Why does onlyAthalia Japonica enter summer diapause among three sympatricAthalia sawflies feeding on crucifers?

Summary This report assesses the primary factor for the evolution of summer diapause of the three species of sawfly,Athalia japonica, A. rosae andA. infumata that feed on cruciferous plants and coexist in the same area.A. japonica has two discrete spring and autumn generations, butA. rosae andA. infumata 5–6 generations. OnlyA. japonica enters summer diapause in response to the long daylengths in spring. Although these three sawflies usually feed on the same cultivated crucifers, they differ markedly in the utilization of wild crucifers. They oviposit only on young leaves.A. japonica mainly usesCardamine plants which sprout in spring and autumn.A. rosae andA. infumata primarily use hosts with new leaves all the year round, i.e. cultivated crucifers andRorippa indica, respectively. The thermal threshold for development is lower inA. japonica than in the other two species. The low heat tolerance ofA. japonica is adapted only to cool shady habitats whereCardamine grows. Presumably, summer diapause ofA. japonica is adaptation to the deterioration of the primary host plants rather than unfavorable climatic conditions. This interpretation is supported by the movement patterns of the threeAthalia sawflies, alternative means to escape from deteriorated habitat conditions.     

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.     

Seasonal life cycle and autogeny in the mosquitoAedes togoi in northern Kyushu, Japan, with experimental analysis of the effects of temperature, photoperiod and food on life-history traits

We studied seasonal changes in the larval population structure, adult size, and autogeny (egg production without a bloodmeal) of the mosquitoAedes togoi on the seacoast of northern Kyushu, Japan. The effects of temperature, photoperiod and food conditions on larval development, adult size and autogeny were studied in the laboratory.Aedes togoi overwintered in both the egg and larval stages and was multivoltine. Adult size was greater in spring and autumn and smaller in summer. Autogeny occurred in spring and autumn but not in summer. Autogenous females were larger than anautogenous females, and larger autogenous females produced more eggs. Laboratory experiments showed that autogeny was promoted under low temperature and short-day conditions which corresponded with spring/autumn conditions in the field. Experimental food limitation in the larval stage greatly reduced adult size and autogenous egg production. In the field, large inter-pool variations in adult size and autogeny rate probably resulted from variations in nutrient and crowding conditions. Seasonal autogeny ofAedes togoi was discussed in terms of life-history strategy under fluctuating environmental conditions where relative advantages of autogenous and anautogenous reproduction alternate seasonally.     

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.     

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.     

The effects of wild cruciferous host plants on morphology, reproductive performance and flight activity in the diamondback moth,Plutella xylostella (Lepidoptera: Yponomeutidae)

The wild crucifers,Rorippa indica andLepidium virginicum, are known to serve as host plants for the diamondback moth (DBM),Plutella xylostella, but they are less suitable than the preferred cultivated cruciferous plant, cabbage, in terms of adult body size and fecundity. The life history traits and flight activity of DBM adults grown on various host plants were investigated. The adults thus reared on each host plant were divided into three size groups (small, medium and large). In general, female adults grown on the wild crucifers were less fecund and lived longer than those reared on cabbage. Flight activity was higher in adults grown on wild crucifers than in those reared on cabbage. Male adults flew longer than females. Fecundity, longevity, flight activity and morphometrical characters of adults were positively correlated with pupal weight in individuals reared on the same host plant. A negative relationship was found between fecundity and flight activity in females of the same size group, but a positive one was observed in females reared on the same host plant.     

Seasonal change of nest survival and related aspects in an aggregation ofLasioglossum duplex (Dalla torre), a eusocial halictine bee (Hymenoptera: Halictidae)

Summary Nest survival in an aggregation of a eusocial halictine bee,Lasioglossum duplex, was censused through an annual cycle. Out of 2,500 nests marked at the beginning of solitary phase in the spring, only 25.5% attained eusocial phase. But 60.5% of 636 nests attaining eusocial phase successfully produced sexual offspring. This shows that solitary phase is the most vulnerable period in the annual cycle. A very low productivity in eusocial phase in the census year was clarified from examination of 99 nests in the autumn. The number of prospective foundresses surviving to the next year was estimated upon the number of brood cells in examined nests and of old females surviving these nests. Deviation between this estimate and the number of nests made in the next spring was 8.8%, confirming a drop of population size to less than one third in the next year. Some nests solitarily made in the summer by dispersed females were examined. Productivity in such nests was extremely low, hence these nests should contribute virtually nothing to the next generation. Possible factors affecting the low productivity in eusocial phase were enumerated though none of these were supported by concrete evidence. Some considerations were given on the relation between obtained results and colony life cycle in eusocial insects, particularly in halictine bees. Bionomics of the eusocial halictine bee,Lasioglossum duplex. VI.     

Seasonal host alternation by the andromeda lace bug,Stephanitis takeyai (Heteroptera: Tingidae) between its two main host-plant species

I studied the seasonal occurrence of the andromeda lace bug,Stephanitis takeyai, on its two main host-plant species. In a secondary forest in Kyoto, this bug altered its hosts seasonally, i.e., from an evergreen shrub,Pieris japonica, in winter to a deciduous shrub,Lyonia elliptica, in summer. In contrast, in Nara park where fewL. elliptica were available, the bug exploited onlyP. japonica. Thus, seasonal host alternation by this bug is not obligate. A comparison of adult longevity and fecundity on the two host-plant species demonstrated the higher quality ofL. elliptica as a food resource. Corresponding to this difference in host quality, there was a dramatic difference in the seasonal population growth in the two study sites. In Nara, the population size at the beginning of the 2nd generation was almost the same as in the overwintered generation, whereas in Kyoto the population size in the 2nd generation was approximately one hundred times as large as in the overwintered generation. Thus seasonal host alternation is adaptive for the bug. In a previous study, I reported that overwintering as eggs in living leaves of their hosts is likely to be common among all the related species of this bug. Thus, this trait can be considered to be a phylogenetic constraint to the group. I speculate that host alternation by this bug has been derived because it is more adaptive from autoecy on an evergreen plant, similar to the pattern currently found in Nara, and that this bug can not only exploit deciduous host due to a phylogenetic constraint.     

Population dynamics of a phytophagous lady-beetle,Epilachna vigintioctopunctata (Fabricius), living in spatio-temporally heterogeneous habitats. III. Effects of habitat structure on population dynamics

Temporal changes in the population size of a phytophagous lady-beetle were analyzed to identify mechanisms affecting lady-beetle population dynamics at different spatial scales. The study area (15 ha) included 18 habitat patches. The major host plants were potato for first generation larvae and eggplant for second generation larvae. The habitat patches were classified into three groups according to the major host plants in each patch: P-E patches (both host plants available), P patches (potato only), and E patches (eggplant only). The winter disappearance of adults in the whole study area, and larval mortality in E patches were apparently the most important factors disturbing the overall population density. Density-dependent movement of females appeared to have the greatest stabilizing effect on the yearly fluctuation of population density. Rate of increase of female adults from the first to the second generation,R, was generally higher on eggplants in E patches than in P-E patches because the adult density of the first generation was much higher in P-E patches. The yearly fluctuation of adult density in each generation tended to be less in patches with all habitat components necessary for the full life cycle (P-E patches). However, such patches were not favorable for first generation females, as indicated by the lower rate of increase from the first to the second generation. The density and stability of lady-beetle populations is discussed in relation to habitat structure.     

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.     

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.     

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.     

Influences of wild crucifers on life history traits and flight ability of the diamondback moth,Plutella xylostella (Lepidoptera: Yponomeutidae)

The larvae ofPlutella xylostella were fed on five wild crucifers,Capsella bursa-pastoris, Lepidium virginicum, Cardamine flexuosa, Rorippa indica, R. islandica and a crop, cabbage. The developmental period of the immature stages, adult longevity, preoviposition period, fecundity and morphometrical characters of the adults were measured. The flight activity of the adults was also measured by the tethered flight method. All the wild plants except forR. islandica were less suitable host plants than cabbage, and larvae which were fed on these less suitable plants emerged as smaller adults with shorter wings. The smaller female adults had lower fecundity but a higher flight activity. Smaller adults measured in terms of their pupal weight among individuals fed on the same host plant had longer wings. These smaller adults with longer wing flew more actively.     

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.     

A population study of the japanese wood mouse,Apodemus speciosus (Mammalia: Muridae), with reference to its social behavior

Summary The role of social behavior on the population regulation of the Japanese wood mouse,Apodemus speciosus, was studied in the Ashu Experimental Forest of Kyoto University by monthly trapping and direct observation on the social behavior at the artificial feeding site. The census was carried out from May to October in 1974 and 1975, and the direct observation in June, August and October, 1975. In the study area, the Japanese wood mouse has two breeding seasons in a year: in spring and in autumn. Minimum survival rate was high in the non-breeding season and low in the breeding season. It was negatively correlated to the number of sexually active adults in both sexes. Home ranges of females tended to be mutually exclusive, and female residents prevented the settlement of female immigrants. On the other hand, home ranges of males overlapped largely and males had a straight social hierarchy. Subordinate males tended to disappear more often than dominant males did, but male residents did not hinder the settlement of male immigrants. Males may regulate their number by driving out the subordinate males, while females by spacing behavior. Contribution from the Laboratory of Animal Ecology, Department of Zoology, Kyoto University, No. 445.     

Extinction of the trochid gastropod,Umbonium (Suchium) moniliferum (Lamarck), and associated species on an intertidal sandflat

On an intertidal sandflat in western Kyushu, Japan in 1979, the trochid gastropod,Umbonium (Suchium) moniliferum (Lamarck), and the thalassinidean ghost shrimp,Callianassa japonica Ortmann, densely inhabited the lower and upper zones, respectvely.Callianassa japonica subsequently expanded its distribution range considerably, having occupied almost the entire sandflat by 1983. Concurrently, theU. moniliferum population gradually declined, becoming extinct in 1986. Furthermore, the populations of 9 species associated withU. moniliferum (predators, an ectoparasite, and subsequent inhabitants of emptyU. moniliferum shells) had disappeared by the end of 1992. The extinction processes ofU. moniliferum and the two other most numerically dominant species, the ectoparasitic gastropod,Odostomia sp., and the hermit crab,Diogenes nitidimanus Terao, were described in detail. It is believed that the bioturbation of sediments byC. japonica was responsible for the extinctions, possible mechanisms involved being discussed. This is the first documented record of the extinction of a large part of a macrobenthic assemblage centering on a filter-feedingUmbonium species which is characteristic of the benthic communities on many intertidal sandflats extending from Japanese to Southeast-Asian waters.     

Population fluctuations and natural mortalities of the pine-moth,Dendrolimus spectabilis

Summary Annual changes in the population of the pine-moth,Dendrolimus spectabilis Butler, were studied in Kashima district. The insect completes two generations in one year, and the adult emerges in early summer and again in autumn. The insect population showed fairly drastic fluctuations in the past years, and a conspicuous peak was recorded in three times, in 1950, 1955 and 1961. It was noted in each of those three years that a great number of adults of the summer generation emerged in autumn. The main reason for this great abundance of the insect population seemed to be high survival rate of the young larvae in summer. It was suggested that the climate in July had powerful influence upon the survival f thos e young larvae. The effects of natural enemies were usually the greatest on the egg population of the pine-moth in either generation. Much greater number of insects in the overwintering generation was destroyed by parasites and diseases than in the summer generation. Finally, a rapid increase of the insect population in autumn was considered to be caused by a combined effect of climatic conditions and of natural enemies.     

Reproductive behavior of the Japanese treefrog,Rhacophorus arboreus (Anura: Rhacophoridae)

Reproductive behavior of the foam-nesting treefrog,Rhacophorus arboreus, is described. Oviposition was made either by an amplectant pair (a female, and an amplectant male) alone or by an amplectant pair and other males (joiners). The snout-vent length of males and females was negatively correlated with the date of the first appearance in the pond. The period of staying in the pond in males was longer than that in females. Physical body contacts of other males, and encounter calls by ampletant males were frequently observed before oviposition with joiners. Snout-vent length of the female and an amplectant male were positively correlated. Encounter calls by amplectant males were likely to precede movement of females. There was no difference of snout-vent length between amplectant males and joiners. Females moved the hindlimbs more frequently than males during oviposition. Amplectant males made encounter cells more frequently in oviposition with joiners, than oviposition without joiners. The patterns of transitions between acts of females and amplectant males were similar between oviposition with and without joiners. Females more frequently responded to amplectant males than joiners. Both the duration of a bout of the movements of hindlimbs of females and the interval between the bouts were longer in oviposition without joiners than that with joiners. The number of the movements per bout in oviposition without joiners was smaller than that with joiners. Encounter calls by amplectant males during oviposition tended to precede the hindlimb movements of females.     

Estimation of prey consumption of a mantid,Paratenodera angustipennis (S.) in a natural habitat

Summary Using a relationship between prey consumption and growth rate, field prey consumption of adults ofParatenodera angustipennis (S.) in a paddy field was estimated. Since a great number of grasshoppers (Oxya japonica) lived in the research area and the mantids had frequently eatenO. japonica, we presumed that they consumed onlyO. japonica. As a result, it was estimated that average prey consumptions for 6 days were 406 mg for the females and 23 mg for the males. From the result of this estimation and the feeding rhythm of the mantids, it is concluded that each female captured one grasshopper on average every a few days. By comparing this actual feeding level ofP. angustipennis with those of other predacious arthropods, characteristics ofP. angustipennis as a predator were discussed.     

Body mass changes inTrimeresurus flavoviridis (Viperidae) on Okinawa Island, Japan

Body masses of wild-caught habu,Trimeresurus flavoviridis, were measured with known error range. Habu larger than mature female size had steeper slope in length-mass regression than smaller individuals. Females outweighed males in most snout-ventlength classes and in early summer. Gravid females outweighed non-gravid ones by about 20% on the average, but the body mass ranges of the two groups overlapped. Through the body mass change in mature females, the proportion of gravid females was estimated to be about 0.5.