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.     

Eradication of the melon fly,Dacus cucurbitae, from Kume is., Okinawa with the sterile insect release method

Summary The sterile insect release method was applied to eradicate the melon fly,Dacus cucurbitae, from the 58.5 km² island of Kume, in the Okinawa Islands group. Weekly releases of 1 to 1.5 million flies irradiated as pupae with 6–7 kR from a cobalt-60 source did not decrease the wild melon fly population. Releases of 1.5–2 million pupae per week made from September, 1975 to January, 1976 decreased the percent egg-hatch of females caught on Kume Is., but did not decrease the percent infestation significantly. The number of pupae released was increased from February, 1976 to accelerate the eradication process. When the number of pupae released exceeded 3.5 million per week, a rapid increase in the ratio of marked (sterile) to unmarked (wild) flies, a remarkable decrease in percent egg-hatch, and a decrease in percent infestation of fruits were observed. There has been no sign of melon fly infestation in wild cucurbit fruits from October, 1976 to the present time (April, 1977), despite the fact that more than 70,000 fruits were carefully examined. The eradication of the melon fly from Kume Is. was thus achieved by April, 1977, after the release of 264 million sterile fly pupae.     

Binomial sampling plans for estimatingBemisia tabaci populations in cantaloupes

Early season infestations of the sweet potato whitefly,Bemisia tabaci (Gennadius), on cantaloupes,Cucumis melo L., were determined by counts of the number of adults per leaf in fields near Yuma, Arizona. We used these data to develop binomial sampling plans based on the relationship between mean densities of whiteflies per leaf,m, and proportion of leaves infested with more thanI whiteflies,P _I, according to the empirical model lnm=a′+b′ ln[−ln(1−P _I)]. The models were developed for the presence-absence approach (I=0) and for a cutoff value of three whiteflies per leaf (I=3). Four independent data sets were used to evaluate the models. Both methods yielded reliable predictions at low infestation levels, but some of the higherm values were overestimated. As the tentative economic threshold forB. tabaci is three adults per leaf, which corresponds to lowP _I values, results of the binomial sampling were satisfactory for pest management purposes.     

Continuous population estimates forDendroctonus frontalis Zimm. (Coleoptera: Scolytidae) occurring in infestations

Summary Infestations ofDendroctonus frontalis Zimm. are often observed to enlarge continuously by the colonization of new hosts in a pattern similar to a forest fire. This pattern of infestation growth presents unique problems in quantitatively estimating populations ofD. frontalis. Beetle populations on each infested tree in an infestation go through five processes: attack, oviposition, reemergence, survivorship, and emergence. These processes, which have been described mathematically in the literature, each take several days for completion. In order to follow the distribution and abundance ofD. frontalis throughout the course of development of a spot, we need a daily estimate of the number of beetles involved in each process on every tree. Since it is not practical to sample each tree daily, we developed a procedure whereby quantitative estimation procedures for within-tree populations were used in combination with the mathematical models for the life processes to produce a daily record of the number of adults successfully attacking trees, the number of eggs oviposited, the number of beetles reemerging, number of beetles surviving within the trees, and the number of beetles emerging. These daily estimates were then summarized for all trees in the spot for the duration of the infestation. The daily record of populations ofD. frontalis, used with information on infestation geometry, were suggested to be of value in describing and elucidating several important facets of population dynamics including dispersal patterns within infestations, between tree beetle loss (mortality), and time lags among the various population processes. The information reported can be used to develop simulation models of population dynamics or to validate existing models. Texas Agric. Experiment Stn. TA No. 14689.     

Inter- and intra-population variations in diapause attribute of the two-spotted spider mite,Tetranychus urticae Koch, in Japan

Summary Populations of the two-spotted spider mite,Tetranychus urticae Koch collected from various localities and from various host plants in Japan showed wide variations in diapause attribute. Diapause percentages at 18°C/9L15D varied from nearly 100% in the north to 0% in the south-west. At intermediate latitudes the mites showed wide inter-population variations. Populations on herbaceous hosts in vinyl- or glass-houses gave significantly lower incidence of diapause than those on roses and deciduous fruit trees. Presence of winter hosts and better host quality under protected environments seemed to favour non-diapausing mites. The temperature threshold for diapause expression also varied widely among local populations. Northern populations consistently had higher and less variable thresholds than populations at intermediate latitudes with thresholds between 15 and 18°C. Inbred lines derived from a population in Kyoto exhibited a wide variation in diapause percentage at 18°C. These results show that diapause inT. urticae is a quantitative threshold trait and that populations in central Japan consist of a variety of genotypes with different diapause traits. This might provide a genetic source for adaptation to local and temporal variations in environmental conditions.     

Population studies ofBrevicoryne brassicae (L.), its parasites and hyperparasites in England

Summary Studies on populations ofBrevicoryne brassicae (L.), its parasites and hyperparasites were carried out by actual counting in the sprouts field and by sticky and water traps.B. brassicae was found to be attacked by one primary parasite,Diaretiella rapae (McIntosh), which in turn is parasitized byAlloxysta brassicae (Ashm.),Asaphes vulgaris Walker,A. suspensus (Nees),Pachyneuron minutissimum (F?rster) andDendrocerus carpenterii (Curtis). The aphid population in the field was started by immigrant alates which were found flying too early to be synchronized with the sprouts plants. SimilarlyD. rapae was not synchronized with the aphids although many individuals could have been carried into a plot through parasitized immigrant alates, of which less than 30% were found parasitized. Because of high hyperparasitism (especially byA. brassicae)D. rapae was not able to maintain a high rate of parasitism to curb the aphid population growth. The maximum percentage mummies being 27.8%, while the maximum, percentage parasitism being 56.6% recorded only during the early 1974 season (mean=12.9%). The decline of aphid population from September onwards was largely due to the cold weather, Syrphid predation and occasionally fungal attack. The high rate of hyperparasitism byA. brassicae is attributed to its better synchronization withD. rapae. The mean percentage of parasite that emerged from mummies collected during 1973–74 wereD. rapae 31.3%,A. brassicae 64.3%,A. vulgaris andA. suspensus 4.3%,D. carpenterii 0.2% andP. minutissimum 0.1%.     

Comparison of male adult population densities of the oriental and artocarpus fruit flies,Dacus spp. (Diptera: Tephritidae), in two nearby villages in Penang, Malaysia

Summary The populations of native male adult oriental fruit flyDacus dorsalis (Hendel) and artocarpus fruit flyD. umbrosus (F.) in two selected site (BU and SD) were estimated weekly by the capture-recapture technique using live traps baited with methyl eugenol. In BU where many varieties of fruit trees were grown, the estimated population densities ofD. dorsalis were between 980 and 3100 male flies per ha between May and July, 1984. During the same period, in SD where there were fewer number and varieties of fruit trees, the estimated population densities were between 300 and 1000 flies per ha. The estimated population densities ofD. umbrosus over the same period were between 570 and 1290 flies per ha in BU; and between 5 and 95 flies per ha in SD. Of a total 6828 markedD. dorsalis flies released only one fly (released 6 weeks earlier in BU) was caught in a different site.     

Behavior and mortality of the southern stink bugNezara viridula in Hawaii

Summary This paper is concerned with some aspects of the ecology of the southern green stink bug,Nezara viridula, in Hawaii. Dispersal studies on adults showed an asymetrical dispersal pattern. The rate of plant to plant to movement by the adults was 3.3–6.7 feet per day. Mortality studies showed that egg predation by ants,Pheidole megancephala, was as high as 80–90 per cent. Egg parasitism byTelenomus basalis was highest during February, 1965, and adult parasitism byTrichopoda pennipes was highest during July, 1964, to January of the following year. The population ofN. viridula declined during May, 1964, to May, 1965. This decline occurred simultaneously with the decline in the number of eggs produced per female. It has been speculated that this reduction in egg production was caused by parasitism byT. pennipes. Published with the approval of the Director of the Hawaii Agricultural Experiment Station as Technical Paper No. 816. It is with great pleasure that I acknowledge with thanks the help that I received in various ways from the following: Dr. WallaceC. Mitchell, Dr. MervinKamran., Mr HarryKaya and Mr. HitoshiKamasaki.     

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.     

Accumulation of phyllodulcin in sweet-leaf plants ofHydrangea serrata and its neutrality in the defence against a specialist leafmining herbivore

Among wild plants ofHydrangea serrata (Hydrangeaceae) in Japan, there are sweet plants whose leave contain a kind of isocoumarin, phyllodulcin, which happens to be 350 times as sweet as sucrose to the human tongue. In a primary beech forest in Ashu, Kyoto, the spatial distribution of sweet plants and temporal and the spatial distribution of phyllodulcin within and among plants were investigated using a high performance liquid chromatograph. The distribution of sweet plants was confined within a valley and was parapatric with non-sweet plants. A plant's characteristic phyllodulcin accumulation did not change, even when transplanted into the different habitats. The phyllodulcin content of the sweet plants varied greatly among plants, and the population mean peaked in July when the plants flowered. Within a plant, phyllodulcin content was elevated by partial defoliation. We examined the possible effect of phyllodulcin on herbivory by a specialist leafmining herbivore,Antispila hydrangifoliella (Lepidoptera: Heliozelidae). We transplanted sweet and nonsweet plants reciprocally between their original habitats, excluded attacks by parasitoids, and compared performance of the leafminer. Leafminer colonization and larval survivorship on transplanted andin situ plants was not significantly different between sites. The fact that accumulation of phyllodulcin did not augment a defensive function, at least against herbivory by the leafminer, and the sporadic distribution of phyllodulcin-accumulating plants, suggest that the genotypes synthesizing phyllodulcin emerged independently at separate localities by mutation, and that the genotypes are almost adaptively neutral in defence against the specialist herbivore.     

Selection and reproductive success in males of the dragonfly,Orthetrum japonicum (Odonata: Libellulidae)

Reproductive success, copulation success, and mating success were measured for a population of male dragonflies,Orthetrum japonicum. Copulation success explained the greatest variation in reproductive success. The proportion of copulations followed by oviposition was positively correlated with the number of oviposited eggs per mating. Directional selection on four morphological characters was estimated. The effect of selection on correlated traits was comparable to that of direct selection. Directional selection varied between traits and between episodes in a single trait. The probability that the observed directional selection on the four morphological traits was expected under the condition of the selective neutrality of traits was not smaller than 5%.     

The role of the brown-eared bulbulHypsypetes amaurotis as a seed dispersal agent

The role of the brown-eared bulbul,Hypsypetes amaurotis, as a dispersal agent for seeds of fruiting plants was studied by field observations for two years, in parallel with laboratory experiments on seed germination. Bulbuls consumed fruits of 53 species from 24 plant families. The fruits of these plants had similar color and size, and these characteristics were likely to enhance the feeding efficiency of the frugivore. Laboratory experiments on 20 food plant species demonstrated that: (1) no seeds were injured by passing through the bulbul's gut; (2) seeds that had passed through the bulbul's gut were still able to germinate and (3) fruit pulp reduced germination ability. When pulp was removed by passing through bulbul's gut, or by hand, germination was improved. An estimate of the home range of six bulbuls suggested that they may transport seeds for at least 300 m.     

Spatial pattern of pine wilt disease caused byBursaphelenchus xylophilus (Nematoda: Aphelenchoididae) within aPinus thunbergii stand

Summary To understand the mechanism of spread of pine wilt disease caused by the pinewood nematode,Bursaphelenchus xylophilus, which is vectored by a cerambycid,Monochamus alternatus, the spatial distribution of trees weakened by the nematode was examined within aPinus thunbergii stand from June to October for 4 years. The weakened trees were distributed in a clumped pattern in 1980 and 1981, at an early stage of infestation. In many cases, they showed a double-clumped pattern. The degree of aggregation was higher in June or July than after August. They were uniformly distributed in June or July 1982 and in June 1983 whereas they showed a double-clumped pattern after August. The trees were frequently weakened in June or July when they were near the trees weakened during the previous year. At quadrat sizes of more than 25 m², spatial overlapping was pronounced between trees weakened during June–July of the current year and those weakened in the previous year. The seasonal changes in spatial distribution of weakened trees were explained by the interaction amongM. alternatus, B. xylophilus andPinus trees.     

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.     

A granulosis virus, possible biological agent for control ofAdoxophyes orana (Lepidoptera: Tortricidae) in apple orchards

Summary Based on the results of ecological surveys ofAdoxophyes orana and its natural enemies in apple orchards sprayed with the granulosis virus and control plots, we constructed working models to simulate the population dynamics in three different experimental plots; those treated with chemical insecticides, those with granulosis virus, and controls. The number of individuals killed by predators, parasitoids and by miscellaneous mortality factors could be calculated on the assumption of imperfect density relations; that is, relation of the number killed by each factor with the initial number of larvae was represented by a curve with an upper asymptote. We could estimate the proportion of virus infection using curves with upper asymptotes. Rate of increase from pupa to middle instar larvae of the next generation was subject to strong density-effect. Simulation we proposed in this paper suggests a possibility that a single spray of the virus at 1st generation can reduce not only the number ofA. orana larvae in the 2nd and 3rd generations but also the degree of fruits injured by this insect. Spray of chemical insecticide is considered to be ineffective in reducing the pest density and the degree of injury to low levels in subsequent generations, as compared with untreated plot, where the density of the 1st generation larvae is low.     

Effects of gall volume on survival and fecundity of gall-making aphidsAdelges japonicus (Homoptera: Adelgidae)

Summary The number of larvae reaching maturity within the gall ofAdelges japonicus was positively related to gall volume, and the relationship between the number of mature larvae and gall volume did not change with different densities of colonized larvae. The population changes in the bud galls ofA. japonicus were surveyed by collecting the galls, which did not suffer predation or parasitism within the galls, from young artificial plantations ofPicea jezoensis over two years. In the year when the density of colonized larvae was high, they suffered a 42% mortality within the galls, whereas mortality was nearly zero in the low density year. The numbers of larvae per gall were positively correlated with gall volume. The regression lines of the number of colonized larvae on gall volume did not differ significantly in the regression coefficients between the two years, whereas the intercept was significantly higher in the year when the density of colonized larvae was higher. However, different within-gall mortality nullified this difference, and the regression lines of the number of mature larvae on gall volume had no significant difference both in the regression coefficients and the intercepts. This suggests that the number of mature larvae per gall was limited by available resources within the gall which were positively related to gall volume. In 25% of the galls in which mature larvae inhabited, the space within the galls were completely filled by the larvae, indicating that space was one of the limiting resources. Gall volume also affected the number of adults that emerged from the gall and the potential number of their progeny.     

Density-dependent predation of the antLasius niger (Hymenoptera: Formicidae) on two attended aphidsLachnus tropicalis andMyzocallis kuricola (Homoptera: Aphididae)

The antLasius niger was observed collecting honeydew and preying on the two aphid speciesLachnus tropicalis andMyzocallis kuricola on the chestnut treesCastanea crenata. Observation determined how the antL. niger controlled their predation on the aphids in response to the density and honeydew-productivity of the aphids.Lachnus tropicalis was a better honeydew source thanM. kuricola forL. niger in terms of the amount of honeydew collected per unit time by the ants. The number of foraging workers on a tree increased with the number ofL. tropicalis on the tree, but not with the number ofM. kuricola. The density ofL. tropicalis perL. niger worker on a tree had a positive effect on the predation activity ofL. niger on both aphids, whereas the density ofM. kuricola per ant did not have any significant effect. The predation pressure by the ant which increased withL. tropicalis density, however, directed toM. kuricola rather than toL. tropicalis. These facts suggest (1) thatL. niger control their predation activities on aphids with regards to the densities of the attended aphids per worker, and (2) that the ants prey on the aphid species producing less honeydew. The effects of the ant predation on aphids and the importance of these predation effect in antaphids interactions were discussed.     

Competitive habitat utilization in the damselfly,Mnais nawai (Zygoptera: Calopterygidae) coexisting with a related species,Mnais pruinosa

Reproductive behaviors related to habitat utilization were studied in males of the damsefly,Mnais nawai, which has two male forms, territorial orange-winged males (nawai) and non-territorial pale-orange-winged males (sahoi), at the upper part of a mountain stream where they partiallycoexist with a related species,Mnais pruinosa, which also has two male forms, territorial orange-winged males (esakii) and non-territorial hyaline-winged males (strigata). These two species showed parapatric distribution; the lower part of the stream was occupied byM. nawai, and the upper part byM. pruinosa. In the present study, cross-matings occurred between bothMnais species, although normal intraspecific matings occurred more frequently than cross-matings. Territorial males of both species copulated with conspecific females that entered their territory and guarded the ovipositing females, probably to avoid sperm displacement resulting from subsequent copulations. Severe competition for oviposition sites by territorial males even occurred between the two species. On the other hand, non-territorial males of both species have alternative mating strategies (including several tactics such as sneaking, takeover and interception). The possible benefits from conflict among territorial males of both species is discussed.     

Nectar secretion pattern of the dish-shaped flower,Cayratia japonica (Vitaceae), and nectar utilization patterns by insect visitors

Summary We studied the relationship between the diurnal nectar secretion pattern of flowers ofCayratia japonica and insect visiting patterns to these flowers. Flower morphology ofC. japonica changed greatly for about 12 hours after flower-opening and the maximum duration of nectar secretion was 2 days. The nectar volume peaked at 11∶00 and 15∶00, and declined at night and at 13∶00 regardless of time elapsed after flower-opening. The nectar volume at the two peaks was, on average, 0.25 μl on bagged inflorescences and 0.1μl on unbagged inflorescences (both, sugar concentration=60%). The flower secreted nectar compensatory when the nectar was removed. This means that insects consume more nectar than the difference of nectar volume between bagged and unbagged flowers. Apis cerana is a primary visitor of this flower, and was the only species for which we confirmed pollen on the body, among many species of flower visiting insects to this flower.Apis cerana visited intensively at the two peaks of nectar secretion. Visits of the other insects were rather constant or intensive only when there was no nectar secretion. Thus flowers ofC. japonica with morphologically unprotected nectaries may increase likelihood that their nectar is used by certain pollinators, by controlling the nectar secretion time in day. In this study the pattern of nectar secretion allowedA. cerana maximum harvest of nectar. Contribution to the ecological and bioeconomoical studies of the pollinator community in Kyoto I.     

Pollination and factors limiting fruit set of chasmogamous flowers of an amphicapric annual,Polygonum thunbergii (Polygonaceae)

Summary We observed phenology and insect visitors of chasmogamous flowers of an amphicarpic annual,Polygonum thunbergii (Polygonaceae) in Kyoto, Japan, and clarified limiting factors for fruit set by bagging and hand-pollination experiments. Flowering season was one month from late September. Flowers were visited by various insect groups (total 30 families and 64 species) and effective pollinators were lower bees (Halictidae) and wasps (Vespidae) in the early flowering season, and middle-sized flies (Syrphidae and Calliphoridae) in the later season. Nectar was secreted at a constant rate (0.013mg sugar/hour) throughout a day and >90% of it was consumed by insects. The average number of flowers per ramet was 122, of which 95% were pollinated in the natural conditions. Bagging experiments showed that 47% of flowers were self-pollinated even under no pollinator visits. Despite of a high probability of cross pollination, the probability of fruit set within the ramet was 0.30 due to resource limitation. We discussed possible adaptive significance of cleistogamous flowers under the condition that seeds could be produced by chasmogamous flowers through self-pollination even under pollinator limited conditions.