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.     

A marking-and-recapture analysis of the adult population of a phytophagous lady-beetle,Epilachna sparsa orientalis

Summary A marking-and-recapture study of a population of overwintered adults ofEpilachna sparsa orientalis was carried out in a small farm containing potatoes and egg-plants. The experiment involved a single release and five recapture samplings during May–June. The data were analysed byLeslie's re-recapture method. Assumption of randomness of sampling proved to be well satisfied. Survival rate of the overwintered adult population was maintained as high as 90 per cent per week or more until early June, and then it declined rapidly towards the end of June. Initially overwintered adults exclusively lived on potato plants, but half of the population alive in mid June migrated to nearby egg-plants. Emigration out of the experimental field, however, seemed not to be occurring. Recapture data of marked individuals in the potato field indicate that majority of the individuals stayed within a radius of 3 m, at least for 2 weeks. Similar data taken from another field also suggest that their mobility was far restricted as compared with random movement within the field. Some comparisons were made between the population behaviours of overwintered adults ofE. sparsa orientalis andE. vigintioctomaculata. Contribution from the Entomological Laboratory, Kyoto University, No. 371.     

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.     

Some beneficial effects of aggregation in young larvae ofPryeria sinica moore (Lepidoptera: Zygaenidae)

Summary The effects of group size on the survival and development of young larvae ofPryeria sinica Moore were investigated by laboratory and field experiments. Under laboratory conditions, about 20% of isolated larvae died of unsuccessful feeding in the first instar, however, larvae survived successfully in aggregations of four or more individuals. In the field, larvae emerge in early spring and wait for new leaves to open before feeding. In this period, the larger the group size of hatchlings the survival rate became higher. The nest-web spun by hatchlings was considered to play an important role in protecting them from desiccation. In the period that larvae began to feed on leaves, more than 36 larvae are necessary to aggregate for the successful establishment of feeding groups. The nest-web played an important role also in the establishment of feeding group. However, the natural group size of the first instar larvae was larger than the minimum group size to spin a sound nest-web in the field experiment. On the other hand, in later stage, larvae in a large group did not have an excess advantage in survival or developmental rate over larvae in a small group. It was found that the experiments on survival and developmental rates could not explain the reason that this species maintain large compact groups in the most part of larval period. The study was partially granted by the Ministry of Education (No. 374205).     

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.     

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.     

The effect of prey size and prey density on the functional response, survival, growth and development of a predatory pentatomid bug,Podisus maculiventris say

Summary Two experiments on the nymphal predation ofPodisus maculiventris were conducted usingSpodoptera litura larvae as prey. First experiment: The predator nymphs divided into three groups were reared individually from second instar to adult in a small vessel. Each nymph in the groups 1, 2 and 3 was allowed to attack the serially growing larvae (these were supplied at the rate of one per day) from 3-, 5- and 7-day old after hatching, respectively. The first prey used for the group 1 was so small that it was not only insufficient to satiate the predator but also was difficult to be searched out. But these disadvantages were soon recuperated due to the rapid growth of the prey and all nymphs could survive to adults. The survival rate of third and fourth instar nymphs in the group 3 was severely affected by vigorous counterattack of older prey larvae. Second experiment: The predator nymphs were individually reared either in a small vessel or in a large one at various rates of food supply (the prey larvae of 7-day old were used). The functional response curves obtained for each instar of the predator took a saturation type within a certain range of the prey density. The saturation level specific to each instar was generally higher for the predator reared in the large vessel than in the small one. The functional response of fourth and fifth instar nymphs was accelerated at a high prey density,viz. 16 larvae per vessel. Even at the low rate of food supply,viz. one larva per day per predator, the predator nymphs could survive to adults, but the size of resultant adults were abnormally small.     

Further improvements to a method for analysing stage-frequency data

Summary An iterative procedure for correcting stage-frequency data is described to allow for situations where the period during which a population is sampled begins after some individuals have entered stage 2 or ends before all individuals are dead. The reason for correcting data in this way is to enableKiritani andNakasuji's method for estimating stage-specific survival rates, with extensions proposed byManly (1976, 1977), to be used to analyse the data. The proposed procedure is illustrated on data obtained by sampling a population of the grasshopperChorthippus brunneus passing through four instar stages to reach the adult stage.     

Pit-relocation of antlion larvae in relation to their density

Of the biotic factors potentially affecting the foraging behavior of pit-building antlion larvae,Myrmeleon bore Tjeder, we focussed on the density effects and examined experimentally influences of larval density on pit construction behavior, spatial distribution and pit relocation rate. Moreover the mechanism of mutual interference among larvae was examined.

Time-specific life tables for the pea aphid,Acyrthosiphon pisum (Harris ), on alfalfa

Summary Time-specific life tables were constructed for three pea aphid,Acyrthosiphon pisum (Harris) (Homoptera: Aphididae), populations using a modification ofHughes' analytical procedure. All populations were studied on second-growth alfalfa (mid-June to mid-July) in south central Wisconsin; data for two populations were collected during 1980, and data for the third population were collected during 1982. The intrinsic rate of increase (r _m) estimated on a physiological time (day-degree) scale under field conditions but in the absence of natural enemies, provided a reliable estimate of potential population growth rate and was used in preference toHughes' approach of estimating potential population growth rates directly from stage structure data. Emigration by adult alatae and fungal disease were the major sources ofA. pisum mortality in each of the three populations studied. These factors were most important because of their impact on reducing birth rates within the local population. Parasitism was never greater than 9 percent. Mortality attributable to predation ranged from 0.0 to about 30.0%; however, even at the highest predator densitiesA. pisum populations increased exponentially.     

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.     

Demographic attributes of an introduced herbivorous lady beetle

Demographic attributes of the adults of an introduced herbivorous lady beetleEpilachna niponica (Coleoptera: Coccinellidae) were investigated from 1975 to 1981 in the Botanical Garden of Kyoto University. Population growth rate varied from 4.8 to 16.8 throughout the study period. Fecundity and mortality in the late larval period contributed most to annual changes in the population growth rate. Population growth rate was negatively correlated with the density of overwintering adults. Adult survival from emergence to the reproductive season, which varied from 0.03 to 0.36 throughout the study, was almost completely determined by survival during the pre-hibernation period. Adult survival to the preproductive season changed in a size- and sex-dependent manner. Larger adults survived better than smaller individuals; male-biased mortality occurred from adult emergence to the reproductive age. Severe intraspecific competition among late instar larvae due to host plant defoliation produced a higher proportion of small-sized adults, resulting in lower adult survival to hibernation. The introduced population had a higher population growth rate and a lower adult survival to the reproductive season than the source population.     

A simulation study of three methods for estimating selective values and survival rates from recapture data

Summary The methods ofManly (1973),Manly (1975) andManly (1977) for estimating survival rates and relative survival rates from recapture data have been compared by computer simulation. In the simulations batches of two types of animal were “released” at one point in “time” and recapture samples were taken at “daily” intervals from then on. The various methods of estimation were then used to estimate, the daily survival rates of type 1 and type 2 animals, and also the survival rate of the type 2 animals relative to the type 1 animals. Simulation experiments were designed to examine (a) the bias in estimates, (b) the relative precision of different methods of estimation, (c) the validity of confidence intervals for true parameter values, and (d) the effect on estimates of the failure of certain assumptions.     

On the nature of errors involved in estimating stage-specific survival rates by Southwood's method for a population with overlapping stages

Summary This paper has examined the effect of within-stage mortality on the estimation of stage-specific survival rates bySouthwood's (1978, p. 358) method. As pointed out bySouthwood, both the severity and timing of mortality affect the mean duration of a life stage, and consequently the estimate of the number of individuals entering that stage. Knowledge of the form of the survivorship curve permits correction of the estimate under certain circumstances. The use ofSouthwood's method with two overlapping stages having different rates and patterns of mortality leads to complex errors in the estimation of survival for the first stage. The nature of these errors is examined analytically and via a simulation model.Southwood's method is fairly robust, with moderate differences in mortality rates leading to acceptable errors in estimating survival for the first stage. When both the rate and pattern of mortality in both life stages are the same, then the survival estimate is made without error. Precise estimates of stage-specific survival will not usually be possible withSouthwood's method because of the errors introduced by the very parameters being measured. Direct measurement of mortality rates and survivorship patterns (seeSouthwood, 1978, p. 309) is strongly advised, at least in preliminary work.     

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.     

A population study ofStenocranus minutus (Fab. ) (Hemiptera: Delphacidae)

Summary The seasonal and annual fluctuations in a population of the delphacid,Stenocranus minutus (Fab.) onDactylis glomerata L. were studied from March 1968 to September 1970. The study involved the comparison of several sampling methods to estimate the egg, nymphal and adult numbers and an investigation into causes of mortality within each stage. A population budget was made for the years 1969 and 1970.     

Effect of avian predation on survival of leaf-rolling lepidopterous larvae

Manipulative field experiments were performed in a northern Japanese deciduous forest to determine the role of avian predation in the survival of three leaf-rolling lepidopterous larvae (Homonopsis foedenratana, Archips viola, and Zeiraphera corpulentana) on Japanese lilac (Syringa reticulata) trees. When all leaf rolls on the lilacs were experimentally removed, the visitation rate of birds declined, indicating that the leaf rolls served as a visual cue for the birds. In a second experiment in which birds were excluded from the lilac trees, the survival of H. foedenratana larvae was improved, whereas the mortality of the other two Lepidoptera stayed the same. This is probably the result of differences in the shape of leaf rolls; larvae of H. foedenratana alter the leaf form to a greater extent, which may facilitate the detection of the rolls by birds. The high predation rate on the tight leaf rolls of H. foedenratana may be offset by enhanced resistance against other predators and better food quality for the enclosed larvae. Received: January 30, 1998 / Accepted: December 2, 1998     

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     

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.