Mating systems in primitive Lepidoptera, with emphasis on the reproductive behaviour of Korscheltellus gracilis (Hepialidae)

Adults of Korscheltellus gracilis (Grote) are crepuscular. Flight is restricted to two 20–40 minute periods each day: the first flight follows sunset during evening twilight, the other precedes dawn. Four to five times as many adults are active during the evening when courtship, mating and oviposition occur. Of these behaviours, only oviposition occurs during the pre-dawn period of activity. Field observations indicate that females release a sex pheromone while fanning their wings. Females usually fly before pairing. There is no obvious female choice or stereotyped courtship behaviour prior to pairing.
In some hepialid genera females attract males, while in others ( Hepialus Fabricius, Phymatopus Wallengren, Sthenopis Packard and Zenophassus Tindale) the calling system is reversed, with males attracting females. Cladistic analysis of the Hepialidae suggests that the plesiomorphic condition for the family is for females to release the long-distance attractant, and that the reversed calling system represents a derived condition restricted to lineages within a single clade.
Available information on the mating systems of other basal lepidopteran lineages and Trichoptera support the hypothesis that ancestral Lepidoptera possessed a female-released, long-range attractant. Support for this position comes from (1) published literature; (2) field observations made by DLW; (3) the apparent absence of male scent structures and recognizable male calling behaviour among the pre-Neolepidoptera; and (4) cases of male antennal elaboration in several basal lepidopteran lineages.     

Circadian Reproductive Rhythms, Pair-Bonding, and Evidence for Sex-Specific Pheromones in Nadezhdiella cantori (Coleoptera: Cerambycidae)

Mating and oviposition rhythms and evidence for sex-specific pheromones of the gray-black citrus longicorn beetle Nadezhdiella cantori were investigated. The reproductive and flight activities of this insect mainly occurred between 1700 and 2400 h, and each activity had two peaks with the first one taking place within the 3 h before sunset (2100 h) and the second occurring within the 3 h after sunset. Circadian rhythms of mating and oviposition were significantly correlated but neither mating nor oviposition was significantly correlated with flight. It is suggested that mating and oviposition repeat in turn during pair-bonding and that flight is an indication of termination of reproductive activities in some individuals of a population. Males differentiated between sexes by touching. Mating behavior of males was stimulated by a contact female sex pheromone on the female body surface. Fighting behavior of males was mediated by a chemical on the male body surface. Females left trails of the contact sex pheromone on the trees when walking, which males used to locate them. Implications of these findings are discussed.     

Rhythmicity of mating and oviposition inCallosobruchus subinnotatus (Pic) (Coleoptera: Bruchidae)

Mating and oviposition behaviors were studied inCallosobruchus subinnotatus. Copulation was most frequent during the late scotophase, 2–3 h before onset of photophase. The females were less willing to mate during photophase, which increased the time to initiate mating while decreasing the duration of mating. Females exhibited increased movement prior to mating, resting immediately after mating, and remained stationary for 6 h when oviposition commenced. Multiple mating by both males and females affected the number of eggs laid, duration of mating, and uncoupling time at the end of mating. Females that mated two or three times laid more eggs than females that mated once or more than three times. Females that remainedin copula for less than 18 min showed greater readiness to remate than those that remainedin copula longer. There was a gradual decrease in the number of eggs females could lay with an increase in the number of previous matings by males.     

Ovipositional Phenology and Behavior of the Spruce Cone Fly, Strobilomyia neanthracina (Diptera: Anthomyiidae)

The behavior of Strobilomyia neanthracina Michelsen, a phytophage infesting spruce (Picea spp.) seed cones, was observed at a field site in northern Ontario and in cages in a greenhouse to investigate spatiotemporal aspects of mating, host location, and oviposition. In the field, adults emerged from 21 to 24 May 1996, which coincided with bud burst of Picea glauca (Moench) Voss seed cones. For 4 days following emergence, Strobilomyia flies could no longer be seen at a monitored P. glauca tree and may have been on a mating or dispersal flight. Subsequently, females but not males were seen again and the oviposition period of ca. 3 weeks began. Mating was observed only in the greenhouse, mostly (i.e., 65%) at age 5–9 days. Although copulations lasted 11–45 min, these females laid infertile eggs only, beginning at age 4 days. No sperm was found in the spermathecal capsules of females, suggesting that no sperm had been transferred during these copulations. In both the field and the greenhouse, ovipositional sequences that resulted in egg deposition occurred throughout the day but few sequences were observed before 1000, probably because flies were not very active at air temperatures below 14°C (most sequences occurred at 25–27°C). In the greenhouse, the typical ovipositional sequence lasted an average of 7 min and consisted of landing on the cone and examining it with the proboscis and sometimes the ovipositor, egg deposition, and postovipositional behaviors such as tapping (touching the cone surface with the flabellum ca. 5 times s ^–1 ), which possibly represents a host marking behavior. In the field, tapping was seen less frequently than in the greenhouse but occurred significantly (P = 0.014) more often after sequences that resulted in egg deposition than after sequences that did not. Eggs hatched after 4–5 days at 20°C. In the greenhouse, the median longevity of females and males was 24 and 17 days, respectively.     

Adult response of the almond seed wasp,Eurytoma amygdali, to chemicals from its host and certain nonhosts

Females of the almond seed wasp,Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae), responded in an olfactometer positively to odours from almond flowers and unripe fruits, but not to almond leaf odours and odours from flowers and unripe fruits of certain other nonhostPrunus species. Males responded to none of these odours. Extracts of undamaged unripe almond fruits (using ethanol, methanol, acetone, hexane, dichloromethane, or petroleum ether) stimulated female aggregation on glass surfaces treated with these extracts; in addition, certain fruit extracts (ethanol, methanol, or acetone) stimulated oviposition. Extracts of undamaged almond leaves (ethanol, methanol, or acetone) and flowers (ethanol or methanol) also stimulated female aggregation and oviposition. Aggregation and oviposition in response to an ethanol extract of almond fruits was intense in females aged 5 to 14 days and from 12∶00 to 18∶00h (photophase between 06∶00 and 20∶00). Certain almond fruit (ethanol, methanol, acetone or hexane) and flower extracts (ethanol or methanol) also provoked female response in the olfactometer. The results strongly suggest that certain chemical stimuli emanating from parts of the almond tree play a major role in host selection and oviposition. Some of the extracts tested may be a good source for the isolation, identification and synthesis of compounds stimulating attraction, aggregation and oviposition in nature. *** DIRECT SUPPORT *** A3414024 00003     

Sex pheromone components ofHeliothis maritima: chemical identification, flight tunnel and field tests

Three sex pheromonal components, (Z)-11-hexadecenal (Z11–16:Ald), (Z)-11-hexadecenol (Z11–16:OH), and hexadecanal (16:Ald), in a ratio of 88.0∶7.2∶4.8, were identified from ovipositor extracts of the fulvous clover moth,Heliothis maritima Grasl. (Lepidoptera: Noctuidae) by gas chromatographymass spectrometry. In addition, trace amounts of (Z)-9-hexadecenal (Z9–16:Ald) were detected in the extracts by GC. A blend of Z11–16:Ald, Z11–16:OH and 16:Ald in a ratio of 100∶6∶3, as well as in combination with 0.1 or 1 part Z9–16:Ald was tested at 0.01, 0.1, 1.0 and 10.0 μg doses in a flight tunnel. In flight tunnel tests male behavioral responses elicited by 0.1 or 1.0 μg doses of the 100∶6∶3∶1 blend were similar to those elicited by an ovipositor extract at 2 female equivalents. Deletion of Z9–16:Ald from the blend at 0.1 μg dose caused a decrease in the male response. In the field test, however, presence or absence of Z9–16:Ald did not significantly influence the number of males trapped in sticky traps with rubber septa containing 100 μg of the respective blends.     

Early outflying and late homeflying in the Indian pygmy bat under natural conditions

The outflying and homeflying activity pattern of a colony of the Indian pygmy bat Pipistrellus mimus occupying a tunnel was studied under natural conditions. Before leaving the tunnel for foraging, the bats made circling flights outside to sample the environmental light conditions. The onset and end of activity was related to the times of sunset and sunrise respectively. The onset of flight during evening commenced after sunset from mid-September to mid-April, before sunset during the rest of the year. Onset of activity occurred at higher light intensities compared to light intensities prevailing during the return flight to the roost in the morning. The duration of activity time showed a curvilinear relationship with the duration of the night. The phase relationship between onset and end of activity, and sunset and sunrise, showed marked seasonal variations in the values of Ψ_onset, Ψ_end, and Ψ_midpoint. However, such changes in the phase angle properties did not obey Aschoff's seasonal rule. Based upon the data obtained on the onset and end of activity patterns for five species of bats, including the data from this study, we report that P. mimus is the earliest to set out for foraging in the evening and among the last to return home in the morning. This might be due to dusk and dawn peaks in activity of the prey insects.     

Mating behavior ofXylotrechus pyrrhoderus Bates (Coleoptera: Cerambycidae) V. Female mounting behavior

Female mounting behavior was observed in the grape borerXylotrechus pyrrhoderus Bates (Coleoptera: Cerambycidae) under laboratory conditions. This behavior was directed towards either females (i.e., female homosexual mounting) or males. The female homosexual mounting was strongly elicited in females which had previously been allowed to fly. The frequency of this behavior was not related to the experience of mating. The investigation of the daily temporal pattern in female homosexual mounting activity indicated that the behavior most frequently occurred between 12∶00 and 16∶00, i.e., when females were searching for the pheromone-releasing males. Oviposition was also observed during this time interval. A detailed analysis revealed female mounting behaviors which are characteristic of male mounting behaviors, such as chasing conspecifics and synchronism of licking and abdominal bending. On the other hand, the frequency of male homosexual mounting was lower than that of females. Taken together with cases of other species previously reported, the expression of mounting behavior in females may be related to the evolution of the reproductive system, in which females approach their mates.     

Attraction of the sexes inFormica lugubris Zett (Hymenoptera: Formicidae)

Summary Sexuals ofFormica lugubris fly to mating places, where females attract males by using a sex pheromone. Females collected on the nest surface before departing on a mating flight are much less attractive than those collected on the mating place after the mating flight, suggesting that the mating flight triggers the release of the sex pheromone. Olfactory cues are essential for males to locate females while they patrol. Males probably use visual cues to locate females once they have alighted nearby them. Males are also attracted by aggregations of other males on the ground, probably because one or several females are likely to be close to male aggregations.     

Reproductive behaviour of female Chorthippus biguttulus grasshoppers

Female grasshoppers of acoustically communicating species assume series of reproductive states that are associated with particular behaviours. Studies on laboratory populations of Chorthippus biguttulus (L.) revealed that females of this species lack the period of ‘passive copulatory readiness’, increase their attractiveness to males by sound production and mate multiple times before their first oviposition. In particular, female Ch. biguttulus display a period of ‘primary rejection’ after their imaginal moult during which they reject male mating attempts followed by a period of ‘active copulatory readiness’ in which they produce acoustic signals and may copulate with courting males. Female stridulation generally stimulated male mating activity and stridulating females attracted more male mating attempts than mute females in the same cage, indicating that males preferentially court females that signal ‘active copulatory readiness’. After receipt of a spermatophore, Ch. biguttulus females displayed periods of ‘secondary rejection’ followed by re-establishment of ‘active copulatory readiness’. Acoustic responses of females to male songs, an indicator of reproductive readiness, were significantly reduced until 2 days after mating and remained slightly reduced in comparison to pre-mating levels. Some females mated multiple times before their first oviposition and cycled between ‘secondary rejection’ and ‘active copulatory readiness’.     

Influence of protein hydrolysate baits on oviposition in the apple maggot, Rhagoletis pomonella

Ovipositional responses of apple maggot (AM), Rhagoletis pomonella (Walsh), females were studied in the laboratory on apples (var: Golden Delicious) treated with different rates of four protein hydrolysate baits in choice and no-choice tests. Protein hydrolysate baits at rates of 0.5 and 1% had no significant effect, but oviposition was greatly reduced at higher rates of 5 and 10%. Apple maggot females exposed to apples treated with protein hydrolysate baits at a rate of 10% made 41–71% fewer punctures and laid 41–73% fewer eggs than in untreated control. No oviposition activity was shown on apples treated with 25 and 100% Nulure®. In no-choice tests the AM females laid 75–96% fewer eggs in apples treated with 10 and 25% Nulure compared to controls and no oviposition occurred in apples treated with 100% Nulure. Apple maggot females arrived in similar numbers on apples treated with 10% Nulure and untreated apples, but only 5% of those arriving on Nulure-treated apples showed ovipositor boring with no egg deposition while 60% of females arriving on untreated apples showed ovipositor boring activity and laid an average of 2.5 eggs per apple. In another experiment, individual AM females displayed similar behavioral responses to 10% Nulure-treated apples; none of the 56 females tested on treated apples displayed ovipositor boring activity, but 59% of the females (N=56) tested on untreated apples displayed ovipositor boring within 5 min of their arrival. Ninetyeight percent of AM females stayed and fed on fruit surfaces for 5 min on Nulure-treated apples without ovipositor boring compared to only 2% on untreated apples. Of the females that arrived on untreated apples, 39% flew away within 5 min without ovipositor boring compared to only 2% of those that arrived on Nulure-treated apples. Results of these two behavioral experiments suggest that upon arrival on a protein bait-treated apple, an apparent change of behavior occurs in AM females and instead of attempting to oviposit, they attempt to feed on fruit surfaces resulting in reduced oviposition activity. These results indicate that the feeding and oviposition-related activities of AM females are probably mutually exclusive and that the feeding behavior preempts oviposition activities on host fruits treated with higher rates of protein hydrolysate baits.     

Mating, cross-mating and related behaviours of Neoseiulus californicus and Neoseiulus fallacis (Acari: Phytoseiidae)

The pairings of Neoseiulus californicus (McGregor) and Neoseiulus fallacis (Garman) from western North America were monitored for tending by adult males, males in the mating position and oviposition and the activity of female deutonymphs and adults. The N. fallacis × N. californicus (♂ × ♀) tests had fewer males tending the deutonymphs but more in the mating position with new females than the reciprocal test. Afterwards, most of the females appeared gravid and approximately 20% produced an egg. Some eggs did not hatch but others became adult males, which mated with their mothers, but no eggs were produced. F1 males tended and mated with new N. fallacis females which had normal offspring. When held with new N. californicus females, F1 males tended the deutonymphs but were not seen mating and no eggs were laid. The pairings of N. californicus× N. fallacis had more males tending, less in the mating position and the females appeared non-gravid and produced no eggs. When same-species males were added to females held with F1 males for 15–20 days, normal levels and sexes of the progeny were produced. The female and male adults of N. fallacis were more active (ambulatory) than those of N. californicus. In within-species tests, the males had a high activity except while tending and mating, the female deutonymphs were inactive and the just mated females were more active than the ovipositing females. The timing of the tending and mating differed in the cross-pairings. Overall, these and other life-history data show that these two mites are distinct species, but that their males are promiscuous in tending and mating. This revised version was published online in November 2006 with corrections to the Cover Date.     

Male-biased sex ratios,female promiscuity,and copulatory mate guarding in an aggregating tropical bug,Dysdercus bimaculatus

The ecological and social bases of the mating system of the seed-feeding bug, Dysdercus bimaculatus(Hemiptera: Pyrrhocoridae), were studied in the lab and in aggregations at the host tree, Sterculia apetala(Malvales: Malvaceae), in Panama. On theoretical grounds, two factors are predicted to be of importance in determining the evolution of male mating tactics in Ms species: the operational sex ratio and the probability that undefended females will mate with other males, subjecting the gametes of deserters to sperm competition. Results of a study of a related species suggested that sperm displacement is probably substantial. Adult sex ratios at numerous sites were significantly male biased, and females whose mates were removed remated before oviposition (i. e., sperm utilization). These results predict that a mate defense tactic is likely to be superior to a nondefense tactic. The biological significance of the parameters is supported by observations that captive pairs often remained in copulafor several days, until just before oviposition. However, substantial variation in copulation duration was also observed, and possible causes of this variation are considered. Causes of male biased adult sex ratios were investigated by monitoring demographic changes within a single aggregation over 2 months. Both female juvenile and adult mortality rates were greater than male. In addition, dissections of reproductive adults showed that the flight muscles of females, but not males, had histolyzed, so that female reproduction is physiologically limited to a single site. Greater rates of immigration among both mature and young males suggests that an excess of males may also be found in the populations of bugs that subsequently colonize other host plants, so that female scarcity is typical of aggregations in all stages of development. The evolution of sex-limtied flight muscle histolysis may be explained by greater patchiness of females than males as mating resources, plus a lower energetic benefit/cost ratio of histolysis for males.     

Field observations of Ostrinia nubilalis eclosion and post‐eclosion activity of females around their natal plants

Abstract The early part of the post‐eclosion, pre‐mating period were examined under field conditions for Ostrinia nubilalis. Post‐eclosion behavior of 25 and 21 females during the first and second flight periods were observed until they left their natal site. Summer generation larvae were reared under field conditions and the timing of adult eclosion was observed. Eclosion occurred at two times during the day, peaking before dawn and before dusk; 46% of females and 56% of males eclosed during the morning period and the rest eclosed during the evening period. After eclosion, females spent 30–60 min expanding their wings. Their typical behavior was to remain calmly on their natal site. None of the females exhibited calling behavior before leaving. All females left their natal sites sometime before dawn. The probability of leaving increased with time. Leaving rates were not significantly different between females of the first and second flight. These field observations indicate that females have several possibilities for pre‐mating movement, which might allow females to move out from their natal field before mating. In addition, we also discuss the influence of pre‐mating movements in relation to the rate of Bacillus thuringiensis (Bt) resistance evolution.     

Field observations of mating,oviposition, and feeding behavior ofAnticarsia gemmatalis (Lepidoptera: Noctuidae) in a soybean field

The temporal occurrence and frequency of mating, oviposition, and feeding of wild velvetbean caterpillar moths, Anticarsia gemmatalis,in a soybean field (Glycine max)were documented. Mating occurred predominately during the first 4 h after sunset and primarily on leaflet bottoms. Oviposition was concentrated within the first 6 hr after sunset. Eggs were laid singly, and in areas of high trichome density, on leaflets, pulvini, and petioles. Feeding occurred primarily at night, with most males feeding during the first half of the night and females during the second half. Males and females shared some food sites but others were visited strictly by males.     

Effect of mating frequency on fecundity and longevity of the predatory mite <Emphasis Type="Italic">Kampimodromus aberrans</Emphasis> (Acari: Phytoseiidae)

The effect of single versus multiple mating on longevity and fecundity as well as the number of matings required to maximize a female’s reproductive success of the predatory mite Kampimodromus aberrans Oudemans were studied under laboratory conditions. Newly emerged adult females of the stock colony of K. aberrans were placed individually on a bean leaf disc, and maintained at 25°C and 16:8 LD. A young male remained with a female for limited periods or continuously. Mating was a requisite for oocyte maturation and oviposition. Females which mated three to four times during their life and females in continuous presence of males, laid significantly and considerably more eggs than single-mated females. Virgin females lived the longest, and those in continuous presence of males the shortest. In all cases and irrespective of the number of matings, the sex ratio of the offspring was male-biased in the first three to four days of oviposition period, and female-biased in later days.     

Behavioural switch of female Mediterranean fruit fly, Ceratitis capitata: mating and oviposition activity in outdoor field cages in Hawaii

1 Laboratory-reared normal, and wild female Mediterranean fruit flies, Ceratitis capitata (Wiedemann), were assayed in outdoor field cages to assess the impact of a mating-induced behavioural switch on mating and subsequent oviposition activity. 2 Virgin females preferred interactions with males leading to mating over attraction to, and oviposition in, artificial yellow spheres containing guava odour or green apples hung in a guava tree. Laboratory-reared females previously mated with either laboratory-reared normal males or laboratory-reared irradiated (sterile) males showed little interest in remating with males and instead, were much more likely to be found arrested on artificial and real fruit and ovipositing. Oviposition on artificial fruit was five times greater by females that had mated with either normal or irradiated males than by virgin females. Wild females showed similar qualitative changes in the mating-induced behavioural switch; however, oviposition activity was significantly less than for laboratory-reared females. 3 These results confirm that mating has a profound effect on the behaviour of female Mediterranean fruit flies and that irradiated males are functionally equal with normal males (lab-reared or wild) in their ability to alter female behaviour. These results are discussed in the context of the sterile insect technique for control of Mediterranean fruit flies in the field.     

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.     

Oviposition behaviour in Paysandisia archon (Lepidoptera: Castniidae)

Paysandisia archon (Burmeister, 1880) (Lepidoptera: Castniidae) is a South American insect developing on palms and recently introduced in Europe where it damages most palm species. Understanding the oviposition behaviour would be decisive for risk assessment and pest management but key points on oviposition behaviour are missing. Using wind tunnel and field experiments, we investigated the oviposition timing, the attraction behaviour of mated females to palms and the different steps of oviposition behaviour. Results showed that oviposition behaviour occurred between 3 pm and 6 pm. In the field, gravid females were significantly more attracted by the palm crown than virgin females. The ovipositing females exhibited distinctive steps before ovipositing. Subsequent to alighting on the crown, pre-oviposition behaviour was characterized by two main behavioural steps: walking and probing the surface with antennae and ovipositor. After the choice of oviposition place, the gravid female remains motionless and the extendible ovipositor is deeply introduced into the upper fibrous part of the crown. About 10 eggs can be lays at the same place. Finally, the female starts to walk again and reinitiates the same behavioural sequences. This paper supports the hypothesis that odours from the crown may play a key role in gravid female attraction. The study assessed that P. archon lays on the palm crown, the part of the palm that should be treated for population monitoring.     

Effects of mating rates on oviposition,sex ratio and longevity in a predatory mite <Emphasis Type="Italic">Neoseiulus cucumeris</Emphasis> (Acari: Phytoseiidae)

Two aspects of mating effects on the fecundity, sex ratio and longevity of Neoseiulus cucumeris (Acari: Phytoseiidae) were examined in laboratory experiments: (1) females mated by one, two or three different males (unmated and 3 days old) at 5-day intervals, and (2) females mated by males with different age/mating status (number of females mated previously by the male). Females allowed to mate with a second or third male at 5-day intervals produced 39 eggs on average, but those mated with a single male produced 28 eggs on average. Matings with additional males 5 or 10 days after the first male increased the duration of the oviposition period of these females by 5–7 days and at the same time reduced the post-oviposition period by about 10 days. Overall, females with additional matings by one or two different males at 5-day intervals survived a few days shorter than females without additional males. Mating with a different female each day, a male of N. cucumeris could mate with 5–8 females, which produced a total of 85–116 eggs: females mated with a male during days 1 and 2 in its adulthood and with a male of the last 2 days of life (days 7 and 8) produced about half as many eggs as females mated with a male during 3–6 days of its adulthood. Females mated with males that are too young or too old had a shorter oviposition period and a longer post-oviposition period and longevity than females mated with middle-aged males. In both experiments, rates of oviposition remained similar in females with high or low fecundity. This indicates that in both cases, the increased fecundity is due to the extension of the oviposition period through additional sperm supplied by the second male and or third male (in experiment 1) or more sperm by males not too young nor too old (experiment 2).