Larval growth dynamics ofHemipyrellia ligurriens (Calliphoridae) andBoettcherisca formosensis (Sarcophagidae) in crowded and uncrowded cultures

Summary Larval growth patterns ofHemipyrellia ligurriens (Calliphoridae) andBoettcherisca formosensis (Sacophagidae) in crowded and uncrowded cultures were compared. Growth of the larvae followed a sigmoid curve. The highest larval growth rates were 0.33 and 0.36 mg h⁻¹ for uncrowded and crowdedH. ligurriens respectively. The corresponding figures were 2.38 and 1.23 mg h⁻¹ forB. formosensis. Larvae of both species attained maximum weight earlier in crowded cultures than in uncrowded cultures, although the final weights attained in crowded cultures were less. The earlier period of most rapid growth in both species was interpreted as a result of intraspecific facilitation at higher larval densities. UncrowdedB. formosensis had a shorter larval to pupal development and an earlier period of most rapid growth than uncrowdedH. ligurriens, suggesting the former may be superior in exploiting carcasses with limiting food.     

Yearly variation in recruitment and its effect on population dynamics inYoldia notabilis (Mollusca: Bivalvia), analyzed using projection matrix model

Summary Long-term variation in recruitment was estimated by constructing projection matrices for a marine bivalve,Yoldia notabilis, at two stations in Otsuchi Bay, northeastern Japan, and the effects of its variation on population dynamics were examined using a simple matrix model. The matrix model was developed from the Leslie matrix, in which the population growth rate λ was expressed as a function of recruitment rater ₀. The equilibrium recruitment rater _s, or the recruitment rate required to maintain population at constant size (λ=1), was expressed by the reciprocal of the reproductive value of a newly recruited individual. The estimates ofr _s for the field population were lower at the shallower station than at the deeper station, reflecting higher survivorship and fecundity. Past recruitment rate estimated both by the field samplings for 3 years and by the back-calculation from the current age structure for over 10 years showed large yearly variation, ranging between 0 and 58.6×10⁻⁴. The estimates were larger thanr _s, and hence, large enough to increase population size (λ>1) only in approximately one-third of the estimated years. This suggests that the population has been maintained by occasional successful recruitment occurring once every few years.     

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.     

Spatial pattern indices based on distances between individuals on a line-segment with finite length

Summary Let us consider a strip-wise habitat of line-segment, like a corridor, to simplify the subject mathematically, and assume that the length of the habitat is γ and there aren individuals. Here, we assume that the spatial pattern of the individuals is random if then distances from the left end of the habitat to each individual follow a uniform distribution on the strip. Under such an assumption, the variance of the distances between any two neighbors is represented by the formulanθ ²(n+1)⁻²(n+2)⁻¹ and the variance betweenn+1 distances betweenn individuals from the left end to the right end to the strip, is represented by the formulanθ ²(n+1)⁻²(n+2)⁻¹. These two kinds of variances can be used for determining (1) the spatial pattern of a population on the strip and (2) the spatial structure within the population, by comparison with the variances calculated from the data. Two examples cited from the literature, a cattle population on a pasture and an aphid population on a sycamore leaf, are presented.     

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.     

Pit-relocation of antlion larvae in relation to their density

Of the biotic factors potentially affecting the foraging behavior of pit-building antlion larvae,Myrmeleon bore Tjeder, we focussed on the density effects and examined experimentally influences of larval density on pit construction behavior, spatial distribution and pit relocation rate. Moreover the mechanism of mutual interference among larvae was examined.

Studies on the distribution pattern of the pine needle gall midge,Thecodiplosis japonensisUchida etInouye (Diptera: Cecidomyiidae) in a pine forest

Summary Numerical changes and distribution patterns of the pine needle gall midge,Thecodiplosis japonensis Uchida etInouye, were studied during the period from 1978 to 1979 in a young plantation ofPinus thunbergii in Shiga Prefecture, Japan. The survivorship curve of this species was characterized by a low mortality of larvae in galls and two high mortalities before the formation of galls and during the overwintering period in soil. The within and between-trees distributions of eggs and larvae in galls were examined by using the regression method. The egg distribution per shoot was aggregative both within and between host plants. The within-tree variations in numbers of eggs per shoot were related to the differences in the abundance of available needles for oviposition per shoot among the canopy layers. The between-tree variations reflected the heterogeneous emergence of adult females in the study plot. The degree of aggregation increased from egg to gall stage in both within- and between-tree distributions and the increase was explained by the different mortality of larvae within trees and the inversely density-dependent mortality between trees. The distribution patterns in the soil habitat stages were examined by the patchness index ( ). This species showed aggregative distributions in soil stages. There was a correlation in spatial patterns of adult emergence between the successive generations. The distribution properties of this species were discussed in connection with the population dynamics and the availability of host plants in the study plot.     

Population dynamics of the green leafhopper,Nephotettix virescens Distant (Hemiptera: Cicadellidae) in synchronized and staggered transplanting areas of paddy fileds in Indonesia

Summary The population dynamics ofNephotettix virescens, a vector of rice tungro virus disease was investigated in a synchronized transplanting area at Jatisari (1984–1986), West Java and in a staggered transplanting area at Sidan (1986–1988), Bali, Indonesia. The FARMCOP suction sampler was employed for population censuses ofN. virescens and its natural enemies. The population growth pattern was affected by transplanting pattern: In the staggered transplanting area, the population density increased from the immigrant generation to the first generation, and sharply decrease thereafter, while in the synchronized transplanting area the population density often reached the highest peak in the second generation. The degree of contageousness in the spatial distribution ofN. virescens was negatively correlated with population density of the immigrant generation. Contribution from Indonesia-Japan Joint Program on Food Crop Protection     

Spatial pattern analysis of the incidence of aster yellows disease in lettuce

The degree of aggregation of lettuce plants infected by aster yellows phytoplasma (AYP) was investigated in 12 fields from three experiments. Position of diseased and healthy plants was mapped in a 6–9×12-m section of each field; for most analyses, fields were divided into 10-plant quadrats. Mean disease incidence (p) ranged from 0.01 to 0.30. The frequency of diseased plants was described by the beta-binomial distribution, with an index of aggregation (θ) ranging from 0 to 0.17, positively correlated withp, and generally increasing over time within a field. Distance-class analysis revealed a core-cluster size of only a few plants. However, spatial autocorrelations ofp between quadrats were not significant, indicating that the scale of spatial pattern was small, generally less than 10 plants. An overall measure of aggregation was given by the slope parameter of the binary form of the power law, in which the log of the calculated variance is regressed on the log of the theoretical variance for a binomial distribution. The slope was 1.18 and significantly different from 1. Results for this “simple-interest” disease are interpreted in relation to the persistent transmission of AYP by its aster leafhopper vector.     

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.     

Ecological studies of collembolan populations in a pine forest soil IV. Comparison of distribution patterns

Summary The distribution pattern of ten species of Collembola was studied during the four years period from July 1971 to May 1975 in a pine forest soil. The distribution patterns were analysed for two scales of distribution, i. e., the distribution over the plot of 10×10 m² and the micro-distribution within a block sample consisting of 36 contigious units each 2×2 cm² in area, by applying the -m regression method. The fundamental pattern which appeared was quite similar for the species examined and individuals were aggregated in response to the heterogeneity of habitat conditions. The causes of aggregations were discussed with regard to some environmental factors. The relative abundances of 10 species within the collembolan community was examined in relation to the habitat utilization and the relative abundance was not related to the degree of aggregation but rather to the area occupied by individuals. This suggests that the more numerically abundant species tend to occupy broader micro-habitat. Biological meaning of aggregation was discussed in connection with the population biology and community organization of collembola.     

A distance measure of spatial pattern in a biological population

Summary Suppose thatn individuals locate independently and randomly on a segment of line of finite length (habitat). Let the theoretical and observed ranges of the sites of the individuals on the segment be μ _n-1 andr _n-1, respectively. Then, the degree of dispersion of the individual sites is measured by the ratio, T _n =n _n-1/μ _n-1, as follows: A random spatial pattern forI _r−1 =1 An aggregated spatial pattern for 0≤I _r <1 A uniform spatial pattern for (n+1)/(n−1)≥I _r >1. Another method was derived. Let the probability that an observed range is less thanr _n−1 beI _p , under the hypothesis of a Beta distribution. Then indicates A random spatial pattern forI _p =1/2 An aggregated spatial pattern forI _p <1/2 A uniform spatial pattern forI _p >1/2. The first index can be used for comparing populations having the same number of individuals, whereas the second one can be used for comparing populations with different numbers of individuals.     

Some strange properties of the logistic equation defined withr andK: Inherent defects or artifacts?

Summary In some situations the logistic equation in the usual expression, dN/dt=r(1−N/K)N, exhibits properties that are biologically unrealistic. For example, whenr≦0 the population can no longer show any normal, negative response in per-capita growth rate to increasing density. Also, when the equation is employed in the Volterra's competition model, a familiar but incredible conclusion is derived which says that the outcome of competition is entirely independent of the reproductive potentialr of each species. It is shown that all such strange properties are mere artifacts arising peculiarly in thisr-K model from its misleading implicit supposition thatK could be independent ofr, and they can be readily removed by alternative use of a plainer, classical form of the model, dN/dt=(r−hN)N.     

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.     

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.     

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.     

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.     

Use of the regression of mean crowding on mean density for estimating sample size and the transformation of data for the analysis of variance

Summary An approximate method for estimating the sample size in simple random sampling and a systematic way of transformation of sample data are derived by using the parameters α and β of the regression of mean crowding on mean density in the spatial distribution per quadrat of animal populations (Iwao, 1968). If the values of α and β have been known for the species concerned, the sample size needed to attain a desired precision can be estimated by simply knowing the approximate level of mean density of the population to be sampled. Also, an appropriate variance stabilizing transformation of sample data can be obtained by the method given here without restrictions on the distribution pattern of the frequency counts. Contribution from the Entomological Laboratory, Kyoto University No. 418. Contributions from JIBP-PT No. 52. Aided in part by a grant from the Ministry of Education for the special project research, ‘Studies on the dynamic status of biosphere’.     

Preference and performance linkage in aPhyllocolpa sawfly on the willow,Salix miyabeana, on Hokkaido

A leaf-folding sawfly in the genusPhyllocolpa (Hymenoptera: Tenthredinidae) attackingSalix miyabeana (Salicaceae) was studied near Sapporo, Hokkaido, along the Ishikari River in 1993. Host plant individuals were young trees 4–7 years old which were growing rapidly, producing some long shoots with large leaves. On a gradient of shoot length classes from 0–5 cm long to over 80 cm long, shoots were much more abundant in the shorter shoot length classes. However, attacks by ovipositing females increased as shoot length increase from 0 attacks on the shortest shoots to 5.17 attacks per shoot on the longest shoots. The frequency of attack per leaf increased from 0 to 0.13 over the same range of shoot lengths. This pattern of attack resulted in a high frequency of larval establishment in feeding sites, between 0.96 and 1.00, in all attacked shoot length categories. However, probability of survival to a late instar larva increased with shoot length and corresponded to the attack pattern, indicating a preference-performance linkage between female ovipositional decisions and larval survival. The patterns found for thisPhyllocolpa species are similar for galling sawflies in North America and Europe, especially in the genusEuura, members of which make stem, bud and leaf midrib galls. Extending the pattern to aPhyllocolpa species broadens identification of pattern and ultimately the generality of the emerging theory on populations of galling sawflies.     

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.