adult, indicating a negative effect of water if received by both males and females. In addition, when the larval diet included glycerol, increased number of eggs laid decreased female longevity, whilst an absence of glycerol in the larval diet resulted in low female longevity that was unlinked with fecundity. Glycerol is used in many artificial insect diets and the fact that it shows a strong effect on key life-history traits (reproductive output and longevity in this species), merits careful re-examination of its effects on these important traits in other laboratory models. We also discuss the possibility that larval diet can affect female reproductive decisions.     

Amino acids in nectar enhance longevity of female Culex quinquefasciatus mosquitoes

Culex mosquitoes feed on a wide range of nectars consisting of mostly carbohydrates and amino acids, however, little is known about the utilization and effects of these different carbohydrates and their accompanying amino acids on longevity. Culex quinquefasciatus larvae were reared on low- and high-quantity food diets to produce adults that were nutritionally representative of wild-caught and laboratory-reared mosquitoes. Emerging adults reared on low- or high-quantity food diets as larvae were then provided Lantana camara nectar mimics containing mixtures of carbohydrates and amino acids to evaluate effects of nectar amino acids on longevity. Carbohydrates (with or without amino acids) were a critical component of the adult diet, and in their absence, adult mosquitoes died within 3-5 days. The nectar mimic that contained both carbohydrates and amino acids did not increase adult longevity of males originating from either poorly or well-fed larvae. However, females receiving adult diets containing both carbohydrates and amino acids lived 5% longer than females fed adult diets with only sugar.     

Flirtation reduces males’ fecundity but not longevity

Theory predicts that due to limited resources males should strategically adjust their investment in reproduction and survival. Based on different conceptual framework, experimental designs, and study species, many studies support while others contradict this general prediction. Using a moth Ephestia kuehniella whose adults do not feed and thus have fixed resources for their lifetime fitness, we investigated whether males adjusted their investment in various life activities under dynamic socio‐sexual environment. We allowed focal males to perceive rivals or additional females without physical contact. We show that males do not adjust the number of sperm they transfer to mates in a given copulation at different immediate or both immediate and mean sperm competition levels. Contradictory to general predictions, our results demonstrate that cues from additional females increase males’ investment in courtship and reduce their lifetime number of copulations and sperm ejaculated, whereas cues from rivals have no effect on these parameters. Males have similar longevity in all treatments. We suggest that the sex pheromone produced by multiple females overstimulate males, increasing males’ costly flirtations, and reducing their lifetime copulation frequency and fecundity. This finding offers a novel explanation for the success of mating disruption strategy using sex pheromones in pest management.     

Life history of Aptesis nigrocincta (Hymenoptera: Ichneumonidae) a cocoon parasitoid of the apple sawfly, Hoplocampa testudinea (Hymenoptera: Tenthredinidae)

Aptesis nigrocincta: Gravenhorst is a bivoltine ectoparasitoid of apple sawfly cocoons, hosts that must be found and parasitized by females at a depth of 10-25 cm in the soil. Females are significantly smaller than males and nearly wingless. After encountering a host, females needed 29.3 min at 20 degrees C and 19. 9 min at 25 degrees C to deposit an egg on the host. Development from egg to adult took 39.6 days for females and 38.0 days for males at 20 degrees C. This small difference was significant. At 20 degrees C, the longevity of females that had no opportunity to oviposit was on average 72.5 days, significantly higher than male longevity (50.6 days). The longevity of females given access to hosts throughout their lifetime averaged 58.6 days. Females were able to mate immediately after emergence and copulation lasted on average 21.7 s. After a pre-oviposition period averaging 5.8 days, females laid 20.2 eggs during their lifetime, thus less than one egg per day. Neither the fecundity nor longevity of individual females was correlated with body size. If females were deprived of food, longevity as well as lifetime fecundity were drastically reduced. Field studies were carried out in one organically managed apple orchard in Switzerland. Aptesis nigrocincta showed parasitism rates ranging from 12.1 to 39.7 % within single parasitoid generations, thereby constituting the most important mortality factor of apple sawfly cocoons.     

Herbivore response to habitat manipulation with floral resources: a study of the cabbage root fly

Biological control of pest insects can be improved by providing natural enemies with additional food resources such as floral nectar within the production field. However, herbivores may also benefit from this practice. The aim of this 3‐year field study was to investigate if dill and buckwheat, aimed as food resources for natural enemies, could increase the densities of the cabbage root fly, Delia radicum L. (Diptera: Anthomyiidae), a severe pest on crucifers. Differences in egg density, numbers of pupae and sex ratio were compared between cabbage plots with or without flowers. Habitat manipulation by intercropping flowering plants with cabbage did not increase the overall D. radicum egg density in our 3‐year study, and there were no significant differences in egg numbers between treatments in any year. No effect on the fecundity of D. radicum was observed, most likely because of the high mobility and feeding behaviour of the female flies, combined with high abundance and diversity of other food sources around the fields during this period. Despite equal egg numbers, fewer pupae were found in plots with flowers than without in one of three studied years. This finding suggests that natural enemies attacking larvae and pupae of D. radicum were either more abundant or efficient in cabbage plots with flowers.     

Bridging Developmental Boundaries: Lifelong Dietary Patterns Modulate Life Histories in a Parthenogenetic Insect

Determining the effects of lifelong intake patterns on performance is challenging for many species, primarily because of methodological constraints. Here, we used a parthenogenetic insect (Carausius morosus) to determine the effects of limited and unlimited food availability across multiple life-history stages. Using a parthenogen allowed us to quantify intake by juvenile and adult females and to evaluate the morphological, physiological, and life-history responses to intake, all without the confounding influences of pair-housing, mating, and male behavior. In our study, growth rate prior to reproductive maturity was positively correlated with both adult and reproductive lifespans but negatively correlated with total lifespan. Food limitation had opposing effects on lifespan depending on when it was imposed, as it protracted development in juveniles but hastened death in adults. Food limitation also constrained reproduction regardless of when food was limited, although decreased fecundity was especially pronounced in individuals that were food-limited as late juveniles and adults. Additional carry-over effects of juvenile food limitation included smaller adult size and decreased body condition at the adult molt, but these effects were largely mitigated in insects that were switched to ad libitum feeding as late juveniles. Our data provide little support for the existence of a trade-off between longevity and fecundity, perhaps because these functions were fueled by different nutrient pools. However, insects that experienced a switch to the limited diet at reproductive maturity seem to have fueled egg production by drawing down body stores, thus providing some evidence for a life-history trade-off. Our results provide important insights into the effects of food limitation and indicate that performance is modulated by intake both within and across life-history stages.