Males of the seed bug Togo hemipterus (Heteroptera: Lygaeidae) use accessory gland substances to inhibit remating by females

In species in which females mate repeatedly, males can adopt several strategies to reduce the risk of sperm competition with future males. The refractory period of females significantly increased as the mating duration increased in the seed bug Togo hemipterus (Heteroptera: Lygaeidae). To elucidate the mechanisms by which mated females are inhibited from remating, we investigated the effects of male-derived substances on the inhibition of mating receptivity of virgin females by injecting the substances into their abdomens. The length of time from injection to mating in virgin females was significantly longer for females injected with accessory gland B solution than for those injected with seminal vesicle, accessory gland A, or control solutions. This is the first report showing that heteropteran males inhibit female remating by using substances from an accessory gland. We discuss and consider the adoption and evolution of this strategy by T. hemipterus males by focusing on female genitalia structures, oviposition habit, and paternity and comparing these traits with those of other heteropterids.     

Evolutionary causes of male-biased sexual size dimorphism from a female perspective in the seed bug Togo hemipterus (Heteroptera: Lygaeidae)

Sexual size dimorphisms (SSDs) in body size are expected to evolve when selection on female and male sizes favors different optima. Many insects show female-biased SSD that is usually explained by the strong fecundity advantage of larger females. However, in some insects, males are as large as or even larger than females. The seed bug Togo hemipterus (Scott) also exhibits a male-biased SSD in body size. Many studies that have clarified the evolutionary causes of male-biased SSD have focused only on male advantages due to male–male competition. To clarify the evolutionary causes of male-biased SSD in body size, we should examine the degree of not only the sexual selection that favors larger males but also natural selection that is acting on female fecundity. The obtained results, which showed higher mating acceptance rates to larger males, implies that females prefer larger males. No significant relationship was detected between female body size and fecundity; body size effects on female fecundity were weak or undetectable. We conclude that male-biased SSD in T. hemipterus can be accounted for by a combination of sexual selection through male–male competition and female choice favoring large males, plus weak or undetectable natural selection that favors large females due to a fecundity advantage.     

Male body size does not affect the refractory period of females in the West Indian sweet potato weevil Euscepes postfasciatus (Fairmaire) (Coleoptera: Curculionidae) and the seed bug Togo hemipterus (Scott) (Heteroptera: Lygaeidae)

Journal of Ethology - Multiple mating by females increases both the risk and intensity of sperm competition for males. The ability to temporarily or permanently inhibit female receptivity to...     

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.     

Effect of Mating Frequency on Female Fitness in Caloglyphus Berlesei (Astigmata: Acaridae)

We studied the effect of mating frequency on female longevity and fecundity in the mite Caloglyphus berlesei. We tested the hypothesis that high promiscuity is selected for in this species because females receive nutrients during copulation. Females were constantly exposed to either one or four males and fed either a standard or poor diet. The prediction that frequent mating benefits females was not confirmed. On the contrary, females kept with four males had significantly lower longevity on both diets and on the standard diet they also had decreased fecundity compared to females exposed to one male. On the poor diet the number of males had no significant effect on fecundity. Thus, increased mating frequency had a detrimental effect on female fitness.     

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.     

黑水虻幼虫的耐饥力及饥饿幼虫复食后的生物学特性

【目的】旨在明确黑水虻Hermetia illucens幼虫的耐饥力及饥饿幼虫复食后的发育、生殖力等生物学特性,为评估饥饿对黑水虻后续发育的影响并指导生产实践提供依据。【方法】将不同日龄或不同体重黑水虻幼虫进行不同时长的饥饿处理,测定和分析其耐饥力与日龄、百头重的关系;对7日龄幼虫进行不同时长的饥饿处理后再复喂,测定不同饥饿时长对其预蛹出现时间、总预蛹率、预蛹重、成虫繁殖等的影响。【结果】黑水虻幼虫饥饿后的半数致死时间(LT₅₀)与日龄成指数关系,与体重呈线性相关。7日龄幼虫的耐饥力较强,LT₅₀超过了50 d,饥饿30 d时存活率仍然达到了95%。百头重小于0.5 g的个体,其LT₅₀小于30 d;百头重在2～2.5 g的个体,其LT₅₀在45～70 d。幼虫超过8日龄时或百头重超过4.5 g时,多数个体在饥饿5 d时就提前进入了后续发育阶段。7日龄幼虫饥饿40 d内复食,其预蛹率未受到影响,但预蛹重显著增加;短期饥饿(0～10 d)对其成虫产卵量无显著影响,长期饥饿(30～40 d)使其成虫产卵量显著下降;饥饿长达30 d以上时,卵孵化率也显著降低。【结论】本研究明确了不同体重或不同日龄的黑水虻幼虫都具有一定的耐饥力;饥饿幼虫复食后预蛹重显著增加。     

Multiple mating reduces male survivorship but not ejaculate size in the polygamous insect Stenomacra marginella (Heteroptera: Largidae)

In polygamous species, successful males should be able to inseminate multiple females, to defeat sperm from previous males, to avoid sperm displacement by other males, and to induce females to use his sperm during fertilization. Since resources are limited, adaptations to perform any of these functions may conflict with each other (and with other life-history traits) and trade-offs are expected to evolve. We studied if males of the polygamous true bug Stenomacra marginella face a trade-off between multiple mating and survivorship, by comparing the survivorship of virgin and multiply mated males. We also looked for physiological costs of ejaculate production by examining ejaculate production in consecutive matings in multiple mated males. Multiply mated males were able to produce ejaculates of similar size in up to six consecutive copulations but they had decreased survivorship in comparison with virgin males. There was no difference in survivorship between males mated three and six consecutive times, suggesting that the negative relation between survivorship and number of copulations is not linear. The decrease in survivorship seems to be a cost of mating and ejaculate production. This cost could favor the evolution of prudence in the allocation of resources to ejaculate production (e.g., cryptic male choice).     

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

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 that C. 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.     

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.     

Sexual receptivity and post-emergence ovarian development in females of Coproica vagans (Diptera: Sphaeroceridae)

Abstract .Males and females of the dung fly species Coproica vagans Haliday 1833 (Diptera: Sphaeroceridae) mate soon after emergence from the puparium. At this time females still have immature ovaries. Therefore, mating precedes vitellogenesis in this species. Data presented here show that mating enhances oogenesis in C. vagans females. Mated females mature their first egg batch sooner and oviposit four days earlier than virgin conspecifics. Mating-related enhancement of oogenesis could be explained either through nutritional benefits to females or male chemical or stimulatory manipulation of the females. Oogenesis was divided into six arbitrary stages, with vitellogenesis beginning in stage 4. Ovarian development beyond stage 4 is rapid compared with pre-vitellogenetic development. Virgin females pause oogenesis in stage 4. The genital opening of mated females is blocked by a mating plug that persists until oviposition begins. The plug seems to ensure the paternity of the last male to mate by preventing females from remating. The operational sex ratio in C. vagans populations is presumed to be strongly male-biased.     

Male Participation in Nest Building in the Dung Beetle Scarabaeus catenatus (Coleoptera: Scarabaeidae): Mating Effort Versus Paternal Effort

The dung beetle, Scarabaeus catenatus, shows not only the rolling but also the tunneling tactic for nest building with bisexual cooperation. Sex roles, however, differed between the tactics. In rolling, the male took the initiative like that of ball-roller species: he rolled a dung ball away and buried it. In tunneling, in contrast, the male usually had a secondary role like that of tunnelers: he was less active in burrow excavation and provisioning. Regardless of the tactics, male participation did not increase female reproductive output measured by the number or size of brood balls in the field, but seemed to function as mate guarding against conspecific males. This suggests that, in both tactics, the male S. catenatus invests primarily in mating effort compared with paternal effort. The relative importance of mating effort in male participation seems to hold true in other dung beetles, irrespective of whether they are ball-roller or tunneler species. In addition, the male mating strategy of S. catenatus is compared with that of other ball-rollers.     

Mating unplugged: a model for the evolution of mating plug (dis-)placement

Mating plugs are male-derived structures that may impede female remating by physically obstructing the female genital tract. Although mating plugs exist in many taxa, the forces shaping their evolution are poorly understood. A male can clearly benefit if his mating plug secures his paternity. It is unclear, however, how plug efficacy can be maintained over evolutionary time in the face of counteracting selection on males' ability to remove any plugs placed by their rivals. Here, I present a game-theory model and a simulation model to address this problem. The models predict that evolutionarily stable levels of mating-plug efficacy should be high when (1) the number of mating attempts per female is low; (2) the sex ratio is male-biased, and (3) males are sperm-limited. I discuss these results in the light of empirical data.     

Precopulation Sexual Selection in Nysius huttoni White (Heteroptera: Lygaeidae) in Relation to Morphometric Traits

Nysius huttoni White is a polygamous bug, endemic to New Zealand, and an important pest of wheat and brassicas. This bug has a female-biased sexual size dimorphism but relative to body length, males have longer antennae, suggesting that the allometric scales of antennal–body relationships may be highly selective in sexual selection. Body weight and most morphometric traits measured have no effect on mating success of either sex. Males significantly preferred mating with females having thicker abdomens, more mature eggs, and longer ovipositors. This result suggests that males may select their mates on the basis of immediate reproductive benefit: fertilizing more eggs and ensuring better survival of these eggs. Males with large genital structures have mating advantages over those with small ones, suggesting that precopulation sexual selection in this species act on male genital traits rather than body weight and nonsexual traits. Finally, females significantly preferred males with greater slopes for the antennal-body relationship for mating. The allometry in the male antennal length may be an indicator of male reproductive fitness.     

Membranes and calcium sequestration during spermiogenesis in the cotton seed bug (Dysdercus intermedius: Heteroptera)

Summary By use of osmium ferricyanide (OsFeCN) staining the fate of cytoplasmic membranes was followed during spermiogenesis in the cotton seed bug (Dysdercus intermedius). During early spermiogenesis interzonal lamellae of endoplasmic reticulum become aggregated as a stack of membranes traversing the entire cell body from the nucleus to the cytoplasmic bridge connecting neighbouring spermatids. Cisternae of endoplasmic reticulum ensheath the acroblast from which vesicles of different sizes are pinched off into the cytoplasm. The oxalate method was used to show that acroblast and associated vesicles are calcium-sequestering sites in spermatids. Membrane profiles with dense calcium oxalate precipitate derived from the acroblast form an uninterrupted membranous sheath at the apical side of the nucleus where the proacrosome will be attached. With further development of the spermatids, the vesicles derived from the acroblast also participate in forming a calciumsequestering sheath enveloping the axoneme and the mitochondrial nebenkern derivatives.     

Relationships among male sexually selected traits in the bean bug,Riptortus pedestris (Heteroptera: Alydidae)

Current concepts of sexual selection suggest that male reproductive success is determined by multiple sexual traits. As expression and production of multiple sexual traits are frequently associated with each other, positive or negative correlations among multiple sexual traits ensue. These relationships among traits associated with male reproductive success may be crucial in the evolution of male reproductive strategies. Here, we investigate phenotypic relationships among sexually selected traits in the armed bean bug Riptortus pedestris. In this insect, males with a larger body and weapon are more likely to win male–male competitions, and males with a larger weapon or higher courtship rate are more attractive to females. There was a significant positive correlation between body size and weapon size, whereas the courtship rate had significant negative correlations with body size and weapon size. Our results suggested that there was a phenotypic trade‐off between courtship rate and male morphology. In this insect, smaller males may make more effort in courtship behavior as an alternative mating tactic.     

Selection of male traits during and after copulation in the seedbug Lygaeus simulans (Heteroptera, Lygaeidae)

In order to evaluate selection of male morphological traits during copulation, a laboratory experiment was performed with the promiscuous seedbug Lygaeus simulans. Three male traits suspected as putative targets of selection were measured: weight, fluctuating asymmetry of a measure on the forewings, and length of a conspicuous genital structure, the processus gonopori. As fitness measures we considered total fecundity (number of fertilized eggs), insemination and fertilization success (established if a female laid fertilized eggs after copulation), and the interval between copulation and oviposition. Eighty-four males were allowed a single copulation with one virgin female each. Out of 67 copulations, 27 (40.2%) resulted in fertilized eggs and the oviposition latency ranged from 6 to 26 days. Regressions of male traits on the fitness measures showed significant phenotypical selection of two male traits: (1) males of average weight are more likely to achieve fertilization and (2) the oviposition latency was shorter for males with lower asymmetry. The copulation-oviposition interval may be especially important for male fertilization success because Lygaeus males perform copulatory mate guarding and the last male copulating with a female fertilizes most of the eggs. No selection of the genitalic trait was detected.     

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.     

Male mate choice based on chemical cues in the cricket Acheta domesticus (Orthoptera: Gryllidae)

1. Males in many animal species exercise mate choice to maximise their reproductive success, assessing females by characteristics related to reproductive potential, such as mating status, body size, and age. The sensory modalities involved in mate choice are often not firmly demonstrated, but only inferred. This is especially true for chemical cues and signals. 2. The present study tests whether males of the cricket Acheta domesticus are able to choose among females based only on chemosensory cues. In A. domesticus, as in many crickets, males call to attract females or roam the habitat silently to search for females. In three‐way choice trials, males were presented with two filter papers that had been placed with females for 24 h prior to the trials and one blank control. Females were either mated or virgin and starved or well‐fed. It was predicted that males would prefer virgin over mated females and those in good condition over starved ones. 3. Males were more likely to contact filters that had been exposed to females. They spent more time examining filter papers from virgin females than those from mated ones, while the condition of the females had no effect. 4. We conclude that males can detect chemical cues from females on substrate and distinguish virgin females from mated ones. Being able to assess sperm competition risk prior to mating or even before further pursuing a trail with chemical cues should confer a considerable benefit to males.     

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.