Female pheromonal chorusing in an arctiid moth, Utetheisa ornatrix

We report an unusual case of communal sexual display in thearctiid moth Utetheisa ornatrix that we designate "female pheromonalchorusing." As in most moths, female U. ornatrix release a long-distancesexual advertisement pheromone during a nightly activity period.We arranged U. ornatrix females in 2 types of signaling conditions:grouped and solitary. When the females were grouped with neighboringsignaling females (grouped), they initiated pheromone releasesooner, continued release with less interruption and over alonger total period, and performed the release with faster abdominalpumping than observed in isolated females (solitary). This differsfrom the usual form of sexual communication in moths: female(chemical) signalers, male receivers, and a general lack ofinteraction among females. At mating, male U. ornatrix transfera large spermatophore that may enhance female reproductive successand which represents either mating effort or paternal investment.This action results in an extended postmating male refractoryperiod leading to a female-biased operational sex ratio. Weargue that this biased sex ratio generates intrasexual competitionamong females, to which they respond by elevating signalingeffort such that the likelihood of at least matching their neighbors'signals is increased. In the field, U. ornatrix are clusteredaround patches of host plants, and we also explore the possibilitythat pheromonal chorusing is driven by cooperation among groupsof related—or nonrelated—females.     

Preference for high concentrations of plant pyrrolizidine alkaloids in the specialist arctiid moth Utetheisa ornatrix depends on previous experience

Secondary metabolites are one the most pervasive defensive mechanisms in plants. Many specialist herbivores have evolved adaptations to overcome these defensive compounds. Some herbivores can even take advantage of these compounds by sequestering them for protection and/or mate attraction. One of the most studied specialist insects that sequesters secondary metabolites is the arctiid moth Utetheisa ornatrix. This species sequesters pyrrolizidine alkaloids (PAs) from its host plant, the legume Crotalaria spp. The sequestered PAs are used as a predator repellent and as a mating pheromone. We used this species to test larval preference for different concentrations of PAs. We purified PAs from plant material and added them at different concentrations to an artificial diet. Larvae of U. ornatrix previously feeding on low and high PA concentration artificial diets were allowed to choose between two new artificial diets with different PA concentrations. The amount of PAs sequestered and larval preference were dependent on their previous exposure to low or high PA content in the diet. Larvae that were pretreated with a low PA diet significantly consumed more diet with the high PA concentration, while larvae that were pretreated with a high PA diet showed no discrimination between future feeding of different PA concentration diets. We discuss our results using mechanistic and evolutionary approaches. Finally, we discuss how these results have important implications on the evolution of plant herbivore interactions and how specialist herbivores may decrease the levels of chemical defenses on plant populations.     

Fate of an alkaloidal nuptial gift in the moth Utetheisa ornatrix: systemic allocation for defense of self by the receiving female

In the moth Utetheisa ornatrix defensive pyrrolizidine alkaloids are sequestered by both sexes as larvae from their foodplants. The adult male transmits some of this alkaloid to the female at mating for eventual incorporation into the eggs. We now show by chemical analyses that the female herself is the first beneficiary of the alkaloid she receives from the male. By the end of mating the male's alkaloid is found already to be generally distributed throughout the female's body (including even the wings), while it is still largely absent from the ovaries. This result is in line with our earlier finding that the male's alkaloidal gift can itself suffice to render a female fully protected against spiders from the moment she uncouples from the male.     

Responses of olfactory receptor neurons in Utetheisa ornatrix to gender-specific odors

We recorded the electrophysiological responses of individual olfactory receptor neurons in sensilla trichodea on the antennae of adult arctiid moths, Utetheisa ornatrix, to stimulation with volatiles associated with both sexes. All trichoid sensilla contain at least two receptor neurons, each with distinct action potential amplitudes and waveforms, that respond dichotomously to male and female odors. Although, neither female neuron responds to extracts of coremata or the male-produced pheromone hydroxydanaidal, they do respond in a gender-specific manner to the volatiles emanating from whole pupae, hemolymph, thoracic froth, and adult animals of several ages. Thoracic froth, which contains pyrrolizidine alkaloids, is thought to play a role in defense. Froth from moths reared on diets, with or without added pyrrolizidine alkaloids, were equally effective in eliciting gender-specific patterns of response. Male trichoid receptor neurons respond to these same materials with similar patterns of activation. These receptor neurons provide information about substances, which we have termed “gender odors,” that are persistently emitted by nearby animals. These substances do not appear to be the same as those already known to be involved in defense or the sexual dialog between individuals. Accepted: 16 March 2000     

Parental Body Mass as a Determinant of Egg Size and Egg Output in an Arctiid Moth (Utetheisa ornatrix)

Females of the moth Utetheisa ornatrix (family Arctiidae) mate preferentially with larger males. As a consequence, females have larger sons and daughters, which have been shown to be more acceptable in courtship and more fecund, respectively. We demonstrate that mating with larger males results in accelerated oviposition by the female on the day after mating and that females are intrinsically prone to lay larger eggs in the first days after mating. Both these additional size-dependent effects have potential adaptive consequences.     

Influence of host plants on development, fecundity and egg hatchability of the arctiid moth Diacrisia casignetum

Measurements were made of the differences between sunflower and climbing hempweed leaves for development, fecundity and egg hatchability of Diacrisia casignetum Kollar. Developmental time and weight gains of sunflower-fed larvae were respectively longer and greater than those of hempweed-fed ones. The development time and weight gain, attributable to differences in host plants, were statistically heterogeneous (P<0.05-<0.10). Relative fecundity and egg hatchability of sunflower reared insects were greater than those of hempweed reared ones, and these were also statistically heterogeneous (P<0.1 and <0.05, respectively). Hempweed leaves contained relatively high titres of phenols and low ratios of proteins to phenols. Ovaries were heavier in insects reared on sunflower than on hempweed, whereas the level of phenols incorporated into the ovary was higher in insects fed on hempweed than those on sunflower. The differences in these parameters may be due to variations in nutritional quality of these hosts and incorporation of ingested phenolic materials to ovary and insect biomass.

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Résumé L'étude a porté sur l'évaluation de l'influence des feuilles de tournesol (Helianthus annuus) et de Mikana cordata sur le développement, la fécondité et l'éclosion des oeufs de D. casignetum Kollaz. La durée de développement et le gain de poids sur tournesol étaient plus importants que sur M. cordata. La durée de développement était statistiquement hétérogène (P<0.05), ce qui peut être dû à des différences entre plantes. La fécondité et le taux d'éclosion chez les insectes élevés sur tournesol étaient plus élevés que ceux observés sur M. cordata, dont les résultats étaient aussi hétérogènes (P<0.1 et<0.05 respectivement). Les feuilles de M. cordata contenaient des taux relativement élevés de phénols et des rapports protéines sur phénols bas. Tout ceci suggère que les différences observées sont attribuables à des variations dans la qualité nutritive de ces hôtes et dans l'incorporation à la biomasse de l'insecte de matériaux phénologiques ingérés.

     

Effect of male and female multiple mating on the fecundity, fertility, and longevity of diamondback moth, Plutella xylostella (L.)

Abstract:  The effect of diamondback moth (DBM), Plutella xylostella (Lep., Plutellidae) male and female multiple mating on fecundity, fertility, and longevity was studied. Males could mate for five times with virgin females during scotophase. The successful copulation rates, fecundity of female, and longevity of both females and males decreased when male mating times increased, whereas copulation duration increased. Correlation coefficient between copulation duration and male mating times was significant ( r  = 0.7358, P = 0.0001, spearman rank-order correlation). There were linear relationships between mating history of males and longevities of males and females, and regression relationships between them were significant. Mated females had similar daily reproductive pattern, which laid the most eggs on the first day after mating in spite of their mates' mating history. Virgin females laid some infertile eggs before they died. Most of the females mated once during their lifespan but 19.9% of females mated twice when one female kept with one male during scotophase. There were no significant differences in the fecundity, fertility and longevity between the single- and twice-mated females. Correlation coefficient between copulation duration and female mating times was not significant ( r  = 0.0860, P = 0.8575). Results suggested that DBM females may be monandrous. Multiple mating did not increase male or female mating fitness.     

Effects of delayed mating on the fecundity,fertility and longevity of females of diamondback moth,Plutella xylostella

Abstract The effects of delayed mating on the copulation duration, female fertility, fecundity, egg fertility, longevity and the number days alive after mating of females of diamondback moth (DBM), Plutella xylostella, were studied. When male mating was delayed, the female fertility, fecundity, egg fertility, longevity and number days alive after mating of DBM decreased, and there was a negative correlation between the age of the moth with those variables except copulation duration. When female mating was delayed, the female fertility, fecundity, percent egg fertility and number days alive after mating of DBM also decreased, but the longevity increased, which also showed a negative relationship between the age of the moth with the variables except copulation duration and longevity. When both males and females delayed mating, the female fertility and fecundity decreased; egg fertility was affected marginally, and the longevity of females increased. The moth age was negatively correlated with those variables.     

Acoustic response to a pheromonal cue in the arctiid moth Pyrrharctia Isabella

ABSTRACT. The acoustic properties of the clicks emitted in response to male courtship pheromone by female Pyrrharctia Isabella (J. E. Smith) (Lepidoptera: Arctiidae) have been investigated. Extracts of male scent organs or synthetic male pheromones can be applied to a glass rod and used to stimulate females to produce these sounds. Power spectra, sound pressure readings, and oscillographic analyses show that the acoustic signals elicited by male extracts or synthetic male pheromones are not distinguishable from those produced in response to disturbance (handling). This is the first reported example of a sound produced by a female moth in a sexual context, and is the first reported example in moths of an acoustic response to a pheromonal stimulus.     

East Atlantic teleconnection pattern and the decline of a common arctiid moth

Teleconnection patterns are large‐scale atmospheric circulation systems and variation in them is often associated with impacts on climate and weather over broad areas. Arctia caja L. is a well‐known, widespread and charismatic tiger moth. In recent decades, the abundance of A. caja in UK has fallen abruptly. The annual abundance of A. caja in UK is known to be affected adversely by wet winter weather and warm spring temperatures. We examined A. caja population dynamics from 1968 to 1999 for weather and climatological effects. Population growth rate displayed endogenous effects of abundance in the previous two seasons. Accounting for this, growth rate in the present season was still affected significantly by winter precipitation and spring temperature. Annual abundance of A. caja was inversely related to winter East Atlantic teleconnection pattern (winter EA index) and annual population growth rate was inversely related to winter EA in the present and previous two seasons. An index of the North Atlantic Oscillation (NAO), commonly used as an indicator of winter climate in northern Europe, did not show a significant relationship with growth rate. We noted, for the first time, that the winter EA index has increased steadily over the past five decades. The model presented here therefore implies a further decline of A. caja population growth rates and abundance in the future. This is the first demonstration of a relationship between EA and population dynamics and indicates the EA and other lesser‐known teleconnection patterns may prove useful in modeling the ecological effects of climate change.     

Aerial application of pheromones for mating disruption of an invasive moth as a potential eradication tool

Biological invasions can cause major ecological and economic impacts. During the early stages of invasions, eradication is desirable but tactics are lacking that are both effective and have minimal non-target effects. Mating disruption, which may meet these criteria, was initially chosen to respond to the incursion of light brown apple moth, Epiphyas postvittana (LBAM; Lepidoptera: Tortricidae), in California. The large size and limited accessibility of the infested area favored aerial application. Moth sex pheromone formulations for potential use in California or elsewhere were tested in a pine forest in New Zealand where LBAM is abundant. Formulations were applied by helicopter at a target rate of 40 g pheromone per ha. Trap catch before and after application was used to assess the efficacy and longevity of formulations, in comparison with plots treated with ground-applied pheromone dispensers and untreated control plots. Traps placed at different heights showed LBAM was abundant in the upper canopy of tall trees, which complicates control attempts. A wax formulation and polyethylene dispensers were most effective and provided trap shut-down near ground level for 10 weeks. Only the wax formulation was effective in the upper canopy. As the pheromone blend contained a behavioral antagonist for LBAM, 'false trail following' could be ruled out as a mechanism explaining trap shutdown. Therefore, 'sensory impairment' and 'masking of females' are the main modes of operation. Mating disruption enhances Allee effects which contribute to negative growth of small populations and, therefore, it is highly suitable for area-wide control and eradication of biological invaders.     

Functional organization of the arctiid moth tymbal (insecta,lepidoptera)

The exoskeletal morphology, muscular organization, and innervation patterns of the tymbals of seven sound-producing species of tiger moths (Arctiidae) were compared with the undifferentiated episterna of two silent species. At least three muscles are involved in sound production: the tymbal muscle, pv2, and the accessory muscles, pvl and/or pv6. All of the tymbal muscles are innervated by the IIIN2a branch of the metathoracic leg nerve, which contains two axons larger than the others. Backfills of the tymbal branch of the IIIN2a reveal a medial sensory neuropil and a population of five ipsilateral motor neurons whose somata are clustered into three groups along the anterior edge of the metathoracic ganglion. The dendritic arborizations of the motor neurons extend to the ganglionic midline but are separate from one part of the auditory neuropil observed in other noctuoids. The study concludes that the arctiid tymbal reveals only minor modifications (e.g., cuticle thinning) of the episterna of silent moths and represents a primitive form of the tymbal compared to those of the Cicadidae.     

Effect of mating status on the fecundity of a cricket,Teleogryllus emma

Abstract Fecundity in some insects is affected by mating status. The effect of mating status on the fecundity and total egg production of Teleogryllus emma (Ohmachi et Matsumura) (Orthoptera: Gryllidae) was examined in this study. The results showed that the pre‐oviposition period was shorter for amphigonic females than that for virgin females. However, no significant difference in pre‐oviposition was found between amphigonic females and those that had mated with a male with either the phallodeum or testes extirpated. There is no difference in adult longevity between the above four groups. The fecundity and total egg production were much higher in amphigonic females than in those controlled under the three non‐amphigonic treatments. The females of T. emma that mated with the testes‐extirpated males produced more eggs (up to two‐fold more) than both the virgin females and those that mated with the phallodeum‐extirpated males, but there was no difference between them. The fecundity‐enhancing substances transferred from male to female can stimulate the female to produce more eggs, but this stimulation has to occur in collaboration with sperm.     

Reproductive biology and function of multiple mating in the mating system of a tree cricket,Truljalia hibinonis (Orthoptera: Podoscritinae)

A tree cricket,Truljalia hibinonis, is known to show a novel sperm removal during copulation. The pattern of copulations and ovipositions showed that the sperm removal functioned to increase reproductive success for sperm removing males. The sperm removal by males evolves under the system in which female accept multiple mating. The possible benefits of multiple mating for females are examined. Multiple mating did not seem to be necessary for avoiding sperm depletion, because females stored huge number of sperm in their sperm storage organ after finishing oviposition. The ingestion of metanotal secretion during copulation also had no effect on increasing fecundity and egg size. However, mating experience may have a positive effect on increasing fecundity slightly, though there were no differences between once- and twice-mated females. The other possible benefits for each male and female are discussed.     

Pyrrolizidine alkaloid deters ant predators of Utetheisa ornatrix eggs: effects of alkaloid concentration,oxidation state,and prior exposure of ants to alkaloid-laden prey

To examine the chemical defense of lepidopteran eggs towards ant predators, eggs of the arctiid moth Utetheisa ornatrix were offered to laboratory colonies of the ant Leptothorax longispinosus. The ants rapidly devoured Utetheisa eggs produced by parents reared on an alkaloid-free diet, but left eggs that had been endowed with parental alkaloid largely unmolested. That defense can be attributed directly to the presence of a pyrrolizidine alkaloid since topical application of monocrotaline in either its free base or N-oxide state conferred protection on otherwise palatable eggs. The N-oxide and free base were not equally effective in conferring protection however; free-base treated eggs were subject to greater predation than N-oxide treated eggs after 24 h. The emergence of differential alkaloid effectiveness only after prolonged exposure is consistent with the notion that chemical protection is seldom absolute. That difference was resolved only when the otherwise food-deprived ants exceeded a certain threshold of hunger. The efficacy of applied monocrotaline as a predator deterrent increased with increasing concentration of N-oxide, but was lowest for eggs treated with an intermediate concentration of free base. The latter concentration effect likely represents an artefact of the behaviour of the ant colonies offered eggs treated with an intermediate concentration, as those colonies were the most voracious predators in reference tests with palatable eggs. In addition to the immediate deterrent value of pyrrolizidine alkaloid, ant colonies that had been exposed to alkaloid-laden eggs subsequently avoided even palatable, unprotected eggs offered 33 days later. Our data provide the first demonstration of such long-term avoidance of chemically protected lepidopteran prey by an invertebrate. The discovery of a mechanism promoting learned avoidance on the part of ant predators has important ramifications to egg-laying strategies of female Utetheisa, both in terms of the dispersion of eggs, and the extent to which eggs are provisioned with alkaloid.     

Interactions between size and temperature influence fecundity and longevity of a tortricid moth,Zeiraphera canadensis

Females of Zeiraphera canadensis Mut. & Free., the spruce bud moth, were reared in the laboratory at constant and alternating temperatures, and in an outdoor insectary, to (1) determine the effects of temperature, age and size on several reproductive parameters and, (2) to test the hypothesis that body size-temperature interactions influence longevity and realized fecundity. Egg maturation was linearly related to age and large moths developed eggs at a higher rate than small ones. Mcan lifetime oviposition rate reached a maximum and remained stable at temperatures 20° C while the mean lifetime rate of egg maturation increased linearly with temperature, indicating that higher temperatures adversely affect oviposition. The production of nonviable eggs increased with age but also with temperature, suggesting high temperature (25° C) reduces egg quality and/or hinders fertilization. The realized fecundity and longevity of females reared under an alternating temperature regime (mean 20° C) was significantly less than that of females reared at constant 20° C. Similar realized fecundity, longevity and mean lifetime oviposition rates for females reared at temperatures alternating between 10 and 25° C (mean 20° C) and those at constant 25° C reflected the inability of females to recover from elevated diurnal temperatures. Longevity was positively related to female body size at constant 15 and 20° C but the relationships were negative for moths exposed to diurnal temperatures equal to or exceeding 25° C. Due to the reduced longevity of large moths at high temperatures, linear regressions between size and realized fecundity were only significant at constant temperatures 20° C. At higher temperatures, the size-fecundity relationship became curvilinear as a result of the diminished reproductive output of large individuals. Reduced fecundity and longevity of large females at high temperatures may have been due to elevated internal temperatures of large-bodied moths. Large females in a controlled-environment chamber maintained at 25° C developed an internal temperature excess (i.e. temperature above ambient) of nearly 2° C while small-bodied females exceeded ambient by only 0.3° C. However, when held at 20° C, the temperature excess of large-bodied moths was much less than 1° C and small-bodied females did not differ from ambient. Such interactions between temperature and body size suggest that there should be stabilizing selection toward moderate-sized individuals and may explain the absence of size-related effects on fecundity and longevity previously reported for several other lepidopterans.     

Asymmetry as a predictor of growth, fecundity and survival

Measures of developmental stability such as fluctuating asymmetry have been assumed to predict individual performance because asymmetry reflects an inability to cope with stressful situations, and because asymmetry hampers locomotion. However, the magnitude of this relationship between important fitness components (growth, fecundity, survival) and asymmetry has never been assessed. Based on a literature survey, estimates of the correlation between asymmetry and the three fitness components are presented. Pearson's correlation coefficients weighted for sample size between asymmetry and growth, fecundity, and survival, respectively, were –0.15, –0.35, and –0.25, respectively, with all three coefficients being highly significant. All three relationships were extremely robust given very large fail-safe numbers. The results were independent of whether studies or species were used as units of analysis. Hence asymmetry is a robust predictor of performance in fitness domains such as growth, fecundity, and survival, although only accounting for 2.1%, 12.3%, and 6.0% of the variance. This may be of importance for studies of sexual selection, but also for ecological and conservation biological studies, where the performance of individuals or groups of individuals are assessed.     

Effect of multiple mating on the reproductive performance of the rice leaffolder moth,Cnaphalocrocis medinalis (Lepidoptera: Crambidae)

To clarify whether multiple mating of females and males affects the reproductive performance of the rice leaffolder moth, Cnaphalocrocis medinalis (Guenée), we examined the effect of the number of matings (once, twice, or three times) for females (female treatment) and males (male treatment) on the incidence of moth mating, number of eggs laid, egg hatchability, and adult longevity. We also compared the effect of multiple mating imposed on males or females separately with the effect of that imposed on both sexes simultaneously (both sexes treatment). The incidence of mating of females and males that mated three times (3-mated females and males) was significantly lower than for females and males that mated twice or once (2-mated or 1-mated females and males). The incidence of mating of 1-mated moths (both sexes) was significantly higher than for 2-mated or 3-mated moths (both sexes). Two-mated or 3-mated females laid significantly more eggs with significantly higher hatchability than 1-mated females. Females that mated with 1-mated males (second male mating) or 2-mated males (third male mating) laid significantly fewer eggs than those that mated with virgin males (first male mating). Females laid significantly more eggs after the second and third matings for moths of both sexes than after the first mating for moths of both sexes. The mechanisms of improvement and decline of female reproductive performance when multiple mating was imposed on males or females are also discussed in relation to the reproductive biology of C. medinalis.     

Gonadotropin release as a function of mating and steroid feedback in the female rat

The aim of this investigation was to study possible relationships between mating-induced and steroid-induced luteinizing hormone (LH) release in spayed Long-Evans rats. Large amounts of LH were released approximately 7 hr following progesterone injection in rats primed with estradiol benzoate (EB). The amount of LH release varied widely depending on (1) the interval between the time of the progesterone injection and the EB priming; (2) the progesterone dose; and (3) the time of day when blood samples were collected. These findings provided confirmation of those of Caligaris, Astrada and Taleisnik (1971a). Females, prepared with estrogen-progesterone treatment in a variety of schedules in which the three above-mentioned variables were altered systematically, were allowed to mate with vigorous males. Mating under these various conditions did not significantly increase plasma LH levels even when the females showed high degrees of sexual receptivity. Sodium pentobarbital prevented the afternoon LH rise resulting from progesterone treatment 3 days after EB priming. Pituitary sensitivity to LRF was not enhanced in the afternoon and the mating did not significantly increase plasma LH in these barbiturate-blocked rats. Following administration of 5 large daily doses of EB without progesterone, however, significant increases in LH were produced by mating on the sixth day. Postcopulatory LH release in these circumstances was dependent on a diurnal factor since the effect of mating was greater in the afternoon than in the morning. Thus, although major LH release can be readily induced by mating in estrogen-treated spayed rats, this effect could not be obtained under conditions of progesterone administration to estrogen primed rats.