Mating Behavior of Trissolcus basalis (Wollaston) (Hymenoptera: Scelionidae): Potential for Outbreeding in a Predominantly Inbreeding Species

The mating behavior of the quasi-gregarious egg parasitoid Trissolcus basalis (Wollaston) was investigated under field conditions. Trissolcus basalis has female-biased sex ratios and is a protandrous species, with males emerging 1–2 days before females. Males competed aggressively for control of the egg mass, with one male assuming dominance and control of the egg mass, although changes in dominance occurred at least once on each egg mass observed. Typical mating behavior involved the dominant male mating his sisters immediately upon their emergence from the egg mass. These behaviors are characteristic of an inbreeding species that manifests local mate competition. However, several aspects of the mating behavior of T. basalis are inconsistent with that of an inbreeding species. Over 18% of emerging females were not mated by the dominant male upon emergence, 13% of females were not observed to be mated at all and may have left their natal site as virgins, 25% of females were mated multiple times and sometimes by multiple males, females remained near the natal site for up to several hours after emergence before emigrating, and males dispersed away from the natal site during female emergence. Trissolcus basalis may be a predominantly inbreeding species but its emergence and mating behavior suggest that low-frequency outbreeding is also likely to occur.     

Uscana lariophaga, West-African egg parasitoid of the cowpea bruchid beetleCallosobruchus maculatus: photoperiod, parasitization and eclosion interactions

The diurnal pattern of parasitization and eclosion of the trichogrammatidUscana lariophaga Steffan, egg parasitoid of the cowpea bruchidCallosobruchus maculatus (Fabricius) was studied for L12:D12 photoperiods. The wasp parasitized eggs throughout each 24-h period with about 70% of the parasitization taking place during the first 12 h regardless whether it was the photophase or the scotophase. More female progeny was produced when the first 12-h period was photophase instead of scotophase. Eclosion of wasps took place over a period of 4 days and occurred during the second half of the scotophase and the first half of the photophase. The number of wasps eclosed during the photophase was similar to that eclosed during the scotophase. Average development time was 8.9 days at 30°C. Male development was completed 6 to 8 h before the females, hence a higher percentage of females emerged in the later eclosion peaks. During scotophases more females eclosed than during photophases. Although the results indicate that the wasp is able to perform under dark storage conditions the effect of permanent low light intensity remains to be studied.     

Hibernation adaptation and eclosion synchrony in leaf-galling sawflies in subarctic Norway

The winter dormancy adaptation in gall-inducing sawflies is poorly known. Diapause termination and the following post-diapause quiescence enhance synchronous eclosion in spring. This is probably the most critical part in the life history in gall-inducing sawflies, as there is only a short phenological window of opportunity for mating and oviposition. In a 2?years’ study, diapause duration, termination, survival and eclosion synchrony were experimentally investigated for three gall-inducing sawfly species (Symphyta: Tenthredinidae: Pontania nivalis, P. glabrifrons and P. arcticornis). Field-collected galls, sampled from willows (Salix spp.) in early autumn, were kept under natural temperatures outdoors until next spring. Subsamples were successively transferred to the laboratory at 14-day intervals from October to April in a test of development time to eclosion and survival ratio. The time to eclosion decreased throughout the experiment, whereas the proportion successfully eclosed and eclosion synchronicity increased, all indicative of prepupae entering a diapause in early autumn. The diapause terminates midwinter, and the prepupae enter a post-diapause quiescence until the temperature in spring allows a direct development and contributes to a nearly synchronous eclosion. In all three species, males eclosed 1–2?days prior to females (protandry). We hypothesize that synchronous eclosion as well as protandry enhance mating and oviposition success. Our finding indicates that gall-inducing sawflies are well adapted to its harsh subarctic and arctic environment.     

Interrelation of mating,flight, and fecundity in navel orangeworm females

The navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae, Phycitini), is an economically important pest of nut crops in California, USA. Improved management will require better understanding of insect dispersal, particularly relative to when mating occurs. A previous study demonstrated a more robust laboratory flight capacity compared to other orchard moth pests, but it was unclear how mating affects dispersal, and how dispersal affects fecundity. In this study, 1‐ and 2‐day‐old females were allowed to fly overnight on a flight mill either before or after mating, respectively, and were then allowed to oviposit. Data on fecundity were compared between treatments to minimally handled or tethered‐only control females. Females that mated before flight flew longer and covered a greater distance than those flying prior to mating. However, timing of flight relative to mating did not affect fecundity, nor did any measure of flight performance. There was no effect on fecundity when females were forced to fly for designated durations from 3 min to 2 h. Together, our data revealed no obvious trade‐off between flight activity and reproductive output. Distances measured on the flight mills (mean ca. 15 km for mated females) may overestimate net displacement in the field where flight tracks are often meandering. The results suggest that most females mate and oviposit in or near their natal habitat, but that some may disperse potentially long distances to oviposit elsewhere.     

Daily rhythms in the sorghum shootfly, Atherigona soccata: oviposition, egg-hatch and adult eclosion

ABSTRACT. Daily rhythms controlling oviposition, egg-hatching and adult eclosion in the sorghum shootfly, Atherigona soccata Rondani, were investigated. Eggs were laid only during the photophase of a LD 12:12 cycle, in two peaks. Under continuous light, this oviposition was considerably attenuated but not made immediately arrhythmic. Egg-hatching and adult eclosion both commenced just before dawn. Some feature of the scotophase during or immediately after black-head formation apparently acts as a signal for hatching. Eclosion was controlled by light but its timing in the field was modified by temperature. The last 2–3 days of the pupal period constituted the most sensitive stage, and light signals received during this period determined the time of eclosion. Ecological advantages of these rhythms to the shootfly are discussed.     

半闭弯尾姬蜂羽化、交配及产卵行为观察

在22℃,相对度60％～70％的实验室条件下,研究了半闭弯尾姬蜂Diadegma semiclausum的羽化、交配与产卵行为。结果表明：半闭弯尾姬蜂蜂蛹从开始羽化到羽化结束历经4d,羽化高峰期出现在羽化的第3d,羽化数量为25头,占全部羽化数量的58．82％;第3d中羽化高峰出现在8：00～13：00,羽化数量为16．5头,占全天羽化数量的66．00％。半闭弯尾姬蜂交配过程大致分为3个阶段：准备阶段、交配阶段、结束阶段;交配后开始产卵,产卵过程大致分为寄主寻找和确定、穿刺和产卵、产卵结束和梳理。     

The dawn flight of the gold swift Hepialus hecta: predator avoidance and the integration of complex lek behaviour (Lepidoptera,Hepialidae)

Predator pressure is a considerable evolutionary force. The evening twilight flight of species in the family Hepialidae Swift moths has been attributed to the moths (which have no hearing and so cannot detect bats) flying in a bat/bird free window. Several species deploy elaborate lek behaviour in this period. The expected flight in the dawn twilight is little reported, except in the non‐leking Hepialus (Korscheltellus) gracilis. A detailed study of the dawn flight in the leking species Hepialus (Phymatopus) hecta shows that it is less extensive than the evening lek flight (both in duration and in the number of moths participating), is confined to a much narrower window around sunrise, involves no reproductive behaviour, and functions only to re‐locate the members of copulating pairs and sessile displaying males from the emergent ground vegetation to less conspicuous roosting sites. Compilation of individual biographies over the 24‐h cycle permits a full construction of the diel activity in time and space. It consists of two twilight flight periods, separated by prolonged roosting during the hours of darkness and daylight. The moths use almost the whole available range of roosting sites, from the base of the ground vegetation to the tree canopy, and cyclically leave and re‐enter the lek site from these positions. The copulating posture and position avoid mammalian predators, and facilitate escape from spiders and wasps by the efficient use of a dead drop. The narrowness of the dawn flight is attributed to the need for this prolonged but conspicuous copulation, which precludes a morning mating, and to a demonstrated asymmetry of twilight activity in birds. The whole integrated spatio‐temporal cycle is attributable to evolutionary pressure to minimize the impact of predators, demonstrated to include birds, bats, wasps, and spiders, and probably also dragonflies, mice and shrews; the pre‐existing dawn flight is permissive to the evolution of sessile male displays during the evening lek. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 305–319.     

Decision-making conditions for intra- or inter-nest mating of winged males in the male-dimorphic ant <Emphasis Type="Italic">Cardiocondyla minutior</Emphasis>

Only winged male and female ants generally mate through nuptial flight during the reproductive season. In the ants of Cardiocondyla, the males show wing dimorphism and their reproductive strategies differ depending on the differences in wing morphology. It has been suggested that wingless “ergatoid” males bearing very similar external morphologies to workers mate within natal nests, whereas winged males bearing typical ant male morphology disperse from their nests to mate. However, some behavioral observations suggest that the winged males of some Cardiocondyla ants such as C. obscurior and C. minutior may mate within natal nests before dispersion. We evaluated the factors affecting the mating behaviors of the winged males of C. minutior under laboratory conditions. We found that (1) the winged males remained and mated with virgin females in natal nests when either virgin winged females or the relatively mature pupae of winged females (i.e., at least 10 days) were present in the nest, (2) the winged males dispersed to adjacent nests with virgin winged females when only mated queens and the relatively young pupae of winged females (i.e., <9 days) were present in the nest, and (3) all winged males were accepted by the workers of non-natal nests irrespective of the distance from the natal nests in the field. Although most ergatoid males were accepted by the workers of close non-natal nests, they were all attacked and killed by the workers of distant non-natal nests. These results suggest that intra-nest mating and the dispersion of the winged males of C. minutior are facultatively determined by the condition of winged females (virginity and relative pupal age) in natal nests. Furthermore, our results suggest that winged males are likely to seek mating partners chemically and to mate with virgin winged females.     

Diel flight periodicity of Chilo partellus

The diel flight periodicity of the nocturnal moth Chilo partellus (Swinhoe) (Lepidoptera;Pyralidae) was measured in the laboratory using an actograph and in the field with suction traps. Females showed almost no flight activity on the night of eclosion. Flight activity of mated females peaked before midnight, the period of peak oviposition activity. Male peak activity occurred after midnight coinciding with female eclosion. Presence or absence of females did not affect when or how long males were active. Data on flight activity and reproductive behaviour are discussed in relation to the use of pheromones to protect maize.     

The Use of the Vibration Signal and Worker Piping to Influence Queen Behavior during Swarming in Honey Bees, Apis mellifera

We investigated worker regulation of queen activity during reproductive swarming by examining the rates at which workers performed vibration signals and piping on queens during the different stages of the swarming process. Worker–queen interactions were first examined inside observation hives during the 2–3 wk that preceded the issue of the swarm (pre‐swarming period) and then inside the swarm clusters during the period that preceded liftoff and relocation to a new nest site (post‐swarming period). Queen court size did not differ between the pre‐ and post‐swarming periods, but workers fed the queens less inside the swarm clusters. Workers performed vibration signals on the queens at increasing rates throughout the pre‐swarming period inside the natal nest, but rarely or never vibrated the queen inside the swarm. Piping was performed on the queens during both the pre‐ and post‐swarming periods and always reached a peak immediately before queen flight. During the final 2–4 h before swarm liftoff, queens were increasingly contacted by waggle dancers for nest sites, some of which piped the queen. The vibration signal may operate in a modulatory manner to gradually prepare the queen for flight from the natal nest, and the cumulative effects of the signal during the pre‐swarming period may make further vibrations on the queen unnecessary when inside the swarm cluster. In contrast, worker piping may function in a more immediate manner to trigger queen takeoff during both the pre‐ and post‐swarming periods. Workers that vibrate and pipe the queen tend to be older, foraging‐age bees. The regulation of queen activity during colony reproduction may therefore be controlled largely by workers that normally have little contact with queens, but help to formulate colony reproductive and movement decisions.     

Relationship of flight and reproduction in beet armyworm, Spodoptera exigua (Lepidoptera: Noctuidae), a migrant lacking the oogenesis-flight syndrome

The beet armyworm, Spodoptera exigua, undertakes long-distance migration. We used flight mills to investigate the interaction between flight and reproduction in this species given the apparent absence of the oogenesis-flight syndrome. This syndrome, common in many migratory insects, is characterized by a suite of traits including migration during the pre-oviposition period followed by a switch to oogenesis. No negative effects of inter-ovipositional flight on lifetime fecundity were observed. Instead, adult reproductive output suffered when female flight was initiated the first day after eclosion and before oviposition, suggesting that migratory flight overlaps with the oviposition period rather than being confined to the pre-oviposition period. Mating status of both females and males had no negative influence on their flight performance except that flight distance and flight duration of 7-day-old mated females were significantly less than in unmated females. Furthermore, the number of eggs produced and mating frequency of females less than 7 days of age were not significantly correlated with flight performance, suggesting reproductive development paralleled and was independent of migratory behavior. This independent relationship between flight and reproduction of adults is consistent with the very short pre-oviposition period in this species, and suggests that resources are partitioned between these activities during pupal development. Together, our results uncovered neither obvious trade-offs nor mutual suppression between flight and reproduction in S. exigua, which indicates the lack of an oogenesis-flight syndrome for coordination of these two energy-intensive processes. We propose a conceptual model of migration for this species based on the current and previous studies.     

Adaptive strategies of overwintering adults: Reproductive diapause and mating behavior in a grasshopper,Stenocatantops splendens (Orthoptera: Catantopidae)

Abstract To understand the adaptive strategies of the overwintering adults of Stenocatantops splendens, the mechanism of maintenance and termination of the reproductive diapause, the variation in mortality between overwintering females and males, and the mating strategy of the males were investigated. The results indicated that the adult reproductive diapause in natural conditions was mainly regulated by photoperiod in the fall – long photoperiods promoted reproductive development and short photoperiods maintained reproductive diapause, and the sensitivity of the overwintering adults to photoperiod was over before the end of the winter. When transferred from natural conditions to controlled laboratory conditions on dates from September through February, pre‐oviposition became increasingly shorter with increasingly deferred transfer dates regardless of photoperiod conditions. The adults treated with low temperature for 30 days in September through November had significantly shorter pre‐oviposition, suggesting that low temperatures in winter had an important role in the termination of reproductive diapause. The female had a significantly lower supercooling point than the male, which was related to their lower mortality after winter. In addition, observations of wild populations of the species indicated that mating behavior prior to winter and the duration of pre‐mating period were not affected by photoperiod; mating and sperm transfer were mostly completed by November. Compared with females only mating before winter, females mating in the spring had shorter life span, longer pre‐oviposition, lower hatching rate and laid fewer egg pods while showing no significant difference with regard to ovipositional interval, per pod number of eggs and nymph dry weight.     

Flight activity of three Spodoptera spp., Spodoptera litura, S. exigua and S. depravata, measured by flight actograph

Flight activities of three Spodoptera species were measured by the aid of flight actograph: S. litura and S. exuiga being regarded as long‐distance migratory insects, and S. depravata being non‐migratory and diapause‐inducible species. In all species tested, flight activities were observed only in scotophase, males showed far higher activities than females, being several times higher at the time of maximum flight activity, which was observed within 2 days after adult eclosion. Total flight activity in males was highest in S. litura, some being flyable even 12 days after eclosion, followed by S. exigua being one‐third compared to the former species, while in S. depravata flight activity was nearly half of that of the second species and most ceased to fly within a week after eclosion. There occurred species‐specific daily rhythms in flight activity during respective scotophase. In S. litura, both females and males exhibited a peak of flight activity shortly after light‐off and exhibited the second flight activity in late scotophase, the females slightly but the males more actively compared to early scotophase. In S. exigua, both sexes did not respond to light‐off, did not show a peak of flight activity in early scotophase, whereas males, but not females prominently increased activity toward the end of scotophase. In S. depravata, both sexes exhibited a peak of flight activity in early scotophase, and the males revived flight activity, being maximum shortly before light‐on, but the females did not show a clear rhythm in flight activity. These features observed in flight activity were discussed in relation with migratory capability.     

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     

Increasing the availability of food and the reproduction ofPoecilus cupreus in a cereal field by strip-management

The spruce bud moth,Zeiraphera canadensis Mutt & Free. (Lepidoptera: Tortricidae), displayed a bimodal activity pattern in two young white spruce,Picea glauca (Moench) Voss, plantations. During the day, when temperatures were highest and relative humidities lowest, most moths remained in the lower crown, presumably in response to the risk of hygrothermal stress and/or predation. Flight activity peaks occurred near dawn and dusk. Moths moved higher in the crowns shortly after sunset and many were observed flying above the tree canopy. This activity peak was associated with oviposition and mating. Oviposition started at sunset and continued for approximately 3 h. Females probed oviposition sites with their ovipositor before ovipositing, suggesting that they use sensillae on the ovipositor to evaluate potential oviposition sites. Most mating occurred between 22∶00 and 04∶00 h. Males hovered adjacent to branches containing females before alighting. After alighting, males usually approached to within 2–5 cm of females and then remained motionless for a short time before attempting mounting, indicating that some short-range communication usually occurs prior to mating. Reduced flight activity between midnight and dawn was not due to low temperatures, which were above the minimal temperatures for flight for males (10°C) and females (12°C) determined in the laboratory.     

Mating behavior and thermoregulation of the reindeer warble fly,Hypoderma tarandi L. (Diptera: Oestridae)

Hypoderma (=Oedemagena) tarandi L. (Diptera: Oestridae) is characterized by a mating strategy in which both sexes meet and mate at two types of distinct topographical landmarks. In the expansive, treeless vidda (= tundra-like) biome, mating places are unique, rocky areas located along rivers and streams or in rocky areas of drying river and stream beds. In wooded valleys below the vidda, flies mated at certain topographical areas along dirt road tracks/paths. Thermoregulatory activities of males occupying perches at mating places included selection of substratum at perch site, orientation of body to sun's rays, crouching, stilting, and flights into upper cooler air. On warm sunny days males perched for just 1–2 min before flying up into cooler air to promote cooling. Laboratory and field studies revealed that flies could not metabolically cool down when held at 25–38°C. Time spent at mating places depended on temperature, duration of sunshine, and wind velocity. Males were very aggressive in pursuing allHypoderma-sized objects that passed by them or that landed near them, but they did not defend specific perch sites. Males either pursued and caught females in flight, or they hopped onto females that landed near them. During 5 years, 74 males and 14 females were seen at mating places. Dissection of six females caught at mating places revealed them to be recently eclosed flies full of fat body and with all eggs intact; two not paired with males were non-inseminated. Three experimentally paired females remainedin copulo for 10, 13, and 19.5 min.     

Local Mating,Dispersal and Sex Ratio in a Gregarious Parasitoid Wasp

Parasitoid sex ratios can be greatly influenced by mating and dispersal behaviour. Many sex ratio models assume that mating is strictly local (only mated females disperse from the natal patch) and that a single male is sufficient to inseminate all females in a brood. Bethylids (aculeate parasitoids) have been used to test predictions of these models, but less attention has been paid to testing their underlying assumptions. We investigated the timing of eclosion, mating and dispersal in mixed-sex and single-sex broods of the bethylid wasp Goniozus nephantidis. In mixed-sex broods, almost all females mate before dispersal and a single male is sufficient to inseminate virtually all females, even when brood sizes are large. Males disperse from both mixed-sex and all-male broods, but males in all-male broods disperse more slowly. Virgin females disperse from all-female broods, which are common. Virgin females can produce a brood, mate with their own sons and subsequently produce mixed-sex broods, but their success rate is very low. Virgin females could potentially circumvent sex allocation constraints by superparasitizing mixed-sex broods, but when presented with hosts bearing mixed-sex broods they destroy all members of the initial brood before ovipositing. Because of the high prevalence of single-sex broods and dispersal of both sexes, the mating structure of G. nephantidis is unlikely to conform to the assumption of strict local mating.     

Reproductive behavior of the honeydew moth,Cryptoblabes gnidiella

Summary

The reproductive behavior of the honeydew moth, Cryptoblabes gnidiella (Millière) (Lepidoptera: Pyralidae), was studied in the laboratory. The sex ratio was 1.1:1, males to females, in both laboratory and field stocks. Most of the females that mated did so during the first night after emergence; males began mating on the following night. Mating occurred 1–2 h before dawn and averaged 100 min. Both sexes mated only once in one night. Most females mated only once in their lifetime, a few mated 2–4 times, whereas males mated up to six times per lifetime. Insects that lived longer also mated more times. When the sex ratio was altered from 3:1 to 1:3, males to females, the percentage of females that mated in one night dropped from 90 to 65, whereas the number of matings per male rose from 0.32 to 2.25. When fresh one-day-old females were provided daily at a ratio of three per male, the males averaged 1.4 matings per lifetime vs. 2.6 with 2- to 3-day-old females. A delay in mating did not affect the percentages of males and females that mated; highest percentages were obtained with 2- to 4-day-old males and females, but a delay in mating resulted in egg fertility dropping from 91 % to 73 %. The preoviposition period lasted a full day after mating, and then most of the eggs were laid during the first night. Average fecundity was 105 eggs per female (maximum: 230).     

Differences in the pre‐diapause and pre‐oviposition accumulation of critical nutrients in adult females of the beetle Colaphellus bowringi

In many insect species, the differentiation of development between diapause and reproduction first becomes obvious during the diapause preparation (pre‐diapause) and pre‐oviposition phases. However, the differentiation of nutrient accumulation between these two phases remains unclear. We compared the weights of pre‐diapause and reproductive adult female Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), and measured their triacylglycerol (TAG), protein, and carbohydrate content from emergence until they had fed for 4 days post‐eclosion. We also compared the ovarian development and accumulation of lipid droplets between pre‐diapause and reproductive adult females in order to determine whether we could visually detect differences in nutrient allocation. The weights of both pre‐diapause and reproductive females increased with duration of feeding. The fresh weight and water content of pre‐diapause females was significantly lower than that of reproductive females after feeding for 3 days post‐eclosion. Pre‐diapause females channeled their reserves into TAG in the fat body, whereas reproductive females converted nutrients into proteins and carbohydrates for egg development. These results quantify differences in nutrient accumulation between pre‐diapause and reproductive adult female C. bowringi, and provide clues for understanding the molecular mechanisms underlying differences in the allocation of nutrients between diapause and reproduction in insects.     

Group changing in free-ranging male rhesus monkeys

Group changing behavior of maleMacaca mulatta was studied over a six-year period at the rhesus monkey colony on two coastal islands at La Parguera, Puerto Rico. Males first left their natal group at a mean age of 47 months and became solitary for the first time at a mean age of 64 months; all had left their natal groups by seven years of age. Age, mating season, sex ratios of adult males and females in the social bands, and geographical barriers all had significant effects on the group shifting. Population size, rank of mother or being an orphan did not significantly affect the changing process. Two factors, age (size) and seniority in the group, were important in determining a male's rank in his new group.