Behavioral patterns that determine the mating rates in a wing dimorphic riffle bug,Microvelia horvathi Lundblad, 1933 (Hemiptera: Heteroptera: Veliidae)

Journal of Ethology - Wing dimorphic insects are known to be a well-established model of the trade-off between flight capability and reproduction. Macropterous morph expends its energy for...     

Cost of mating in male Menochilus sexmaculatus (Fabricius) (Coleoptera: Coccinellidae)

Mating and reproduction not only provide benefits but can also incur substantial costs. An increasing number of recent studies have indicated that mate searching, mating, and ejaculate production requires large amount of energy which can result in high cost to males. Here, we examine the effect of number of matings of males and mating duration on mating and reproductive success in Menochilus sexmaculatus (Coleoptera: Coccinellidae) by subjecting them to 3 consecutive matings of 30, 60, 90 min, and complete mating (a) with no interval and (b) with interval of 24 hr between the matings. The results reveal a reduction in reproductive performance for both the setup and reduction is more when mating is consecutive without gap. This shows that between subsequent matings males require time to recuperate the sperm/ejaculate. As a male engages in a greater number of matings, the time it takes to begin mating when paired with a female increases and the overall duration of mating decreases. However, frequent mating attempts made by males result in reduction in their body weight. Thus, in this ladybird beetle mating and related physiological process consume lots of energy and the body weight of males tend to decrease which further modifies their reproductive performance. The study helps us to identify the cost of mating from male's perspective and indicates that males need to rejuvenate between the matings.     

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

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

转Bt基因水稻对两种弹尾虫及尖钩宽黾蝽捕食作用的影响

转Bt基因水稻KMD1、KMD2和对照水稻XS11稻田主要有两种弹尾虫：灰橄榄长角跳虫 Entomobryagriseoolivata (Packard) 和钩圆跳虫 Bourletiella christianseni Banks。两种Bt稻田中灰橄榄长角跳虫种群密度均显著高于对照XS11稻田;在以KMD1和KMD2腐烂 茎叶为食的灰橄榄长角跳虫成虫中检测到微量Cry1Ab杀虫蛋白。室内测定结果表明,不管是单 头捕食还是多头协同捕食,尖钩宽黾蝽Microvelia horvathi Lundblad 成虫对用3种供试 水稻残体饲养的灰橄榄长角跳虫的捕食量和功能反应均符合HollingⅡ型方程,其日捕食量、瞬 时攻击率（a）和处理时间（T_h）均无显著差异。     

Relative male and female contributions to the supercooling point of their offspring in Microvelia reticulata (Heteroptera: Veliidae)

Cold hardiness is a key life history trait in temperate and polar ectothermic species, as it affects survival during overwintering, but its evolution is poorly understood. While many studies of cold hardiness in insects have shown differences between species, populations or developmental stages, data on the relative contribution of individual genotypes to cold hardiness are scarce and mainly limited to drosophilid fly species. We used a sib‐analysis (paternal half‐sib/full‐sib breeding design) to estimate the relative contributions of parental generation to the supercooling point (SCP) of the offspring of a heteropteran non‐model insect species, Microvelia reticulata (Heteroptera: Gerromorpha: Veliidae). We found that parent identity affected significantly SCP values of their offspring. Magnitudes of estimated sire and dam variance components were similar but the distributions of individual contributions to SCP differed between sires and dams, which points to sex‐specific genetic or parental effects on SCP in this species. The animal model failed to find a realistic estimate of heritability (h²) of SCP, suggesting that the underlying genetics of SCP in M. reticulata can not be characterized by purely additive effects.     

Remating, sperm transfer, and sperm displacement in the cactophilic species Drosophila buzzatii Patterson & Wheeler (Diptera: Drosophilidae)

The mating system of Drosophila buzzatii is characterized by short copulation duration, frequent remating in both males and females, and male ejaculate partitioning. Additional features of the system are strong sperm displacement and a high frequency of sterile matings. Remating frequencies and the effects of remating on various mating parameters were studied. In order to characterize variation, five isofemale lines from geographically distant localities in Australia (three localities), Brazil and the Canary Islands were used. Mating parameters studied were: premating time, copulation duration, interval between successive matings, and progeny number as a measure of sperm transfer. Variation for sperm displacement was studied in crosses between laboratory stocks and a number of isofemale lines from Australia. There were significant between‐line differences in female remating frequencies, premating time, copulation duration, interval between successive matings, and progeny numbers, indicating genetic variation for these traits. Females from the five lines mated on average 1.6 to 3.1 times in 4 h, with a maximum of eight matings for one female. The males were given a maximum of ten virgin females in sequence and more than one‐third of the males mated all ten females in the 2 h observation period. Copulation duration decreased and interval between matings increased with copulation number in multiply mated males. Mean copulation duration was c. 2 min. Sperm transfer, measured as the average number of progeny from a single mating, was low (c. 25) and multiply mated females gave more progeny than single mated females, although with much lower progeny numbers than observed in wild‐caught non‐virgin females. A surprisingly high proportion of observed matings gave no progeny, i.e. they were sterile matings. Sperm displacement was strong in most crosses and remained strong in multiply mated females. The results are discussed in relation to the evolution of mating patterns in Drosophila.     

Postmating sexual selection: the effects of male body size and recovery period on paternity and egg production rate in a water strider

The role of male body size in postmating sexual selection wasexplored in a semiaquatic insect, the water strider Gerris lateralis.To separate effects of male size per se from those due to numericsperm competition, male recovery period (shown here to be proportionalto ejaculate size) was manipulated independently of body sizein a factorial experiment where virgin females were mated firstwith sterile males and then with focal males. Both relativemale fertilization success and female reproductive rate were measured.The number of sperm transferred increased with male recoveryperiod, an effect that was mediated by longer copulation duration,but there were no effects of body size on ejaculate size. Neithermale size nor recovery period had any significant direct effectson male fertilization success. However, copulation durationinfluenced relative fertilization success, suggesting that malesable to transfer more sperm also achieved higher fertilizationsuccess. Females exercised cryptic female choice by modulatingtheir reproductive rate in a manner favoring large males andmales that were successful in terms of achieving high relativefertilization success. Thus, successful males gained a twofoldadvantage in postmating sexual selection. This study has important implicationsfor previous estimates of sexual selection in this group of insectsbecause pre- and postmating sexual selection will be antagonisticdue to limitations in male sperm production: males mating frequently(high mating success) will on average transfer fewer sperm ineach mating and will hence tend to fertilize fewer eggs permating (low fertilization success).     

Variability in the life history of the water strider Gerris lacustris (Heteroptera: Gerridae) across small spatial scales

Abstract.  1. Variability in life-history traits of the common pond skater Gerris lacustris (L.) (Heteroptera: Gerridae) was investigated on a small geographical scale in Bavaria, Germany (≈ 49 °N). In an area of approximately 3.5 × 3.0 km, voltinism and wing length data of G. lacustris were recorded from 65 ponds located either in an open field or surrounded by forest.
2. Life history differed significantly between populations in the two habitat types. Gerris lacustris showed a bivoltine life-cycle in most of the field ponds with a high proportion of long-winged individuals in the diapause generation and a large increase of the short-winged morph in the midsummer generation. In contrast, almost all populations inhabiting forest ponds were univoltine. Only a few forest populations produced any second generation larvae. The long-winged morph dominated during the whole season in forest ponds.
3. The appearance of first-instar larvae was delayed by 3–4 weeks in forest ponds compared with field ponds. Presumably, low temperatures led to this reduction in the developmental rate of eggs and larvae. Consequently, 'field larvae' of the first generation developed under increasing daylight conditions (triggering direct reproduction and brachyptery) but 'forest larvae' developed under a shortening photoperiod (triggering diapause and macroptery).
4. Offspring raised from 'forest parents' in the laboratory showed the same variability in life history when reared in a cool (16.9 ± 2.4 °C) or warm (23.1 ± 1.8 °C) temperature regime.
5. Given the small spatial scale of the study, it is concluded that the variability in the life history of G. lacustris has a strong phenotypic component.     

Life history adaptation along a latitudinal cline in the water strider Aquarius remigis (Heteroptera: Gerridae)

We addressed the general hypothesis that life history differences among eastern populations of the North American water strider, Aquarius remigis (Heteroptera: Gerridae), along a north-south gradient are manifestations of genetic differentiation due to natural selection. We raised offspring of two field-caught populations from each of three latitudes in a common laboratory environment at 20° C and two photoperiods. Nearly all Quebec (PQ) individuals (ca. 46° N) entered diapause to reproduce the following spring (univoltine life cycle), while intermediate proportions of New York (NY; ca. 43° N) and New Jersey (NJ; ca. 41° N) individuals reproduced directly, producing a second generation (bivoltine life cycle). PQ females were smaller, developed faster, and laid smaller eggs than NY and NJ individuals; NY and NJ populations differed little in these variables. NY females had longer life spans than either PQ or NJ females, but lower oviposition rates. Total reproductive output did not differ across latitudes. Photoperiod affected body length, development time, and reproductive pathway, resulting in a latitude by environmental interaction. PQ individuals reproduced directly under 15L : 9D (summer) conditions only, while the NY and NJ populations exhibited more direct reproduction under 13L : 11D (spring or fall) conditions. Some life history characters of the NY and NJ populations displayed the higher variability indicative of phenological transition zones. These results indicate local adaptation of populations to long-term climatic patterns. Water striders appear to adapt to longer seasons by extending development, growing larger, and breeding directly. Larger body size and extended or rapid development are associated with bivoltinism and increase in egg size, but not necessarily with higher fecundity or oviposition rate. The phenological transition zone appears to be unrelated to a transition zone a little further south established by allozyme data and morphology, as all populations studied here could be electrophoretically identified as northern "type".     

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.     

Copulation, the dynamics of sperm transfer and female refractoriness in the leafhopper Balclutha incisa (Hemiptera: Cicadellidae: Deltocephalinae)

Abstract.  Mature sperm of the leafhopper Balclutha incisa (Matsumara) (Cicadellidae: Auchenorrhyncha: Hemiptera) are stored as a series of sperm bundles within seminal vesicles prior to ejaculation. During transfer, sperm are pumped from the vesicles into the ejaculatory duct to the complex aedeagus. Sperm transfer is marked by a c . 30-fold expansion of the spermatheca to accommodate both sperm and seminal fluid. Sperm number increases exponentially with male age, reaching a maximum of 700 000 after 14 days, while the number of sperm available on days 2–5 is between 70 000 and 100 000. During mating, maximum sperm transfer occurs after 7 min and mating is complete after about 10 min. Ejaculate size, defined by both sperm and associated accessory gland fluid, is influenced by male mating status and the interval since the previous mating. There is a positive correlation between duration of copulation and both ejaculate and the time to subsequent mating. Sperm are more likely to be retained in the testes during mating by males of 2–5 days post-emergence than older males. The number of sperm received by the female can be manipulated experimentally by mating males once (medium ejaculate) or twice (small ejaculate) immediately after their first mating. Females that receive small ejaculates from sperm-depleted males have a far shorter refractory period than females receiving medium to large ejaculates. Both ejaculate size and the time after males have mated influence the female post-mating refractory period as measured by the female's responsiveness to male sexual signalling.     

Morphology and neurophysiology of tarsal vibration receptors in the water strider Aquarius paludum (Heteroptera: Gerridae)

Substrate vibratory information receptors are extensively studied in insects and spiders, however for water surface dwelling species little data is available. We studied the vibration receptive organs in tarsi of the water strider Aquarius paludum, using light, transmission and scanning electron microscopes, and recorded the neural activity of the organs in response to vibrational stimuli, which were afterwards analysed with a custom made spike sorting program.We found that the tarsal chordotonal organ has one set of three scoloparia: one in the tarsomere I and two in the tarsomere II, all of which consisted of a few scolopidia. The chordotonal organ clearly responded to vibratory stimulation. Furthermore, we found that a pair of large subapical emergent dorsal setae, which had been deemed mechanosensory by previous authors, are not so. In turn, four ventral subapical trichobothria that are in direct contact with the water surface during locomotion, proved to be mechanosensory. The anatomical and ultrastructural observations support these electro-physiological results.     

Infection rates and pathogenicity of trypanosomatid gut parasites in the water strider Gerris odontogaster (Zett.) (Heteroptera: Gerridae)

Summary Trypanosomatid flagellates are common protozoan gut parasites of a wide range of insect species. Water striders (Gerridae) harbour the trypanosomatid Blastocrithidia gerridis. Three different populations of the water strider Gerris odontogaster in northern Sweden were sampled to assess the infection rate dynamics of trypanosomatids. The initially very low infection rates (0%–15%) early in the season were followed by a rapid increase during the reproductive period of the water striders, reaching very high levels (80%–90%). The pathogenic effects of trypanosomatids on G. odontogaster adults were studied in laboratory experiments. The parasites caused a general reduction of host vigour. Male skating endurance was negatively correlated with the intensity of the trypanosomatid infection. However, infection of trypanosomatids increased the mortality among adults only when the water striders were subjected to food stress. The trypanosomatids did not reduce the fecundity of females provided with food. We suggest that trypanosomatid gut parasites may be an important mortality factor in water strider populations. Since the pathogenicity of the parasites is enhanced by stress, parasitism by trypanosomatids may contribute to the regulation of host populations.     

Avoidance of egg parasitism through submerged oviposition by tandem pairs in the water strider, Aquarius paludum insularis (Heteroptera: Gerridae)

Abstract.  1. Females of the water strider Aquarius paludum insularis (Motschulsky) (Heteroptera: Gerridae) carry males on their backs and oviposit under water after copulation. This study focuses on the benefit  A. paludum insularis receives by ovipositing in tandem.
2. Males guarded females in tandem through to the end of oviposition in 85% of copulations. Females in tandem dived deeper than single females, and the density of A. paludum insularis eggs increased with water depth. The proportion of eggs parasitized by a scelionid wasp, Tiphodytes gerriphagus Marchal (Hymenoptera: Scelionidae) decreased with increasing water depth.
3. These results suggest that during oviposition guarding by males is beneficial for females, because it enables pairs to dive and lay eggs deeper and in oviposition sites where the risk of egg parasitism is lower.     

Decoy presentations as a means to manipulate the risk of extrapair copulation: an experimental study in a semicolonial raptor, the Montagu's harrier (Circus pygargus)

Mate guarding and frequent copulations are two alternative paternity assurance strategies found in birds. In species with intensecourtship feeding, like raptors, the "frequent copulation"strategy is expected because male food provisioning conflictswith mate guarding. We evaluated experimentally the paternityassurance behavior of a semicolonial raptor, the Montagu'sharrier Circus pygargus, using decoy presentations to simulateterritorial intrusions. Breeding pairs were exposed to maleand female decoys at different periods during the female's reproductive cycle. Agonistic responses to decoys were intra-sexual,and the timing and intensity of male attacks toward male decoyssupported responses related to the risk of extrapair copulation(EPC): Male aggression peaked during the presumed fertile periodand almost disappeared after clutch completion. During thefertile period, copulation rate was significantly higher, andcopulations lasted longer, during male decoy presentations than during controls. Males also spent more time close to the femaleduring male decoy presentations compared to controls, bothduring the early prelaying and fertile periods, but not duringincubation. In the fertile period, males also increased presencetime close to the female in the hour following the removal of the male decoy. Conversely, female decoy presentations hadno significant effect on copulatory behavior or male presencetime. These results showed that the risk of EPC can be experimentallymanipulated by the means of decoy presentations, simulatingmale territorial intrusions, and that male Montagu's harriersincrease their short-term copulation frequency and female surveillancewhen they perceive themselves at an increased EPC risk.     

Phylogeny of the water strider genus Gerris Fabricius (Heteroptera: Gerridae) based on COI mtDNA, EF-1α nuclear DNA and morphology

Phylogenetic relationships between water striders (Heteroptera: Gerridae) of genus Gerris Fabricius were examined using molecular and morphological characters. The molecular dataset was 820 bp DNA from the 3′ half of the mitochondrial gene encoding cytochrome oxidase subunit I and 515 bp DNA from the nuclear gene encoding elongation factor 1 alpha. The morphological dataset was a slightly modified version of a previously published dataset. Representatives from all eight recognized species groups of Gerris, as well as six species from three related genera, including Gigantometra gigas, Limnoporus esakii, L. rufoscutellatus, Aquarius najas, A. conformis and A. paludum, were included. Unweighted parsimony analyses of the COI sequences gave a topology with strong support for only those nodes that were already recognized as closely related based on morphological characters. Similar analyses of EF‐1α gave a cladogram with a topology quite different from that based on morphology and COI. Unweighted parsimony analyses of the ‘total evidence’ dataset largely supports the traditional view of Gerris phylogeny. Finally, the implications of the reconstructed phylogeny in relation to biogeography and ecological phylogenetics of Gerris is discussed.     

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.     

Segregation products of male mice doubly heterozygous for the RB(6.16) and RB(16.17) translocations: Influence of sperm karyotype on fertilizing competence under varying mating frequencies

The meiotic segregants of male mice heterozygous for Rb(6.16)24Lub and Rb(16.17)7Bnr were viewed, for the first time, at first cleavage metaphase. Chromosomes were analyzed after G-banding, C-banding, and karyotyping. To study sperm aging effects, chromosomes of 202 one-cell zygotes derived from males mating at intervals of approximately 3,14, and 21 days were examined. At least 89.6% of sperm-derived complements were products of 2:2 segregation; at most, a possible 6.4% were 3:1 segregants. The six expected types of 2:2 segregants, both balanced and unbalanced, were equifrequent in the total zygote population derived from sperm of all ages. When the data were analyzed according to mating frequency, the 3-day sperm population considered most likely to be fresh showed a deficiency of the segregant nullisomic for chromosome 6 and disomic for chromosome 17, when compared with the reciprocal segregant (P < 0.025) as well as to all other 2:2 segregants (P < 0.05). However, these sperm fertilized in greater numbers (P < 0.01) than their reciprocal segregant (disomic for 6 and nullisomic for 17) in the 14-day sperm population. While sperm with chromosomal abnormalities are capable of fertilization, the competence of segregants nullisomic for 6 and disomic for 17 apparently depends on the prior storage period in the male. Further, the results suggest that the effect of aneuploidy on sperm function is dependent on the specific chromosome(s) involved.