Genetic variation of density responses in relation to wing polymorphism in the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae)

Responses to nymphal density in the determination of wing form were compared between the offspring from brachypter x brachypter crosses and those from macropter x macropter crosses of the oriental chinch bug, Cavelerius saccharivorus. In the offspring from crosses between macropters, there was a strong tendency for macropters to increase to a rather high level with increasing nymphal density in both sexes. In contrast to this, in the offspring from crosses between brachypters, the appearance rate of macropters attained a maximum value in moderately crowded conditions and conversely decreased to a lower level in more crowded conditions in both sexes. Thus density response patterns concerning the determination of wing form were quite different between the offspring from different crosses in the wing form, indicating that there is a genetic basis underlying wing polymorphism in this species. As for the body size of emerged adults, macropters tended to be larger than brachypters in the same crowded condition. Moreover, the rate of decrease of body size with nymphal density was lower in the offspring from crosses between macropters than in the offspring from crosses between brachypters. This indicated that the former offspring are more tolerant of nymphal crowding than the latter. The difference in such a tolerance against nymphal crowding between the offspring from different crosses was considered to be related to the difference in the appearance of macropters in the crowded conditions between them.     

Reproductive properties of the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae), in relation to its wing polymorphism

Reproductive properties of the oriental chinch bug, Cavelerius saccharivorus, were investigated for three types of female adults: brachypters produced in both high and low density populations and macropters produced in high density populations. With respect to egg-laying potential in terms of clutch size, clutch number, and total fecundity, macropters were superior to brachypters produced in the same high density population, although inferior to brachypters produced in the low density population. The mean fecundity of female adults in each type was positively correlated with mean body length. Macropterous females markedly prolonged their initiation of both copulation and oviposition and had a much longer life span compared with both types of brachypters. However, there was no fundamental difference in reproductive schedules between the two types of brachypters.     

Genetic correlation of wing polymorphism between females and males in the oriental chinch bug,Cavelerius saccharivorus Okajima (Heteroptera: Lygaeidae)

Genetic correlations of the wing form and the relative wing length between females and males were estimated in the oriental chinch bug, Cavelerius saccharivorus, by calculating the correlation between the sexes of the proportion macropterous or the mean relative wing length in full-sib families obtained from different wing forms of parents emerged in a high density population. There was a significantly positive genetic correlation between the sexes in both the proportion macropterous and the mean relative wing length. However, the appearance rate of macropters tended to be much lower in males than in females under the rearing conditions which promote the appearance of macropters. This was evident especially in the offspring of brachypterous parents. These indicated that in C. saccharivorus the wing polymorphism of males is not a simple result of the genetic correlation of wing morphology between the two sexes. It was considered that both of the female and male fitness advantages to wing reduction, as well as the genetic correlation between the sexes, would influence the evolution of wing polymorphism in this species.     

Population studies ofCavelerius saccharivorus Okajima (Heteroptera: Lygaeidae): Adult dispersal in relation to the density

Summary Two adjacent stations (H and L) were set up to study movement ofC. saccharivorus adults in sugarcane fields. At the beginning of the study the density ofC. saccharivorus including all stages of development was quite different between the two. The density of the first generation adult on Station H was about 5 times that on Station L. The number ofC. saccharivorus on both the stations became almost the same one month after the beginning of the study. At the beginning of the study macropterous adults were more numerous in Station H than in Station L. However percentage of macropterous adults on Station L increased after one month whereas that on Station H declined. About 2,000 marked adults were released on each station during the early period of the emergence of the first generation adult. Marked insects were recaptured on both the stations one month after the release. The adults released on the dense population (H) tended to disperse more actively than those on the scarce population (L). Marked macropterous adults moved more actively than brachypterous ones. The density related dispersal of adults was considered to be an important factor to regulate the population density.     

Population studies ofCavelerius saccharivorus Okajima (Heteroptera: Lygaeidae): Adult dispersal in relation to the density

Two adjacent stations (H and L) were set up to study movement of C. saccharivorus adults in sugarcane fields. At the beginning of the study the density of C. saccharivorus including all stages of development was quite different between the two. The density of the first generation adult on Station H was about 5 times that on Station L. The number of C. saccharivorus on both the stations became almost the same one month after the beginning of the study. At the beginning of the study macropterous adults were more numerous in Station H than in Station L. However percentage of macropterous adults on Station L increased after one month whereas that on Station H declined. About 2,000 marked adults were released on each station during the early period of the emergence of the first generation adult. Marked insects were recaptured on both the stations one month after the release. The adults released on the dense population (H) tended to disperse more actively than those on the scarce population (L). Marked macropterous adults moved more actively than brachypterous ones. The density related dispersal of adults was considered to be an important factor to regulate the population density.     

Population studies ofCavelerius saccharivorus Okajima (Heteroptera: Lygaeidae): A few findings on population interchange

The change in population density of Cavelerius saccharivorus was studied in the sugar cane field and the Miscanthus habitats around it. This species spent 2 (or partly 3) generations a year. It was suggested that the density rose in the sugar cane field where the density had been low enough, when the population density increased in the nearby sugar cane fields. Likely, the density in the Miscanthus habitat increased with the density in the nearby sugar cane field. It seems that these are mainly due to flying movement of adults, and that the movement takes place in every season. It was found through the investigation in the sugar cane field that the percentage of long-winged adults increased with the population density. These adults seemed to emmigrate to the nearby sugar cane fields and Miscanthus habitats. The remarkable increase in the percentage of long-winged adults in the sugar cane field was probably due to the immigration into there. In the Miscanthus habitat the adult population was consisted mainly of long-winged ones. Through all the observations, it was suggested that the adult movement was associated with the population regulation and performed chiefly by long-winged adults.     

A male fitness advantage to wing reduction in the oriental chinch bug,Cavelerius saccharivorus okajima (heteroptera: Lygaeidae)

A male fitness advantage to wing reduction was investigated for the oriental chinch bug, Cavelerius saccharivorus, which is wing dimorphic. Field surveys for the frequency of matings between wing morphs showed that the mating probability was much higher in brachypterous males than in macropterous males in the early breeding season. Brachypterous males copulated with not only brachypterous females but also macropterous females in this season. This led to a considerable rate (30%) of insemination of macropterous females just before emigration even in the early breeding season. A rearing experiment for the pre-reproductive period of females revealed that females mated with brachypterous males copulated and oviposited earlier than females mated with macropterous males. These results indicate that there is a fitness advantage to wing reduction in the males of C. saccharivorus in terms of the earlier sexual maturation.     

Reproduction and egg diapause of the oriental chinch bug,Cavelerius saccharivorus okajima (Heteroptera: Lygaeidae), in the subtropical winter season in relation to its wing polymorphism

Reproduction and egg diapause of the oriental chinch bug, Cavelerius saccharivorus, in the subtropical winter season were investigated in relation to its wing polymorphism. Macropterous females collected from the southern part of Okinawa Is. in the autumn season delayed their oviposition and were less fecund early in the adult life period, but survived much longer than brachypterous females collected from the same locality. The total fecundity was not significantly different between wing morphs. The diapause of eggs laid by brachypters tended to be terminated more easily at a high temperature than that of eggs laid by macropters. This indicated that the eggs laid by macropters were more intense in their diapause than those laid by brachypters. However, irrespective of the parental wing form, diapause showed considerable variation in its intensity within and between clutches.     

Phenological studies on the wing dimorphism of a semi-aquatic bug,Microvelia douglasi (Heteroptera: Veliidae)

Seasonal changes in the percentage of macropterous individuals in natural population of Microvelia douglasi in permanent habitats were investigated in Shimane, western part of Japan. Macropterous forms of 5th instar nymphs occurred most frequently from late June to early July. However, the seasonal appearance of macropterous adults differed between habitats and years. There were significant effects of density, temperature, photoperiod and food availability on the percentage of macropters under laboratory conditions. The percentage of macropterous forms generally increased with density, but the mode of the response was modified by temperature, photoperiod and food availability. When insects were reared under long day photperiod (16L8D) with abundant food (0.5–1.0 g/container/day), the percentage of macroptery varied most among the different density treatments at the intermediate temperature (24°C) and the difference was reduced at higher temperature (28° and 32°C). Effects of temperature were small under the short day photoperiod (12L12D). Under 12L12D photoperiod sensitivity of density responses were higher at 32°C and lower at 24°C than those under 15L8D photoperiod. When the insects were supplied with food every other day, the density response became obscure. Ecological significance of environmental responses in the wing morph determination was discussed in relation to the heterogeneous environment of water surface.     

Lawn parameters influencing abundance and distribution of the hairy chinch bug (Hemiptera: Lygaeidae)

Management of lawns that promotes conditions detrimental to the development of insect pests may represent a valuable environmentally benign turfgrass management strategy. In the cool-humid region of Quebec, Canada, we investigated 45 lawns infested with hairy chinch bug, Blissus leucopterus hirtus Montandon, to identify lawn parameters related to its distribution and abundance. Kentucky bluegrass, creeping bentgrass, and perennial ryegrass, respectively, accounted for 55.8, 19.6, and 9.3% of the grass species. Chinch bug population density was associated positively with abundance of perennial ryegrass, whereas it was marginally negatively related with the abundance of creeping bentgrass. An index of the severity of chinch bug infestation was obtained for each lawn by combining estimates of number of infested patches per lawn, average size of the patches, and chinch bug number per patch. The index was associated positively with abundance of Kentucky bluegrass and perennial ryegrass. There was evidence that abundance of creeping bentgrass was associated negatively with the number of infested patches per lawn, area of the patches, and number of chinch bugs within those patches. The number of infested patches increased, whereas patch area and chinch bug number per patch tended to decrease, when broad-leaf weeds were more abundant on a lawn. No significant relationship was found between thatch thickness and patterns of chinch bug abundance and distribution. These results suggest that management of lawns to respectively increase and decrease abundance of creeping bentgrass and perennial ryegrass could facilitate control of hairy chinch bug populations in cool-humid regions.     

Seasonal changes in wing dimorphism of the lygaeid bug Dimorphopterus japonicus (Heteroptera: Lygaeidae) in relation to environmental factors

The lygaeid bug Dimorphopterus japonicus Hidaka, which mainly feeds on a eulalia, Miscanthus sinensis, shows marked wing dimorphism of brachyptery and macroptery. Its production of macropters is stimulated in rearing conditions such as high temperature, long photoperiod and crowding during the nymphal stage. In this study, we investigated the seasonal prevalence in occurrence and the seasonal change in incidence of macroptery in D. japonicus for field populations in Okayama, western Japan. The results demonstrated that nymphal density was a key factor in determining the proportion of macropters, indicating an escape strategy from crowded populations. The field surveys also revealed that this bug has a univoltine life cycle in Okayama. There was a seasonal change in the incidence of macroptery in the new generation. The combined effects of temperature and photoperiod on wing‐form determination explained this seasonality trait.     

Evidence of Environmental and Vertical Transmission of Burkholderia Symbionts in the Oriental Chinch Bug,Cavelerius saccharivorus (Heteroptera: Blissidae)

The vertical transmission of symbiotic microorganisms is omnipresent in insects, while the evolutionary process remains totally unclear. The oriental chinch bug, Cavelerius saccharivorus (Heteroptera: Blissidae), is a serious sugarcane pest, in which symbiotic bacteria densely populate the lumen of the numerous tubule-like midgut crypts that the chinch bug develops. Cloning and sequence analyses of the 16S rRNA genes revealed that the crypts were dominated by a specific group of bacteria belonging to the genus Burkholderia of the Betaproteobacteria. The Burkholderia sequences were distributed into three distinct clades: the Burkholderia cepacia complex (BCC), the plant-associated beneficial and environmental (PBE) group, and the stinkbug-associated beneficial and environmental group (SBE). Diagnostic PCR revealed that only one of the three groups of Burkholderia was present in ∼89% of the chinch bug field populations tested, while infections with multiple Burkholderia groups within one insect were observed in only ∼10%. Deep sequencing of the 16S rRNA gene confirmed that the Burkholderia bacteria specifically colonized the crypts and were dominated by one of three Burkholderia groups. The lack of phylogenetic congruence between the symbiont and the host population strongly suggested host-symbiont promiscuity, which is probably caused by environmental acquisition of the symbionts by some hosts. Meanwhile, inspections of eggs and hatchlings by diagnostic PCR and egg surface sterilization demonstrated that almost 30% of the hatchlings vertically acquire symbiotic Burkholderia via symbiont-contaminated egg surfaces. The mixed strategy of symbiont transmission found in the oriental chinch bug might be an intermediate stage in evolution from environmental acquisition to strict vertical transmission in insects.     

Induction of diapause in a migratory seed bug,Neacoryphus bicrucis (Say) (Heteroptera: Lygaeidae)

Summary Neacoryphus bicrucis hibernates in the adult stage and exhibits a facultative reproductive diapause. Constant photoperiods of the durations encountered in the field do not evoke the diapause response but Shortening of the photoperiods does, providing the decrease in daylength is of a certain length and occurs within a certain photoperiodic interval. The sensitive stages are larval instars II to V. The percentage of diapausing bugs increases as temperature is lowered.It is suggested that the response to daylength shortening in N. bicrucis is an adaptation to ensure seasonal synchronization in a highly migratory species where successive generations may breed at different latitudes.     

Pharmacological observations on body fluids from the milkweed bug,Oncopeltus fasciatus (Dallas) (Heteroptera: Lygaeidae)

A report is given on the pharmacological activity of body fluids from milkweed bugs maintained on a diet of sunflower seeds, together with observations on the toxin accumulating system of the adult insect.     

Mating experience weakens starvation tolerance in the seed bug Togo hemipterus (Heteroptera: Lygaeidae)

Organisms are exposed to various stresses caused by environmental fluctuations. One of the most common stresses is the shortage of food. Individuals of many species must survive periods of starvation. There appears to be a trade‐off between reproduction and survival. When residual reproductive value declines for an individual, life‐history theory predicts an increase in current reproductive investment. Current reproductive investment differs between virgin and mated individuals. It is likely that mating experience influences starvation tolerance. However, few studies have investigated sex differences in the effect of mating experience on starvation tolerance or clarified the causes of reductions in starvation tolerance in both sexes. In the present study, these questions are investigated using the seed bug Togo hemipterus (Heteroptera: Lygaeidae).The results of the present study demonstrate that mating is costly for both sexes. Mated males show very short survival times and a daily reduction in weight, and daily energy expenditures are significantly greater in mated males than in virgin males. It is possible that starvation increases the mating effort of males, such as behavioural activities and the amount of time spent searching for females. A trade‐off between survival duration and lifetime fecundity is found in virgin females. However, there is no trade‐off in mated females, which have very short survival times. Whether male seminal substances contribute to the short survival times of mated females is considered. This is the first report demonstrating the influence of sex and mating experience on starvation tolerance. Sex‐specific causes for reductions in starvation tolerance are discussed.     

Mating aggregation of a stink bug,Megacopta punctissimum (Montandon) (Heteroptera: Plataspidae)

Summary Adults of a stink bug,Megacopta punctissimum, form aggregations on stems of the bush-clover,Lespedeza crytobotria, in spring. They do not oviposit there, but feed and mate within aggregations. Oviposition is made on other leguminous plants such as the kudzu-vine,Pueraria lobata. Mean size of aggregations (groups of two or more individuals which sit on a stem at distance less than 3 cm from their nearest neighbors) consisted of 4.4 individuals. The sex ratios within aggregations were similar to overall sex ratio of the population including solitary individuals (0.58). More than 50% of females found in aggregations were copulating. Percentage of femalesin copula in larger aggregations was higher than that in smaller aggregations, and this difference was considered to be caused by the higher chance of sexual enconters in larger aggregations. It was suggested that the aggregation of this species on the bush-clover is not a ‘harem’ (a male monopolizes a group of females reported in some other stink bugs) but is mating aggregation to raise the chance of mating. This work was partially supported by Grant-in-Aid (No. 56480039) from the Ministry of Education, Science and Culture.     

Seasonal wing length dimorphism in a tropical seed bug: ecological significance of the short-winged form

Summary Short-winged forms of the bug, Jadera aeola, were discovered during the late dry season at a study site on Barro Colorado Island, Panama. They were smaller than long-winged adults not only in elytron and hindwing sizes but also in head, thorax, rostrum and hind femur dimentions. When newly emerged adults were kept under natural photothermal conditions, short-winged females started ovipositing earlier, produced more eggs, and died earlier than did long-winged females. This was partly due to the fact that long-winged adults usually enter diapause after adult energence (Tanaka et al. 1987b). Egg size was another factor contributing to the high egg production in the short-winged form; it was significantly smaller in the short-winged form than that in the long-winged form. Wing form was apparently environmentally induced because all off-spring of short-winged adults became long-winged. No short-winged form was obtained in the laboratory when nymphs were reared on different species of seeds at different photoperiods. Although the factor(s) responsible for determination of the wing form remains unknown, the ecological significance of this short-winged form seems clear, i.e. exploitation of the food resource left in the late dry season. Wing length dimorphism in J. aeola could be a response to unpredictable availability of the food resource determined by seed production of the host plants (Sapindaceae) and by the timing of the onset of a wet season.     

Nosema blissi sp. n. (Microsporida: Nosematidae), a pathogen of the chinch bug,Blissus leucopterus hirtus (Hemiptera: Lygaeidae)

Nosema blissi sp. n. is described from the Malpighian tubules of adults of Blissus leucopterus hirtus. Spores measured 6.5 ± 0.3 × 2.5 ± 0.1 μm in Giemsa-stained preparations. The polar filament lay in 37 to 40 coils, arranged in a single layer in the posterior portion of the spore, and in several layers in the anterior portion.     

Life cycle and morphology of Anisops sardeus Herrich-Schaeffer, 1849 (Heteroptera: Notonectidae)

The life cycle of Anisops sardeus was studied by rearing individuals from egg to adult stage in laboratory conditions at a water temperature of 23.2 ± 1.4 °C during the wet season (May-June) and 19 ± 1.8 °C during the dry season (December-February). The incubation period averaged 8 ± 0.8 and 11.5 ± 1.7 days during the wet and dry seasons, respectively. Duration of the five instars averaged 3.4 ± 0.5, 4.4 ± 0.5, 4.8 ± 0.8, 5 ± 0.7, and 6.9 ± 0.7 days, respectively during the wet season, and 4.9 ± 0.7, 6.5 ± 1.1, 7.5 ± 1.1, 8.1 ± 0.7, and 9.4 ± 1.1 days, respectively during the dry season. Total developmental time averaged 32.5 ± 2 and 47.9 ± 2.8 days in the wet and dry seasons, respectively. The average period of incubation and developmental time of the five instars were shorter in the wet season as compared to those in the dry season. But individuals were larger in the dry season. The variations in morphometric ratios of different characteristic features of laboratory reared specimens among different developmental stages in both the wet and dry seasons, and field collected specimens in the wet season were highly significant as revealed by one-way MANOVA (F = 95.45, p < 0.001; F = 124.38, p < 0.001; F = 5022.85, p < 0.001, respectively). Five instars are described in detail with emphasis on 29 morphometric ratios. This study discerned six morphometric ratios such as length of wing pad/ width of wing pad (WL/WW), length of wing pad/body length (WL/BL), length of head/length of body (HL/BL), length of meso femur/length of meso tibia (FE2L/TI2L), synthlipsis/width of head (S/HW), and vertex/synthlipsis (V/S) which can be used for discriminating instars I-V.