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.     

Competition between laboratory populations of green leafhoppers,Nephotettix spp. (Homoptera: Cicadellidae)

Summary Intra- and interspecific competition between laboratory populations of four green leafhoppers,Nephotettix spp. was studied in the laboratory under three different temperature regimes of 24°C, 27°C and 30°C. For the single-species population of the three tropical species, the equilibrium density increased as the temperature increased. On the other hand, for the temperature speciesN. cincticeps, the highest equilibrium density was at the intermediate temperature and the lowest at high temperature. Interspecific interactions between two tropical (N. virescens vs.N. nigropictus), a tropical and a temperature (N. virescens vs.N. cincticeps) and a rice-feeding and a grass-inhabiting (N. virescens vs.N. malayanus) Nephotettix species were also studied in the laboratory at the three temperature regimes. Temperature differentially affected the outcome of competition between twoNephotettix species. BetweenN. virescens andN. nigropictus, the latter was more successful over the former at low and intermediate temperatures, while the former was more successful at high temperature. BetweenN. virescens andN. cincticeps, the temperate species inhibited the growth of the tropical species at low temperature while the tropical species inhibited the growth of the temperate species at high temperature. At intermediate temperature, the population ofN. virescens persisted at a slightly higher density over the population ofN. cincticeps. Between the rice-feedingN. virescens and the grass-inhabitingN. malayanus, regardless of temperature the population density of the latter was greatly reduced and later became extinct while the population of the former continued its growth. These consequences of competition between twoNephotettix species conformed fairly well to those predicted by theLotka-Volterra model using demographic parameters specified for each species.     

Population dynamics of the whitebacked planthopper,Sogatella furcifera (Hemiptera: Delphacidae) with special reference to the relationship between its population growth and the growth stage of rice plants

A population census was conducted to describe the effects of the growth stage of rice on the population dynamics ofS. furcifera, in particular, on immigration, seasonal abundance, population growth rate, and wing-form expression. The number of immigrants was highest on rice plants 17 to 30 days after transplanting (DAT), which suggested that immigrants prefer to settle or remain more on rice plants at the tillering stage (approximately 20–30 DAT). Population growth rate from immigration to the 1st generation decreased with an increase in rice plant age. In contrast, population growth rate from the 1st to 2nd generation was not influenced by rice plant age and was negatively density-dependent. The percentage of macropters (flyers) was positively related to the growth stage of rice when rice was in the vegetative or early part of the reproductive stage, and reached 100% at about 10 days before heading (booting stage). Threafter, most adult females molted into macropters regardless of population density. The roles of host plant age and crowding effect on the population dynamics ofS. furcifera are discussed.     

Dynamics of buried seed population and seedling cohorts of two dominant weeds in a hill agroecosystem of the humid subtropics of India

Dynamics of the buried seeds and plant population of two dominant weeds, viz.,Emilia sonchifolia (Linn.) DC. andRichardsonia pilosa HBK were studied in the crop fields of Meghalaya, north-east India during radish and maize cropping and intervening fallow periods. The total buried seed population ofR. pilosa was always larger than that ofE. sonchifolia, but the germinable fraction was invariably greater in the latter. A major portion (39–41%) of the viable (germinable+dormant) seed population in both weeds was confined to the surface soil layer (0–5 cm). The viable seed population ofE. sonchifolia peaked during April, while that ofR. pilosa showed two peaks (during August and December). The survival pattern and half-lives of seedling cohorts showed, some differences in the two weed species, but both being summer annuals, their populations behaved in a similar manner by showing higher seedling recruitment (K) and survivorship (p) rates in the summer crop (maize) than in the winter crop (radish). However, the density of plants that could attain adulthood was significantly higher inE. sonchifolia thanR. pilosa which might have resulted in greater seed input of the former to the soil leading to its greater abundance in the crop fields. Supported by the University Grants Commission, New Delhi (Grants No. F. 3-37/87 SR II)     

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.     

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.     

Population studies ofBrevicoryne brassicae (L.), its parasites and hyperparasites in England

Summary Studies on populations ofBrevicoryne brassicae (L.), its parasites and hyperparasites were carried out by actual counting in the sprouts field and by sticky and water traps.B. brassicae was found to be attacked by one primary parasite,Diaretiella rapae (McIntosh), which in turn is parasitized byAlloxysta brassicae (Ashm.),Asaphes vulgaris Walker,A. suspensus (Nees),Pachyneuron minutissimum (F?rster) andDendrocerus carpenterii (Curtis). The aphid population in the field was started by immigrant alates which were found flying too early to be synchronized with the sprouts plants. SimilarlyD. rapae was not synchronized with the aphids although many individuals could have been carried into a plot through parasitized immigrant alates, of which less than 30% were found parasitized. Because of high hyperparasitism (especially byA. brassicae)D. rapae was not able to maintain a high rate of parasitism to curb the aphid population growth. The maximum percentage mummies being 27.8%, while the maximum, percentage parasitism being 56.6% recorded only during the early 1974 season (mean=12.9%). The decline of aphid population from September onwards was largely due to the cold weather, Syrphid predation and occasionally fungal attack. The high rate of hyperparasitism byA. brassicae is attributed to its better synchronization withD. rapae. The mean percentage of parasite that emerged from mummies collected during 1973–74 wereD. rapae 31.3%,A. brassicae 64.3%,A. vulgaris andA. suspensus 4.3%,D. carpenterii 0.2% andP. minutissimum 0.1%.     

Humid microenvironment prerequisite for the survival and growth of nymphs of the rice brown planthopper,Nilaparvata lugens (St?l) (Homoptera: Delphacidae)

Nymphs ofNilaparvata lugens were experimentally reared from the 2nd instar in a cage covering part of the leaf sheath of an individual rice plant grown in a Wagner pot. Plants were covered with the cage from the water surface of the pot to 10 cm above the surface (lower cage-group) or from 10 cm to 20 cm above the surface (upper cage-group). Temperatures measured at three different parts of the cage remained fairly constant in both groups at around 25°C (23.7–25.2°C in mean value). In the lower cage-group, relative humidities measured at the three heights in the cage in (76.3–90.5% in mean value) markedly increased with the approach to the water surface. The nymphs of this group, particularly during the molting period, aggregated close to the surface. Eighty-two percent of the released nymphs emerged in this group. Relative humidities measured at three heights of the upper cage-group were 69.5–72.7% in mean value, and all the nymphs in this group died within 3 days after their release although half of them stayed on the rice plants within 6 h after their release. The role of relative humidity as a limiting factor on the range of the microhabitat and the population density ofN. lugens in rice fields was discussed on the basis of the results.     

Behavior and mortality of the southern stink bugNezara viridula in Hawaii

Summary This paper is concerned with some aspects of the ecology of the southern green stink bug,Nezara viridula, in Hawaii. Dispersal studies on adults showed an asymetrical dispersal pattern. The rate of plant to plant to movement by the adults was 3.3–6.7 feet per day. Mortality studies showed that egg predation by ants,Pheidole megancephala, was as high as 80–90 per cent. Egg parasitism byTelenomus basalis was highest during February, 1965, and adult parasitism byTrichopoda pennipes was highest during July, 1964, to January of the following year. The population ofN. viridula declined during May, 1964, to May, 1965. This decline occurred simultaneously with the decline in the number of eggs produced per female. It has been speculated that this reduction in egg production was caused by parasitism byT. pennipes. Published with the approval of the Director of the Hawaii Agricultural Experiment Station as Technical Paper No. 816. It is with great pleasure that I acknowledge with thanks the help that I received in various ways from the following: Dr. WallaceC. Mitchell, Dr. MervinKamran., Mr HarryKaya and Mr. HitoshiKamasaki.     

Life tables and population parameters of the green leafhopper,Nephotettix cincticeps (Uhler), in the southwestern district of Japan with special reference to the first generation on foxtail grass

Summary The three year (1988–1990) life tables ofNephotettix cincticeps were constructed, and the daily survival rate and longevity of female adults were estimated by Hokyo and Kiritani’s (1967) method for the overwintering and the first-generations on the foxtail grass in Okayama, southwestern Japan. The life tables and the population parameter values estimated were compared with those in the other generations on rice. The FARMCOP suction sampler was employed to survey the population density. The durations of pre-ovarial maturation of female adults of the 1st generation on foxtail grass and rice seedling were similar. Longevity of adults of the overwintering and the first generations which emerged on the wild host was longer than that of the other generations (2nd and 3rd generations) on the rice plants. Fecundity of females decreased successively as the generation proceeded and it became lowest in the final 3rd generation. Only about 3.5 percent of first-instar nymphs of the 1st generation emerged as adults in the fallow field. The survival rate of nymphs on foxtail grass was always lower in comparison with that on rice plants. However, the survival rates of nymphs on foxtail grass and rice seedling were not significantly different from each other under laboratory conditions. In the fields, senescence of foxtail grass occurred in the midst of nymphal period of the 1st generation. The survival rate of nymphs on foxtail grass decreased with the increasing in the nymphal density. Abundance of spiders during the 1st generation was higher than that in the early stage of rice plants.     

The effects and limits of territoriality on population regulation in grey red-backed voles,Clethrionomys rufocanus bedfordiae

Summary The effects of breeding territoriality on the stability of grey red-backed vole (Clethrionomys rufocanus bedfordiae) populations were investigated on a control grid and a grid on which the voles were fed, in an outdoor enclosure in Hokkaido, Japan. Vole populations were monitored by live trapping from 1984 to 1986: (1) Population density was 2–7 times greater on the experimental grid to which food was added than on the control grid. Reproductive output was more closely associated with the difference in density between grids than survival or dispersal (immigration and emigration) rates. (2) The number of adult females and pregnancy rate of the experimental population were significantly greater than those of the control one. The difference in the number of adult females between the populations was greater than that in pregnancy rate. (3) The proportion of successful litters and the number of weanlings per litter were not significantly different between the control and experimental population. (4) Adult females held territories on both the control and experimental grid; they were spaced out more than would be expected from random occupation. The territories overlapped more on the experimental grid than on the control grid. (5) Mean territory size of adult females on the experimental grid was about half of that on the control grid. The territory size was correlated negatively with population density. (6) The proportion of trap sites that were used by adult females was significantly greater on the experimental grid than on the control grid. This suggests that adult females on the experimental grid used the area more extensively. This factor, in association with territory size and overlapping of territory, was also important in causing the difference in the number of adult females between the grids. (7) These results call into question the hypothesis that territoriality stabilizes the density in populations ofClethrionomys.     

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.     

Comparative studies on the seasonal occurrence, nymphal development and food menu in two giant water bugs,Diplonychus japonicus vuillefroy andDiplonychus major esaki (Hemiptera: Belostomatidae)

Summary We compared the seasonal changes in population density and nymphal development at different water temperatures and under different food conditions between two giant water bugs,Diplonychus japonicus andD. major, in Okayama, Japan. D. japonicus produced 1–2 generations a year, whileD. major was strictly inivoltine. The developmental velocity was higher inD. japonicus than inD. major. The thermal constant ofD. japonicus was less than that ofD. major. These results suggest thatD. japonicus is adapted, to higher water temperature thanD. major. In the field,D. japonicus preyed predominately on Lymnaeidae and Physidae snails, whileD. major preyed on aquatic insects such as dragonfly nymphs.     

Studies on the adult polymorphism in the southern green stink bug,Nezara viridula (hemiptera: Pentatomidae)

Summary Polymorphism in adult colour pattern ofNezara viridula is determined on the genetic basis. The basic colour patterns of adult are classified into four types, i. e. G, O, R and F. No appreciable differences between these types were observed in respect to various physiological traits of nymphs and adults, except that type G seems to be superior in reproductive ability but to be inferior in ability of surviving winters at least to types R and F. Inter-generation changes in percentage frequency of G type were examined from 1959 to 1967 covering more than 30 generations. Percentage frequencies of G types prior to the severe winter of 1962–3 fluctuated greatly around a mean of 87.9%, while they did to a lesser extent than before with a mean of 85.0% after the winter. Unexpected high percentages of G type were recorded frequently in summer generations, viz. 1st and 2nd. On the other hand, the relative frequencies of G and O types decreased after hibernation in contrast to the increases in those of F and R. This sort of changes in genetic composition related to the winter of 1962–3 was observed in several populations segregated from each other. This seasonal alternation of selective activity in the environments is considered to be responsible for retention of the polymorphism. Alternative possible causes, i. e. difference in habitat preference, non-random mating and selective predation by predators among polymorphs, may safely be rejected as irrelevant to the mechanism in maintaining polymorphism. The polymorphism of this insect seems to be in a transient state rather than balanced one contributing little to population regulation, but the persistence ofN. viridula in the periphery range may be assisted by retention of the polymorphism.     

Natural mortality at different population densities of a pine shoot moth,Evetria cristata

Summary A field population ofEvetria cristata was studied in 10 plots in 1962 and in 6 plots in 1963. These plots were divided into 2 or 3 groups of different population levels of the shoot moth in respective years. The survival of the insect was then analysed in these different groups of plots. The survival rate ofE. cristata from eggs to adults in the first generation was found always higher in the group with low population density, which indicates the existence of some factors that affect the population more severely when the insect is more abundant.Lissonota evetriae andPediobius sp. seemed to have killed more proportion of the hosts where the shoot moth density was high. However, the total effect of the all natural enemies was not always great in the plots with high density of the moth. The survival of the second generation of the moth in 1963 was observed to be much higher at any population level than in the other generations.     

Intraspecies competition in a field population ofGregopimpla himalayensis (Hymenoptera: Ichneumonidae) parasitic onMalacosoma neustria testacea (Lepidoptera: Lasiocampidae)

Summary Intraspecies competition in a field population ofGregopimpla himalayensis (Hym.: Ichneumonidae) parasitic on the prepupae ofMalacosoma neustria testacea (Lep.: Lasiocampidae) was investigated. The parasite oviposits the sufficient number of progeny (5 individuals/0.1 g dry weight of host) to exhaust a single host in a single attack. However, at the intensity less than 22–26 individuals/0.1 g d.w. of host, all individuals can emerge, i.e. density-dependent mortality does not occur. Within this range of intensity, survival of parasite larvae is guaranteed by diminution in body size and decreasing sex ratio. In contrast, total biomass of parasites showed a peak at 5 individuals/0.1 g d.w. of host at which a single host is exhausted. Above the intensity of 22–26, extraordinary minute individuals appeared and they died before maturation. If intraspecies competition play a role in regulation ofG. himalayensis population in the field, the process is usually not through density-dependent mortality but through decreasing reproductive rate caused by decrease in the sex ratio, adult longevity and fecundity. Contribution Ser. 2, No. 275. Entomological Laboratory, Faculty of Agriculture, Kyushu University.     

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.     

Population behaviour of the southern green stink bug,Nezara viridula, with special reference to the developmental stages of early-planted paddy

Summary Population behaviour of adults and 5th-instar nymphs ofNezara viridula L. was analysed by means of the marking-and-recapture method in an early-planted paddy field. The field contained five varieties of rice which differend in growth states. It was estimated that a total of more, than 7,000 adults of the first generation, in which at least 3,000 were females, invaded the field from early July to middle August. Egg-mass census data, however, indicated that only 10 per cent or less of the females participated in egg-laying. This was largely due to the, low rate of adult survival. The adults preferred younger plants, for both feeding and oviposition. The method described byIwao et al. (1966) permitted estimate that 3,300–3,400 of the 5 th-instar nymphs and 1,100–1,200 of the adults of the second generation were produced from 298 egg-masses (25, 700 eggs); while 95–6 per cent of the individuals were thought to have died before reaching adulthood. Most of the 5 th-instar nymphs moved less than 4 m in three days as long as the condition of food plants remained suitable, but they tended to move more towards younger plants when those on which they lived became too mature. The apparent survival rate of the second generation adults was very low, probably due both to a rapid emigration and a high mortality of newly-emerged adults.     

Functional response and searching efficiency inPseudogonatopus flavifemur Esaki and Hash. (Hymenoptera: Dryinidae), a parasite of rice planthoppers

Summary The functional response ofPseudogonatopus flavifemur E & H (Hym., Dryinidae) was investigated by offering hosts (brown planthopper) at densities ranging from 8 to 160 per cage. The response curve was found to be sigmoid, i. e.Holling's (1959) Type III curve. In experiments involving 310 hosts per cage distributed unevenly in 5 densities (10, 20, 40, 80 and 160 per hill), and a different female parasite density each time (viz. 1, 2, 4, 8 or 16 per cage), the behavioral response was described well by the “random predator equation” ofRoyama (1971) andRoger (1972), which is a convex exponential curve. The area of discovery (a) decreased with an increase in female parasite density (P), and the relationship was described by the equation: loga=−1.0099−0.3638 logP. There was an apparent increase in handling time per host as the number of female parasites increased. Superparasitism, a rare phenomenon under natural conditions, was often observed in the laboratory. The potential ofP. flavifemur as a biocontrol agent of the brown planthopper is discussed.     

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.