Why doLuehdorfia butterflies lay eggs in clusters?

Summary Luehdorfia butterflies lay eggs in clusters. Clones of their host plants (Asiasarum andHeterotropa) are distributed pacthily among the understory of deciduous forests. Groups ofLuehdorfia larvae often exhaust the clones and may wander over the forest floor seeking new clones. The highest mortality observed is during this wandering period. To elucidate whyLuehdorfia butterflies lay eggs in clusters, a simulation experiment was made for hypothetical populations which lay eggs in clusters or singly. Field data on larval mortality, consumption, density of host clones and leaf weight forLuehdorfia japonica were incorporated into the model. The predictions of the simulation were: (1) When the egg density is low, the single egg type could leave many more pupae than the egg clustering type, but when the egg density is high, the former might leave smaller number of pupae than the latter; and (2) There are optimal sizes of egg clusters for different egg densities and the optimal size becomes larger as the egg density increased. This work was supported in part by Grant-in-Aid No. 439017 and No. 56480039 from the Ministry of Education, Science and Culture, Japan.     

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.     

Spatial distribution and mortality process ofAnoplophora malasiaca (Coleoptera: Cerambycidae) eggs in citrus groves

Summary The study was carried out in two (A and B) citrus groves to clarify the spatial distribution patterns of eggs and larvae, and to analyse the mortality process of eggs. From the analysis by using the mean density and the mean crowding, it was clarified that the distributions of eggs were contagious and that larvae were more contagiously distributed than eggs. The τ andz indices showed that the operation of egg mortality was inversely density-dependent in both groves, and that the degree of inverse density-dependence was greater in A than in B grove. The spatial correlations between the emergence holes and the eggs or larvae in each tree, which were analysed by using ω index, showed that the distributions were more overlapping between the emergence holes and the larvae than the eggs. As the result of dividing trees into several groups according to the number of emergence holes, it was clarified that the survival rates of eggs were positively correlated with the number of emergence holes. In conclusion, inversely density-dependent mortality process was considered to be caused by lower mortality rates of eggs in the trees with more emergence holes. Especially in A grove, because the trees with more emergence holes were larger in diameter and more egg oviposition, the inversely density-dependent mortality process was considered to be detected more conspicuously than in B grove.     

Contest competition inDrosophila subobscura

The role of larval intraspecific competition in laboratory populations ofDrosophila subobscura was investigated. Mortality is density-independent during the first 3 days after hatching but becomes density dependent as development proceeds to pupation. Although total biomass per patch was independent of initial egg density, competition betweenDrosophila larvae leads to the formation of smaller pupae. This resulted in a population that was dominated by suppressed individuals. Development rate ofD. subobscura larvae was not affected by high larval densities. Smaller pupae give rise to females with fewer eggs in their ovarioles. A simple simulation model, predicting the effects of intraspecific competition on the fecundity of the nextDrosophila generation is described.     

Effects of host and parasitoid age on search behaviour and oviposition rates inLariophagus distinguendusF?rster (Hymenoptera: Pteromalidae)

Summary Oviposition rates and related behaviours were quantified forLariophagus distinguendus F?rster attackingCalosobruchus chinensis (L.) andC. maculatus (F.). Oviposition rates varied with parasitoid age; parasitoids aged 1–7 days laid approximately twice as many eggs per day as those aged 8–14 days. Similar differences were noted in search rates and handling times; younger parasitoids had higher attack rates and lower handling times than older parasitoids. Search rates and handling times also varied with the host stage available for attack. Search rates were higher and handling times were lower on larger stages. The results are discussed with reference to their impact on the dynamical behavior of insect parasitoid-host populations.     

Biology ofHyphantria cuneaDrury (Lepidoptera: arctiidae) in Japan. V. Preliminary life tables and mortality data in urban areas

Summary Studies on the population dynamics of the fall webworm,Hyphantria cunea have been carried out at three survey stations and along selected roads in the urban area of Tokyo since 1966. Twelve survivorship curves obtained during two years and 8 life tables show that the mortality rate in early developmental stages of the fall webworm is remarkably low as compared with that of other lepidopterous defoliaters and the mortality rate in later developmental stages is compensatory high. The low mortality rate in early stages is considered to be due to the protective role of the nest-web and the lack of egg and larval parasites. All but one parasitic species emerge from prepupae and pupae. Spiders living in the nest-web of the fall webworm play an important role in reducing the number of young larvae. Direct observations and caging experiments showed that relatively high mortality during later larval stages is mainly due to predation by birds (in the first generation) and wasps (in the second generation). The generation mortality in the survey stations always exceeded the level where the population is kept at the steady state, and the outbreak of this moth is considered to be continued by the immigration of adults from large trees growing in gardens on which the larvae can escape from predation pressure. Contributions from JIBP-PT No. 51. A part of this study was supported by the special project research, ‘Studies on the dynamic status of biosphere’, sponsored by the Ministry of Education.     

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.     

Density-dependent growth and reproduction of the apple snail, Pomacea canaliculata : a density manipulation experiment in a paddy field

To examine density dependence in the survival, growth, and reproduction of Pomacea canaliculata, we conducted an experiment in which snail densities were manipulated in a paddy field. We released paint-marked snails of 15–20 mm shell height into 12 enclosures (pens) of 16 m² at one of five densities – 8, 16, 32, 64, or 128 snails per pen. The survival rate of released snails was 95% and was independent of snail density. The snail density had a significant effect on the growth and egg production of individual snails. This density dependence may have been caused by reduced food availability. The females at high density deposited fewer and smaller egg masses than those at low density, and consequently produced fewer eggs. The females at densities 8 and 16 deposited more than 3000 eggs per female, while the females at density 128 oviposited only 414 eggs. The total egg production per pen was, however, higher at higher snail density. The survival rates of juvenile snails were 21%–37% and were independent of adult density. The juvenile density was positively correlated with the total egg production per pen and hence was higher at higher adult density. However, the density of juveniles larger than 5 mm in shell height, i.e., juveniles that can survive an overwintering period, was not significantly different among density treatments. These results suggest that snail density after the overwintering period is independent of the density in the previous year. Thus, density dependence in growth and reproduction might regulate the population of P. canaliculata in paddies. Received: October 23, 1998 / Accepted: July 16, 1999     

Delayed maturation in the colonial coralGoniastrea aspera (Scleractinia): whole-colony mortality, colony growth and polyp egg production

The present study examines (1) the cost of reproduction on colony growth, and (2) relationships among sexual maturity, whole-colony mortality rate and colony growth rate inGoniastrea aspera free from external influences by macrobenthos. Survival of colonies in permanent plots was followed for two years. Egg production by polyps in colonies collected just before the first spawning of a year was estimated by dissecting the polyps. Growth of the colonies (increase in number of polyps) was followed over one annual reproductive cycle. The cost of egg production on colony growth was apparent through colony ontogeny: (1) immature colonies had a greater annual growth rate than mature colonies, but produced almost no eggs; (2) in mature colonies, growth rate was negatively correlated with NE/PV (number of eggs per polyp volume mm^-3). Annual whole-colony mortality was high in colonies with fewer than11 polyps in initial colony size, while mortality was extremely low once a colony grew beyond this size. This critical size for low whole-colony mortality was much smaller than the colony size (40 polyps) which would attain maturity one year later. Age at maturity was estimated as six years. While survival to maturity may be a selective force for the evolution of delayed maturation, the present data suggest that high colony fecundity, achieved after a long growth period as an immature colony, and an abrupt decrease of colony growth rate after maturation are the crucial forces.     

Elimination ofSitotroga cerealella (Olivier) bySitophilus zeamais Motschulsky from mixed laboratory cultures on maize

Summary This paper describes a series of experiments conducted to determine whySitophilus zeamais Mots. andSitotroga cerealella (Oliv.) could not survive together in maize cultures in the laboratory. The effect ofS. zeamais on different developmental stages ofS. cerealella was investigated. The presence of adultS. zeamais slightly affected moth copulation, egg laying and moth eggs in a mixed culture, but large numbers of developing moths inside maize grains were killed by the adult weevil through feeding on the grains. The major cause of elimination ofS. cerealella byS. zeamais from mixed cultures was therefore found to be damage to the immature moths in grain and such moth mortality increased as the developing moths became bigger in the grains. A weevil: grain ratio of approximately 1.4∶1 was found to be the critical weevil density at which the moth disappeared from the mixed cultures.     

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.     

Estimatingdendroctonus frontalis (Coleoptera: Scolytidae) daily infestation dynamics

Mathematical procedures are given to estimate infestation totals and daily life stage arrivals, departures, and mortality ofDendroctonus frontalis Zimmermann for an infested tree in the field. These estimates are based on minimal sample data and are designed to utilize all available information. Daily arrival estimates for larvae, pupae, and callow adults are obtained by indirect analysis without direct observation of these stages. The procedures are applied to 147 infested trees, and the results are transformed to a common time basis to obtain daily expectations by life stage for an “average” tree. These expectations suggest optimal times for field sampling or relative times of sampling when optimal times are missed. Expected daily arrival distributions by life stage for a single egg and a single attacking adult are given. Procedures are given for utilizing collateral information to obtain an infestation total and daily arrival estimates for a boundary life stage. The results of this study are applicable to anyD. frontalis field study, and the procedures given are applicable to any bark inhabiting insect having similar habits.     

Evaluation of the role ofMicrovelia douglasi atrolineata (Bergroth) (Heteroptera: Veliidae) as predator of the brown planthopperNilaparvata lugens (St?l) (Homoptera: Delphacidae)

Summary The biology of the veliid bugM. d. atrolineata, its predatory behavior, and the effects of plural hunting were studied to evaluate its role as a predator of the brown planthopperNilaparvata lugens in the Philippines. The probability of planthoppers falling onto the water surface and provision of habitat continuity was measured by a sticky trap placed at the base of rice hills in a greenhouse and in paddy fields. The developmental period of immature stages combined was 21 days. If given prey, females laid 25 eggs on the average during an adult life span of 18 days. Starved adults could survive for only 3–5 days. The functional response to prey density was sigmoid, and the maximum number of prey killed was 7 per day. Prey feeding was completed in 12–36 min. The percentage of successful prey attacks averaged 5–8%, decreasing with higher (and larger) developmental. stages of prey, but adult prey were found the soonest. Plural hunting increased the probability of capturing prey by as much as 2.5 times that by individual hunting. Late-instar nymphs, which may be more active, fell from rice hills in a greenhouse more than early-instar nymphs, and the number falling increased with density. In the field the percentage of planthoppers falling to the water in 1 day varied considerably, from 1% for nymphs in one field to 67% for adults in another field. On the basis of work described above and given the high density of veliid predators in flooded paddy fields of tropical Asia,M. d. atrolineata is considered one of the most important natural enemies of the brown planthopper.     

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.     

A laboratory study on the predatory miteTyphlodromus pyri (Acarina: Phytoseiidae): I. The effect of temperature and food consumption on the rate of development of the eggs and immature stages

Summary Development periods were determined for eggs and juvenile stages of the predatory phytoseiid miteTyphlodromus pyri at four constant temperatures and three feeding levels. For the non-feeding stages (eggs and larvae) power curve functions were fitted to development rate in terms of temperature. The results are compared to the general arthropod response to temperature. A non-linear model was fitted to nymphal development rate in terns of temperature and food, in which the weight gain over an instar is assumed to be a linear function of daily food consumption. Examples of the same shape development rate—food relationship are given for two more phytoseiid mites and for other arthropods. The results are discussed with respect toT. pyri as an integrated control agent.     

Adult population parameters and life tables of an epilachnine beetle (Coleoptera: Coccinellidae) feeding on bitter cucumber in Sumatra

Summary The population dynamics of an epilachnine beetle, which is closely related toEpilachna sparsa Dieke (henceforth called “sp. C”) and feeds on bitter cucumberMomordica charantia, was studied by mark-recapture of adults and the construction of life tables. The study was repeated three times, i.e., March–May, July–September and October–December in 1982, in Padang, Sumatra, Indonesia. After the establishment of the host plants, adults of “sp. C” soon colonized, and each study period ended in the death of the plants due to defoliation by the larvae and adults. The estimated mean length of residence of adults ranged from 6–11 days, but this was probably much shorter than the actual longevity, because the adults were so active that they flew away, or dropped off the plants, when they were approached or slightly disturbed. Life tables indicated that egg mortality ranged from 17.8–53.9%, and a parasitic waspTetrastichus sp. B made up 41.1–64.2% of egg mortality. Two wasps,Tetrastichus sp. C andPediobius foveolatus killed 1.2–19.4% (7.6–100%)^* of 4th instars and only the latter species attacked the pupae, killing 24.6–59.1% (45.1–72.4%). Parasitism and starvation by overcrowding contributed most to the total mortality from egg to adult emergence, which ranged from 89.4–99.5%. “Sp. C” had a higher diversity and level of parasitism than the Japanese species,E. vigintioctopunctata. The high dispersal power of “sp. C”, coupled with the prolongedl _x−m_x schedules shown under laboratory conditions, was advantageous for exploiting the food plant which was available throughout the year, but was rather patchily distributed in space.     

Analysis of contagiousness in the action of mortality factors on the western tent caterpillar population by using the \mathop m* - m\mathop m\limits^* - m relationshiprelationship

Summary The contagiousness in the operation of mortality processes on the colonies of the western tent caterpillar,Malacosoma californicum pluviale, was analyzed from two different aspects: successive changes in the frequency distribution of the number of surviving individuals per colony in the course of development, and the distribution pattern of the individuals killed by some biotic mortality factors. Also, for a tachinid parasite,Tachinomyia similis, the analysis was made on the egg-laying pattern on colonies as well as on individual larvae. The methods of these analyses were all based on the relation of mean crowding (m) on mean . A braconid parasite,Rogas sp., tended to kill few individuals together once it attacked a colony, but its effect on host colonies was rather equivalent to the random removal of individuals from all the colonies. Diseases in the late-stage larvae before cocooning was contagious in their action. Nuclear polyhedrosis virus seemed to have no basic contagiousness in its action, but it caused highly contagious distribution of deaths among the colonies when its average incidence was high. A spore-formingBacillus had a tendency to kill several individuals once it appeared in a colony, but the distribution of its incidence (no. of times it appeared per colony) was considered to be nearly at random. The female ofTachinomyia tended to lay more than one egg successively on the same colony. It also attacked individual larvae with a definite tendency for aggregation, which seemed to be resulted from the parasite’s preference to large hosts. When the number of eggs laid on prefered hosts exceeded a certain threshold, however, the fly seemed to change its attention to less attractive, smaller individuals.     

Population biology of the broad-horned flour beetle,Gnathocerus cornutus (F.) II. Crowding effects of larvae on their survival and development

Summary Crowding effects of larvae on survival and development were examined for the broad-horned flour beetle,Gnathocerus cornutus (F.). The larvae matured about 3 weeks after hatching regardless of their densities, but pupation was severely hindered by crowding. There existed an upper limit for the number of the pupae produced and its mechanism was studied by a statistical analysis of the distribution patterns of pupal cells and the experiment in which glass tubes were artificially supplied in addition as pupation site. These studies show thatG. cornutus larvae have a habit to construct cells for pupation and this habit leads to a contest competition for pupation site at high densities. The significance of the contest competition for population regulation was discussed comparing the results onTribolium confusum Jacqueline duVal.     

The effects of age and weather on egg laying inDanaus plexippus L. (Lepidoptera: Danaidae)

Summary The effects of age and weather conditions on egg laying inD. plexippus were determined for caged females. Age (measured in physiological time), temperature and solar radiation influence egg laying in this species of butterfly. An algorithm taking these factors into account in presented and accounts for 88% of the daily variation in egg laying. CagedD. plexippus begin to lay eggs six—seven days after emergence, peak egg production (about 60 eggs/♀) occurs about 15 days later. Females continue to lay eggs throughout their adult life, which in a flight cage was about 40 days. This egg laying pattern is compared with other published fecundity schedules. The effect and importance of a female being prevented from laying her eggs, on her life-time egg production, is also discussed.