A study of natural populations of the celery leaf-miner,Philophylla heraclei L. (Diptera, Tephritidae) II. Importance of changes of mines for larval populations

Summary The leaf-mining larvae of the celery fly,P. heraclei, have the ability to leave their primary mine to bore a secondary mine in another leaflet or leaf. This phenomenon is always numerically significant under natural conditions (depending on the plots and the years, 38 to 97% of the larvae observed made such a change), although it is not an obligatory behavior. The distribution of the secondary mines on the leaflets and on the leaves varies with the importance of the migrations, with the total unoccupied and healthy leaf area and with the relative position of different leaves on the plant. These migrations are due to the insufficiency of healthy food available for the larvae: either because the quantity of parenchyma offered by a leaflet is too small for a larva, or because there is intraspecific competition following multiple egg-layings on the same leaflet, or because there is a deterioration of the quality of the parenchyma, particularly following the development of celery leaf-spot. The possibility for the larvae ofP. heraclei to bore secondary mines appears to be an extremely important factor in the population dynamics of this species. This permits the reduction of the negative effects of intraspecific competition and celery leaf-spot, and permits the colonization by the larvae of young leaves not used by the females at the time of egg-laying.     

Climatic change and migration from Oceania: Implications for Australia,New Zealand and the United States of America

It is possible that climatic change may stimulate population movements as people turn to migration as one strategy of adaptation. This paper attempts to assess possible migration flows which may occur, in response to climatic shifts over the next thirty years, from small island states in the south-west Pacific ocean region to the United States, Australia and New Zealand. It is argued that the small island states appear vulnerable to climatic change, with low coral atolls being most at risk. Adverse impacts of climatic change will be one extra pressure on small island states, many of which are already struggling to cope with sustainable management of their natural resources and with the demands of their rapidly growing populations for education, housing and employment. The migration strategy is likely to entail significant medium-term health, psychological and social costs for some Pacific island migrants as they try to move or cope with life in western industrialised societies.     

Factors and consequences of a non-functional alary polymorphism inPyrrhocoris apterus (Heteroptera: Pyrrhocoridae)

Functional alary polymorphisms have been studied rather extensively in several insect species. This review article deals with factors controlling wing polymorphism in a flightless species,Pyrrhocoris apterus (L.), and discusses its adaptive significance and mechanisms for their persistence under natural conditions. The macropterous morph is determined by a recessive allele whose penetrance depends on photoperiod and temperature. Natural populations of this species contain a small fraction of flightless macropters. The disadvantages of being a macropter (increase of development time, decrease of fecundity) are minimal, while the benefit may consist in the tendency to prereproductive arrest of ovarian development in teneral females. It prevents establishing a second generation which would mostly die during the next winter. The mechanism of alary morph regulation may be an ancestral trait linkingP. apterus with other polymorphic Heteroptera, while its decreased penetrance may be a derivative character. Variation in fitness due to alary morphs is small compared to the one associated with differences in body size. The latter is environmentally determined, and not linked to the genetic basis of wing polymorphism. In the “mosaic” of phenotypes of various size the significance of the genetic macroptery may be close to neutral.     

Pollination and factors limiting fruit set of chasmogamous flowers of an amphicapric annual,Polygonum thunbergii (Polygonaceae)

Summary We observed phenology and insect visitors of chasmogamous flowers of an amphicarpic annual,Polygonum thunbergii (Polygonaceae) in Kyoto, Japan, and clarified limiting factors for fruit set by bagging and hand-pollination experiments. Flowering season was one month from late September. Flowers were visited by various insect groups (total 30 families and 64 species) and effective pollinators were lower bees (Halictidae) and wasps (Vespidae) in the early flowering season, and middle-sized flies (Syrphidae and Calliphoridae) in the later season. Nectar was secreted at a constant rate (0.013mg sugar/hour) throughout a day and >90% of it was consumed by insects. The average number of flowers per ramet was 122, of which 95% were pollinated in the natural conditions. Bagging experiments showed that 47% of flowers were self-pollinated even under no pollinator visits. Despite of a high probability of cross pollination, the probability of fruit set within the ramet was 0.30 due to resource limitation. We discussed possible adaptive significance of cleistogamous flowers under the condition that seeds could be produced by chasmogamous flowers through self-pollination even under pollinator limited conditions.     

Seasonal change of nest survival and related aspects in an aggregation ofLasioglossum duplex (Dalla torre), a eusocial halictine bee (Hymenoptera: Halictidae)

Summary Nest survival in an aggregation of a eusocial halictine bee,Lasioglossum duplex, was censused through an annual cycle. Out of 2,500 nests marked at the beginning of solitary phase in the spring, only 25.5% attained eusocial phase. But 60.5% of 636 nests attaining eusocial phase successfully produced sexual offspring. This shows that solitary phase is the most vulnerable period in the annual cycle. A very low productivity in eusocial phase in the census year was clarified from examination of 99 nests in the autumn. The number of prospective foundresses surviving to the next year was estimated upon the number of brood cells in examined nests and of old females surviving these nests. Deviation between this estimate and the number of nests made in the next spring was 8.8%, confirming a drop of population size to less than one third in the next year. Some nests solitarily made in the summer by dispersed females were examined. Productivity in such nests was extremely low, hence these nests should contribute virtually nothing to the next generation. Possible factors affecting the low productivity in eusocial phase were enumerated though none of these were supported by concrete evidence. Some considerations were given on the relation between obtained results and colony life cycle in eusocial insects, particularly in halictine bees. Bionomics of the eusocial halictine bee,Lasioglossum duplex. VI.     

Application of a marking-and-recapture method for the analysis of larval-adult populations of an insect,Nezara viridula (Hemiptera: Pentatomidae)

Summary Difficulty arises in applying marking-and-recapture methods to insects when the probability of recapture of marked individuals is changed with advancing age, either due to detachment of the mark by moulting (in the case of larvae) or to changes in their survival rate or their behaviour. A modification of the re-recapture method (Leslie et al., 1953) has been devised to analyze the capture-recapture data of the 5th-instar larvae and adults ofNezara viridula L. Estimation of the rate of moulting to the adult stage is made with the aid of additional information on larval survival. Migration rates of the larvae between the two halves of the census field is estimated byIwao's (1963) method. Through these analyses, the dynamic feature of the population during transition from the 5th instar to, adult is revealed. Several problems involved in the application of marking-and-recapture methods to insect populations are discussed. Contribution from the Entomological Laboratory, Kyoto University No. 392.     

Mining pattern of the honeysuckle leaf-minerPhytomyza lonicerae

Summary The mining pattern of the honeysuckle leaf-minerPhytomyza lonicerae which feeds onLonicera gracilipes was analyzed with special reference to the avoidance of parasitoids' attacks. The leaf-miner makes the linear mine with branches and crosses by turning and branching, off the backtracking mine. Both average numbers of turnings and branchings were about 4. The speed of mining was 1.5 times faster in backtracking than in advancing. The mine width increased as the mining larva grew and total volume of consumed plant tissue was nearly constant independent of variability of the number of turnings and other characteristics of the mine. The majority of larval mortality were due to attacks by parasitoid wasps (Chrysochairs pentheus andDiglyphus minoeus), which locate the present position of leaf-miners, by tracing mine tracks, and wasps attack leaf-miners at a higher rate in backtracking than in advancing. The adaptive significance of branching and crossing by leaf-miners was discussed in terms of the reduction of parasitoids' attacks.     

Rapid evolution of larval development time during mass-rearing in the melon fly, Bactrocera cucurbitae

A quantitative genetic analysis of rapid evolution of a life history trait has been conducted on the first 24 generations of mass-rearing in the melon fly Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). The phenotypic variance of larval development time in each generation was divided into genetic and residual components. Mean and phenotypic coefficients of variation of larval development time decreased gradually as generations proceeded as a result of artificial selection for shorter larval period in the mass-rearing procedure. There was a trend that additive genetic coefficients of variation in larval development time decreased with generations. These changes are entirely attributed to genetic responses to laboratory selection under the mass-rearing environment because the population was maintained at a very large size so as to exclude random genetic drift and inbreeding depression, which would be other factors responsible for the observed genetic changes. The residual coefficients of variation in larval development time did not change with generations. Realized heritability of larval development time was low. The heritabilities for larval development time estimated from parent–offspring regression at generations 60 and 70, when the evolutionary plateau was asymptotically reached, were not significantly larger than 0. Received: April 22, 1999 / Accepted: September 20, 1999     

Coherent mortality forecasts for a group of populations: An extension of the lee-carter method

Mortality patterns and trajectories in closely related populations are likely to be similar in some respects, and differences are unlikely to increase in the long run. It should therefore be possible to improve the mortality forecasts for individual countries by taking into account the patterns in a larger group. Using the Human Mortality Database, we apply the Lee-Carter model to a group of populations, allowing each its own age pattern and level of mortality but imposing shared rates of, change by age. Our forecasts also allow divergent patterns to continue for a while before tapering off. We forecast greater longevity gains for the United States and lesser ones for Japan relative to separate forecasts.     

Factors influencing development and survival ofCulex pipiens pallens larvae (Diptera: Culicidae) in polluted urban creeks

Summary Factors influencing development and survival ofCulex pipiens pallens immatures in polluted urban creeks were studied in Saga, Japan. Addition of food shortened developmental durations and increased pupal size in floating cages at only the least polluted site out of 4 study sites. There was no evidence for developmental delay due to overcrowding in natural populations. Survival was not increased by artificial feeding at any site. Using the developmental parameters obtained for caged cohorts, survival in natural populations was estimated from differences between the actual number and the expectation if there were no mortality. Pupation rates ranged from 1> to >10% by site and season. Mortality due to predation, evaluated from the difference in pupation rates between caged and natural populations, exceeded 30% in 4 out of 6 cases (twice at 2 sites and once at the other 2 sites). Lethal factors in the creek water caused the predominant mortality in 2 other cases. Regulation ofC. pipiens pallens population breeding in eutrophic, open creeks was discussed in the context of larval dispersal through behavioral interference.     

Analytical studies of light trap records in the Hokuriku district, II the green rice leafhopper,Nephotettix cincticeps

Summary Light-trap records on the green rice leafhopper,Nephotettix cincticeps, were dealt with to study its population fluctuations in the Hokuriku district. Crude data were modified for distinguishing years of the low intensity of infestation by the insect from the rest of years. It is then clearly demonstrated that the low intensity of infestation were ordinarily preceded by heavy snowfall, although heavy snowfall could not be regarded as the only factor checking the vigorous multiplication of the populations.     

A model for analyzing insect stage-frequency data when mortality varies with time

Summary A model is developed for the analysis of insect stage-frequency data which may be applied to populations with age-dependent mortality. The analysis of stage-frequency data is divided into two steps. In the first step, the number of different mortality rates and their values are estimated. The second step provides estimates of developmental rates and variances for each developmental stage and in addition provides estimates of the number of recruits to each stage. The model may be used both in analysis and prediction of insect stage frequencies. Hence, in addition to estimating developmental and mortality rates from stage-frequency data, it may also be used as a simulation model for an insect population. The model is applied to two populations ofHemileuca oliviae Cockerell, a lepidopterous pest of New Mexico grasslands. The model identifies, in the two populations, different mortality rates that are related to plant productivity.     

Effects of parasitoid community structure upon the population dynamics of the honeysuckle leafminer,Chromatomyia suikazurae (Diptera: Agromyzidae)

Population dynamics of a leafminer,Chromatomyia suikazurae (Agromyzidae, Diptera) and its parasitoid community were studied for ten years at seven natural populations along an altitudinal gradient in Japan. This species which mines leaves of a forest shrub,Lonicera gracilipes (Caprifoliaceae), was attacked by 25 hymenopterous parasitoid species. Annually, the parasitoid community structure varied less within a population than among populations. The seven parasitoid communities were clustered into three groups corresponding to the altitudinal gradient: (a) lowland communities dominated by late-attacking, generalist pupal idiobiont eulophids and with highest species diversity, (b) hillside communities dominated by an early-attacking, specialist larval-pupal koinobiont braconid and (c) highland communities dominated by an early-attacking, generalist larval idiobiont eulophid. Annual changes of the host larval densities among the local populations were largely synchronous rather than cyclic. Among these populations, host density levels and mortality patterns greatly varied. By analyzing these inter-populational differences of host mortality patterns, the following conclusions were drawn: (1) The host mortality patterns were determined by the host utilization patterns of the locally dominant species. (2) The host pupal mortality but not larval mortality was related to species diversity but not to species richness itself of each parasitoid community. (3) Density dependence was detected only in pupal mortality at a lowland population dominated by late-attacking pupal parasitoids. These results suggest that interspecific interactions of parasitoids add additive effects to host population dynamics dissimilarly among local populations with different parasitoid communities.     

Estimation of the individual number entering each development stage in an insect population

Summary A method for estimating the number entering each development stage from data obtained by regular sampling through one generation of an insect population was described. This method is consisted of the following two procedures: The provisional estimates are calculated on the assumption that each stage has a common mortality in a sampling interval. Then these estimates are corrected on another assumption that the mortality is different in each stage but constant during a stage. The result of testing its validity with two laboratory populations of the common cabbage butterfly,Pieris rapae crucivora, showed the availability of the present method.     

Seasonal host alternation by the andromeda lace bug,Stephanitis takeyai (Heteroptera: Tingidae) between its two main host-plant species

I studied the seasonal occurrence of the andromeda lace bug,Stephanitis takeyai, on its two main host-plant species. In a secondary forest in Kyoto, this bug altered its hosts seasonally, i.e., from an evergreen shrub,Pieris japonica, in winter to a deciduous shrub,Lyonia elliptica, in summer. In contrast, in Nara park where fewL. elliptica were available, the bug exploited onlyP. japonica. Thus, seasonal host alternation by this bug is not obligate. A comparison of adult longevity and fecundity on the two host-plant species demonstrated the higher quality ofL. elliptica as a food resource. Corresponding to this difference in host quality, there was a dramatic difference in the seasonal population growth in the two study sites. In Nara, the population size at the beginning of the 2nd generation was almost the same as in the overwintered generation, whereas in Kyoto the population size in the 2nd generation was approximately one hundred times as large as in the overwintered generation. Thus seasonal host alternation is adaptive for the bug. In a previous study, I reported that overwintering as eggs in living leaves of their hosts is likely to be common among all the related species of this bug. Thus, this trait can be considered to be a phylogenetic constraint to the group. I speculate that host alternation by this bug has been derived because it is more adaptive from autoecy on an evergreen plant, similar to the pattern currently found in Nara, and that this bug can not only exploit deciduous host due to a phylogenetic constraint.     

The role of medium conditioning in the population dynamics of the housefly

Summary The role of larval medium conditioning on the behavior of larvae and of ovipositing adults of the housefly,Musca domestica L., was investigated through behavioral preference tests. Larvae were strongly attracted to medium conditioned by themselves or other larvae, while adult females overwhelmingly oviposited in fresh medium. Medium conditioning occurs within a few hours after hatching of an egg cohort and is effective in preventing overcrowding of a single site by shutting off further oviposition within 24 hours after eggs are initially deposited. A model of medium conditioning optimizes density for developing larvae and could also provide for regulation of local larval populations. This study was carried out during the tenure of grants from the National Science Foundation (GB-35473) and The National Institutes of Health (1-R01-AI-10988-01) to the senior author.     

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.     

Contest competition inDrosophila subobscura

The role of larval intraspecific competition in laboratory populations ofDrosophila subobscura was investigated. Mortality is density-independent during the first 3 days after hatching but becomes density dependent as development proceeds to pupation. Although total biomass per patch was independent of initial egg density, competition betweenDrosophila larvae leads to the formation of smaller pupae. This resulted in a population that was dominated by suppressed individuals. Development rate ofD. subobscura larvae was not affected by high larval densities. Smaller pupae give rise to females with fewer eggs in their ovarioles. A simple simulation model, predicting the effects of intraspecific competition on the fecundity of the nextDrosophila generation is described.     

Net delay of next conception by contraception: A highly simplified case

Abstract A probability model, building on the work of Perrin and Sheps, is presented and applied. The model makes it possible to follow a cohort of women from a pregnancy outcome to next conception. Principal simplifying assumptions are homogeneity among women and constant fecundity. Cases treated include no contraception; a single contraceptive method practised; and a second contraceptive method practised following a first one. Expected durations to next conception may be calculated in relation to the following factors: outcome of previous pregnancy, length of anovulation, natural fecundability, time first contraceptive method is initiated, characteristics of first contraceptive (effectiveness and continuation rate), characteristics of second contraceptive, and gap between termination of first and commencing second contraceptive. By suitably pairing runs of the model, one can construct experiments in which the only differentiating factor is use and non-use of a specified contraceptive, and, by differencing the corresponding mean durations to next conception, compute the net delay of next conception produced by that contraceptive.