Evolutionary character changes and population responses in an insect host-parasitoid experimental system

In an insect host (the cowpea weevilCallosobruchus maculatus)- parasitoidHeterospilus prosopidis) experimental system, the population densities of the component species oscillated for the first 20 generations and then abruptly stabilized as the parasitoid density decreased. Examination of the host and parasitoid after the 40th generation in the long-term experiment showed that (1) host larvae exhibited contest-type competition (killing other larvae inhabiting the same bean), in contrast to the founder population being scramble-type competitors and (2) the parasitoid attack rate on the host did not change. There was also an evolutionary trade-off between body size and the rates of larval survival and development, suggesting a cost of contest competition on larval survivorship and development. I tested model predictions (Tuda and Iwasa 1998) that (1) host equilibrium population size should gradually decrease as the proportion of the contest type increases and that (2) random attacks of the parasitoid on the host should reduce the rate of increase in proportion of the contest type, and the effect should become manifest especially during the first 20 generations. Two of three host-only replicates showed significant decrease in population sizes. Although the density of emerging adults per bean did not differ between replicates of the host-only and host-parasitoid systems, comparison of the host body size between them on day 270 (at the 13th generation) showed that the host was more contest-type in the host-only system than in the host-parasitoid system, as the model predicted, and later on day 650 the effect of the parasitoid had disappeared.     

Inter- and intraspecific interactions among larvae of specialist and generalist parasitoids

Intra- and interspecific larval interactions that take place in a host body were investigated for two tachnid fliesEpicampocera succincta andCompsilura concinnata (Diptera: Tachinidae) parasitizingPieris butterfly larvae.E. succincta, a specialist onPieris butterflies, showed contest-type intraspecific competition, eliminating all the other conspecific larvae. On the other hand, an extreme generalist parasitoidC. concinnata exhibited scramble-type competition, sharing the host with other conspecifics and suffering reduced body size as a result. However, when these two species occurred together in a single host,C. concinnata had a much higher chance of survival. Moreover,C. concinnata could often survive in the presence of a parasitoid waspCotesia glomerata (Hymenoptera: Braconidae) whileE. succincta could not. The high tolerance ofC. concinnata could be attributable to its being an extreme generalist: To attack and survive on many different hosts, one has to be able to deal with various competitors. The competitive inferiority of the specialistE. succincta, on the other hand, may be a result of relatively recent encounter with, those competitors.     

Effects of interspecific competition on development and reproduction in two giant water bugs,Diplonychus japonicus vuillefroy andDiplonychus major esaki (Hemiptera: Belostomatidae)

Summary We studied the interpecific competition between 2 species of predatory aquatic bugs,Diplonychus japonicus andD. major by conducting a field experiment. We set up 3 types of experimental plots in the paddy fields whereD. major predominated. The two plots contained single species of eitherD. japonicus orD. major, respectively, and one plot had both species in equal number. We compared the development and the reproductive performance between plots in each species. InD. japonicus, the number of eggs and early instar nymphs were significantly smaller in the plots containing both species than in the monospecific plots. However, the numbers of late instar nymphs and newly emerged adults were not significantly different between plots. The proportions of starved nymphs in both plots were larger than those in theD. japonicus's natural habitats. The final densities of adults in both plots were lower than those in the natural habitats. These results suggest that lower density ofD. japonicus in these paddy fields is due to the lack of available food for nymphs rather than the effects of interspecific competition withD. major. InD. major, significant differences were not found in the number of eggs, each instar nymphs and adults. These results suggest that the effects of interspecific competition did not affect the reproductive performance ofD. major.     

The effects of wild cruciferous host plants on morphology, reproductive performance and flight activity in the diamondback moth,Plutella xylostella (Lepidoptera: Yponomeutidae)

The wild crucifers,Rorippa indica andLepidium virginicum, are known to serve as host plants for the diamondback moth (DBM),Plutella xylostella, but they are less suitable than the preferred cultivated cruciferous plant, cabbage, in terms of adult body size and fecundity. The life history traits and flight activity of DBM adults grown on various host plants were investigated. The adults thus reared on each host plant were divided into three size groups (small, medium and large). In general, female adults grown on the wild crucifers were less fecund and lived longer than those reared on cabbage. Flight activity was higher in adults grown on wild crucifers than in those reared on cabbage. Male adults flew longer than females. Fecundity, longevity, flight activity and morphometrical characters of adults were positively correlated with pupal weight in individuals reared on the same host plant. A negative relationship was found between fecundity and flight activity in females of the same size group, but a positive one was observed in females reared on the same host plant.     

Principal component analysis of interrelations in stored-wheat ecosystems infested with multiple species of insects

Summary Principal component analysis (PCA) was used to determine the effects of infestation of bulk stored wheat by multiple species of insects at 30±2°C for 60 wk. Eight 204-liter drums containing wheat at 15.5% moisture content were used as three distinct man-made ecosystems: (a) Control system (2 drums), insect-free; (b) RST system (3 drums), artificially infested withRhyzopertha dominica (F.),Sitophilus oryzae (L.), andTribolium castaneum (Herbst); and (c) COT system (3 drums), infested withCryptolestes ferrugineus (Stephens),Oryzaephilus surinamensis (L.), andTribolium castaneum. The variables measured tri-weekly within each system included carbon dioxide, oxygen, temperature, grain moisture, seed damage, grain weight and volume, dust weight and volume, fat acidity values (FAV) of the wheat, seed germination, microflora includingAlternaria alternata (Fr.) Keissler,Aspergillus glaucus group,Aspergillus candidus Link, and bacteria, insects and the miteTarsonemus granarius Lindquist. PCA provided multivariate synopsis of the data quantifying several important relationships among the variables monitored. Tri-weekly and cumulative 60-wk analyses of each system showed that high bacterial counts were associated with high FAV levels;Tarsonemus numbers were positively related toAspergillus; Alternaria and seed germination were negatively related to FAV, bacteria and grain damage; and that the number of insects was related to the presence ofAspergillus and negatively related to the presence of bacteria. Seed germination andAlternaria infection often decrease rapidly presumably because of infection by fungi of theAspergillus glaucus group. The combined action ofR. dominica andAspergillus spp. enhanced seed damage and increased grain moisture content thus promoting bacterial growth which in turn inhibited insect and mold growth. Fat acidity values increased with time unless seed damage and bacterial infection were extensive as in the RST system. Contribution No. 903, Agriculture Canada, Research Station, 195 Dafoe Road, Winnipeg, Manitoba, R3T 2M9.     

Population studies ofCavelerius saccharivorus Okajima (Heteroptera: Lygaeidae): Adult dispersal in relation to the density

Summary Two adjacent stations (H and L) were set up to study movement ofC. saccharivorus adults in sugarcane fields. At the beginning of the study the density ofC. saccharivorus including all stages of development was quite different between the two. The density of the first generation adult on Station H was about 5 times that on Station L. The number ofC. saccharivorus on both the stations became almost the same one month after the beginning of the study. At the beginning of the study macropterous adults were more numerous in Station H than in Station L. However percentage of macropterous adults on Station L increased after one month whereas that on Station H declined. About 2,000 marked adults were released on each station during the early period of the emergence of the first generation adult. Marked insects were recaptured on both the stations one month after the release. The adults released on the dense population (H) tended to disperse more actively than those on the scarce population (L). Marked macropterous adults moved more actively than brachypterous ones. The density related dispersal of adults was considered to be an important factor to regulate the population density.     

Distribution patterns of and sampling plans forTetranychus urticae Koch (Acarina: Tetranychidae) on roses

Summary I sampledTetranychus urticae Koch (Acarina: Tetranychidae) regularly from four rose gardens in Kyoto and Nara Prefectures in 1988–1990. When mite density was low,T. urticae showed an uneven vertical distribution, being more abundant in the lower third and absent from the upper third of the plants. Mite density was less variable within than between plants, suggesting that a sampling plan which includes more leaves from different plants rather than from different levels of a plant is favourable. Spatial distribution ofT. urticae was nonrandom and followed the negative binomial distribution. In addition, both the Taylor’s power law and the Iwao’s patchiness regression described the distribution well. An empty-sample method for estimating mite density from the proportion of empty sampling units was developed. Sampling plans for determining the sample size required to reach a predetermined precision level, based on this method and by directly counting the mites, were designed. The counting method was more accurate than the empty-sample method. However, when the time factor was taken into account, the latter was more favourable, because it was faster than the former at a density range of 1.5–300 mites leaf⁻¹, which was most commonly encountered in the field.     

Influences of wild crucifers on life history traits and flight ability of the diamondback moth,Plutella xylostella (Lepidoptera: Yponomeutidae)

The larvae ofPlutella xylostella were fed on five wild crucifers,Capsella bursa-pastoris, Lepidium virginicum, Cardamine flexuosa, Rorippa indica, R. islandica and a crop, cabbage. The developmental period of the immature stages, adult longevity, preoviposition period, fecundity and morphometrical characters of the adults were measured. The flight activity of the adults was also measured by the tethered flight method. All the wild plants except forR. islandica were less suitable host plants than cabbage, and larvae which were fed on these less suitable plants emerged as smaller adults with shorter wings. The smaller female adults had lower fecundity but a higher flight activity. Smaller adults measured in terms of their pupal weight among individuals fed on the same host plant had longer wings. These smaller adults with longer wing flew more actively.     

Effects of gall volume on survival and fecundity of gall-making aphidsAdelges japonicus (Homoptera: Adelgidae)

Summary The number of larvae reaching maturity within the gall ofAdelges japonicus was positively related to gall volume, and the relationship between the number of mature larvae and gall volume did not change with different densities of colonized larvae. The population changes in the bud galls ofA. japonicus were surveyed by collecting the galls, which did not suffer predation or parasitism within the galls, from young artificial plantations ofPicea jezoensis over two years. In the year when the density of colonized larvae was high, they suffered a 42% mortality within the galls, whereas mortality was nearly zero in the low density year. The numbers of larvae per gall were positively correlated with gall volume. The regression lines of the number of colonized larvae on gall volume did not differ significantly in the regression coefficients between the two years, whereas the intercept was significantly higher in the year when the density of colonized larvae was higher. However, different within-gall mortality nullified this difference, and the regression lines of the number of mature larvae on gall volume had no significant difference both in the regression coefficients and the intercepts. This suggests that the number of mature larvae per gall was limited by available resources within the gall which were positively related to gall volume. In 25% of the galls in which mature larvae inhabited, the space within the galls were completely filled by the larvae, indicating that space was one of the limiting resources. Gall volume also affected the number of adults that emerged from the gall and the potential number of their progeny.     

Genetic control ofHylemya antiqua III. Differences in pupation ability between two strains

Summary Some relevant traits of a wild (L) and a laboratory (C) strain ofHylemya antiqua (Meigen), determining differences in their pupation ability under experimental conditions have been investigated in relation to genetic control. The wild strain showed an intrinsic higher pupation ability than the laboratory strain. The minimum feeding period was 0.6 days longer for the C strain. The minimum larval dry weight was different for the two strains. With a normal feeding period C larvae pupated on average 1.63 days later than L larvae. By the shortening of the larval feeding period an acceleration of the larval development of both strains was observed: the acceleration of development was more marked for the wild strain. This result has been contrasted with published works onDrosphila. The consequences of these differences as far as the competitive ability of each strain is concerned, have been discussed in relation to genetic control.     

Anopheles subpictus Grassi: Observations on survivorship and population size using mark-release-recapture and dissection methods

Summary A mark-release-recapture experiment to estimate population survivorship and absolute size was performed with wild-caughtAn. subpictus adults at the village of Khano-Harni, Lahore District, Punjab Province, Pakistan during September 1978, the end of the monsoon rainy season, when temporal population abundance was maximized. Daily survival rate estimated from the recapture sequence of marked adults was low, males=0.192 and females=0.343. Survivorship for females estimated by several vertical age-grading procedures ranged from 0.347 to 0.628. Both stage- and age-specific life tables were calculated from vertical age-grading data determined by the dilatation method. Female and male population size was estimated byBailey’s modification of theLincoln Index and was found to average 4478.4 and 6106.8, respectively. The bionomics, survivorship and population size ofAn. subpictus in the Lahore are indicated that this species was probably not important in the transmission of human malaria.     

How small can the difference among competitors be for coexistence to occur

Closely related competitors comprising ofEscherichia coli strains having the same metabolic system and differing only with a few bases on the glutamine synthetase gene in the plasmid pKGN were previously shown to coexist in a chemostat. The differences among these closely related competitors can be considered large enough to allow coexistence as the level of enzyme activity is different. To bring the difference among competitors to the slightest possible, the mutation was introduced on the noncoding region of the plasmid pKGN harbored in the wild-type strain (strain W). The new strain, strain W’, carries the plasmid pKGN’ with a 4-base insertion at theHind III site in the polycloning site of pKGN. As the noncoding region is a nucleotide segment that is not translated into amino acids, the relatedness between strains W and W’ is the closest possible from the genetic point of view. Interestingly, though both strains are almost identical, they can coexist stably in a chemostat irrespective of the initial population size. These experimental results suggest that in the natural ecosystem, no matter how akin competitors are, coexistence is not impossible.     

Genetic control and evolution of acoustic signals in planthoppers (Homoptera: Delphacidae)

Acoustic signals are part of the specific mate recognition system of planthoppers. The genetic control of acoustic signal characters was studied in the planthopperRibautodelphax imitans. Artificial selection for interpulse interval in the female call revealed a large additive genetic component for this polygenic character. Other female call characters showed a correlated response. Some male call characters also appeared to be genetically correlated with the female character selected for, despite the rather different structure of male and female calls. Parent-offspring regression provided significant heritability estimates for those male call characters that also responded to artificial selection in the female call, one of which appeared to be influenced by sex-linked genes. It is argued that the differentiation of this mate recognition system in planthopper populations and species could be the result of founder effects, enabled by the genetic plasticity of the call characters and the existence of a wing length dimorphism in these animals.     

Comparison of dispersal ability between the green leafhoppers,Nephotettix virescens andN. cincticeps (hemiptera: Cicadellidae) by tethered flight

Summary Dispersal propensity of green leafhoppers was compared between a tropical species,Nephotettix virescens, and a temperate species,N. cincticeps. The flight ability was measured with tethered flight technique under laboratory conditions (25°C, 16L-8D). The pre-flight period was shorter and the flight duration was longer inN. virescens than inN. cincticeps in both sexes. No significant correlations were found between the flight activity and morphometric characters for either of the two species. The results suggested that dispersal propensity ofN. virescens is higher than that ofN. cincticeps.     

Egg cluster size variation in relation to the larval food abundance inLuehdorfia puziloi (Lepidoptera: Papilionidae)

Summary Mean egg cluster size ofLuehdorfia puziloi yessoensis varied among habitats. The mean egg cluster size tended to be large when abundance of the larval food leaves expressed as the fresh weight of leaves per unit area at a given habitat was high. Since this variation was observed among closely located study plots (butterflies can easily move between study plots), the egg cluster size variation among habitats is likely to be a result of flexible response by females to varying food abundance for larvae.     

A galling aphid with extra life-cycle complexity: Population ecology and evolutionary considerations

Summary In most gall-forming aphids, only the fundatrix is able to induce a gall on the host plant. InSmynthurodes betae Westw. (and a few other species), F₂ descendants emerge from the mother gall and induce their own, morphologically different galls. This constitutes an added complexity to the already very complex life cycle of gall-forming aphids. We investigated the ecology ofS. betae on marked trees and shoots at four sites in Israel. Gall initiation, gall distribution and density, and temporal changes in clone size within the galls were investigated during two consecutive years. We discuss the possibility that the two-gall life cycle evolved from the typical one-gall system of most gall aphids, and the possible selective advantage of this added complexity in the life-history strategy of gall aphids. Although the total reproductive output ofS. betae is not higher than in related species with a single gall per life cycle, there seems to be an advantage in the subdivision of each aphid clone into several galls, thus reducing the risk of the accidental extinction of the clone (genotype) by environmental factors, including parasites and predators.     

Seasonal life cycle and autogeny in the mosquitoAedes togoi in northern Kyushu, Japan, with experimental analysis of the effects of temperature, photoperiod and food on life-history traits

We studied seasonal changes in the larval population structure, adult size, and autogeny (egg production without a bloodmeal) of the mosquitoAedes togoi on the seacoast of northern Kyushu, Japan. The effects of temperature, photoperiod and food conditions on larval development, adult size and autogeny were studied in the laboratory.Aedes togoi overwintered in both the egg and larval stages and was multivoltine. Adult size was greater in spring and autumn and smaller in summer. Autogeny occurred in spring and autumn but not in summer. Autogenous females were larger than anautogenous females, and larger autogenous females produced more eggs. Laboratory experiments showed that autogeny was promoted under low temperature and short-day conditions which corresponded with spring/autumn conditions in the field. Experimental food limitation in the larval stage greatly reduced adult size and autogenous egg production. In the field, large inter-pool variations in adult size and autogeny rate probably resulted from variations in nutrient and crowding conditions. Seasonal autogeny ofAedes togoi was discussed in terms of life-history strategy under fluctuating environmental conditions where relative advantages of autogenous and anautogenous reproduction alternate seasonally.     

Correlated effects of the response to conditioned medium in the flour beetle,Tribolium castaneum

Summary Dispersal rates were measured in lines selected for high and low response of egg production to conditioned media, and responsivness of egg production was measured in lines selected for high and low dispersal. A positive correlation was found between these two traits, each of which had previously been found to have a simple genetic basis. It is suggested that inTribolium castaneum the sensitivity to environmental conditions is mediated through a Sensor gene, which can activate either response, according to circumstances.