Effects of plant density on the survival rate of cabbage pests

The population density of herbivores depends on the spatial scale as well as the temporal scale. In a small-scale, short-term experiment, the number of individuals entering from the surrounding area will be most influential in determining the herbivore density. In large-scale, long-term experiments, however, the density of herbivores will rather be influenced by the survival rate of individuals inside the field because most of the herbivorous population derives from the parents that developed inside the field. If we want to predict the large-scale long-term density of herbivores, therefore, emphasis should be placed on the estimation of survival rate. To elucidate the effects of plant density on the large-scale long-term abundance of cabbage pests, we examined the survival rates of three lepidopterous pests, the small white butterfly Pieris rapae crucivora Boisduval (Pieridae), the beet semi-looper Autographa nigrisigna (Walker) (Noctuidae), and the diamondback moth Plutella xylostella (Linnaeus) (Yponomeutidae) under two levels of plant spacing (sparse plot, 2 m × 2 m interval; dense plot, 0.5 m × 0.5 m interval). The experiment with four blocks was repeated in two seasons. The number of eggs per plant was larger in the sparse plots than in the dense plots for all species. The survival rate of eggs and larvae, on the contrary, was lower in the sparse plots than in the dense plots. The lower survival rate of eggs in the sparse plots was mainly caused by the density dependency, while the lower survival rate of larvae in the sparse plots was mainly caused by the direct effects of plant density. It was thus suggested that the density of herbivores may become lower in the sparsely planted field in the long run because of the higher mortality of larvae. Received: September 16, 1998 / Accepted: March 22, 1999     

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.     

Heritability estimates of life history traits in small white butterflyPieris rapae crucivora

Summary Heritabilities and genetic correlations of life history characters (pupal weight, age-specific fecundities, and egg weight) of small white butterflyPieris rapae crucivora are estimated by a quantitative genetic method (sib analysis). The results indicate moderate or high heritabilities and a largely negative genetic correaltion in age-specific fecundities.     

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

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

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.     

Genetic variance and covariance patterns of larval development in the small white butterflyPieris rapae crucivora Boisduval

Summary Quantitative genetic theory indicates that genetic covariance patterns among life history characters should have played an important role as genetic constraint in life history evolution. Highly positve (and negative) genetic correlations between larval development time (or larval growth rate) and adult size characters were detected by means of sib analysis for the small white butterflyPieris rapae crucivora. The genetic associations suggested that evolution of developmental characteristics and adult phenotypic traits were constrained by pleiotropy. The positive genetic correlations between development time and adult body size may be compatible with the trade-off between them, but the negative genetic correlations between larval growth rate and adult body size are not predicted from theories of optimal energy allocation. That phenotypic correlations drastically differed from the genetic correlations indicates limitations of evolutionary inferences based only on phenotypic variation.     

Dispersive behaviour of larvae of the common cabbage butterflyPieris rapae crucivora

Summary To investigate the aggregative nature of the larvae ofPieris rapae crucivora, each 50 individuals of the 4th and 5th instar larvae collected from cabbage farms and reared under crowded and solitary conditions were released on an experimental arena and their dispersive behaviour was observed with the lapse of time. Both the 4th and 5th instar larvae showed the trend to approach toward random distribution when they were released under clumped condition, and they maintained random distribution when they were released at random. Therefore, it may be concluded that the larvae have not any aggregative nature caused by the mutual attraction among individuals. However, as the larvae reared in crowds sometimes showed the slight aggregative behaviour, it seems that the larval dispersal is different between densely and sparsely populated plants in field.     

The learning abilities of the white cabbage butterfly,Pieris rapae, foraging for flowers

This study examines the role of learning and memory in the butterflyPieris rapae crucivora Boisduval during foraging for flowers. In an outdoor cage with 6 flower species,P. rapae showed various visiting patterns: some visited only one species, while others visited several species in a day. The foraging process for flowers ofErigeron annuus (L.) Pers. could be divided into two successive steps: (1) landing on the nectaring caputs, and (2) finding the source of nectar in the caput. Butterflies learned to proceed through the two steps more efficiently with successive attempts: they gradually decreased landings on nectarless caputs and probings on the nectarless petals of ligulate flowers respectively. As a result, handling time per unit caputs became shorter, and apparent rewards per unit time, i.e. the efficiency of collecting nectar, increased. In addition, once learned,P. rapae could remember a rewarding flower color for 3 days, which was not interfered with by learning another flower color. This indicates thatP. rapae keeps memory for a period longer than 3 days, and that they can remember at least two flower species as suitable flower resources. Furthermore, data indicated that they sometimes can apply the foraging skills obtained on other flower species to a novel one. These abilities could enable butterflies to easily switch flower species, or to enhance labile preference. It has been known thatP. rapae also shows flower constancy, which may be due to memory constraints. Therefore, they may appropriately use two foraging tactics: visit consistency and labile preference, to get enough nectar according to their circumstances.     

Spatio-temporal distribution patterns of two tachinid flies,Epicampocera succincta andCompsilura concinnata parasitizingPieris

Summary Larvae of genusPieris in the northern part of Kyoto City are parasitized by two tachinid flies:Epicampocera succincta, a specialist on genusPieris, andCompsilura concinnata, a generalist with very wide host-range. We surveyed the parasitism rates ofPieris by both flies for two years at six study areas. In these study areas, there lived three host species in the genusPieris: P. rapae, P. melete, andP. napi, but neither tachinid parasitizedP. napi to any significant extent. In the mountainous district,P. rapae andP. melete coexisted and their populations were relatively continuous, while in the lowland, onlyP. rapae larvae were abundant in spring and autumn, but even they disappeared in summer. Parasitisms byE. succincta occurred mainly in mountainous district and never in the lowland.C. concinnata parasitizedPieris in all the areas, but its parasitisms occurred mainly in autumn. We analyzed the factors affecting the spatial and temporal patterns of parasitism rates and presumed that the temporal discontinuity of host population restricted the distribution of the specialist parasitoid.     

Host plant conspicuousness and the distribution of eggs and larvae in the butterfly,Anthocharis scolymus (Lepidoptera: Pieridae)

Summary We investigated how the distribution pattern of eggs and larval on the host plant,Turritus glabra, was influenced by the oviposition behavior of the pierid butterflyAnthocharis scolymus. Females searched for the host plants visually and they frequently approached taller host plants with sparse surrounding vegetation. After encountering host plants, oviposition behavior of females was independent of host plant characteristics such as height, density, and type of surrounding vegetation. A female laid eggs singly on a host plants. Most females appeared to lay their eggs regardless of the presense of eggs on the host plant. Consequently egg and larva tended to be abundant on conspicuous host plants as measured by height or relative isolation from other plants. However, overcrowding of eggs on an individual host decreased the survival rate of larvae.     

Patterns of prey utilization in a web of orb-weaving spiderAraneus pinguis (Karsch)

Summary The pattern of prey utilization of the orb-weaving spiderAraneus pinguis was studies by comparing between arthropods restrained in the empty webs (spiders were removed) and those unattacked in the intact webs (spiders were not removed). The number of arthropods was larger in the empty webs than in the intact webs. In the empty webs, web area, mesh width, number of radii, and signal thread length were presumed to affect the number of arthropods left in a single web. As for the intact webs, web area, mesh width, and web-exposure time were important factors. In the empty webs, the density of arthropods decreased away from the hub. On the other hand, the density of unattacked arthropods in the intact webs was the same throughout the web. Arthropods in the empty webs were larger than those in the intact webs. The upper limit in size of unattacked arthropods increased along with the distance from the hub in the intact webs, but not in the empty webs. These results indicate increase in the minimum size of eaten arthropods increased in the former. This positive sizedistance relation may have resulted from the adaptive switching of spiders’ alternative foraging methods (i.e., the rapid attack at encounter and the later eating during web deconstruction) on the basis of the prey profitability.     

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.

Sibling and non-sibling cannibalism by larvae of a lady beetleHarmonia axyridisPallas (Coleoptera: Coccinellidae) in the field

Summary Sibling (cannibalism among siblings) and non-sibling cannibalism (cannibalism among non-siblings) were studied in a natural population of a lady beetle,Harmonia axyridis Pallas. Of all the eggs laid (n=2269), 24.76% (n=562) were killed by sibling cannibalism and 36.10% (n=819) were killed by non-sibling cannibalism. Sibling cannibalism occurred constantly and intensively in most egg batches throughout the entire oviposition period. On the other hand, non-sibling cannibalism was more intense in the middle and late oviposition periods, and when the egg batches were close to an aphid colony. This may be due to the high density ofH. axyridis larvae relative to aphid density in the middle and late oviposition periods and also the larvae searching intensively near an aphid colony.     

The role of parasitoids in evolution of habitat and larval food plant preference by three Pieris butterflies

This article attempts to explain that parasitoids provide the evolutionary pressure responsible for relationships between habitat use and larval food plant use in herbivorous insects. Three species of butterflies of the genus Pieris, P. rapae, P. melete, and P. napi use different sets of cruciferous plants. They prefer different habitats composed of similar sets of cruciferous plants. In our study, P. rapae used temporary habitats with ephemeral plants, P. melete used permanent habitat with persistent plants, although they also used temporary habitats, and P. napi used only permanent habitat. The choice experiment in the field cages indicated that each of the three butterfly species avoided oviposition on plants usually unused in its own habitat, but accepted the unused plants which grew outside its own habitat. Their habitat use and plant use were not explained by intrinsic plant quality examined in terms of larval performance. Pieris larvae collected from persistent plants or more long lasting habitats were more heavily parasitized by two specialist parasitoids, the braconid wasp Cotesia glomerata and the tachinid fly Epicampocera succincta. The results suggest that Pieris habitat and larval food plant use patterns can be explained by two principles. The evolution of habitat preference may have been driven by various factors including escape from parasitism. Once habitat preference has evolved, selection favors the evolution of larval food plant preferences by discriminating against unsuitable plants, including those which are associated with high parasitism pressures. Received: December 3, 1998 / Accepted: January 20, 1999     

A comparative study of the searching efficiencies of a parasite and a hyperparasite

Summary The searching efficiencies of a primary parasite (Diaeretiella rapae (McIntosh)) and a hyperparasite (Alloxysta brassicae (Ash.)) were investigated and compared. In both species, at all parasite densities, there was a curvilinear relationship (P<0.001) between the number of hosts parasitised and the host density. A linear regression (loga=logQ−m logP) was fitted for log area of discovery against log parasite density (P<0.001). The area of discovery for its immediate (i.e. primary) host (viz.Diaeretiella for the hyperparasite and aphid forDiaeretiella) is lower in the hyperparasite than in the primary parasite. InDiaeretialla both the searching efficiency and the mutual interference constant increased (but not significantly,P>0.05) in the presence of its males.     

Reproductive properties of the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae), in relation to its wing polymorphism

Summary Reproductive properties of the oriental chinch bug,Cavelerius saccharivorus, were investigated for three types of female adults: brachypters produced in both high and low density populations and macropters produced in high density populations. With respect to egg-laying potential in terms of clutch size, clutch number, and total fecundity, macropters were superior to brachypters produced in the same high density population, although inferior to brachypters produced in the low density population. The mean fecundity of female adults in each type was positively correlated with mean body length. Macropterous females markedly prolonged their initiation of both copulation and oviposition and had a much longer life span compared with both types of brachypters. However, there was no fundamental difference in reproductive schedules between the two types of brachypters.     

Canonical correlation between groups of acarine, fungal and environmental variables in bulk grain ecosystems

Summary Interrelations among acarine, fungal, and environmental components of bulk grain ecosystems were determined by canonical correlation analyses. Twenty-seven variables were measured monthly in samples collected from 2 identical grain bulks in a granary in Winnipeg during the years 1959–67. The relationships between 9 kinds of arthropods and 6 environ mental variables, and between the same arthropods and 12 kinds of actinomycetes and fungi were examined. The maximum canonical correlation between arthropods and environmental factors was 0.35, and between arthropods and microorganisms was 0.28; both are highly significant (p<0.001). In the first analysis correlations of the variables with the canonical variates revealed that correlations of the variables with the canonical variates revealed that sampling location, depth, and temperature are the primary environmental antecedents involved, and the criterion is primarily composed of mitesTarsonemus spp.,Tydeus interruptus and the psocid,Lepinotus reticulatus. In the second analysis the fungiNigrospora sphaerica, Aspergillus spp., andCochliobolus sativus are involved with the mitesCheyletus eruditus andAcarus siro. Generally, the results of these analyses complement the findings of factor and regression analyses of the same data reported earlier. Contribution No. 337 from the Research Station, Canada Department of Agriculture, Winnipeg, Manitoba.     

Female size and age-specific fecundity in the small white butterfly,Pieris rapae crucivora Boisduval (Lepidoptera: Pieridae)

Summary Individual differences in several reproductive parameters of femalePieris rapae were investigated in a controlled laboratory condition. Lifetime and age-specific fecundity showed considerable variability between individuals. Larger females began oviposition at an earlier age than smaller ones, and larger females were more fecund than smaller ones. Larger females laid a larger proportion of their eggs in the early stages of their reproductive lifetime, whilst smaller females laid the larger proportion of their eggs later in their reproductive lifetime. The significance of the variance in age-specific fecundity associated with female size is discussed with respect to the seasonal change in size and habitat utilization of this species.     

Comparison of male adult population densities of the oriental and artocarpus fruit flies,Dacus spp. (Diptera: Tephritidae), in two nearby villages in Penang, Malaysia

Summary The populations of native male adult oriental fruit flyDacus dorsalis (Hendel) and artocarpus fruit flyD. umbrosus (F.) in two selected site (BU and SD) were estimated weekly by the capture-recapture technique using live traps baited with methyl eugenol. In BU where many varieties of fruit trees were grown, the estimated population densities ofD. dorsalis were between 980 and 3100 male flies per ha between May and July, 1984. During the same period, in SD where there were fewer number and varieties of fruit trees, the estimated population densities were between 300 and 1000 flies per ha. The estimated population densities ofD. umbrosus over the same period were between 570 and 1290 flies per ha in BU; and between 5 and 95 flies per ha in SD. Of a total 6828 markedD. dorsalis flies released only one fly (released 6 weeks earlier in BU) was caught in a different site.