Tritrophic interactions among coumarin, the herbivore Spodoptera litura and a gregarious ectoparasitoid Bracon hebetor

The secondary metabolites that play a defensive role in plants not only affect the growth and development of herbivores, but they can also influence their natural enemies. In the present study, the effect of coumarin was evaluated on a braconid parasitoid, Bracon hebetor (Say) (Hymenoptera: Braconidae) using polyphagous pest, Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae). Reproductive potential of the parasitoid declined significantly due to parasitization of host larvae feeding on coumarin supplemented diet. Total development period of B. hebetor extended significantly (2.17–2.38 days) when the host larvae were reared on diet amended with higher concentrations (625–3,125 parts per million) of coumarin as compared to unamended diet. Ingestion of coumarin supplemented diet by S. litura adversely affected the emergence and parasitization potential of B. hebetor. As compared to the control a significant decline in hatching of eggs was recorded when parasitoid wasps developed on host larvae fed on diet containing higher concentrations of coumarin (625–3,125 ppm). Negative effects of coumarin were also recorded from the next generation of the parasitoid.     

Influence of Dietary Experience on the Induction of Preference of Adult Moths and Larvae for a New Olfactory Cue

In Lepidoptera, host plant selection is first conditioned by oviposition site preference of adult females followed by feeding site preference of larvae. Dietary experience to plant volatile cues can induce larval and adult host plant preference. We investigated how the parent’s and self-experience induce host preference in adult females and larvae of three lepidopteran stem borer species with different host plant ranges, namely the polyphagous Sesamia nonagrioides, the oligophagous Busseola fusca and the monophagous Busseola nairobica, and whether this induction can be linked to a neurophysiological phenotypic plasticity. The three species were conditioned to artificial diet enriched with vanillin from the neonate larvae to the adult stage during two generations. Thereafter, two-choice tests on both larvae and adults using a Y-tube olfactometer and electrophysiological (electroantennography [EAG] recordings) experiments on adults were carried out. In the polyphagous species, the induction of preference for a new olfactory cue (vanillin) by females and 3^rd instar larvae was determined by parents’ and self-experiences, without any modification of the sensitivity of the females antennae. No preference induction was found in the oligophagous and monophagous species. Our results suggest that lepidopteran stem borers may acquire preferences for new olfactory cues from the larval to the adult stage as described by Hopkins’ host selection principle (HHSP), neo-Hopkins’ principle, and the concept of ‘chemical legacy.’     

Feeding patterns in grasshoppers (Orthoptera: Acrididae): Factors influencing diet specialization

Summary Diets of grasshopper species from two arid grassland communities in Trans-Pecos, Texas, were determined by gut analysis. Species-specific food plant choice and niche breadths are presented for each of these species. As a group, grasshoppers range from monophagous to polyphagous feeders although most species fall in the oligophagous to polyphagous group. Phylogenetic constraints are evident such that gomphocerinae are primarily grass feeders while melanoplinae feed predominantly on forbs; the oedipodinae show less clearcut tendencies.Feeding patterns are remarkably constant from site to site and overall, community niche breadth distributions between sites do not differ greatly. Individual species tend to eat the same plant species at various sites and maintain similar niche breadths. Species with relatively specialized diets tend to feed on predictable plant species such as grasses and long-lived perrenial forbs.Grasshopper feeding patterns present some problems to the current theory of herbivore diet specialization since forb feeding melanoplines tend to be polyphagous (contrary to predictions). Life history patterns unrelated to tracking host plants may explain some aspects of diet breadth since diet selectivities are presumably adjusted according to the probability of finding suitable food plants.     

Feeding patterns of monophagous,oligophagous, and polyphagous insect herbivores: The effect of resource abundance and plant chemistry

Summary Leaf tissue preferences of monophagous, oligophagous, and polyphagous insect herbivores were determined using young and mature leaf tissue abundances and herbivore feeding observations. Larvae of monophagous and oligophagous herbivores preferred young leaf tissues while, overall, larvae of polyphagous species preferred mature leaves of their various host plants. Even though a species is often polyphagous over its geographical range, larvae from local populations may be very specialized in their diet. When this occurs these specialized larvae prefer the more nutritious and perhaps more toxic young leaves of some of their host plants. Resource abundance and plant chemistry are discussed as major factors influencing herbivore feeding patterns.     

Influence of sex and dietary fat on the group effect in larvae of Acheta domesticus (L.)

Larvae of Acheta domesticus show the group effect when reared in pairs on an artificial diet with the opposite sex, but not when reared with the same sex. When wheat-germ oil is added to the artificial diet, all paired larvae exhibit the group effect, regardless of sex. Dietary fat also improves survival and growth, but the identity of the component(s) producing these results has not been determined. The possible roles of linolenic acid and linolenic acid synthesis in this insect are discussed.     

Experience-based behavioral and chemosensory changes in the generalist insect herbivore Helicoverpa armigera exposed to two deterrent plant chemicals

Behavioral and electrophysiological responses of larvae of the polyphagous moth species Helicoverpa armigera to two plant-derived allelochemicals were studied, both in larvae that had been reared on a diet devoid of these compounds and in larvae previously exposed to these compounds. In dual-choice cotton leaf disk and pepper fruit disk arena assays, caterpillars reared on a normal artificial diet were strongly deterred by strychnine and strophanthin-K. However, caterpillars reared on an artificial diet containing strychnine were insensitive to strychnine and strophanthin-K. Similarly, caterpillars reared on an artificial diet containing strophanthin-K were also desensitized to both deterrent chemicals. Electrophysiological tests revealed that the deterrent-sensitive neurons in taste sensilla on the maxillae of caterpillars reared on each deterrent-containing diet displayed reduced sensitivity to the two chemicals compared with the caterpillars reared on normal diets. We conclude that the experience-dependent behavioral plasticity was partly based on the reduced sensitivity of taste receptor neurons and that the desensitization of taste receptor neurons contributed to the cross-habituation to the two chemicals.     

Larval diet influence on oviposition behaviour inSpodoptera littoralis

Females ofSpodoptera littoralis Boisd. (Lepidoptera: Noctuidae) with different feeding experiences during their larval development were tested for their ovipositional response to methanol extracts of larval frass and semisynthetic diets. The effect of the following frass, diet and diet component extracts was tested: (a) frass fromS. littoralis orAgrotis segetum larvae fed on a potato-based diet; (b) frass fromS. littoralis larvae fed on a wheat germ-based diet; (c) potato and wheat germ-based diets; and (d) potatoes and wheat germ. Ovipositing females without prior experience of the potato diet were deterred by extracts of: (1) larval frass from either species fed on potato diet; (2) the potato-based diet; (3) potato. Also females with experience of the potato diet during only a part of their larval development were deterred from oviposition by frass of larvae reared on the potato diet and by the diet itself. However, for females reared on the potato diet for their entire larval development, oviposition was no longer deterred by either of the three extracts listed above. Extracts of: (1) frass from larvae of either species reared on wheat germ diet: (2) the wheat germ diet; or (3) wheat germ did not significantly affect oviposition. Females with ablated antennae were still deterred by frass extracts from larvae fed on potato diet, when they had been reared on the wheat germ diet. In feeding experiments, larvae of larval stage one and of larval stage three-four reared on either of the two diets preferred to feed on the wheat germ diet. However, the preference was significantly stronger for larvae with no prior contact with the potato diet. The effect of larval experience on the loss of oviposition-deterring activity by extracts of larval frass, diets and diet components is discussed in view of induction and selection.     

Intra- and interspecific predation on four life stage groups by the adult females of Metaseiulus occidentalis, Typhlodromus pyri, Neoseiulus fallacis and Amblyseius andersoni

Do adult females of oligophagous species such as Neoseiulus fallacis (Garman) and Metaseiulus occidentalis (Nesbitt) show less intra- and interspecific predation on phytoseiids when other foods are scarce than polyphagous species such as Amblyseius andersoni Chant and Typhlodromus pyri Scheuten? We caged single adult females of each species without food with ten of their own eggs or larvae, with ten eggs or larvae of the other species or with ten nymphs or adult females of M. occidentalis (T. pyri for M. occidentalis). We assessed the ambulatory activity, survival time, egg levels and prey loss in each test. Polyphages (in particular T. pyri) lived longer than oligophages (in particular N. fallacis) without food. The small T. pyri detected its own stages and benefited most by feeding on small active stages of other species. Amblyseius andersoni, the largest mite, fed and gained the most of any species when held with nymphs and female adults. Metaseiulus occidentalis fed on eggs of all four species to enhance survival. The large hyperactive N. fallacis gained the least from these behaviours. Each mite seemed uniquely adapted to survive conditions of scarce prey and these behaviours may explain their roles in phytoseiid mite complexes. Overall, oligophagous adult females fed less and gained less by feeding on phytoseiids than did polyphagous adult females.     

Dietary diversification and variations in the number of labrum sensilla in grasshoppers: Which came first?

The diversity of the diet of grasshoppers (Acrididae, Orthoptera) is related to multiple factors, including the chemoreceptors on the antennae, palps and on the epipharyngeal face of the labrum. In the present study, we sought to understand the nature of the diet of 12 Moroccan acridian species and to try to relate various aspects of their diet to the number of labrum sensilla. If the effect of the labrum size on the number of sensilla is removed, four groups of species are recorded: (i) polyphagous species with a broad diet and numerous sensilla; (ii) polyphagous species with a graminivorous diet and numerous sensilla; (iii) oligophagous species feeding exclusively on Poaceae and with a medium number of sensilla; and (iv) strictly monophagous species feeding on a single plant species and with the smallest number of sensilla. These observations show the close relationship between the diet and the number of labrum sensilla. However, Sphingonotus rubescens, a polyphagous species, is an exception to this trend as it harbours a medium number of sensilla. We propose that the modification in the number of labrum sensilla is a result of a progressive adaptation to a different diet and does not represent its cause.     

Dependence on host constituents controlling food acceptance by Manduca sexta larvae

The host range of Manduca sexta L. (Lepidoptera: Sphingidae) is restricted in nature to plants in the family Solanaceae. However, naive hatchling larvae often accept and continue their development on foliage from a wide spectrum of unrelated plants. In contrast, solanaceous-experienced larvae refuse to feed on other plants. Experiments were designed to explore the role of constituents of various plants in this behavioral phenomenon. Fourth instar larvae reared on solanaceous hosts: tomato (Lycopersicon esculentum Mill.), potato (Solanum tuberosum L.), or tobacco (Nicotiana tabacum L.), and the leguminous non-host cowpea (Vigna sinensis Savi), or on artificial diet based on wheat germ were tested for their acceptance or preference when offered a new diet in both choice and no-choice situations. Under no-choice conditions, acceptability of cowpea foliage was strongly affected by the larval dietary experience. Most larvae reared on solanaceous foliage did not feed, whereas those larvae reared on non-solanaceous food readily accepted cowpea. Moreover, solanaceous foliage, leaf discs and extracts were readily accepted by larvae regardless of their dietary experience. Larvae reared on any of the solanaceous plants strongly preferred cowpea discs that were treated with solanaceous extract, while larvae reared on non-solanaceous diets did not discriminate between treated and control discs. Assays of cowpea extracts as well as acceptance of cowpea discs treated with solanaceous extract indicated a lack of detectable deterrent in cowpea. Our results suggest that the mechanism for induced host specificity in M. sexta larvae involves development of a dependence on solanaceous chemical constituents. We argue that this dependence on host plant chemistry has adaptive significance.     

Development and Leaf Consumption by Spodoptera cosmioides (Walker) (Lepidoptera: Noctuidae) Reared on Leaves of Agroenergy Crops

Spodoptera cosmioides (Walker) (Lepidoptera: Noctuidae) is a polyphagous pest that threatens more than 24 species of crop plants including those used for biodiesel production such as Ricinus communis (castor bean), Jatropha curcas (Barbados nut), and Aleurites fordii (tung oil tree). The development and leaf consumption by S. cosmioides reared on leaves of these three species were studied under controlled laboratory conditions. The egg-to-adult development time of S. cosmioides was shortest when reared on castor bean leaves and longest when reared on tung oil tree leaves. Larvae reared on castor bean and Barbados nut leaves had seven instars, whereas those reared on tung oil tree leaves had eight. Females originating from larvae reared on castor bean and Barbados nut leaves showed greater fecundity than did females originating from larvae reared on tung oil tree leaves. Insects fed on castor bean leaves had shorter life spans than those fed on tung oil tree and Barbados nut leaves although the oviposition period did not differ significantly. The intrinsic and finite rates of increase were highest for females reared on castor bean leaves. Total leaf consumption was highest for larvae reared on tung oil tree leaves and lowest for those reared on Barbados nut leaves. We conclude that castor bean is a more appropriate host plant for the development of S. cosmioides than are Barbados nut and tung oil tree.     

Starvation resistance of gypsy moth,Lymantria dispar (L.) (Lepidoptera: Lymantriidae): tradeoffs among growth,body size,and survival

Summary Survival and body composition of starving gypsy moth larvae initially reared on aspen foliage or artificial diet differeing in nitrogen (N) and carbohydrate concentration were examined under laboratory conditions. Diet nitrogen concentration strongly affected starvation resistance and body composition, but diet carbohydrate content had no effects on these. Within any single diet treatment, greater body mass afforded greater resistance to starvation. However, starving larvae reared on 1.5% N diet survived nearly three days longer than larvae reared on 3.5% N diet. Larvae reared on artificial diet survived longer than larvae reared on aspen. Differences in survival of larvae reared on artificial diet with low and high nitrogen concentrations could not be attributed to variation in respiration rates, but were associated with differences in body composition. Although percentage lipid in larvae was unaffected by diet nitrogen concentration, larvae reared on 1.5% N diet had a higher percentage carbohydrate and lower percentage protein in their bodies prior to starvation than larvae reared on 3.5% N diet. Hence, larger energy reserves of larvae reared on low nitrogen diet may have contributed to their greater starvation resistance. Whereas survival under food stress was lower for larvae reared on high N diets, growth rates and pupal weights were higher, suggesting a tradeoff between rapid growth and survival. Larger body size does not necessarily reflect larger energy reserves, and, in fact, larger body size accured via greater protein accumulation may be at the expense of energy reserves. Large, fast-growing larvae may be more fit when food is abundant, but this advantage may be severely diminished under food stress. The potential ecological and evolutionary implications of a growth/survival tradeoff are discussed.     

Studies on the group effect in Acheta domesticus (L.) using artificial diets

Larvae of Acheta domesticus were reared on artificial diets both singly and in groups of six. The growth of group-reared larvae was significantly faster, and survival of singly reared larvae was greatly reduced. Survival of singly reared larvae was raised to that of grouped larvae when 1% wheat-germ oil was added to the diet. Wheat-germ oil also improved the growth of grouped larvae and of singly reared female larvae. The fatty acid composition of body lipid was similar, except for linolenic acid, whether the larvae were singly or group-reared, or whether wheat-germ oil was fed or not. These results are discussed from the point of view of the mechanism of the group effect.     

Gypsy moth (Lymantria dispar) larval development and survival to pupation on diet plus extractables from green ash foliage

Gypsy moth, Lymantria dispar L., rate of larval development, molting, pupal weight and survival were studied on an artificial diet containing different concentrations of green ash ethyl acetate extractables (EtOAc Exts). Insects were reared on experimental diets from egg to pupa. Addition of EtOAc Exts to artificial diet significantly prolonged larval development and reduced their survival compared to larvae reared on control diet. Weights of pupae were significantly reduced when larvae were reared on diet containing the lowest dosage of EtOAc Exts (i.e., 0.01%) versus on control diet. EtOAc Exts in diet (e.g., 0.01, 0.06 and 0.2%) frequently caused incomplete ecdysis which invariably resulted in larval death. Impaired feeding, locomotion and excretion are likely causes of death. The combination of these results with our earlier findings of repellents and feeding deterrents against gypsy moth larvae (GML) in green ash foliage shows that the non-host status of green ash to the highly polyphagous GML involves three orders of chemical defense: repellents, feeding deterrents and inhibitors of nutritional and developmental physiology. As the insect becomes sequentially exposed to these orders of defense, it incurs higher costs because the adverse effects become less reversible.     

Nutritional and developmental factors in larval growth and retrogression of Trogoderma glabrum

Diet is an important determinant of the developmental pattern of Trogoderma glabrum (Herbst). Two nutritionally adequate diets were compared. The average weight of larvae of the same age was greater when cultured on meridic diet than on standard crude diet. The expected pattern of instars, as determined by head capsule widths, was observed for larvae reared on standard diet. On meridic diet, head capsules of each instar were significantly wider, and both sexes pupated after 5 stadia.Survival during retrogression correlated positively with weight at the time of isolation. Larvae reared on meridic diet survived longer than larvae of equal culture age or weight from standard diet. Survival and maintenance of weight were greatly improved by direct contact with nutritional media, some of which were incomplete (powdered cellulose plus 10% lactalbumin, fructose, or glucose). Cellulose alone, however, was more detrimental to survival than no medium.A moult after transfer to isolation tubes produced 3 morphologically distinct forms, determined by the extent of maturation. Immature larvae retromoulted to smaller larvae; fully mature larvae pupated; and larvae that had initiated but not completed pupal differentiation became prothetelic.     

Dietary cholesterol requirements of housefly, Musca domestica, larvae.

Quantitative analyses have been made of the dietary cholesterol requirement for the growth of the larvae of Musca domestica. The larvae will not grow on diets to which no cholesterol is added, a few pupae and adults are obtained when the concentration of cholesterol is 0·05 μmol/g of diet, but the concentration has to be raised to 0·36 μmol/g of diet before the maximum numbers of pupae and adults are obtained. Further addition of cholesterol above 0·36 μmol/g diet did not have any significant effect on the weight and growth of the larvae. However, the ratios of the cholesterol to phospholipid fractions recovered from the larvae increased rapidly when the concentration of cholesterol in the diet was raised from 0·05 to 0·56 μmol/g of diet. Above this concentration only a slight increase in the ratios was observed. Larvae reared on diets containing 0·05 μmol cholesterol/g of diet contain only 25 per cent of the cholesterol content of the larvae reared on the diets containing more than 0·28 μmol of cholesterol/g of diet, the cholesterol content being expressed relative to the weight of the larvae,The absence of cholesterol synthesis has been demonstrated in the larvae by feeding [4-¹⁴C] cholesterol. The specific activity of the cholesterol recovered from the larvae is the same as that of cholesterol added to the diet. Irrespective of the cholesterol concentration of the larval diet, approximately 97 per cent of the radioactivity recovered from the larvae behaved as free cholesterol, less than 1 per cent as cholesterol esters and the rest as unidentified ‘polar sterols’. The results are compared with those from similar studies on other insects.     

Butterfly community structure in fragmented habitats

We analysed effects of habitat fragmentation on the diversity, abundance, and life history traits of butterflies on 33 calcareous grasslands. Diversity of butterflies was positively correlated with habitat area (as was plant diversity), but not with habitat isolation. In contrast to expectations, butterfly densities of polyphagous and oligophagous species declined with habitat area whereas densities of monophagous species increased. The z -values, i.e. the slope of species–area relationships, increased with food plant specialization, from 0.07 in polyphagous, 0.11 in oligophagous, 0.16 in strongly oligophagous to 0.22 in monophagous species, and were 0.14 in plant species. Significant z -values were not only found for total species richness, based on a sample size adjusted to fragment area ( z  = 0.12), but also for the local density of butterfly species richness, based on equal sample size across all habitat fragments ( z  = 0.06). To our knowledge, this is the first study to show differential responses of monophagous, oligophagous and polyphagous species to area with respect to species richness and population density.     

Diet-switching by gypsy moth: effects of diet nitrogen history vs. switching on growth, consumption, and food utilization

Gypsy moth (Lymantria dispar (L.) (Lepidoptera: Lymantriidae)) larvae were reared from hatch on 1.25% N or 3.5% N artificial diet (previous diet) and switched reciprocally to the other diet (current diet) after molting into the second, third, fourth, or fifth instar. The nitrogen concentration of food consumed during previous instars had a strong residual effect on the growth rate in subsequent instars when a diet switch was made during instars two through four, but did not affect growth rate of fifth-instar larvae despite effects on food consumption and utilization. In early instars, larvae reared on 1.25% N artificial diet and then switched to 3.75% N diet had lower mass-adjusted growth rates than larvae continuously reared on 3.75% N diet. Conversely, larvae reared on 3.75% N diet and switched to 1.25% N had higher mass-adjusted growth rates than larvae reared continuously on 1.25% N diet. Relative to larvae previously reared on 1.25% N diet, fifth-instar male larvae previously reared on 3.75% N diet had slightly lower consumption rates, higher net growth efficiency (ECD), and higher gross growth efficiency (ECI). Larvae previously reared on 3.75% N diet tended to have lower food assimilation efficiency (AD) and lower nitrogen assimilation efficiency (AD(N)). Although both previous and current diet nitrogen concentration strongly affected larval growth and food utilization, the interaction term between these was not significant for any response variables except ECD and ECI. Because the interaction term reflects the effect of switching per se, the results indicate that there was a metabolic cost associated with switching, but no inherent net cost or benefit of diet-switching to growth.     

Spatial and temporal variation in the parasitoid assemblage of an exophytic polyphagous caterpillar

Abstract 1. Over 3400 larvae of the polyphagous ground dwelling arctiid Grammia geneura were sampled and reared over seven generations in order to characterise its parasitoid assemblage and examine how and why this assemblage varies over time and space at a variety of scales.
2. The total parasitoid assemblage of 14 species was dominated both in diversity and frequency by relatively polyphagous tachinid flies.
3. Both the composition of the parasitoid assemblage and frequency of parasitism varied strikingly among and within sampling sites, seasons, and years.
4. Overall rates of parasitism increased consistently over the duration of caterpillar development.
5. Within sampling sites, parasitism rates were non-random with respect to habitat structure and caterpillar behaviour for the most abundant parasitoid species.
6. The large variability in parasitoid assemblage structure over space and time in this system may be a function of local host population abundance, habitat-specific parasitism, and indirect interactions between G. geneura and other Macrolepidoptera through shared oligophagous and polyphagous parasitoids.     

Dietary complementation across life stages in the polyphagous lady beetle Coleomegilla maculata

We investigated the life history consequences of changes in diet between larval and adult life stages in the polyphagous lady beetle Coleomegilla maculata DeGeer (Coleoptera: Coccinellidae). Beetles were reared on three larval diets: greenbug, Schizaphis graminum Rondani (Homoptera: Aphididae), eggs of the flour moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), and bee pollen. The reproductive performance of females was then evaluated on an adult diet of either greenbug or moth eggs. Moth eggs appeared to be the most suitable diet for larvae, yielding the largest adults, and pollen the least suitable, resulting in the smallest adults and greatly extended developmental time. Pollen‐reared beetles tended to have lower fecundity and fertility than those reared on animal protein, regardless of adult diet. Female fitness was generally increased by a change in diet upon emergence to the alternative source of animal protein, suggesting that dietary complementation occurred across life stages. Among females reared on greenbug, a change of diet to moth eggs reduced the period required for production of 12 clutches and increased egg fertility compared to continued feeding on greenbug. Among females reared on moth eggs, a change of diet to greenbug increased fecundity compared to continued feeding on moth eggs. Among females fed an adult diet of greenbug, those fed moth eggs as larvae had faster production of 12 clutches and higher fecundity. We discuss these novel results in the context of coccinellid life history and ecology and their potential implications for other insects that are predatory as both larvae and adults.