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.     

Wing dimorphism and the migratory syndrome: Correlated traits for migratory tendency in wing dimorphic insects

The hypothesis that the morphological, physiological, and behavioral traits comprising the migratory syndrome in insects are genetically correlated through pleiotropic effects of genes controlling the titre of a common hormonal determinant is explored. Evidence that juvenile hormone (JH) influences the component traits of the migratory syndrome is presented, and thus JH is assumed to be the underlying, common determinant. However, readers are cautioned that this does not imply that JH is solely responsible for these traits, nor is this necessary for the arguments presented. For wing dimorphic taxa, the “correlated traits hypothesis” predicts covariance within wing morphs between JH titre and the proportion winged. Four simple genetic models for wing-morph determination are considered: single-locus with short-winged (SW) dominant; single-locus with long-winged (LW) dominant; polygenic, fixed threshold, shifting distribution; and polygenic, shifting threshold, fixed distribution. In each case, wing morphology is assumed to be a threshold trait with the liability being JH titre at some critical stage of development. All models predict covariation between %LW and the mean JH titre of at least one of the wing morphs, but the form and direction of the relationship depends critically on the genetic model used. The results suggest that we should expect the traits associated with the migratory syndrome, and hence the trade-offs associated with the evolution of wing dimorphism, to be correlated with proportion winged and, in this sense, to be frequency-dependent.     

Hybridization and speciation in the carabid beetles of the subgenusOhomopterus (Coleoptera, Carabidae, genusCarabus)

Natural hybridization among wingless carabid beetles of the subgenusOhomopterus (Carabidae, genusCarabus) is reviewed, and its significance in the evolution of this subgenus discussed. Natural hybridization occurs between parapatric species of similar size. Two case studies of natural hybridization suggest that natural hybridization could have affected the evolution of this subgenus in different ways. When there is a large difference in genital morphology between hybridizing species, interspecific copulation often results in genital injuries that causes mortality of copulating individuals, and hence reduces the fitness of hybridizing individuals greatly. In such a case, hybridization may be effective in maintaining the parapatric distribution of the two species, and in the long term, may promote reinforcement selection for traits which are effective in prezygotic reproductive isolation. When the morphological difference in genitalia is not so large as to cause genital injury, a hybrid population may be established at the intermediate zone between two parental species, provided that the immigration rates of the two species into the intermediate zone are small. Thus, natural hybridization may have contributed to both divergence and reticulate evolution in this subgenus.     

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.     

Population dynamics of the whitebacked planthopper,Sogatella furcifera (Hemiptera: Delphacidae) with special reference to the relationship between its population growth and the growth stage of rice plants

A population census was conducted to describe the effects of the growth stage of rice on the population dynamics ofS. furcifera, in particular, on immigration, seasonal abundance, population growth rate, and wing-form expression. The number of immigrants was highest on rice plants 17 to 30 days after transplanting (DAT), which suggested that immigrants prefer to settle or remain more on rice plants at the tillering stage (approximately 20–30 DAT). Population growth rate from immigration to the 1st generation decreased with an increase in rice plant age. In contrast, population growth rate from the 1st to 2nd generation was not influenced by rice plant age and was negatively density-dependent. The percentage of macropters (flyers) was positively related to the growth stage of rice when rice was in the vegetative or early part of the reproductive stage, and reached 100% at about 10 days before heading (booting stage). Threafter, most adult females molted into macropters regardless of population density. The roles of host plant age and crowding effect on the population dynamics ofS. furcifera are discussed.     

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.     

Contest and scramble competitions in two bruchid species,Callosobruchus analis andC. Phaseoli (Coleoptera: Bruchidae)

Summary We performed multiple-generation competition experiments betweenCallosobruchus analis andC. phaseoli with different bean sizes. In each system, we supplied 5 g of mung beans (Vigna radiata) every 10 days. We examined three types of bean conditions: 5 g of large beans, 5 g of small beans, and a mixture of 2.5 g of large and small beans. In small bean condition,C. analis dominatedC. phaseoli in all three replicates andC. phaseoli was extinct by the 260th day. On the contrary,C. phaseoli overcameC. analis within 250 days in large beans in all three replicates. In mixed beans condition the two bruchid species coexisted more than 500 days in two out of the three replicates. Even in the exceptional case, both species coexisted for 460 days. These results were examined in the light of the predictions from short-term larval competition experiments and a game theoretical model by Smith and Lessells (1985). The density and frequency dependent results during larval competition inside a bean was concluded to be a main factor to produce the above long-term competition results.     

Differences in adult and reproductive lifespan in the two male forms ofMnais pruinosa costalis selys (Odonata: Calopterygidae)

The damselflyMnais pruinosa costalis (Odonata: Calopterygidae) is unusual in that males are dimorphic, existing as clear-winged non-territorial ‘sneaks’ and orange-winged territorial ‘fighters’. Here we report the results of population census data and behavioural observations in the field and laboratory, and present estimates of emergence period, reproductive period, total lifespan, and reproductive success of each morph. Clear-winged males are smaller and have lower daily reproductive success than orange-winged males, but live for longer in the field and laboratory. We accounted for the difference in the ‘operational reproductive life’ of the two morphs and estimated lifetime reproductive success: there was no difference between clear-winged and orange-winged males. We discuss possible mechanisms for the maintenance of the two forms.     

Contest and scramble competitions inCallosobruchus maculatus (Coleoptera: Bruchidae) II. Larval competition and interference mechanisms

Summary Competition between contest and scramble strategists was examined using two strains ofCallosobruchus maculatus, the contest strain (iQ) and the scramble strain (tQ). The direct larval interference experiment within each strain showed that the contest strain performed strong direct interference throughout its larval stage against the conspecific opponent(s). The scramble strain also performed its interference, but only during the 3rd and the 4th larval stages. The inter-strain larval competition experiments inside a large and small mung bean (Vigna radiata) showed that the contest strain was, in general, superior to the scramble strain in competition, but the competition results were density- and frequency-dependent. In the large beans, the proportion of adult emergence of the contest strain increased with the increase of its own initial density but decreased as the initial density of the scramble strain increased. The shape of the proportion of adult emergence became nonlinear in the small beans. The cause of the complexity of inter-strain competition results was discussed in the light of the difference in larval interference ability between the two strains.     

Effects of spatio-temporal intervals between newly-hatched larvae on larval survival and development inMonochamus alternatus (Coleoptera: Cerambycidae)

The effects of distance between hatching larvae on survival and development were investigated inMonochamus alternatus. Two newly-hatched larvae were inoculated intoPinus densiflora bolts at a distance of 2.5 cm or 10 cm, simultaneously or at an interval of 2 weeks. Some larvae were inoculated singly as a control. When larvae were inoculated simultaneously, mortality of the closely-inoculated larvae was significantly higher than that of distantly-inoculated larvae. Such high mortality was identified as due to conspecific bites. When the two larvae were inoculated asynchronously, the first-inoculated larvae killed some second-inoculated larvae but were never killed by them. Consequently, mortality was higher in second-inoculated larvae than in first-inoculated larvae. In particular, there was a significant difference in mortality between them when the larvae had been inoculated closely. The mortality of second-inoculated larvae was higher in the closely-inoculated group than in the distantly-inoculated group although there was no significant difference between them. In the case of two simultaneously-inoculated larvae, the initial distance between them had no significant effect on the development and growth in the early larval stage. When the larvae were inoculated asynchronously, the first-inoculated larvae grew more quickly than singly-inoculated control larvae.     

The role of vole populations in prevalence of the parasite (Echinococcus multilocularis) in foxes

Effects of population fluctuation of the gray-sided vole(Clethrionomys rufocanus) on the prevalence (infection rates) of the parasiteEchinococcus multilocularis in red fox(Vulpes vulpes) populations was investigated from 1985 to 1992 in eastern Hokkaido (Abashiri, Nemuro, and Kushiro area), Japan. This parasite needs two hosts to complete its life cycle; the gray-sided vole as its intermediate host and the red fox as its final host. We found that: (1) Infection rates in foxes depended on the current-year abundance of voles in all three study areas, particularly in Abashiri. (2) In addition to this direct density-dependence, delayed density-dependence between the infection rate and the prior-year abundance of voles was detected in Nemuro and in Kushiro. (3) The regional differences in density-dependence pattern were related to regional differences in the winter food habits of red foxes: in Abashiri the proportion of voles in the fox’s diet greatly decreases in winter, while the proportion remains high in winter in Nemuro and in Kushiro, probably because of shallower snowpack. These results suggest that infection rates in foxes in Abashiri were less influenced by the prior-year prevalence, since the infection cycle might be interrupted in winter, when voles became less important in fox’s diet. In contrast, the state of the prevalence may carry over from year to year in Nemuro and in Kushiro, because red foxes continue to eat a considerable amount of voles throughout year. The regionally contrasted results for the relationship between infection rate in foxes and vole abundance were parallel to the regional difference in fluctuation pattern of vole populations, which are highly variable in Abashiri area, but less variable in Kushiro-Nemuro area. Drastic change in vole populations appears to affect the host-parasite system.     

Reproductive strategy of the sugi bark borer,Semanotus japonicus (Coleoptera: Cerambycidae) on Japanese cedar,Cryptomeria japonica

The inner bark of Japanese cedar,Cryptomeria japonica D. Don., is the main food of the sugi bark borer,Semanotus japonicus Lacordaire, but may also be involved in resistance to attack by the borer. I used newly hatched larvae to inoculate cedar logs that had been cut 2-weeks to 12-months earlier, and undamaged living cedar trees. On living trees, all larvae were killed by resin flow from the traumatic resin canals between the outer- and inner bark or between the inner bark and cambium. In logs that were cut more than 4 months prior to inoculation, larvae were unable to complete development. In logs cut 2 weeks prior to inoculation, almost all larvae were able to reach the adult stage but they were smaller in size than adults from living trees damaged by this borer. These results suggest that poor nutrient conditions in the inner bark of logs affected development and survival of the borer. Because of its nutritional advantage, living cedars provide a better environment for the sugi bark borer. However, all larvae are killed by resin flow, suggesting that this insect is “in between” being primary and secondary with respect to living trees, or a “weak” primary insect. The sugi bark borer seems to develop early in the season so that early instar larvae encounter reduced amounts of resin flow and so that late-instar larvae feed mostly in summer when nutrient levels in the inner bark are at their highest.     

A simple evolutionary model of dormancy and dispersal in heterogeneous patches with special reference to phytophagous lady beetles: I. Stable environments

Summary A simple evolutionary model of dormancy and dispersal is presented with special reference to phytophagous lady beetles. In order to investigate spatially heterogeneous environments, we assume the simplest patch structure, that is, there are only two patches, main and sub. Environments are also assumed to be temporally constant. The main patch is superior to the sub patch, but density effect at the main patch is higher than at the sub patch. Optimal dormancy and dispersal are obtained at the same time by the method of evolutionarily stable strategy (ESS). In the univoltine life cycle, dormancy strategy vanishes because dormant individuals do not reproduce at all but suffer from a certain mortality rate during winter hibernation. In the bivoltine life cycle, the dormancy and dispersal rates constitute a trade-off: the rates change together with a negative correlation when the mortality rate during dispersal or during winter hibernation changes. When suitability of the main patch gradually deteriorates, the optimal strategy changes as follows: neither dormancy nor dispersal is adopted at the most suitable condition, the dispersal rate is increased without dormancy in the intermediate condition, and then the dormancy rate is increased with a constant dispersal rate. We discuss the field observation data of lady beetles in the light of results of our model.     

Prey preference of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand)

Summary Functional responses of the wolf spider,Pardosa pseudoannulata (Boesenberg et Strand) attacking the rice brown planthopper,Nilaparvata lugens (St?l.), and the mirid predatorCyrtorhinus lividipennis Reuter were both those of Holling Type II. The attack rate was higher and handling time lower forC. lividipennis. However, when caged with the two prey, the wolf spider showed a significant preference forN. lugens at a lower prey proportion. Proportions of prey attacked were significantly different from the expected ratios of prey available as well as from the predicted preferences derived from the functional response parameters. As proportions ofN. lugens attacked changed from greater to less than expected as the proportions ofN. lugens available increased, a “reverse switch” behaviour seems to be evident.     

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.     

Host plant conspicuousness and the distribution of eggs and larvae in the butterfly,Anthocharis scolymus (Lepidoptera: Pieridae)

Summary We investigated how the distribution pattern of eggs and larval on the host plant,Turritus glabra, was influenced by the oviposition behavior of the pierid butterflyAnthocharis scolymus. Females searched for the host plants visually and they frequently approached taller host plants with sparse surrounding vegetation. After encountering host plants, oviposition behavior of females was independent of host plant characteristics such as height, density, and type of surrounding vegetation. A female laid eggs singly on a host plants. Most females appeared to lay their eggs regardless of the presense of eggs on the host plant. Consequently egg and larva tended to be abundant on conspicuous host plants as measured by height or relative isolation from other plants. However, overcrowding of eggs on an individual host decreased the survival rate of larvae.     

Effects of interspecific competition on development and reproduction in two giant water bugs,Diplonychus japonicus vuillefroy andDiplonychus major esaki (Hemiptera: Belostomatidae)

Summary We studied the interpecific competition between 2 species of predatory aquatic bugs,Diplonychus japonicus andD. major by conducting a field experiment. We set up 3 types of experimental plots in the paddy fields whereD. major predominated. The two plots contained single species of eitherD. japonicus orD. major, respectively, and one plot had both species in equal number. We compared the development and the reproductive performance between plots in each species. InD. japonicus, the number of eggs and early instar nymphs were significantly smaller in the plots containing both species than in the monospecific plots. However, the numbers of late instar nymphs and newly emerged adults were not significantly different between plots. The proportions of starved nymphs in both plots were larger than those in theD. japonicus's natural habitats. The final densities of adults in both plots were lower than those in the natural habitats. These results suggest that lower density ofD. japonicus in these paddy fields is due to the lack of available food for nymphs rather than the effects of interspecific competition withD. major. InD. major, significant differences were not found in the number of eggs, each instar nymphs and adults. These results suggest that the effects of interspecific competition did not affect the reproductive performance ofD. major.     

Genetic control and evolution of acoustic signals in planthoppers (Homoptera: Delphacidae)

Acoustic signals are part of the specific mate recognition system of planthoppers. The genetic control of acoustic signal characters was studied in the planthopperRibautodelphax imitans. Artificial selection for interpulse interval in the female call revealed a large additive genetic component for this polygenic character. Other female call characters showed a correlated response. Some male call characters also appeared to be genetically correlated with the female character selected for, despite the rather different structure of male and female calls. Parent-offspring regression provided significant heritability estimates for those male call characters that also responded to artificial selection in the female call, one of which appeared to be influenced by sex-linked genes. It is argued that the differentiation of this mate recognition system in planthopper populations and species could be the result of founder effects, enabled by the genetic plasticity of the call characters and the existence of a wing length dimorphism in these animals.     

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%.     

Comparison of dispersal ability between the green leafhoppers,Nephotettix virescens andN. cincticeps (hemiptera: Cicadellidae) by tethered flight

Summary Dispersal propensity of green leafhoppers was compared between a tropical species,Nephotettix virescens, and a temperate species,N. cincticeps. The flight ability was measured with tethered flight technique under laboratory conditions (25°C, 16L-8D). The pre-flight period was shorter and the flight duration was longer inN. virescens than inN. cincticeps in both sexes. No significant correlations were found between the flight activity and morphometric characters for either of the two species. The results suggested that dispersal propensity ofN. virescens is higher than that ofN. cincticeps.