Cumulants of random sum distributions

Consider a random variable S being the sum of a number N of independent and identically distributed random variables X_j (j = 1, 2, ...) where the number N is itself a non-negative integer-valued random variable independent of the X_j An explicit expression of the r-th cumulant of S is given in terms of the cumulants of N and X_j, Asymptotic properties of the distribution of S are also discussed.     

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.     

Male mating behavior in relation to spermatophore transfer in the white cabbage butterfly

The lifetime mating frequency of female butterflies is believed tobe dependent on the reproductive status of the males which they have mated. This report assesses those status usingPieris rapae L. Multiple mating females mated males with a short time interval after the last mating or males with many mating records. Such males, like small ones, produced small spermatophores during copulation, which may have resulted in high mating frequency of those females. The males with short time interval after the last mating or those with many mating records also showed a long mating duration. Alternative interpretations of the adaptive significance of this behavior for males are discussed.     

Ecological studies on a dragonfly,Nannophya pygmaea ramber (Odonata: Libellulidae) I. seasonal changes of adult population and its distribution in a habitat

Summary Adult population of a dragonflyNannophya pygmaea Ramber inhabited in a damp ground was investigated with mark-and-recapture method in 1975. The following results about the seasonal changes of the population size and distribution in the habitat were obtained. Adults emerged from late May to mid August. The number of the adults was most abundant in early June, but that of matured males in early July. From the recapture data, the estimate of daily survival rate was 0.82, and the length of immature stage in males was estimated as 5 days or so. Total number of post-teneral adults emerged in the habitat was estimated as about 9,000. The mean crowding-mean density regression method was applied for the analysis of the distribution pattern of the adults. Matured males showed a spaced-out distribution, while females and immatured males distributed themselves rather aggregatively. Such a distribution pattern of the matured male would be attributed to their territorial behaviour. The territorial behaviour was considered to force the matured males to extend the distribution area in July when they were most abundant. From the above-mentioned results and some observations, the meaning of the territoriality in this species was discussed.     

Inter- and intraspecific interactions among larvae of specialist and generalist parasitoids

Intra- and interspecific larval interactions that take place in a host body were investigated for two tachnid fliesEpicampocera succincta andCompsilura concinnata (Diptera: Tachinidae) parasitizingPieris butterfly larvae.E. succincta, a specialist onPieris butterflies, showed contest-type intraspecific competition, eliminating all the other conspecific larvae. On the other hand, an extreme generalist parasitoidC. concinnata exhibited scramble-type competition, sharing the host with other conspecifics and suffering reduced body size as a result. However, when these two species occurred together in a single host,C. concinnata had a much higher chance of survival. Moreover,C. concinnata could often survive in the presence of a parasitoid waspCotesia glomerata (Hymenoptera: Braconidae) whileE. succincta could not. The high tolerance ofC. concinnata could be attributable to its being an extreme generalist: To attack and survive on many different hosts, one has to be able to deal with various competitors. The competitive inferiority of the specialistE. succincta, on the other hand, may be a result of relatively recent encounter with, those competitors.     

Comparative population studies of threePieris butterflies,P. rapae, P. melete andP. napi, living in the same area

Summary Utilization of patchy habitats by adult populations of threePieris butterflies,P. rapae, P. melete andP. napi was studied throughout the flight season in an area of their coexistence, about 3×1.5 km, in a farm village in the mountains in Inabu, Aichi Prefecture. Field study was by the mark-recapture method. Results were analyzed by dispersal distances and recapture duration decay curves for adults of different age-classes estimated on the basis of physical condition of their wings, together with supplementary information of daliy egg-laying rate of females, obtained in field cages. Sexually immature, mated femals ofP. rapae after teneral stage showed a migratory flight. On the other hand, reproductive females and all males ofP. rapae were strongly resident within suitable habitats, and reproductive females begun to lay eggs abundantly at sunny places of newly suitable areas within a short period.P. melete seemed to disperse gradually from emerged stites and females of this species continued to lay some constant numbers of eggs for more than ten days over a wider area.P. napi appeared more likeP. melete thanP. rapae. The habitats of the three species can be characterized as follows:P. rapae, temporary, continued for pre-reproductive females but localized for reproductive females and all males, and unstable;P. melete, permanent, widespread, and stable;P. napi, permanent, localized, and stable. The numbers of generations ofP. rapae, P. melete andP. napi were estimated to be about six, three and three, respectively. Seasonal fluctuations in the number of adults were influenced by the stability of their habitats, i. e., the population size fluctuated sharply inP. rapae, but it was much more stable inP. melete andP. napi. In view of these results, it can be said thatP. rapae fits the general characteristics of a r-strategist whereasP. melete andP. napi are more K-strategic thanP. rapae.     

Warning coloration in sawflyAthalia rosae larva and concealing coloration in butterflyPieris rapae larva feeding on similar plants evolved through individual selection

Summary Green larvae of the butterflyPieris rapae and black larvae of the sawflyAthalia rosae feed on green leaves of the same cruciferous plants. To demonstrate thatP. rapae has concealing coloration and thatA. rosae has warning coloration, the larvae of the two species were supplied to naive chicksGallus gallus on white, green or black backgrounds.P. rapae larvae were palatable and their green body color acted as a concealing coloration. On the other hand,A. rosae larvae were unpalatable and their black body color acted as a warning coloration. There is a general consensus that warning coloration is an altruistic character which needs victims, and thus can evolve through kin selection or green beard selection. However, blackA. rosae larvae were seldom injured by chicks' attack, in particular, on the green background. Therefore, the warning coloration ofA. rosae larvae can be a selfish character and hence can evolve through individual selection as well as concealing coloration ofP. rapae.     

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     

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.