Sex ratio of Nysius huttoni White (Hemiptera: Lygaeidae) in field and laboratory populations

Abstract

We studied the parasitoids of three insect seed predators of the mast-seeding genus Chionochloa spp. (snow tussocks) at Mt Hutt, New Zealand. Megacraspedus calamogonus (Lepidoptera: Gelechiidae) is parasitised by four species, three Hymenoptera (Zealachertus tortriciphaga [Eulophidae], Diadegma sp. [Ichneumonidae] and Dolichogenidea sp. [Braconidae]) and one Diptera (Uclesiella sp. [Tachinidae]). Diplotoxa similis (Diptera: Chloropidae) is parasitised by Callitula sp. (Hymenoptera: Pteromalidae). Eucalyptodiplosis chionochloae (Diptera: Cecidomyiidae) is parasitised by two Hymenoptera, Gastrancistrus sp. (Pteromalidae) and Zelostemma chionochloae (Platygastridae); all three species have extended diapause. Overall parasitism was 68.5% in M. calamogonus, 1% in D. similis, and 41% in E. chionochloae. Such parasitism in M. calamogonus and E. chionochloae may reduce seed predation in Chionochloa and alter the selective benefit of mast seeding (predator satiation) to the plant. However, seed predation is still high at many sites, so some of the seed predators and parasitoids may be food-limited (bottom-up regulation).     

Effect of day length on development and reproductive diapause in Nysius huttoni White (Heteroptera, Lygaeidae)

Abstract:  Nysius huttoni White is endemic to New Zealand and an important pest of wheat and brassica crops. To provide critical information for pest forecast, management and quarantine inspection, we investigated the effect of day length on the growth, development and reproductive diapause of this pest under a series of photoperiodic regimes: 16 : 8, 14 : 10, 12 : 12 and 10 : 14 h [light : dark (L : D)]. Long day lengths [16 : 8 and 14 : 10 h (L : D)] promote a continuous lifecycle while short day lengths [12 : 12 and 10 : 14 h (L : D)] slow up the growth and development, prolong the pre-mating period, and induce the reproductive diapause. The absence of oviposition for approximately 30 days is recognized as the criterion for N. huttoni reproductive diapause definition. When all life stages are kept under the short day length conditions, only 60–73% of females enter reproductive diapause; if the exposure to short day lengths starts in late instar nymphs, 100% of females enter reproductive diapause. If only adults experience short day lengths, does diapause incidence fall in between, with up to 26% of females laying a few eggs before entering diapause. The critical photoperiod for the induction of reproductive diapause falls between 13.3 : 10.7 and 13.5 : 10.5 h.     

Thermal requirements for the development and reproduction of Nysius huttoni White (Heteroptera: Lygaeidae)

Nysius huttoni White is an economically important pest of wheat and brassica crops in New Zealand. Because of its frequent presence in export fruit packages, it is also considered an important quarantine pest to countries that trade with New Zealand. To provide critical information for the pest risk analysis, forecast and management of N. huttoni, we investigated the effect of five consistent temperatures (10, 15, 20, 25, and 30 degrees C) on its development, survival and reproduction. At 10 degrees C both eggs and nymphs did not develop but the latter grew. Nymphs could survive 10 degrees C for >1.5 mo, with the fifth instar nymphs surviving for up to 145 d. Adults could live for at least 100 d at this temperature. This species could not complete its lifecycle at or below 15 degrees C. Between 15 and 30 degrees C, fifth instar stage was significantly longer than other nymphal stages. Egg hatch rate and total survival rate for all stages were significantly higher at 20 degrees C than at other test temperatures. The developmental rate of different life stages increased linearly with the increase of temperatures from 15 to 30 degrees C. The estimated low temperature threshold for the completion of lifecycle was 11.9 degrees C, and that for mating and oviposition was 12.3 and 16.8 degrees C, respectively. The thermal requirement for completing a life cycle of N. huttoni was 625 DD. The time needed for completing a life cycle was similar for both sexes. Temperature had little effect on adult body weight and sex ratio. Implications of the above findings are discussed.     

Function of Prolonged Copulation in Nysius huttoni White (Heteroptera: Lygaeidae) Under Male-Biased Sex Ratio and High Population Density

Prolonged copulation as a response to sex ratio and population density has been demonstrated for a number of species but the functions behind it remain controversial. It is difficult to determine the function of prolonged copulations without examining the mechanisms of sperm transfer and storage, and fertilization success–copulation duration relationships. Here, we report our work on a polygamous seed bug, Nysius huttoni White (Heteroptera: Lygaeidae), where we manipulated the population density and sex ratio for a series of mating trials, quantified the relationship between copulation duration and ejaculate amount as well as reproductive outputs, and examined the sperm transfer and storage mechanisms. Our results suggest that males prolong copulations to increase their fertilization success relative to their rivals in response to high sperm competition intensity under male-biased sex ratio and high population density. The mechanism behind the positive correlation between copulation duration and fertilization success may be attributed to the characteristic structures of aedeagus and spermatheca in this insect, which allow direct transfer of sperm to a storage site of fixed size during copulation.     

Evaluation of potential trap plant species for the wheat bug Nysius huttoni (Hemiptera: Lygaeidae) in forage brassicas

1. The wheat bug Nysius huttoni is a major pest of brassica seedlings. Management of this insect currently relies on seed treatment with neonicotinoids and spraying with chlorpyrifos and pyrethroid insecticides. These practices can generate severe external costs, including human health, the environment and biodiversity. Trap cropping is one alternative option to protect brassica seedlings from N. huttoni.
2. Trap crop species evaluated in field cage experiments were: alyssum (Lobularia maritima L. Desvauxcv. Benthamii White), wheat (Triticum aestivum L. cv. Morph), coriander (Coriandrum sativum L. cv. Santo) and clover (Trifolium repens L. cv. Nomad). These were compared with kale (Brassica oleracea L. cv. Kestrel). In open‐field experiments, alyssum (L maritima), wheat (T. aestivum) and a mixture of alyssum (L. maritima) and wheat (T. aestivum) were used. All of these were compared to kale (B. oleracea).
3. Alyssum and wheat were the most favoured potential trap plants for N. huttoni. Results indicated that two treatments: alyssum (used as a single trap crop) or ‘alyssum plus wheat’ (a multiple trap crop), may be useful in brassica fields to protect the seedlings from N. huttoni damage.
4. Such a trap cropping protocol potentially reduces pesticide use in forage brassicas and can also deliver multiple ecosystem services such as biological control of insect pests.

     

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.     

Possible Female Choice During Copulation in Ozophora baranowskii (Heteroptera: Lygaeidae): Female Behavior, Multiple Copulations, and Sperm Transfer

The remarkable behavior of female Ozophora baranowskii of lightly tapping the male with the hind legs during copulation was studied in relation to multiple copulations, copulation length, and sperm transfer. Multiple copulations did not affect female fecundity, fertility, or longevity. The incidence of tapping was higher in shorter copulations, which did not result in sperm transfer to the female. Female tapping behavior may be a mechanism for choosing between males after copulation has begun.     

Sexual Selection of Zorion guttigerum Westwood (Coleoptera: Cerambycidae: Cerambycinae) in Relation to Body Size and Color

Zorion guttigerum is a flower-visiting longhorned beetle endemic to New Zealand. Sexual selection of this species in relation to the body size and color form of different sexes was investigated in the field. The population sex ratio, based on censuses of feeding and mating sites (flowers), is male-biased. Females are significantly larger than males. Both sexes have antennae of similar length but the antennal length relative to the elytral length is greater in males than in females, and the antennal length of males increases more with an increase in body size than that of females. Both sexes have dark blue (DB) and yellowish-brown (YB) individuals. Both pair-bonded and solitary males are similar in elytral and antennal length. In pair-bonded males, DB individuals are significantly more numerous than YB ones, but in solitary males, the number of both color forms is similar. Males tend to have territory protection behavior, fighting with and chasing away rival males from feeding and mating sites. Larger males usually win the fight but the size-dependent fighting advantage does not translate into mating success. Male color plays an important role in mating success, with DB males having a significantly better chance to mate than YB males. Furthermore, male body size and color also have interactions in mating success: males of DB color morph obtain a greater mating advantage according to body size. Pair-bonded females are significantly larger and have longer antennae than solitary females, suggesting that males prefer larger females for mating. In addition, females of DB color morph with longer antennae are also preferred by males for mating. The significance of these findings is discussed.     

Proximate Mechanisms of Sexual Selection in the Firefly Photinus pyralis (Coleoptera: Lampyridae)

Male competition in courtships of the firefly Photinus pyralis is intense. Morphometric analysis revealed that male size is variable, possibly exhibiting alternative body plans. We examined the outcomes of courtships to determine the proximate mechanisms of selection that may have led to male allometry. Two distinct selective events exist within mate competition: (1) searching and (2) scramble competition. Although many courtships failed, those without rivals were shorter and successful males spent less time in the mate-guarding position than when several males were in competition. Winners outflashed rivals, but flash dialogs occurred at similar frequencies across rival densities, indicating that females require some quantum of information before mating. Female flash preference was examined in choice tests to evaluate female discrimination ability. Females preferred flashes of greater intensity and precedence. This suggests that flash synchronization is a competitive display rather than a cooperative behavior to maintain female responsiveness. Selection for male size was density dependent: females preferred large males in searching, while small males were favored in scrambles. Simulated scrambles showed that small males have superior locomotor abilities, allowing them to reach the female rapidly. Comparison with a species lacking scrambles, P. macdermotti, revealed that selection in P. pyralis is disruptive, targeting traits alternately, and may have led to the evolution of higher allometric ratios in a species with direct male competition. Both male competition and female choice are important determinants of the outcome of P. pyralis courtships.     

Chromosomal Polymorphism, Morphometric Traits and Mating Success in Leptysma argentina Bruner (Orthoptera)

The South-American species Leptysma argentina Bruner is polymorphic for a centric fusion between chromosomal pairs 3 and 6 (fusion 3/6). Cytogenetic and morphological studies revealed that fusion 3/6 significantly increases some morphometric variables in males. A selection components study showed that the fusion is positively selected for longevity, the direct effect being exerted on thorax height. Moreover, a unisexual approach revealed that sexual male selection acts by increasing third femur length. In the present study the effect of fusion 3/6 on morphometric variables and the effect of body size and karyotype on mating success in both males and females were analysed through a bisexual approach. Total body length (TL) and third femur length (FL) were significantly larger amongst the fusion carrier females. Besides, the largest females and the carriers of the fusion have increased mating success. The selection differential suggests that directional sexual selection favours the females with increased TL and fusion dosage; however, the selection gradient revealed that none of the variables here analysed is a direct target of sexual selection. The male sexual selection study agrees with previous results, showing the importance of the third pair of legs in male mating success among grasshoppers.     

First report on C-banding and fluorescent banding in species of Dieuches (Rhyparochrominae: Lygaeidae: Heteroptera)

Although the karyotypes of twelve species of Dieuches Dohrn, 1860 belonging to Rhyparochrominae have been described so far, there is no information about heterochromatin and its characterization in terms of base composition for any of the species. In the present paper, C-banding and fluorescent banding have been applied for the first time to three species of Dieuches : D. uniguttatus, D. insignis (2n = 12 = 8A + 2m + XY) and D. coloratus (2n = 14 = 10A + 2m + XY). Dieuches uniguttatus and D. insignis show distinct terminal C-bands along with a few interstitial bands in all the autosomal bivalents, whereas in D. coloratus , one autosomal pair is almost completely heterochromatic, three show C-positive bands while one is totally euchromatic. The sex chromosomes too show heterogeneity in distribution of C-heterochromatin among three Dieuches species. Characterization of heterochromatin in D. uniguttatus and D. insignis using DAPI/CMA₃ staining reveals that in D. uniguttatus , C- heterochromatin blocks of all the autosomal bivalents, which are predominantly A–T rich, whereas in D. insignis , these are rich both in A–T and G–C. In D. uniguttatus, sex chromosomes X and Y have localized G–C rich regions whereas in D. insignis , these are scattered in X and absent in Y. As variations in the heterochromatin represent the main source of karyological differentiation among and within species, it seems that there occurred extensive redistribution of heterochromatin within the complement as the three species evolved. There is need for cytological details of more species to understand evolutionary aspects in the genus Dieuches .     

Effect of Body Weight on Reproductive Performance in Cnephasia jactatana (Lepidoptera: Tortricidae)

The larger-the-better theory predicts that fitness is positively linearly associated with body size or weight. We used the kiwifruit pest, Cnephasia jactatana Walker, to test whether larger insects perform better reproductively. We divided our insect population into three weight groups: light, average, and heavy, and assessed the reproductive performance of 9 breeding treatments (3 male weights × 3 female weights). Female fecundity is positively correlated with female body weight in low and average weight groups. There is no such correlation in the heavy weight group, suggesting that further weight increase has no fitness gain for females. The positive linear relationship between fertility and female weight in all weight groups may be attributed to the fact that permanently paired heavy females are more likely to remate, gaining more sperm and thus higher fertility. However, the previous study also indicates that mated females are less likely to be mated again when males have a choice. Therefore, in the natural environment the realized fertility may still follow an asymptotical pattern similar to the fecundity in relation to female weight. Males' beneficial effect on female reproductive outputs increases linearly with their body weight in all weight groups, indicating that male reproductive performance fits the larger-the-better theory. Fertility rate is not affected by the body weight of either sex. Heavy and average females lay eggs earlier and have higher daily fecundity and fertility than light females. Females of all weight groups have similar oviposition and postoviposition periods. Male weight and female–male weight interactions have no effect on oviposition parameters.     

Morphometric variation in theAlonsoa meridionalis complex (Scrophulariaceae)

Morphological field studies of 27 populations of theA. meridionalis complex were conducted in Ecuador. It is possible to distinguish four different form series by the position of the stamens in relation to the style. This trait was congruent with the overall morphometric variation, shown by the results of a canonical discriminant analysis (CDA). The taxonomy of the species complex is discussed based upon the results of the CDA. The degree of outcrossing varied in different form series, from a comparatively high degree of selfing to predominantly outcrossing, as indicated by the variation within relative to among populations. This is probably due to the varying potentials for self-deposition owing to differences in the degree of herkogamy.     

Vitellogenesis inhibition in Oncopeltus fasciatus females (Heteroptera: Lygaeidae) exposed to cadmium

Newly moulted females of the insect Oncopeltus fasciatus were exposed to cadmium (Cd) dissolved in the drinking water (50-400 mg l(-1) Cd) for 5 days. Cd exposure delayed ovarian maturation and inhibited egg production. Exposure to Cd, moreover, decreased hemolymph levels of the two major vitellogenin polypeptides of O. fasciatus, VG1 and VG2, in a concentration-dependent way, probably by a reduction in their synthesis. The ovarian levels of VG1 and VG2 were also decreased in Cd-exposed females. It was next investigated whether Cd effects might be a consequence of the endocrine disruption of vitellogenin synthesis, which is controlled by juvenile hormone (JH). JH replacement therapy did not restore VG1 or VG2 levels in Cd-exposed females, but did so in starved females. Our results do not therefore support a disturbance of JH production or a reduction in feeding as the cause of the reduced vitellogenin polypeptide levels, but rather point to the site of action of JH, the JH receptor, as the target of Cd effects.     

Male Sexual Signals and Female Choice in Drosophila grimshawi (Diptera: Drosophilidae)

Sexual selection and sexual signaling have been prominent topics in recent behavioral studies, but limited data have led to controversy regarding these topics. For example, the Hawaiian Drosophila are often cited as examples in which female choice has resulted in the evolution of elaborate male courtship signals, but relatively few data exist to test these claims adequately. We studied D. grimshawi, a lek-forming Hawaiian Drosophila, to determine whether there was evidence for female choice without male competition and to elucidate the possible cues females use to discriminate. Male mating success was found to be nonrandom and males that courted females intensely and deposited many pheromone-containing streaks on the substrate were the most successful. Hence, multiple cues seem to be involved in male mating success in this species. Some males performed only one display, however, and may represent an alternate male mating tactic. The protein content of the adult male diet significantly influenced the level of pheromone streak deposition, and thus, foraging environment may affect the outcome of sexual selection.     

Changes in the haemocyte profile ofSpilostethus hospes (Fab) (Heteroptera: Lygaeidae) in relation to eclosion,sex and mating

Five types of haemocytes have been identified in the haemolymph ofSpilostethus hospes. Their morphology and micrometric measurements have been provided. Changes in the total and differential haemocyte population [total haemocyte count (THC) and differtial haemocyte count (DHC)] as well as in the absolute number of haemocytes in circulation have been assessed in relation to eclosion, sex and mating. The haemogram profile was studied prior to and immediately after eclosion and also prior to and after copulation. Though the THC was not significantly different immediately before and after eclosion, there was a significant increase in total count prior to copulation. Mated females registered an increase in total count but there was no appreciable change in the mated males. Granulocytes were the most abundant of the haemocyte types in both the sexes and mating caused a significant increase in the plasmatocyte count in females. Changes in the blood volume as well as the mitotic activity of the haemocytes is also discussed     

Mating Behavior, Sexual Selection, and Copulatory Courtship in a Promiscuous Beetle

The function of male movements during copulation is unclear. These movements may be a result of the necessary mechanics of insemination, or they may also have further function, for instance, stimulating or courting a female during mating, perhaps influencing female mate choice. We present data from three experiments exploring the mating behavior and copulatory movements of the highly promiscuous beetle Psilothrix viridicoeruleus. Male mating success in the struggle over mating was not related to male or female size (measured by weight) but successful males were more vigorous in terms of copulatory movements. These males took longer to mount females but copulated longer and remained mounted longer. We discuss these results in terms of the mating system of Psilothrix and also in terms of observations of the timing of insemination during copulation. We suggest that copulatory movements in this species are best understood as copulatory courtship.     

Oviposition and development response of perilla seed bugs (Nysius sp.) (Heteroptera: Lygaeidae) to five crop seeds

The perilla seed bugs, Nysius plebeius Distant and Nysius hidakai Nakatani, are emerging pests of perilla crop in Korea. Here, we investigated the life-history variables of N. plebeius and N. hidakai on five commonly grown crop seeds: perilla [Perilla frutescens (L.) Britt.], sesame (Sesamum indicum L.), sorghum [Sorghum bicolor (L.)], proso millet (Panicum miliaceum L.) and foxtail millet (Setaria italic L.). Bug oviposition, total developmental period, adult emergence, adult longevity and adult weight of both Nysius species were determined in both choice and no-choice tests. The oviposition and development periods of N. plebeius and N. hidakai varied significantly among seed species. The most preferred seed species for oviposition were perilla seeds, followed by sesame, sorghum and foxtail millet, whereas proso millet seeds were the least preferred. The bugs development period was faster in perilla seeds, with higher adult emergence, greater adult longevity and heavier adult weight. The higher oviposition preference and faster development with longer longevity of both Nysius species on perilla seeds is probably due to the physicochemical and nutritional attributes of the seeds. This information may enable the exploration of resistant genetic materials and chemical traits associated with seeds for breeding programme. Further, the outcomes of this study are discussed in terms of bug's behavioural response to crop seeds and management strategy of Nysius species as alternative to chemical management.     

Genital and body allometry in two species of noctuid moths (Lepidoptera: Noctuidae)

One‐size‐fits‐all and related hypotheses predict that static allometry slopes for male genitalia will be consistently lower than 1.0 and lower than the slopes for most other body parts (somatic traits). We examined the allometry of genitalic and somatic morphological traits in males and females of two species of noctuid moths, Spodoptera exigua (Hübner, [1808]) and Helicoverpa armigera (Hübner, [1808]). The relationship between genitalic traits and body size was generally strongly negative‐allometric in males but with no significant differences from 1.00 in females of the two species examined. However, in females, the slope of genital traits was also lower than the slopes for somatic traits. The relationship between somatic traits and the body size indicator was approximately isometric in most cases in males, except in four traits in S. exigua, in which the slopes showed slight negative allometry, and the hind tibia in H. armigera, in which the slope had positive allometry. However, in females, some somatic traits showed isometric and some other showed negative allometry in both species. The coefficients of variation (CV) for all structures in the males were low, not exceeding 10%. Genitalic traits showed significantly lower CV than somatic traits in males. In females, somatic traits showed lower CV than genitalic traits but with no significant difference in the H. armigera. Our observations of strongly negative allometry for genitalic traits in males are consistent with stabilizing selection on genital size and we suggest that male performance in interactions with females is the source of selection on male genital allometry. The difference in the degree of phenotypic variation between genitalic and somatic traits in the two studied species is attributed to the different developmental‐genetic architectures of these traits. Female genitalia showed a similar trend to the males, although the difference between genital and somatic traits was not significant in females. This finding suggests that selection is acting differently on male and female genitalia. Positive allometry of hind tibia in H. armigera may be a result of secondary sexual function.     

Functional morphology of the hind legs as weapons for male contests inLeptoglossus australis (Heteroptera: Coreidae)

Functional morphology of the hind legs as weapons in male contests was analyzed in the leaf-footed plant bug, Leptoglossus australis Fabricius. Measurement of some metrical tratis of the hind femur and tibia indicated that the weapon was the hind femur. Sexual dimorphism in the femoral length and width, and in the number of spines on the lower side of the femur, indicated that these parts play a significant role in male contests. It was also suggested that the length from the base to the widest part of the femur had a functional significance for male-male combat behaviors ofL. australis.