Flight of male Cadra cautella along plumes of air and pheromone superimposed on backgrounds of pheromone

Male Cadra cautella were presented with five heterogeneous pheromone clouds (created from source doses of 0, 0.01, 1, 100, and 10 000 ng) with and without superimposed plumes of either clean air or sex pheromone in a wind tunnel. Moths provided with the lowest doses of background clouds without a superimposed plume did not fly upwind. Moths provided with higher doses of background clouds, with or without superimposed air plumes, increased their track, course, and drift angles (i.e., their zigzags headed more towards crosswind) with increased dose, but slowed their velocity. No differences in flight track parameters were observed for moths provided with a superimposed pheromone plume, regardless of the background cloud dose. Moreover, moths were able to locate the source of superimposed air plumes in the highest background dose, and of superimposed pheromone plumes in any background dose. The significance of these results is discussed in the context of mating disruption.     

Regulation of Wolbachia density in the Mediterranean flour moth,Ephestia kuehniella,and the almond moth,Cadra cautella

The Mediterranean flour moth, Ephestia kuehniella, is infected with A-group Wolbachia (wKue), and the almond moth, Cadra cautella, is doubly infected with A- and B-group Wolbachia, which are designated as wCauA and wCauB, respectively. In both insects, the Wolbachia populations increased greatly during embryonic and larval stages. The Wolbachia population doubled every 3.6 days on average in E. kuehniella larvae, whereas those of wCauA and wCauB doubled every 2.1 days in C. cautella larvae. The populations of wCauA and wCauB that had been transferred into the E. kuehniella background increased at similar rates to that of wKue in the natural host E. kuehniella, suggesting that the host genetic background influences Wolbachia proliferation. To examine whether the populations of the two Wolbachia variants in double infection is regulated collectively or independently, we measured the infection load in the ovaries of three transfected E. kuehniella lines in different infection states: single infection with wCauA, single infection with wCauB, and double infection. The density of each Wolbachia variant did not differ significantly between the singly and doubly transfected hosts, suggesting independent regulation.     

Genetic architecture of sensory exploitation: QTL mapping of female and male receiver traits in an acoustic moth

The evolution of extravagant sexual traits by sensory exploitation occurs if males incidentally evolve features that stimulate females owing to a pre‐existing environmental response that arose in the context of natural selection. The sensory exploitation process is thus expected to leave a specific genetic imprint, a pleiotropic control of the original environmental response and the novel sexual response in females. However, females may be subsequently selected to improve their discrimination of environmental and sexual stimuli. Accordingly, responses may have diverged and the original genetic architecture may have been modified. These possibilities may be considered by studying the genetic architecture of responses to male signals and to the environmental stimuli that were purportedly ‘exploited’ by those signals. However, no previous study has addressed the genetic control of sensory exploitation. We investigated this question in an acoustic pyralid moth, Achroia grisella, in which a male ultrasonic song attracts females and perception of ultrasound likely arose in the context of detecting predatory bats. We examined the genetic architecture of female response to bat echolocation signals and to male song via a cartographic study of quantitative trait loci (QTL) influencing these receiver traits. We found several QTL for both traits, but none of them were colocalized on the same chromosomes. These results indicate that – to the extent to which male A. grisella song originated by the process of sensory exploitation – some modification of the female responses occurred since the origin of the male signal.     

Genetic variance and phenotypic plasticity in a component of female mate choice in an ultrasonic moth

Abstract.— Female response to male advertisement signals in lesser waxmoths showed substantial genetic variation, phenotypic plasticity across rearing environments, and genotype-by-environment interactions resulting in crossing reaction norms. These results represent two previously underemphasized means by which genetic variation may be maintained in sexually selected traits: genetic variation in female response to male traits, and variation in the selection acting on both males and females. Genotype-by-environment interactions and reaction norms that cross indicate that divergent selection may act on male and female sexual traits if the level of environmental change is high. The processes that contribute to the maintenance of genetic variation may thus also contribute to population differentiation.     

Sexual selection in a moth: effect of symmetry on male mating success in the wild

Sexual selection is generally caused by female choice and male–malecompetition. In female choice process, female preference isfavored indirectly and/or directly by sexual selection. In indirectselection, females expressing the preference might gain indirectgenetic benefits. In direct selection, females expressing thepreference might gain direct benefits or avoid male-imposedcosts. The white-tailed zygaenid moth Elcysma westwoodii ismonandrous, and males often gather around a female to mate withher, suggesting a high opportunity for sexual selection on maletraits. We quantified phenotypic selection on male morphologyin this species in the field. The morphological characters analyzedincluded body weight, antenna length, forewing length, hindwing length, hind wing tail length, genital clasper length,and the fluctuating asymmetry (FA) of these bilateral traits.In E. westwoodii, selection favored males with more symmetricgenital claspers, as well as longer and more symmetrical hindwings and antennae. Negative correlations between FA and sizewere also detected in the clasper and the antenna. Our resultssuggest that FAs of male traits, in particular the genital clasper,may have indirect and direct influences on mating success. Duringa copulatory attempt, an E. westwoodii male will try to graspthe female's abdominal tip with his claspers but often failto do so because of the female's reluctance to mate. The femaleabdominal tips are smooth and strongly sclerotized and couldthus be difficult for males to grasp. We hypothesize that moresymmetrical male claspers are more efficient in overcoming femalereluctance.     

Influence of plume structure and pheromone concentration on upwind flight of Cadra cautella males

Abstract. The effects of pheromone plume structure and its concentration on the pheromone-mediated flight of male Cadra cautella (Lepidoptera: Phycitinae) were investigated in a laminar-flow wind tunnel. When two C. caurella males flew simultaneously along a ribbon plume of mixed smoke and pheromone, their inflight behaviour was dependent on the instantaneous structure of the plume they encountered. When a male intercepted an intact ribbon filament, he sustained a crosswind course, whereas when he intercepted a turbulent filament (created by an upwind male fragmenting the ribbon plume), he adopted a flight course more due upwind. These results indicate that C. cautella males altered their in-flight manoeuvres in response to instantaneous changes in the fine structure of the pheromone plume. We also demonstrated that differences in the fine structure of the plume had more influence on the flight pattern of C. cautella males than a 1000-fold range in pheromone dose. The size of the plume was increased by adding wind deflectors upwind of the pheromone source, independent of source dosage, males following ribbon plumes flew slow zigzag tracks, whereas males following large, turbulent plumes flew directly to the source in fast, straight tracks with less counterturning.     

Isolation, identification and stimulatory activity of a second component of the sex pheremone system (complex) of the female almond moth, Cadra cautella (Walker)

$\text{Cis}$-9-tetradecen-l-ol acetate was isolated from both extracts of female abdominal tips and of filter paper exposed to female $\text{Cadra}$$\text{cautella}$ (Walker). It is the second sex stimulant isolated from this species and, in stimulatory bioassays, it is ca. 1/100, 000 as active as the stimulant isolated previously, $\text{cis}$-9, $\text{trans}$-12-tetradecadien-l-ol acetate. According to attractancy tests, the natural sex pheromone system of complex contains other components in addition to these 2 stimulants.     

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.     

Sexual signal loss,pleiotropy, and maintenance of a male reproductive polymorphism in crickets

Pleiotropy between male signals and female preferences can facilitate evolution of sexual communication by maintaining coordination between the sexes. Alternatively, it can favor variation in the mating system, such as a reproductive polymorphism. It is unknown how common either of these scenarios is in nature. In Pacific field crickets (Teleogryllus oceanicus) on Kauai, Hawaii, a mutation (flatwing) that segregates as a single locus is responsible for the rapid loss of song production in males. We used outbred cricket colonies fixed for male wing morph to investigate whether homozygous flatwing and normal-wing (wild-type) females differ in responsiveness to male calling song and propensity to mate when paired with either a flatwing or normal-wing male in the presence or absence of courtship song. Flatwing females were less likely to mount a male than normal-wing females. Females of both genotypes showed a preference for normal-wing males and were more likely to mate in the presence of courtship song; normal-wing females were particularly likely to mate with song. Our results show that negative pleiotropy between obligate male silence and female mating behavior can constrain the evolution of sexual signal loss and contribute to the maintenance of a male reproductive polymorphism in the wild.     

Temperature coupling as an emergent property: parallel thermal effects on male song and female response do not contribute to species recognition in an acoustic moth

Temperature coupling exists when changes in male signal production with temperature are paralleled by changes in female response. Such thermal effects have been observed in various ectothermic animals producing acoustic, visual, and electric signals in which the signal rate may be subject to stabilizing selection imposed by female preference. Often, coupling was considered as an adaptive function wherein male and female thermal effects coevolved under selection pressure favoring species recognition, although this assumption has not been tested definitively. We investigated thermal effects on pulse-pair rate in male song and female acceptance threshold for male song rate in an acoustic moth, Achroia grisella, in which male song rate is subject to directional selection. Male song rate and female acceptance threshold do exhibit parallel increases as temperature rises from 18 degrees C to 30 degrees C, but female thresholds are much lower than male song rates and the thermal effect on female response cannot augment species recognition. In further investigations using inbred lines of A. grisella we found that the male and female thermal effects are genetically correlated, and we discuss the likely sources of this covariance. We consider several explanations for the occurrence of temperature coupling in this species and suggest that it represents an emergent property arising from the neuromuscular responses to temperature that are common to several physiological systems.     

Flight behaviour of males of two moths, Cadra cautella and Pectinophora gossypiella, in homogeneous clouds of pheromone

Abstract It is thought that orientation by male moths along pheromone plumes is guided by interception of filaments of pheromone along that plume and that clean air gaps are required for upwind progress. Given that several investigations have determined that cells sensitive to pheromone can resolve only low rates of encounter with pheromone filaments, generally up to 10 pulses/s, it would appear that individual filaments encountered at higher rates would not be resolved by the insects' sensory system and therefore the stimulus would be perceived as a non-flickering signal. Behaviourally, this has been thought to be expressed as the cessation of upwind progress. Previous studies with Cadra cautella (Walker) (Lepidoptera: Pyralidae) demonstrated that upwind flight by these males is not inhibited in rapidly pulsed plumes. Therefore, to determine whether a flickering signal is required for upwind progress by C. cautella , males were introduced to homogeneous clouds of pheromone in a wind tunnel and their behaviour recorded. For comparison, Pectinophora gossypiella (Saunders) (Lepidoptera: Gelechiidae), a species from a long-diverged lineage, was also used. Upwind progress by C. cautella is not impeded by the constant olfactory signal provided by a homogeneous cloud of pheromone, but this is not true for P. gossypiella . Furthermore, although C. cautella directs its flight upwind in a homogeneous cloud, its heading is not always due upwind. Potential mechanisms are discussed. It is suggested that C. cautella does not require a flickering signal to progress upwind.     

Pheromone-mediated behaviour of male lightbrown apple moth, Epiphyas postvittana, correlated with adaptation of pheromone receptors

ABSTRACT. Two pheromone components are required to elicit close-range precopulatory behaviour in male lightbrown apple moths, Epiphyas postvittana (Walker) (Lepidoptera). The receptor cells which respond to the major component (I) ( E )-11-tetradecen-1-yl acetate, have a fast disadaptation rate with recovery occurring within 5 s after stimulation, while the cells responding to the second component (II), ( E, E )-9, 11-tetradecadien-1-yl acetate, recover after approximately 300 s.
Studies on the behaviour of males in the laboratory show a close correlation between the duration of a memory effect, during which males will respond to compound I alone after receiving an initial exposure to I and II, and the time-course for disadaptation of component II-responding cells. These results suggest several possibilities for mechanisms of integration of sensory input by the CNS.     

Genetic analysis and population survey of sex pheromone variation in the adzuki bean borer moth, Ostrinia scapulalis

Sexual communication in many moths occurs between females emitting a sex pheromone and males responding to it. Females of Ostrinia scapulalis (Lepidoptera: Crambidae) show a large variation in blend ratios of the two sex pheromone components (E)‐ and (Z)‐11‐tetradecenyl acetates. E type females produce a pheromone with a high percentage of (E)‐11‐tetradecenyl acetate, whereas Z type females produce the opposite blend. We established laboratory cultures of E and Z types. Females of the F₁ generation produced an intermediate blend (I type) in both reciprocal crosses of the E and Z cultures. Results of further crossing experiments suggested that the three pheromone types are primarily controlled by a single autosomal locus with two alleles. Also, analyses of the variation in pheromone blend within F₁, backcross and F₂ families suggested that other genetic factors modify the pheromone blend of the I and Z types. Investigation of the pheromone variation in natural populations at 14 localities in Japan has shown that the E type was predominant in northern Japan, whereas the pheromone was highly polymorphic in central Japan. At a locality in central Japan, the pheromone was constantly polymorphic for several years, and the pheromone type frequencies did not deviate from Hardy–Weinberg expectations, providing no evidence of selection or assortative mating between the pheromone types. Analyses of pheromone variation within families derived from feral females indicated that matings between a pair with different genotypes for pheromone production was occurring in natural populations. Overall, this study showed that the genetic basis of the pheromone variation in O. scapulalis is very similar to that in its sibling species Ostrinia nubilalis although the state of pheromone polymorphisms in natural populations appears to differ between the two species. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 84 , 143–160.     

Visual cues collimate the trajectories of almond moth Cadra cautella males flying in wind and still air within a wind‐formed plume of pheromone

Male Cadra cautella (Walker) moths are videotaped in three dimensions in a 3‐m long wind tunnel as they fly within a 65‐cm wide plume of pheromone. Moths are presented two floor patterns, either ‘aligned’, a 25‐cm wide ‘trail’ of solid red circles along the tunnel's midline, or ‘offset’, in which the trail veers 25 cm to the left at the tunnel's midpoint. These visual patterns are presented either in a continuous airflow or airflow that is stopped before moths reach the tunnel's halfway point. Moths fly relatively straight paths over the aligned pattern in still air after the wind is stopped. With the offset pattern in wind and when the wind is stopped, moths swerve towards the offset pattern before again progressing along the plume. Prominent visual cues appear to ‘collimate’ (i.e. align with a directional cue) the moth's course as long as the moth remains in contact with pheromone. In wind, these moths appear to favour trajectories that enhance visual feedback, even if the path taken is not directly upwind. During wind lulls, this manoeuvre may enable moths to continue progress towards calling females along a visually set course. The centring of trajectory over prominent visual cues suggests that these moths favour a route that enhances visual feedback.     

Elevation of pheromone response threshold in almond moth males pre-exposed to pheromone spray

Abstract. High percentages of naive Cadra cautella (Walker) (Lepidoptera: Pyralidae) males not pre-exposed to pheromone flew upwind to sources containing 50 ng (83%) and 500 ng (97%) of pheromone, but not to sources containing 5 μg (23%) and 50 μg (4%).Of the naive males that flew upwind in response to 50 ng sources, 67% located and landed on the source, whereas fewer than 19% of the naive males that flew upwind in response to higher doses located and landed on the sources.A 2-minute pre-exposure of C.cautella males to a spray cloud containing 50 ng, 500 ng, 5 μg or 50 μg of pheromone, induced shifts in response levels such that in wind-tunnel bioassays performed 1 h later, there was an increase in the doses that optimally elicited upwind flight and landing on the source that was proportional to the pre-exposure dose.Few of the pre-exposed males flew upwind to (10–43%) and landed on (0–33%) 50 ng sources, whereas they now perferentially flew upwind to(58–81% and 52–73%) and landed on (33–68% and 55–60%) pheromone sources of doses of 500 ng and 5 μg, respectively.Therefore pre-exposure to pheromone promoted a shift of threshold for response, and not an overall reduction in responsiveness to pheromone.