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

Summary 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)

Summary 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 inC. 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.     

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

Summary 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.     

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

Summary 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 ofC. saccharivorus in terms of the earlier sexual maturation.     

Ecological significance of the wing polymorphism of the oriental chinch bug,Cavelerius saccharivorusOkajima (Heteroptera: Lygaeidae)

Summary The wing polymorphism of the oriental chinch bug,Cavelerius saccharivorus, was studied in relation to its reproductive strategy. The frequency distribution of wing form in terms of the relative wing length (R.W.L.) changed with increase in the population density from a single modality biased toward the brachypterous form to bimodality with both macropterous and more extreme brachypterous forms. Since some evidences in the field showed that fliers are limited to macropters, such a bimodality of wing form indicated thatC. saccharivorus employs a mixed strategy with some portion of adults staying on in the same habitat, whereas the rest disperse. Moreover, macropters were larger in body size than brachypters emerging in high density populations, although smaller than brachypters emerging in low density populations. Larger body size as well as the delay of ovarian maturation in macropters was considered to be a beneficial trait for migratory behaviour.     

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): A few findings on population interchange

Summary The change in population density ofCavelerius saccharivorus was studied in the sugar cane field and theMiscanthus 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 theMiscanthus 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 andMiscanthus 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 theMiscanthus 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.     

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

Summary Seasonal changes in the percentage of macropterous individuals in natural population ofMicrovelia 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.     

Spatio-temporal variation of natural and sexual selection in breeding populations of the coreid bug,Colpula lativentris (Heteroptera: Coreidae)

Spatio-temporal variations of lifetime reproductive succes (LRS) of both male and female individuals of a coreid bugColpula lativentris were measured and analyzed using the multiple regression method of Arnold and Wade (1984a, b). The standardized variance of LRS was larger in males than that in females as males often to secure mates for a long period whereas females could easily find mates and oviposit simply dependent on ovarial maturation. LRS was partitioned into 4 consecutive fitness components: (1) reproductive lifespan, (2) copulating efficiency, (3) guarding efficiency (for males) or oviposition efficiency (for females), and (4) number of eggs per clutch. In males copulating efficiency was the largest determining factor of LRS, whereas in females reproductive lifespan was the most important factor. Such tendencies were stable on both a yearly and local basis. Patterns of relative contribution of natural selection (reproductive lifespan and number of eggs per clutch) and sexual selection (copulating efficiency and guarding or oviposition efficiency) to LRS were clearly different between males and females. This sexual difference is, at least to some extent, thought to be brought about by sexual selection among males for mating opportunity, though no physical fight was observed among males. Directional selection on body length was found only in relation to the clutch size of females because large females tended to lay larger clutches. No significant directional selection was found in other fitness components.     

Studies on the adult polymorphism in the southern green stink bug,Nezara viridula (hemiptera: Pentatomidae)

Summary Polymorphism in adult colour pattern ofNezara viridula is determined on the genetic basis. The basic colour patterns of adult are classified into four types, i. e. G, O, R and F. No appreciable differences between these types were observed in respect to various physiological traits of nymphs and adults, except that type G seems to be superior in reproductive ability but to be inferior in ability of surviving winters at least to types R and F. Inter-generation changes in percentage frequency of G type were examined from 1959 to 1967 covering more than 30 generations. Percentage frequencies of G types prior to the severe winter of 1962–3 fluctuated greatly around a mean of 87.9%, while they did to a lesser extent than before with a mean of 85.0% after the winter. Unexpected high percentages of G type were recorded frequently in summer generations, viz. 1st and 2nd. On the other hand, the relative frequencies of G and O types decreased after hibernation in contrast to the increases in those of F and R. This sort of changes in genetic composition related to the winter of 1962–3 was observed in several populations segregated from each other. This seasonal alternation of selective activity in the environments is considered to be responsible for retention of the polymorphism. Alternative possible causes, i. e. difference in habitat preference, non-random mating and selective predation by predators among polymorphs, may safely be rejected as irrelevant to the mechanism in maintaining polymorphism. The polymorphism of this insect seems to be in a transient state rather than balanced one contributing little to population regulation, but the persistence ofN. viridula in the periphery range may be assisted by retention of the polymorphism.     

Receptivity of female remating and sperm number in the sperm storage organ in the bean bug,Riptortus clavatus (Heteroptera: Alydidae)

This study examines the relationship between the number of sperm in the seminal receptacle (spermatheca) and the receptivity of female remating in the bean bugRiptortus clavatus Thunberg. On the 21 st day after the first mating when receptivity to remating was > 70%, females receptive to remating had significantly fewer sperm ( < 40 on average) in the spermathecae than females reluctant to do (about 150 on average). However, averages of the number of eggs laid by receptive and reluctant females within 21 days were almost same. The proportion of fertilized eggs for receptive females at 15–21 days after copulation was significantly lower than that for reluctant females. Spermatozoa transferred from a male to a female’s spermatheca were detected 5 min after copulation and then increased continuously to about 500 with the first hour. When copulation durations were manipulated artificially, the shorter the copulation period (=females had less sperm in their spermathecae), the higher the remating rate became. Females may perceive the number of sperm in their seminal receptacles and then determine whether they copulate or not. These results support the hypothesis that females mate multiply in order to replenish inadequate sperm supplies to fertilize all eggs produced.     

Factors and consequences of a non-functional alary polymorphism inPyrrhocoris apterus (Heteroptera: Pyrrhocoridae)

Functional alary polymorphisms have been studied rather extensively in several insect species. This review article deals with factors controlling wing polymorphism in a flightless species,Pyrrhocoris apterus (L.), and discusses its adaptive significance and mechanisms for their persistence under natural conditions. The macropterous morph is determined by a recessive allele whose penetrance depends on photoperiod and temperature. Natural populations of this species contain a small fraction of flightless macropters. The disadvantages of being a macropter (increase of development time, decrease of fecundity) are minimal, while the benefit may consist in the tendency to prereproductive arrest of ovarian development in teneral females. It prevents establishing a second generation which would mostly die during the next winter. The mechanism of alary morph regulation may be an ancestral trait linkingP. apterus with other polymorphic Heteroptera, while its decreased penetrance may be a derivative character. Variation in fitness due to alary morphs is small compared to the one associated with differences in body size. The latter is environmentally determined, and not linked to the genetic basis of wing polymorphism. In the “mosaic” of phenotypes of various size the significance of the genetic macroptery may be close to neutral.     

Seasonal host alternation by the andromeda lace bug,Stephanitis takeyai (Heteroptera: Tingidae) between its two main host-plant species

I studied the seasonal occurrence of the andromeda lace bug,Stephanitis takeyai, on its two main host-plant species. In a secondary forest in Kyoto, this bug altered its hosts seasonally, i.e., from an evergreen shrub,Pieris japonica, in winter to a deciduous shrub,Lyonia elliptica, in summer. In contrast, in Nara park where fewL. elliptica were available, the bug exploited onlyP. japonica. Thus, seasonal host alternation by this bug is not obligate. A comparison of adult longevity and fecundity on the two host-plant species demonstrated the higher quality ofL. elliptica as a food resource. Corresponding to this difference in host quality, there was a dramatic difference in the seasonal population growth in the two study sites. In Nara, the population size at the beginning of the 2nd generation was almost the same as in the overwintered generation, whereas in Kyoto the population size in the 2nd generation was approximately one hundred times as large as in the overwintered generation. Thus seasonal host alternation is adaptive for the bug. In a previous study, I reported that overwintering as eggs in living leaves of their hosts is likely to be common among all the related species of this bug. Thus, this trait can be considered to be a phylogenetic constraint to the group. I speculate that host alternation by this bug has been derived because it is more adaptive from autoecy on an evergreen plant, similar to the pattern currently found in Nara, and that this bug can not only exploit deciduous host due to a phylogenetic constraint.     

Egg cluster size variation in relation to the larval food abundance inLuehdorfia puziloi (Lepidoptera: Papilionidae)

Summary Mean egg cluster size ofLuehdorfia puziloi yessoensis varied among habitats. The mean egg cluster size tended to be large when abundance of the larval food leaves expressed as the fresh weight of leaves per unit area at a given habitat was high. Since this variation was observed among closely located study plots (butterflies can easily move between study plots), the egg cluster size variation among habitats is likely to be a result of flexible response by females to varying food abundance for larvae.     

Scramble competition polygyny in the coreid bug (Colpula lativentris)

Summary The mating system of the coreid bug,Colpula lativentris, was studied, and compared with those in other hemipteran bugs. Copulating pairs formed a compact aggregation on a shoot of the Japanese knotweed,Polygonum cuspidatum, and there continued copulation and feeding for more than one day. This aggregation was defined as “a shoot aggregation”. Other than this aggregation composed of copulating pairs, there were loose aggregations on the ground composed of males and females not in copula. This loose aggregation was defined as “a ground aggregation”. Males searched for receptive females mostly in a ground aggregation. Sex ratio was constant and variance of sex ratio was also very small in any shoot aggregations and in large ground aggregations. However, in small ground aggregations, sex ratio was strongly male biased and variance of sex ratio was fairly large. This was because receptive females tended to leave a small ground aggregation and join a large one. These results suggest that a male can attain higher mating success in a larger ground aggregation.     

Mating and diapause in hibernating adults ofMenida scotti puton (Heteroptera: Pentatomidae)

InMenida scotti aduts which mate during hibernation, we examined the effect of photoperiod on diapause induction in terms of the developmental degree of reproductive organs and corpora allata. In a hibernating population, mating season and physiological changes related to dipause were investigated. When newly emerged adults were reared under long- or short-day conditions, reservoir of the ectodermal accessory gland and corpora allata in males developed, while the developments of ovary and corpora allata in females were suppressed under both conditions. This suggests that diapause of this species is induced only in females independently photoperiodic conditions during adult stage. In a hibernating population, mating was observed from early to late November and from late March to early May. Observations of the development of reproductive organs and corpora allata suggest that diapause is induced only in females in the former period, while both sexes are not in diapause in the latter period.     

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

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

Fertility schedules ofunaspis yanonensis (Hemiptera: Diaspididae) in relation to daily temperature

Summary Fertility schedules ofUnaspis yanonensis on citrus leaves were investigated from 1962 to 1971 for the first generation and from 1964 to 1971 for the second generation, at the Okitsu Branch of the Fruit Tree Research Station in Shizuoka Prefecture. The fertility curves were commonly characterized by a first major peak and a few subsequent minor peaks, although their shapes differed from year to year. Models estimating these fertility curves of each generation were constructed in relation to the fluctuations in average daily temperature, on the assumption that the curve was composed of two peaks, i.e., bimodal. Estimated curves were well fitted to observed ones. Contribution B-173, Fruit Tree Res. Stn.