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.     

How three rocky shore snails coexist on a limited food resource

Summary Competition theory poses a major problem when several species coexist on what appears to be one resource. ThreeThais species living on the Pacific Northwest Coast provide an example: at many sites, all three depend primarily on one barnacle species. Growth rates of three species were measured for 3 years and these provide an indirect means to assess how these snails use their common food resource. Major temporal differences were observed:T. lamellosa grew 0–1 mm/mo during the spring and 2–3 mm/mo during the summer, whileT. emarginata andT. canaliculata grew 2–3 mm/mo during the spring and 0–1 mm/mo during the summer. However, all species are opportunists when food is available, and seasonal and interspecific differences disappeared when all three species were kept well fed together in the laboratory. Therefore, temporal differences arise from spatial segregation rather than from intrinsic differences in activity, and must arise because barnacle abundance patterns differe consistently from one area of the shore to another. Species-specific activity patterns lead each species to a food intake that is independent of the food supply on the shore as a whole and is also independent of food intake by the other two species. Where two snail species depend upon a single food species, their use of the food supply appears to make it function as two different resources. This resource use is possible because prey quality is markedly dependent on shore level.     

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.     

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.     

Postmetamorphic growth, survival, and age at first reproduction of the salamander,hynobius nebulosus tokyoensis Tago in relation to a consideration on the optimal timing of first reproduction

Summary The postmetamorphic growth and survival of the salamanderHynobius nebulosus tokyoentis Tago were surveyed in the study site located in Habu village of Hinodemachi, a suburb of Tokyo City, during 1975–1981. A laboratory experiment on the growth rate of juveniles was conducted in parallel with the field survey. The result indicated that this salamander grew at the rate of 8,mm in s.v.l. per year during the juvenile stage, but its growth rate decreased markedly as low as 1.8 mm for males and 1.1 mm for females, once it had attained sexual maturity. According to the “capture-recapture” procedure the annual survival rate after metamorphosis was found to be quite high; that is, approximately 0.7. By using the growth rate of juveniles and the difference between the sizes at metamorphosis and sexual maturity, the age at first reproduction was estimated to be 4 year for males and 5 year for females. From the data obtained in this study, the intrinsic rates of increase (r) were calculated for various values of age at first reproduction under different survival schedules, and the relationship between the age at first reproduction and fitness as measured byr was examined. The result indicated that an optimal age maximizing fitness always existed under respective survival schedules, and the observed age at first reproduction of this salamandei was found to coincide well with the predicted optimal age.     

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.     

Differences in egg wastage by superparasitism, contrastingVenturia (=Nemeritis) canescens searching singly versus searching in groups

Summary A statistical and graphic study is presented of the wastage of eggs byVenturia (=Nemeritis) canescens when searching singly and in groups of 10 among hosts at four different host densities in laboratory universes as described byHuffaker andMatsumoto (preceding paper of this journal). The host insect was the fluour mothAnagasta kühniella and the host densities used were 10, 30, 100 and 200 per universe. Intensity of egg wastage due to superparasitim varied significantly according to host density, and between the two parasite densities employed, 1 and 10, using bothF-tests and chi-square tests. Plots ofk-factor analysis on this egg wastage showed high negative correlations with host density, and the raw data for single parasites was well represented by a parabola while that for the grouped parasites departed from this relationship only at the lowest host density. These studies were conducted as a part of a general investigation into the processes operating in the population dynamics of arthropods under grants from the National Institutes of Health, U. S. Public Health Service (#A10-1611), and the National Science Foundation and Environmental Protection Agency (NSF DEB7504223) to the University of California. The findings, opinions and recommendations expressed herein are those of the authors and not necessarily those of the University of California, the National Institutes of Health, the National Science Foundation or the Environmental Protection Agency.     

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.     

Life table studies on the paddy field population of the green rice leafhopper,Nephotettix cincticeps Uhler (Hemiptera: Cicadellidae), with special reference to the mechanism of population regulation

Summary The population growth of the green rice leafhopper,Nephotettix cincticeps, in the paddy field was analyzed based on the life table data accumulated for six years. The paddy field population, which stems from the invading adults of the first generation (G-I), repeats two complete generations, and the hatchlings of the fourth generation (G-IV) enter diapause and overwinter as the fourth instar nymphs in fallow paddy fields. It was clarified that the density dependent reduction in the mean longevity and oviposition rate of adult females in G-II and III played a primary role in stabilizing the annual population densities. The annual average of the mean longevity of G-II females (3.9 days) was much shorter than that of G-III ones (7.7 days) and thus the density dependent reduction in the mean longevity induced a more prompt regulatory effect on the oviposition of G-II females compared with G-III ones. As the result, two equilibrium densities of eggs were obtained,e.g., ca 100 and 700 eggs per hill in G-III and IV, respectively. Density dependent decrease in the proportion of mature females in the adult population was especially conspicuous in G-II, and this was closely associated with the density dependent reduction in the mean longevity and fecundity. Thus, the density dependent dispersal (emigration) of the adult females by flight in G-II and III was the most convincing factor in the process of population regulation. The density dependent dispersal of the adult females is effective in avoiding the deleterious effects of nymphal crowding in a breeding habitat unit (a paddy field), and may result in a more even distribution of the population over a continuous habitat units in a locality than otherwise.     

Host-size-dependent sex ratio in a parasitoid wasp

Charnov's host-size model explains parasitoid host-size-dependent sex ratio as an adaptive consequence when there is a differential effect of host size on the offspring fitness of parasitoid males versus females. This article tests the predictions and the assumptions of the host-size model. The parasitoid wasp Pimpla nipponica Uchida (Hymenoptera: Ichneumonidae) laid more female eggs in larger or fresher host pupae when choice among hosts of different sizes or ages was allowed. Then, whether an asymmetrical effect of host size and age on the fitness of females versus males existed in P. nipponica was examined. Larger or fresher host pupae yielded larger wasps. Larger females lived longer, whereas male size did not influence male longevity. Large males mated successfully with relatively large females but failed with small females, whereas small males could mate successfully either with small or with large females. Thus, small-male advantages were found, and this held true even under male–male competition. Ovariole and egg numbers at any one time did not differ among females of different sizes. Larger females attained higher oviposition success and spent less time and energy for oviposition in hosts. Larger females produced more eggs from a single host meal. Taken together, females gained more, and males lost more, by being large. Host size and age thus asymmetrically affected the fitness of offspring males versus females through the relationships between host size or hast age and wasp size, which means the basic assumption of the host-size model was satisfied. Therefore, sex ratio control by P. nipponica in response to host size and age is adaptive. Received: November 13, 1998 / Accepted: January 18, 1999     

Density dependence depends on scale; at larval resource patch and at whole population

Summary A previous study (Tuda and Shimada, 1993) has shown that the equilibrium population size of the azuki bean beetle was lower at 32°C than at 30°C and that this difference was due to a reduced maximum population size of emerged progeny through inside-bean process. In this paper, these results were analyzed further on the scale of the individual bean where interaction among larvae took place. Per-bean numbers of deposited eggs, hatched eggs, and emerged adults have been recorded at seven different parental densities under the two temperature conditions. Three individual-bean-scale process hypotheses that may explain the reduced maximum emergence density on the whole population scale are suggested: (1) a lower maximum emergence per bean at 32°C than at 30°C, if the bean scale and the wholepopulation scale share the same density-dependent pattern in adult emergence, (2) a limited range of hatched egg number per bean at 32°C, resulting from the adult oviposition process outside beans, and (3) different patterns of density-dependent emergence between the two different scales. This study showed that the inside-bean pattern of responses on the bean scale was a simple saturated curve at 30°C, but one with a discontinuous decline at higher hatched egg densities at 32°C. On the contrary, during outside-bean process, the peak number of hatched eggs decreased on this scale as observed on the wholepopulation scale. I discuss why the extracted factor of inside-bean process on the whole-population in the previous study could not be applied to the bean-scale pattern.     

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 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.     

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.     

Relation between plant density and arthropod density in cabbage fields

The relationship between plant density and the abundance of arthropods was examined by planting cabbages in four densities (0.25, 1, 4, or 8 plants per square meter). Four herbivorous species were examined: the small white butterfly Pieris rapae crucivora Boisduval, the diamondback moth Plutella xylostella (Linnaeus), the beet semi-looper Autographa nigrisigna (Walker), and the green peach aphid Myzus persicae (Sulzer). The number of spiders and the number of eggs of syrphid flies were also examined. The number of individuals per unit ground area increased curvilinearly with increasing plant density for most arthropods. To clarify the mechanism causing such a curvilinear increase, the oviposition behavior of Pieris rapae crucivora was examined. The oviposition process of a female was divided into three components: (1) entering the field, (2) alighting on a plant to bend her abdomen, and (3) attaching an egg on the leaf. The first and the second components curvilinearly increased with increasing plant density, while the third component was not influenced by the plant density. The analysis of the flight path of P. rapae crucivora indicated that the curvilinear increase in the frequency of abdominal bending behaviors occurs since a female flies at least about 1.3 m between successive abdominal bending behaviors, irrespective of the plant density, when the plant density is sufficiently high. Received: October 23, 1998 / Accepted: February 15, 1999     

The outbreak mechanisms of the green rice leafhopper,Nephotettix cincticeps Uhler, in northern Japan

Intraspecific regulatory processes keep the population ofNephotettix cincticeps stable at a low density in southern Japan. In northern Japan, however, the yearly population density of the insect fluctuates violently, and large outbreaks occasionally take place. To clarify the difference in the population dynamics between the two regions, we analyzed light-trap and sweep-net sampling records from prefectural and national agricultural experimental stations. The survival rate of the overwintering population decreased with increases in the period of continuous snow cover (PCSC) in the north, and initial population densities in the years of long PCSC were too low for populations to reach equilibrium density by the end of the active breeding season. This made yearly population fluctuations in the north much larger than in the south. The equilibrium density in the north was higher than in the south. The higher equilibrium density presumably permits the higher population density and larger yearly population fluctuations in the north. A major factor responsible for the difference in equilibrium densities between the two regions is the difference in heading dates of the host plant (rice). Qualitative differences among rice plant varieties, and among biotypes ofN. cincticeps, may also be important.     

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.     

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     

Population dynamics ofTelenomus fariai (Hymenoptera: Scelionidae), a parasite of Chagas' disease vectors IX. Larval competition and population size regulation under laboratory conditions

Summary A series of experiments were carried out with the endophagous egg parasiteTelenomus fariai on its hostTriatoma phyllosoma pallidipennis to determine the possible role of intraspecific competition by the parasite progeny in population regulation of the parasite. Eight parasite densities (1, 3, 5, 10, 20, 30, 40, and 50 individuals per vial) were used, and the design of sequentially sacrificed replicates applied. Survivorship curves for each density indicated smaller number of progeny per host at higher densities, and the shapes of the curves suggested a relatively early mortality process.Morris' linear regression technique for determining within-generation density-dependence was used, and the results showed that only larval mortality could be identified as density dependent. The same technique applied within the larval stage proved that only mortality of larvae in their second, third, and fourth day of development were responsible for population regulation. The applicability of the technique, as well as the relevance of the results for natural population, is discussed.