Spatial aggregation across ephemeral resource patches in insect communities: an adaptive response to natural enemies?

Although an increase in competition is a common cost associated with intraspecific crowding, spatial aggregation across food-limited resource patches is a widespread phenomenon in many insect communities. Because intraspecific aggregation of competing insect larvae across, e.g. fruits, dung, mushrooms etc., is an important means by which many species can coexist (aggregation model of species coexistence), there is a strong need to explore the mechanisms that contribute to the maintenance of this kind of spatial resource exploitation. In the present study, by using Drosophila-parasitoid interactions as a model system, we tested the hypothesis whether intraspecific aggregation reflects an adaptive response to natural enemies. Most of the studies that have hitherto been carried out on Drosophila-parasitoid interactions used an almost two-dimensional artificial host environment, where host larvae could not escape from parasitoid attacks, and have demonstrated positive density-dependent parasitism risk. To test whether these studies captured the essence of such interactions, we used natural breeding substrates (decaying fruits). In a first step, we analysed the parasitism risk of Drosophila larvae on a three-dimensional substrate in natural fly communities in the field, and found that the risk of parasitism decreased with increasing host larval density (inverse density dependence). In a second step, we analysed the parasitism risk of Drosophila subobscura larvae on three breeding substrate types exposed to the larval parasitoids Asobara tabida and Leptopilina heterotoma. We found direct density-dependent parasitism on decaying sloes, inverse density dependence on plums, and a hump-shaped relationship between fly larval density and parasitism risk on crab apples. On crab apples and plums, fly larvae benefited from a density-dependent refuge against the parasitoids. While the proportion of larvae feeding within the fruit tissues increased with larval density, larvae within the fruit tissues were increasingly less likely to become victims of parasitoids than those exposed at the fruit surface. This suggests a facilitating effect of group-feeding larvae on reaching the spatial refuge. We conclude that spatial aggregation in Drosophila communities can at least in part be explained as a predator avoidance strategy, whereby natural enemies act as selective agents maintaining spatial patterns of resource utilisation in their host communities.     

Patterns of resource distribution among conspecific larvae in two fruit fly species: Anastrepha fraterculus and Ceratitis capitata (Diptera: Tephritidae)

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Competition between Bufo larvae in a eutrophic pond

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Larval stop,delayed development and survival in overcrowded cultures of Drosophila melanogaster: Effect of urea and uric acid

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Egg size affects larval performance in a coleopteran parasitoid

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Multiple infestation by seed predators: the effect of loculate fruits on intraspecific insect larval competition

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Infection of Drosophila melanogaster by Tubulinosema kingi: stage-specific susceptibility and within-host proliferation

Despite its importance as a model organism very little is known about the interaction between Drosophila and its microsporidian pathogens. Here we report on the relative susceptibility of Drosophila melanogaster life history stages to infection by Tubulinosema kingi, and on patterns of pathogen proliferation. We find that only larvae can be infected, and that this susceptibility decreases with larval age. Following infection, the pathogen shows little subsequent proliferation in larvae, a limited amount in pupae while it replicates greatly in adults. We present evidence that the host launches a cellular immune response after infection with the pathogen, although its effectiveness remains to be demonstrated.     

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Some interrelations between density and genetic structure of aDrosophila population

The importance of relative growth rates and their variances in larval populations is discussed. In competition between wild type andBar mutant larvae ofDrosophila melanogaster, an influence of density on the rate of selection was demonstrated experimentally. A further experiment using two wild stocks also showed that relative fitnesses vary with the degree of competition, and indicated that this may be connected with differences in variance of growth rates.     

Optimization of breeding output for larval stage of Anopheles gambiae (Diptera: Culicidae): prospects for the creation and maintenance of laboratory colony from wild isolates

Domesticating anopheline species from wild isolates provides an important laboratory tool but requires detailed knowledge of their natural biology and ecology, especially the natural breeding habitats of immature stages. The aim of this study was to determine the optimal values of some parameters of Anopheles gambiae larval development, so as to design a standard rearing protocol of highland isolates, which would ensure: the biggest fourth instars, the highest pupae productivity, the shortest duration of the larval stage and the best synchronization of pupation. The density of larvae, the size of breeding water and the quantity of food supplied were tested for their effect on larval growth. Moreover, three cheap foodstuffs were selected and tested for their capability to improve the breeding yield versus TetraMin? as the standard control. The larval density was a very sensitive parameter. Its optimal value, which was found to be ≈1?cm-2 surface area, yielded a daily pupation peak of 38.7% on day 8 post-oviposition, and a global pupae productivity of 78.7% over a duration range of three days. Anopheles gambiae's larval growth, survival and developmental synchronization were density-dependent, and this species responded to overcrowding by producing smaller fourth instars and fewer pupae, over elongated immature lifetime and duration range of pupae occurrence, as a consequence of intraspecific competition. While shallow breeding waters (<3?cm) produced a higher number of pupae than deeper ones, no effect of the breeding habitat's absolute surface area on larval development was observed. Increasing the daily food supply improved the pupae productivity but also boosted the water pollution level (which was assessed by the biological oxygen demand (BOD) and the chemical oxygen demand (COD)) up to a limit depending on the food quality, above which a rapid increase in larval mortality was recorded. The food quality that could substitute the manufactured baby fish food was obtained with weighed mixture of 1 wheat+1 shrimp+2 fish. On establishing an anopheline mosquito colony in the laboratory, special care should be taken to design and maintain the appropriate optimal values of larval density, water depth, daily diet quantity and nutritional quality.     

Predation byXylocoris flavipes [Hem.: Anthocoridae]: Influence of stage,species and density of prey and of starvation and density of predator

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Integrating aspects of larval lepidopteran behavior that enhance survival into conservation plans could increase the overall impact of the efforts. We previously recommended that where possible, maintaining 2–4 ramets of closely-spaced common milkweed would support the development of at least one monarch through pupation, based on a seemingly innate behavior in which monarch larvae (Danaus plexippus) abandon their natal milkweed ramet (Asclepias sp.). Here, we explored the impact of intraspecific competition on larval ramet abandonment behavior and fitness of monarch larvae in small artificial milkweed patches.

We observed larvae reared under direct and indirect intraspecific competition, and larvae reared alone.

We found no influence of intraspecific competition; however, our study provides further support that milkweed ramet abandonment is a seemingly innate behavior. This behavior occurs before all of the available leaf biomass on a ramet is consumed and prior to the pre-pupal wandering stage.

Implications for insect conservation: Results from our study suggest that in the absence of predation, parasitism, and interspecific competition, and when sufficient plant biomass is present to support larval development, the presence of an intraspecific competitor does not influence larval behavior or fitness. Based on milkweed ramet abandonment behavior, we continue to suggest maintaining small patches of 2–4 milkweed ramets when possible.

     

Differential response to larval crowding of a long‐ and a short‐lived medfly biotype

Response of endophytic fruit fly species (Tephritidae) to larval crowding is a form of scramble competition that may affect important life history traits of adults, such as survival and reproduction. Recent empirical evidence demonstrates large differences in adult life history traits, especially longevity, among Mediterranean fruit fly (Ceratitis capitata; "medfly") biotypes obtained from different regions of the world. However, whether the evolution of long lifespan is associated with response to stress induced by larval crowding has not been fully elucidated. We investigated, under constant laboratory conditions, the response of a short‐ and a long‐lived medfly biotypes to stress induced by larval crowding. Survival and development of larvae and pupae and the size of resulting pupae were recorded. The lifespan and age‐specific egg production patterns of the obtained adults were recorded. Our findings reveal that increased larval density reduced immature survival (larvae and pupae) in the short‐lived biotype but had rather neutral effects on the longed‐lived one. Only larvae of the long‐lived biotype were capable of prolonging their developmental duration under the highest crowding regime to successfully pupate and emerge as adults. Response of emerging adults to larvae crowding conditions was similar in the two medfly biotypes. Those individuals emerging from high larval density regimes had reduced longevity and fecundity. Long‐lived biotype individuals, however, appeared to suffer a higher cost in longevity compared with the short‐lived one. The importance of our findings to understand the evolution of long lifespan is discussed.     

幼虫密度对二点委夜蛾生长发育及繁殖的影响

【目的】在不同幼虫密度饲养条件下,研究二点委夜蛾Athetis lepigone生长发育及繁殖的情况,明确幼虫密度对该害虫的室内种群增长的影响。【方法】本实验设置5个幼虫饲养密度即1,5,10,20和30头/瓶(750 mL),分别观察5个饲养密度下该虫的各个龄期及整个幼虫发育历期及存活率、蛹重、蛹期以及成虫生殖情况。【结果】幼虫密度对该虫幼虫各龄期及整个幼虫发育历期及存活率、蛹重、蛹期以及成虫生殖情况均有显著性影响。整个幼虫发育历期随着密度的增加而缩短,10头/瓶达到最短(18.27 d),之后随着幼虫密度的增加而显著延长;幼虫至蛹的存活率随着密度增高而显著下降,30头/瓶最低(39.37%)。蛹期随着密度的增加而延长(10头/瓶除外)。蛹重和每雌产卵量均以1头/瓶最高,随着幼虫密度的增加而显著下降。雌雄蛾寿命均以10头/瓶最长,与1和5头/瓶没有显著性差异。生命表分析显示:二点委夜蛾的种群增长指数以5头/瓶最高,幼虫密度过低或者过高均不利于种群增长。【结论】幼虫密度是影响二点委夜蛾种群增长的重要因子之一。     

THE ECOLOGY OF ANISOPUS FENESTRALIS SCOP. (DIPTERA) IN SEWAGE BACTERIA BEDS

The assessment of populations of Anisopus fenestrate (Scopoli, 1763) in sewage bacteria beds by the trapping of adults at the surface has been previously proved to be unsatisfactory. Methods were therefore developed for determining the numbers of egg masses, larvae and pupae as well as the total organic matter in a unit volume of bed. The relative larval populations of three areas under investigation were related to the loading of the beds as measured by the sewage strength and rate of application. In the upper 2 ft. 6 in. the numbers of larvae and pupae per unit volume of bed decreased with depth. There was no evidence of any vertical migration of the larvae before pupation. The horizontal distribution of larvae was found to be affected by the method of distribution. In beds served with fixed spray jets larvae are more abundant nearer the jets than at some distance away. In beds served with travelling distributors the larvae are more abundant in the zones below the jets than in the drier intermediate zones. In both cases relatively higher percentages of pupae in the drier zones might indicate a horizontal migration of larvae before pupating or of the pupae themselves.
The incidence of the larvae throughout 1949 in all three beds showed recurring peaks. In the following year a more intensive investigation on one area showed that this was due to successive generations of the fly; the proximity of the peaks is determined mostly by temperature and their size by the amount of food available in the bed during the larval grazing phase.
In the area not treated with insecticide intraspecific competition during a period when food was limiting, due to depletion by the previous generation, resulted in a natural reduction in the population. In the area treated with insecticide the food was retained and this natural control was thereby delayed.     

Life history variability of a grazing stream insect (Liponeura cinerascens minor; Diptera: Blephariceridae)

SUMMARY 1. Fourteen populations of Liponeura cinerascens minor from different sites in the Swiss Alps were investigated. Our goals were to describe the life history of this blepharicerid species and to analyse how it was influenced by temperature and food availability. 2. Temperature regimes and periphyton density at the sampling sites varied considerably. Mean annual temperature ranged from 3.8 to 6.5 °C, with annual amplitudes (i.e. the difference between the mean of the coldest and the warmest month) between 3.7 and 12.0 °C. Averaged periphyton density, which was assessed semiquantitatively on a scale from 0 to 5, ranged from 1.1 to 3.3. 3. Larvae or pupae of L. c. minor were found between January and October. Presence of larvae or pupae at individual sites ranged from 100 to 224 days and appeared to be mainly a result of hatching patterns. Individual larval development required between 7 and 20 weeks. Our findings suggest that L. c. minor is univoltine and undergoes an extended (≥4 months) egg dormancy during late summer, autumn and winter. 4. Development of L. c. minor was observed from <1 to 15.8 °C, indicating it is a cold‐stenotherm. Its geographical distribution seems to be constrained by high summer temperatures. Within the tolerated temperature range (0–16 °C), however, temperature had no apparent effect on hatching, larval growth rate or pupal size. 5. Larval growth rate and size of pupae were significantly correlated with food availability. Completion of larval development required 300–400 degree‐days at sites where periphyton was readily available, but >800 degree‐days where periphyton availability was low. 6. No evidence for intraspecific food competition was found, whereas food competition by other blepharicerid species, mainly Hapalothrix lugubris, appeared to be a major reason for the observed food limitation. However, as L. c. minor is much more widespread than H. lugubris, food competition between these two species is likely to occur infrequently. We therefore contend that variations in the life history of L. c. minor are mainly a consequence of its thermal constraints.     

Biochemical and genetic studies on alkaline phosphatase ofCeratitis capitata

Two forms of alkaline phosphatase exist in the integument of the “white pupae” (wp) and dark pupae (dp) mutant strains ofCeratitis capitata, during transition from larvae to pupae. They were separated by DEAE-cellulose chromatography. Both isoenzymes have a molecular weight of approximately 180,000 and two pH optima, at 9.4 and at 11.0. The isoenzymes of the “dark pupae” mutant catalyze the hydrolysis of phosphotyrosine and β-glycerophosphate but not phosphoserine, phosphothreonine, ATP, and AMP. In contrast, the isoenzymes of the white pupae mutant hydrolyze all the substrates tested. The ALPase 1 of the dark pupae mutant was inhibited byL-tyrosine, butL-phenylalanine had no effect on either isoenzyme. The effects of divalent cations, EDTA, temperature, urea, and 2-mercaptoethanol were also investigated. Electrophoretic analysis did not reveal any variants of the larval and pupal isoenzymes, but ALPase A, an adult stage-specific isoenzyme, was found to be polymorphic. The electrophoretic variants were shown to be controlled by three codominant alleles located on the third chromosome ofCeratitis capitata. Since we found no hybrid enzyme, we conclude that ALPase A is monomeric.     

Coexistence of three differentDrosophila species by rescheduling their life history traits in a natural population

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