Some beneficial effects of aggregation in young larvae ofPryeria sinica moore (Lepidoptera: Zygaenidae)

Summary The effects of group size on the survival and development of young larvae ofPryeria sinica Moore were investigated by laboratory and field experiments. Under laboratory conditions, about 20% of isolated larvae died of unsuccessful feeding in the first instar, however, larvae survived successfully in aggregations of four or more individuals. In the field, larvae emerge in early spring and wait for new leaves to open before feeding. In this period, the larger the group size of hatchlings the survival rate became higher. The nest-web spun by hatchlings was considered to play an important role in protecting them from desiccation. In the period that larvae began to feed on leaves, more than 36 larvae are necessary to aggregate for the successful establishment of feeding groups. The nest-web played an important role also in the establishment of feeding group. However, the natural group size of the first instar larvae was larger than the minimum group size to spin a sound nest-web in the field experiment. On the other hand, in later stage, larvae in a large group did not have an excess advantage in survival or developmental rate over larvae in a small group. It was found that the experiments on survival and developmental rates could not explain the reason that this species maintain large compact groups in the most part of larval period. The study was partially granted by the Ministry of Education (No. 374205).     

One-host one-parasitoid system: Population dynamics of a zygaenid mothPryeria sinica Moore in an undisturbed habitat

Summary The population dynamics ofPryeria sinica was investigated in an undisturbed area in 1976–1979. We analyzed the process stabilizing the local population by the life table approach for immature stages and the mark-recapture method for the adult stage. Females usually layed about 130 eggs in an egg-mass. The shape of the survivorship curve was convex and was characterized by a relatively low mortality in the egg and larval stages and by a relatively high mortality in the prepupal and pupal stages. The low mortality in the early stage seemed to be not only due to the peculiar life cycle of this species (larvae develop in early spring when natural enemies are not active) but due to their protective nest-webs, larval warning coloration and repellent smell. The high mortality after cocooning was caused by severe parasitization byAgrothereutes minousubae. The number of adult in the population varied by 2.10-fold, which was less than that of other gregarious moths. The life table data and field observations suggest that adult female dispersal would have acted as a stabilizing factor, andA. minousubae as a conditioning factor on the dynamics of the moth population.     

Population dynamics of a migrant skipper butterflyParnara guttata (Lepidoptera: Hesperiidae) II. Survival rates of immature stages in paddy fields

Summary Survival rates and mortality factors of a migrant skipperParnara guttata were censused in paddy fields in 4 localities of central and western Japan during 1975–1980, and 10 life tables were developed for 3 generations. Mortality rates of eggs, older larvae and pupae were high but those of younger larvae were very low. The high mortality rate was caused by parasitoids. Ten primary and 6 secondary parasitoids were recorded. Out of three egg parasitoids,Telenomus sp. was predominant. Larval parasitoids,Apanteles baoris andPediobus mitsukurii which are specific to genusParnara were predominant in wet land habitat, i.e. paddy land. On the other hand, tachinid flies which have a wide host range and a high searching ability were predominant in dry land habitats. Mortalities of eggs and pupae contribute more to the change in the total mortality of immature stages than those of other stages. egg parasitoids which are the major mortality factor of eggs acted density-dependently but larval and pupal parasitoids did not do so. The total mortality of immature stages occurred more or less density-dependently. Larval density on rice plants scarecely affected the survival rate of larvae.     

Population dynamics of a phytophagous lady-beetle,Epilachna vigintioctopunctata (Fabricius), living in spatio-temporally heterogeneous habitats. III. Effects of habitat structure on population dynamics

Temporal changes in the population size of a phytophagous lady-beetle were analyzed to identify mechanisms affecting lady-beetle population dynamics at different spatial scales. The study area (15 ha) included 18 habitat patches. The major host plants were potato for first generation larvae and eggplant for second generation larvae. The habitat patches were classified into three groups according to the major host plants in each patch: P-E patches (both host plants available), P patches (potato only), and E patches (eggplant only). The winter disappearance of adults in the whole study area, and larval mortality in E patches were apparently the most important factors disturbing the overall population density. Density-dependent movement of females appeared to have the greatest stabilizing effect on the yearly fluctuation of population density. Rate of increase of female adults from the first to the second generation,R, was generally higher on eggplants in E patches than in P-E patches because the adult density of the first generation was much higher in P-E patches. The yearly fluctuation of adult density in each generation tended to be less in patches with all habitat components necessary for the full life cycle (P-E patches). However, such patches were not favorable for first generation females, as indicated by the lower rate of increase from the first to the second generation. The density and stability of lady-beetle populations is discussed in relation to habitat structure.     

A life table ofMogannia iwasakii (Homoptera: Cicadiidae) in sugarcane field of Okinawa

Summary A preliminary life table of the sugarcane cicada,Mogannia iwasakii, which became a pest of sugarcane in Okinawa recently, was presented based on the results of field survey. Mortality rates during egg and later nymphal stages were low but a remarkable reduction of the number of individuals in a cohort took place at the 1st nymphal instar, mainly due to predation by ants. Ants also killed the cicada during and just after the final ecdysis and birds concentrated their feeding on adults. Prenatal mortality of adults was, however, low in the infested area, suggesting the ‘escape’ of cicadas from the pressure of predators. This espace is considered to be a factor responsible to the maintainance of outbreak for a long time. Increase in the acreage of ratton canes and wide-use of organochlorine insecticides during the 1960’s were assumed to be the factors responsible to the initial increase of this species.     

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.     

Age-specific survivorship analysis ofHeliothis spp. Populations on cotton

Summary Population dynamics ofHeliothis virescens (F.) andHeliothis zea (Boddie) (Lepidoptera: Noctuidae) eggs and larvae were studied for two years in a small plot of cotton,Gossypium hirsutum (L.). Due to morphological and ecological similarities, the pooledHeliothis population was considered for most of the analyses. Two generations ofHeliothis eggs and larvae were completed during each year. Stage recruitment was estimated for the eggs and larval instars 2–6, and recruitment variances were estimated by a Monte Carlo method. A modified form of the Weibull distribution was developed and used as a model to characterize survivorship curves for each of the fourHeliothis generations. A Type I survivorship curve (mortality rate increasing with age) was inferred for both Generation 1 (early season) data sets, whereas a Type II survivorship curve (mortality rate constant and thus independent of age) was inferred for both Generation 2 (late season) data sets. The shapes of the survivorship curves for the individualH. virescens andH. zea populations were inferred to be the same as those for the pooled populations. Analysis of the contributions of various factors toHeliothis stage-specific mortality indicated that natural enemies (predators and parasites) and the availability of food for larvae were responsible for between-generation differences in survivorship patterns.     

Studies on the spatial distribution pattern of larvae of the mosquito,Anopheles sinensis, in rice fields

Summary The spatial distribution patterns of the population ofAnopheles sinensis larvae were studied in the rice field area in the suburb of Urawa city in Japan, during the summer seasons in 1973 and 1974. The distribution pattern of the larval population within the field, analysed by the m−m regression method, indicated that the basic component of larval distribution was not a group of individuals but a single individual and such components were distributed contagiously over the field. This basic pattern did not change significantly according to developmental stage, census date or field. Therefore, we could describe the distribution pattern of the population in a rice field by the single linear regression, x=0.021+1.339x(r²−0.912). Also, the relation for the whole population in the field area including the five fields could be shown by the linear regression, x=0.049+1.749x(r²−0.959). The value of α remained to be nearly equal to zero, but the value of β became larger than the value for the single-field relation. Such a change in distribution pattern seemed to reflect the greater heterogeneity in conditions among the fields than within individual field. Using the information on the distribution patterns mentioned above, some considerations were given on the sampling plans for mosquito larvae, including samplesize determination and application of sequential methods to estimate population size as well as to classify population level.     

Life tables for worker honeybees

Summary Life tables for worker honeybees covering all life span, and those for adults, were prepared for three seasonal cohorts,June bees, July bees andwintering bees. Survivorship curves forJune andJuly bees show a convex type being exceptional for insects, with relatively high mortality at egg and feeding larval stages and at later adult stage after most bees became potential foragers. Adult longevity greatly lengthens inWinteriing bees and survivorship curve drops approximately with the same rate. A remarkable similarity of survivorship curves for men and honeybees was demonstrated, apparently due to highly developed social care in both. Some comments were given on mortality factors. The importance of life tables for population researches was shown by applying our result to the population growth curve made byBodenheimer, based upon the data byNolan. At the asymptote of the uncorrected curve, the ratio of total population estimated by uncorrected curve to that by corrected curve reaches about 3∶2. Contribution No. 821 from the Zoological Institute, Faculty of Science, Hokkaido University, Sapporo, Japan. Contributions from JIBP-PT No. 45. This study was in part supportod by a grant in aid from the Ministry of Education for the special project research, “Studies on the dynamic status of biosphere.” Population and bioeconomic studies on the honeybee colonies. II. We express our sincere thanks to Dr. YosiakiIt?, National Institute of Agricultural Sciences, Tokyo, for his kind stimulation and advices to the present work.     

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.     

The dynamics of laboratory populations ofCallosobruchus chinensis andC. Maculatus (Coleoptera: Bruchidae) in patchy environments

Summary Experiments are described showing the long-term dynamics of two species of bruchid beetles (Callosobruchus chinensis andC. maculatus) in arenas in which the resource of 50 black-eyed beans is divided between 5, 10 or 50 ‘patches’. Both species of adult beetles exhibit clumped distributions between patches. Within a patch there is a tendency for a density dependent reduction in (1) eggs laid per female, (2) the proportion of eggs hatching per bean (C. chinensis only) and (3) larval survival which is strongly overcompensating (particularly inC. maculatus). A discrete generation model is used as a framework to draw these results together and show how the different factors affecting natality and mortality can influence the population dynamics. Finally, the importance of the resource renewal interval in influencing the period of the population cycles is discussed.     

An introduction experiment of an herbivorous lady beetle: Characteristics of the adult population

In May 1971, 45 adults of an herbivorous lady beetleEpilachna niponica (Coleoptera: Coccinellidae) from Asiu Experimental Forest were introduced into a botanical garden of Kyoto University, where is 10 km south of the southern limits of its distribution with being 3–5°C warmer than the original site. The introduced population of the lady beetle was thus investigated from 1975 to 1981. Mark-release-recapture experiments were applied to individual adult beetles, to estimate population size and daily survival rate. Overwintering adults emerged from hibernation around early April, reaching peak numbers in late April to early May, then gradually declined to late June. No adults remained at the end of June. Adult survival was maintained at a high level to early May, and declined consistently until late in the reproductive season. New adults began to emerge in late June and quickly reached a peak in early July; thereafter they decreased in number and had entered hibernation by late October. In spite of seasonally deteriorating food resources and heat stress in summer, new adults showed moderately high survival during the inimical period. New adults which emerged later in the season tended to be smaller in body size than those that emerged early. The proportion of females in the new adult population gradually increased throughout the pre-hibernating period, suggesting that male-biased mortality occurred during this period. When compared to the source population, the introduced population had a higher rate of population growth. Coupled with the improved population growth, heavy leaf damage during the larval period suggested that intensive intraspecific competition was most likely to occur among larvae in the introduced population.     

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.     

Life table analysis of the green rice leafhopper,Nephotettix virescens (distant) (Hemiptera: Cicadellidae), and efficient vector of rice tungro disease in asynchronous rice fields in Indonesia

Summary Population dynamics ofNephotettix virescens was studied in 17 paddy fields transplanted at intervals of about 1 month in 1988–1990. The adult density was highest either in the immigrant or the 1st generation and sharply decreased to the 2nd generation. The survival rate of the 1st generation was lowest in the transition season when areal population density increased. Key factor analysis revealed that the nymphal and adult mortality of the 1st generation (kn) was the principal source of population fluctuations. No significant correaltion was found between kn and natural enemy density, natural enemy density/healthy egg density, or the precipitation during the nymphal period. On these bases adult emigration was suspected to be the key factor. Areal population build-up ofN. virescens in the transition season was considered to occur as a result of increasing immigration to young stages of rice. Contribution from Indonesia-Japan Joint Program on Food Crop Protection (ATA 162), which was implemented by the Directorate of Food Crop Protection, Ministry of Agriculture, Indonesia and Japan International Cooperation Agency, Japan.     

Population dynamics ofTelenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas' disease vectors IX. Larval competition and population size regulation under laboratory conditions

Summary A series of experiments were carried out with the endophagous egg parasiteTelenomus fariai on its hostTriatoma phyllosoma pallidipennis to determine the possible role of intraspecific competition by the parasite progeny in population regulation of the parasite. Eight parasite densities (1, 3, 5, 10, 20, 30, 40, and 50 individuals per vial) were used, and the design of sequentially sacrificed replicates applied. Survivorship curves for each density indicated smaller number of progeny per host at higher densities, and the shapes of the curves suggested a relatively early mortality process.Morris' linear regression technique for determining within-generation density-dependence was used, and the results showed that only larval mortality could be identified as density dependent. The same technique applied within the larval stage proved that only mortality of larvae in their second, third, and fourth day of development were responsible for population regulation. The applicability of the technique, as well as the relevance of the results for natural population, is discussed.     

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.     

A study of natural populations of the celery leaf-minerPhilophylla heraclei L. (Diptera, Tephritidae)

Summary The study of the population dynamics of the celery leaf-miner,Philophylla heraclei, must take into account the fact that the host-plant is not present permanently and that it develops between the two annual generations of the insect. This development affects in particular the quality and quantity of the leaves fed upon by the mining larvae. It is possible to know the numerical change of the populations from one annual generation to the next (bivoltine insect) and from one year to the next by counting the populations in the various larval instars, responsible for the injury caused to celery. An accurate method of counting of larval numbers, taking into account their change parallel with that of the host-plant, is described in this text. It makes it possible to know with accuracy the duration and importance of the infestations in relation to external factors and to factors inherent to the host-plant itself. Certain traits of larval behaviour, particularly the transfer of a larva from its original mine to a secondary mine as a result of food shortage can thus be established and estimated quantitatively.     

A granulosis virus, possible biological agent for control ofAdoxophyes orana (Lepidoptera: Tortricidae) in apple orchards

Summary Based on the results of ecological surveys ofAdoxophyes orana and its natural enemies in apple orchards sprayed with the granulosis virus and control plots, we constructed working models to simulate the population dynamics in three different experimental plots; those treated with chemical insecticides, those with granulosis virus, and controls. The number of individuals killed by predators, parasitoids and by miscellaneous mortality factors could be calculated on the assumption of imperfect density relations; that is, relation of the number killed by each factor with the initial number of larvae was represented by a curve with an upper asymptote. We could estimate the proportion of virus infection using curves with upper asymptotes. Rate of increase from pupa to middle instar larvae of the next generation was subject to strong density-effect. Simulation we proposed in this paper suggests a possibility that a single spray of the virus at 1st generation can reduce not only the number ofA. orana larvae in the 2nd and 3rd generations but also the degree of fruits injured by this insect. Spray of chemical insecticide is considered to be ineffective in reducing the pest density and the degree of injury to low levels in subsequent generations, as compared with untreated plot, where the density of the 1st generation larvae is low.     

Comparison of life tables between the solitary eumenid waspAnterhynchium flavomarginatum and the subsocial eumenid waspOrancistrocerus drewseni to evaluate the adaptive significance of maternal care

Summary I compared life tables between the solitary eumenid waspAnterhynchium flavomarginatum Smith and the subsocial eumenid waspOrancistrocerus drewseni Saussure in Kyoto, Japan, during 1980–1983. The subsocial eumenid is parthenogenetic in this study area. There were 9 identified mortality factors in the solitary eumenid and 7 in the subsocial eumenid, 6 of which were common to the two eumenids. The important differences of mortality between the two eumenids were seen in the egg, larval, and prepupal stages. In the egg stage, mortality by the phorid flyMegaselia sp. was much lower in the subsocial eumenid (1.4%) than in the solitary eumenid (15.0%) likely because of the matenal care of the subsocial eumenid (progressive provisioning and other related behavior), which reduced predation pressure. In the larval stage, mortality by the miltogrammine flyAmobia distorta was also lower in the subsocial eumenid (8.1%) than in the solitary eumenid (23.8%) also probably because of the maternal care of the subsocial eumenid. A comparison of mortality in the two eumenids between the stable, long continuing natural nest sites and the additional temporal ones showed that the phorid fly remained near its birth place and parasitized stable nest sites. The miltogrammine fly followed returning eumenid wasps and parasitized those nest sites that have a high host density. In the prepupal stage, mortality by endogenous death was higher in the subsocial eumenid than in the solitary eumenid. Mortality due to the rhipiphorid beetle was also higher in the subsocial eumenid probably due to more frequent flower-visits by the subsocial eumenid. The defense mechanism of the subsocial eumenid was discussed in relation to the evolution of subsociality. Contribution to the ecological studies of the eumenid wasps. I.     

Below-replacement fertility: Determinants and prospects in South Asia

Although 10 countries and two of China’s special administrative areas, totalling 1,528 million people or 44 per cent of Asia’s total population, are now characterized by fertility rates below long-term replacement levels, no such countries are yet found in South Asia. This paper first examines the characteristics of 12 Asian administrations with very low fertility at various stages of their fertility declines and then compares the findings with the present situation in three South Asian countries, Sri Lanka, India and Bangladesh. This allows a prediction of when the South Asian countries will reach replacement fertility in accord with the trends in two key criteria, the percentage of girls in secondary school and the infant mortality rate. These conclusions are then buttressed for each country by the findings of anthropological demographic research programs in which the authors were involved. The predictions are that all three countries will attain a total fertility rate of 2.1 within the next 30 years and that the UN2000 Revision of the medium population projection is plausible in that regard. However, the authors part company with the UN projection in their assessment that the nature of these societies means that they will all subsequently fall to still lower fertility levels.