Experimental studies on the distribution of the aphid counts

Summary The present paper dealt with the sequential changes of the distribution pattern of apterous females aphid populations, that were artificially settled at the beginning on the experimental barley ‘field’. The aphids were settled at random or even with a fixed denisty per plant. For five or six days after the settling, the number of individuals followed the negative binomial distributions in all cases while the parametersk andp were varying. The estimated values ofk were rather small for the first one week after the settling, which may suggest that the number of moving aphids between plants was relatively small and the degree of concentration expressing the intrinsic increase was high. After that, as the number of individuals increased, the number of moving aphids between plants would be considered to be increased. It was found that with the lapse of time the degree of concentration decreased ork became larger. The distribution of aphids per blade in a plant was also described briefly.     

Influence of parental density on the distribution pattern of eggs in the common cabbage butterfly,Pieris rapae crucivora

Summary To investigate the relation between the distribution pattern of eggs and the parental density in the common cabbage butterfly,Pieris rapae crucivora, the countings of egg number per plant were made on both cabbage plants cultivated in the farm and planted in the net house in which the female butterflies were released at various densities. The frequency distribution of eggs fits well to the negative binomial excepting the cases where they agree withPoisson series, and the degree of aggregation expressed as the reciprocal of the parameter, 1/k, tends to decrease as the egg or parental density increases. At the same parental density, however, the distribution of eggs can be described by the negative binomial with a common parameter,k _c, regardless of the difference in the density of laid eggs. In the case where a single female butterfly lays eggs, the spatial pattern of egg distribution is always lean, while its frequencies conform toPoisson or the negative binomial series. This lean changes toward patchy with increasing the parental density. From these results, it is concluded that the degree of aggregation in the distribution of eggs decreases with the increase of the parental density.     

Prey capture tactics of spiders: An analysis based on a simulation model for spider's growth

Summary The prey capture tactics of spiders was analyzed, considering the energy gained by the capture of prey and that required for it. For the purpose of it, a growth model of spiders was constructed, expressing the flow rate of prey biomass to the spider's body by differential equations. Solving these equations under the differing values of three parameters, growth curves of spiders was obtained. These three parameters are the amount of prey biomass supplied daily to spiders,x ₀, the rate of prey capture of spiders, α, and a coefficient of the respiration rate required for the capture of prey,k. When the value ofk increased, spiders could grow only at high value ofx ₀. These results suggest that habitats with small prey biomass are preferred by spiders adopting a sit-and-wait tactics for prey capture, which requires small values ofk. Wolf spiders are one of these spiders showing that tactics. On the other hand, web-builders which require large amount of energy for spinning webs (namely, take large value ofk), are able to grow only in the habitats with large prey biomass. Each species of spiders are considered to locate in a certain point between both extremes of these tactics for the capture of prey.     

An analysis of the predator-prey system in the paddy field

Summary A predator-prey system involvingLycosa pseudoannulata as predators and both the green rice leafhopper,Nephotettix cincticeps and the brown planthopper,Nilaparvata lugens as prey was studied in an experimental paddy field. Daily predation rates on the prey and weekly disappearance rates ofLycosa females were related to densities of both predators and prey by multiple regression. During the off-season of reproduction for predators and prey, it proved possible to simulate the interaction betweenLycosa and its prey.     

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.     

Prey preference of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand)

Summary Functional responses of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand) attacking the rice brown planthopper,Nilaparvata lugens (St?l.), and the mirid predatorCyrtorhinus lividipennis Reuter were both those of Holling Type II. The attack rate was higher and handling time lower forC. lividipennis. However, when caged with the two prey, the wolf spider showed a significant preference forN. lugens at a lower prey proportion. Proportions of prey attacked were significantly different from the expected ratios of prey available as well as from the predicted preferences derived from the functional response parameters. As proportions ofN. lugens attacked changed from greater to less than expected as the proportions ofN. lugens available increased, a “reverse switch” behaviour seems to be evident.     

The spatial distribution pattern of the brown planthopperNilaparvata lugens St?l (Homoptera: Delphacidae) in west Java, Indonesia

Summary Spatial distribution pattern of the brown planthopper (BPH) was analyzed at 9 experimental fields in the northern part of West Java during two consecutive rice cropping seasons, i.e., wet and dry seasons. The population of each developmental stage and wing form of BPH at each location showed consistent departure from the random (Poisson) distribution, the variances of the densities in most cases exceeding their means. Namely, the distribution pattern of BPH per hill of rice plant was found to have a general tendency to be aggregated or contagious and to fit fairly well to the negative binomial model. The tendency for aggregation was further confirmed by both the β-values of -m regression being larger than unity and theC _A -values being larger than zero for each developmental stage. Although significant variations in the distribution pattern as measured by β- orC _A -value were observed between different developmental stages, between wing forms and among locations, the degree of aggregation for a given developmental stage at each experimental field remained fairly stable throughout the crop period, despite wide temporal changes in population density. Possible factors to explain these characteristics of the spatial distribution pattern of the BPH in West Java were discussed with reference to the process generating it.     

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.     

Modelling the effects of population aggregation on the efficiency of insect pest control

Summary A methodology is developed to assess the effects of spatial distribution on the efficiency of insect pest control. This methodology is especially applicable to pest control methods whose efficiency of action depends either positively or negatively on pest density It is applied here to the sterile insect technique and pheromone trapping for male annihilation, which both depend negatively on density. This methodology relies on quantifying clumps of various size and then relating this to efficiency of control and predicting the total pest production given the information on clump sizes and efficiency of control for each clump size. It is found that control is about four times as difficult for a population that is highly clumped (k of the negative binomial distribution=0.25) as for a regularly dispersed population.     

Process generating the ditribution pattern of eggs of the common cabbage butterfly,Pieris rapae crucivora

Summary The process generating the negative binomial in the distribution pattern of eggs of the common cabbage butterfly,Pieris rapae crucivora, was investigated by releasing the female adults in a net house where cabbages were planted. The distribution of butterflies visited and laid an egg or more per plant followed thePoisson series under the uniform light condition, while that of eggs laid per visit conformed to the logarithmic distribution. From these results, it may be concluded that the negative binomial arises from compounding of thePoisson and the logarithmic distribution. The observed frequency of eggs found per plant fitted to the negative binomial with parameter thus computed theoretically. The change in the degree of aggregation with the increase of the parental density was considered in connection with the above results. Aided by a grant from Scientific Research Expenditure of the Ministry of Education.     

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.     

Inconstancy of Taylor'sb: Simulated sampling with different quadrat sizes and spatial distributions

Summary Taylor's power law,s ² =am ^b , provides a precise summary of the relationship between sample variance (s ² ) and sample mean (m) for many organisms. The coefficientb has been interpreted as an index of aggregation, with a characteristic value for a given species in a particular environment, and has been thought to be independent of the sample unit. Simulation studies were conducted that demonstrate that the value ofb may vary with the size of the sample unit in quadrat sampling, and this relationship, in turn, depends on the underlying spatial distribution of the population. For example, simulated populations with hierarchical aggregation on a large scale produced values ofb that increased with the size of the sample unit. In contrast, for a simulated population with randomly distributed clusters of individuals, the value ofb eventually decreased with increasing quadrat size, as sample counts became more uniform. A single value ofTaylor'sb, determined with a particular sample unit, provides neither a fixed index of aggregation nor a complete picture of a species' spatial distribution. Rather, it describes a consistent relationship between sample variance and sample mean over a range of densities, on a spatial scale related to the size of the sample unit. This relationship may reflect, but not uniquely define, density-dependent population and behavioral processes governing the spatial distribution of the organism. Interpretation ofTaylor'sb for a particular organism should be qualified by reference to the sample unit, and comparisons should not be made between cases in which different sample units were used. Whenever possible, a range of sample units should be used to provide information about the pattern of distribution of a population on various spatial scales.     

Evaluation of statistical precision and design of efficient sampling for the population estimation based on frequency of occurrence

Summary The binomial sampling to estimate population density of an organism based simply upon the frequency of its occurrence among sampled quadrats is a labour-saving technique which is potentially useful for small animals like insects and has actually been applied occasionally to studies of their populations. The present study provides a theoretical basis for this convenient technique, which makes it statistically reliable and tolerable for consistent use in intensive as well as preliminary population censuses. Firs, the magnitude of sampling error in relation to sample size is formulated mathematically for the estimate to be obtained by this indirect method of census, using either of the two popular models relating frequency of occurrence (p) to mean density (m), i.e. the negative binomial model,p=1−(1+m/k) ^−k, and the empirical model,p=1−exp(−am ^b). Then, the equations to calculate sample size and census cost that are necessary to attain a given desired level of precision in the estimation are derived for both models. A notable feature of the relationship of necessary sample size (or census cost) to mean density in the frequency method, in constrast to that in the ordinary census, is that it shows a concave curve which tends to rise sharply not only towards lower but also towards higher levels of density. These theoretical results make it also possible to design sequential estimation procedures based on this convenient census technique, which may enable us with the least necessary cost to get a series of population estimates with the desired precision level. Examples are presented to explain how to apply these programs to acutal censuses in the field.     

Relative humidity as an environmental factor determining the microhabitat of the nymphs of the rice brown planthopper,Nilaparvata lugens (St?l) (Homoptera: Delphacidae)

Summary Microhabitat of the nymphs and the adults of the rice brown, planthopper,Nilaparvata lugens is known to be the lower parts of rice plants. In this study, possible environmental factors determining the range of the microhabitat were experimentally analyzed by using the nymphs under laboratory conditions. Thirty individuals of the 1st or the 3rd instar nymphs were released to a potted rice plant covered with a transparent acrylic cylinder. When the top of the cylinder was kept opening (open condition), the temperatures in the cylinder were almost constant, and relative humidities in the cylinder decreased with the increase of the height from the water surface of the pot. In the open condition, most nymphs and all exuviae were found on the basal parts of rice plants where the humidity was more than ca. 90% r.h. When the top of the cylinder was kept closing with parafilm (closed condition), the temperatures in the cylinder were almost constant, and relative humidities in the cylinder were more than 95% r.h. In the closed condition, the nymphs and the exuviae were distributed sparsely to the whole parts of rice plants. In both of the open and the closed conditions, patterns of nymphal distributions on rice plants during the dark regime were the same as those during the light regime under 25±2°C and 16L∶8D. Four different temperatures ranging from 20°C to 35°C did not influence on the microhabitat in the open condition. It was concluded from the results that relative humidity is the important environmental factor to determine the microhabitat of the nymphs ofN. lugens which showed to prefer very humid condition more than ca. 90% r.h.     

Stabilization effects of spatial aggregation of vectors in plant disease systems

Stabilization effects of spatial aggregation of vectors were examined in insect-borne plant disease systems by constructing a model that describes the yearly dynamics of rice stripe virus disease transmitted by the small brown planthopperLaodelphax striatellus (Fallén). Two transmission paths between vectors were considered: vertical transmission from parents through eggs, and horizontal transmission from infected plants by acquisition feeding. In this model, a paddy field was divided into quadrats and horizontal transmission was assumed to occur within each quadrat. A negative binomial distribution was used to describe the frequency distribution of vectors per quadrat. The parameters of the model were estimated using field data collected in Ibaraki Prefecture, Japan. The model showed that (1) the disease cannot invade into an epidemiological system if the mean crowding of vectors is less than a critical value, (2) the proportion of infected vectors is maintained at about 30% irrespective of the vector density if vectors are highly aggregated, and (3) the proportion of infected plants is maintained at a low level irrespective of the vector density if vectors are highly aggregated. It was also shown that these stabilization effects of aggregation in this epidemiological system come from a mechanism that is common to other systems such as single-species systems and competition systems.     

Coexistence and weak amensalism of congeneric gall-forming aphids on the Japanese elm

Closely related species of gall-forming aphids are often associated with a single host species. SixTetraneura species coexist on the Japanese elm,Ulmus davidiana, in Sapporo, northern Japan. This paper describes the probabilities of coexistence on macro- and microgeographic scales (i.e., on host trees and host leaves) and examines whether coexistence with conspecific or heterospecific galls on leaves or shoots has any effect on the fecundity of each aphid species using multiple regression. A Monte Carlo simulation showed that the frequency distribution of the numbers of species on individual host trees differed significantly from that expected from the null model. There were significantly positive or negative associations between species.Tetraneura radicicola andsorini always coexisted with other species on trees they infested. Multiple regression revealed that the coexistence of conspecific or heterospecific galls on individual leaves had no influence on the fecundity ofradicicola andsorini, but had a negative influence on that of sp. O. On average,radicicola andsorini produced a smaller number of offspring in galls than sp. O, and obviously consumed only a small part of resources available on the galled leaves. Evidence available suggests that although amensalism does arise between sp. O and other species, its influence in not so strong as to exclude sp. O competitively from theTetraneura community.     

Predator-prey system structure in patchy and ephemeral phytotelmata: Aquatic communities in small aroid axils

Mechanisms allowing the persistence of an aquatic predator-prey system in tiny pools (the mean ca. 0.5 ml) held by taro axils were analyzed from the viewpoint of temporal and spatial patterns of habitat use. Predatory larvae of a mosquitoTopomyia tipuliformis concentrated in young axils, while 9 other taxa utilized both young and old axils or concentrated more in older axils. The total prey density was lower in axils with the predator but the density of each prey taxon (except for a few cases) and the number of prey taxa did not differ between axils with and without predators. This indicates thatT. tipuliformis is a general predator and does not influence prey community organization through selective removal of particular prey taxa. Inter-axil distribution ofT. tipuliformis was aggregated in the first instar but uniform in the third and fourth instars due to intraspecific cannibalism, which assures the survival of a single individual under short food supply. Distributions of prey taxa were mostly aggregated, fitting the negative binomial distribution. Thirty seven out of 45 associations of 10 taxa were independent with 3 negative (between the predator and some late-colonizing prey) and 5 positive (between some prey taxa) associations. Probabilistic refuges (produced by independent aggregated distributions) reduced interspecific encounters which may result in competition and predation, and thus probably facilitated prey coexistence. There was no evidence for the importance of predation in structuring the prey community. This system may be prey-dominated in that predator persistence depends on prey community existence but prey community structure depends less on predation.     

Niche modification and stability of competitive systems. II. Persistence of interspecific competitive systems with parasitoid wasps

Summary Effects of niche shift in ecological time scale on the population dynamics of competing species were studied in the experimental populations of two parasitoid wasp species,Anisopteromalus calandrae andHeterospilus prosopidis (both are solitary parasites), on a host, the azuki bean weevil,Callosobruchus chinensis. Four resource conditions were set up with combination of kind of bean (azuki or black eye), and host distribution (uniform or clumped). In each resource condition, four developmental stages of hosts were provided as a resource spectrum for parasitoid wasps. Population dynamics of the two wasp populations were investigated in each resource condition in Multi-Generation Competitive Systems (MGCS), in which fresh hosts of four developmental stages were periodically introduced and were parasitized competitively by the two wasp species. Competitive coexistence of both wasps occurred in the azuki-clumped condition, where the peaks of the resource utilization curves separated in the two species; pupae inA. calandrae and the early or late fourth instar inH. prosopidis, A. calandrae was eliminated in the azuki-uniform condition andH. prosopidis went extinct in two black eye conditions irrespective of host distributions. The degrees of overlap of the resource utilization patterns of the two wasp species during population dynamics were not significantly different among resource conditions irrespective of the results of coexistence or extinction. Even in the azuki-clumped condition, however, extinction ofA. calandrae was observed when resource partitioning could not be realized with only the late fourth instar larvae available to wasps. Further analytical experiments showed that parasitizing ability ofA. calandrae increased with host density per bean with azuki beans, butA. calandrae could express higher parasitizing ability with black eye beans thanH. prosopidis irrespective of host density per bean. The flexibility in parasitizing ability byA. calandrae for various host stages under different resource conditions was thought to be the major factor in determining the competitive coexistence or the extinction of either species under different resource conditions. The present experiments also suggested that different second-best host stages between competitors could be a major contributing factor to competitive coexistence.     

Crucial factors determining the spatio-temporal distribution patterns of threeAthalia sawflies feeding on common cruciferous plants

Summary ThreeAthalia sawflies,A. japonica, A. rosae andA. infumata, feeding on cruciferous plants, coexist in Japan. However, it is not known what ecological strategies they use and what environmental factors are crucial to such strategies. I attempted to explain these questions by examining the relationship between the spatio-temporal distribution patterns of threeAthalia sawflies and their habitats in three districts (Lowland, Intermediate and Mountain). The three sawflies have different spatio-temporal distribution patterns, though they usually used common cruciferous plants.A. japonica was abundant in spring and autumn but disappeared during summer in all the districts. In the Lowland, populations ofA. rosae andA. infumata, like that ofA. japonica, crashed in summer. HoweverA. rosae occurred mainly in summer in the Intermediate and Mountain. AlthoughA. infumata occurred in the same seasons asA. rosae in all districts, population levels ofA. infumata were always lower than those ofA. rosae. The crucial factors controlling their population patterns were the availability of host plants and temperature. Population crashes ofA. rosae andA. infumata were due to food depletion, and those ofA. japonica were due to heat stress. On the other hand, their population patterns may be interpreted as phenological synchronization with their original host plants, though they all existed on common cruciferous plants. The three sawflies may have evolved different strategies to escape from unfavorable habitat conditions. Such strategies are speculated to be summer diapause inA. japonica, long distance migration inA. rosae, and local dispersal inA. infumata.