Repeated larval diapause and diapause-free development in geographic strains of the burnet moth Zygaena trifolii Esp. (Insecta,Lepidoptera)

Summary Zygaena trifolii is a long-day insect with temperature-dependent photoperiodic responses. All larval instars are sensitive to photoperiod; however, diapause may occur at the third larval stage or any subsequent larval instars. There were quantitative differences within populations in the threshold photoperiod for diapause induction. The diapause response was polymorphic, so that larvae might enter diapause at different instars under the same culture conditions. Furthermore, decreasing photoperiods below a critical daylength shifted the diapausing instar towards earlier stages. Geographic strains of Z. trifolii showed discontinuous clinal variation. Near the northern edge of the distribution [Cologne (Köln), FRG], there is first an obligatory diapause, mainly during early instars, and additional facultative (repeat) diapauses during later larval instars in subsequent years. In the southern part of its distribution, this burnet moth is partially bivoltine in the field with a facultative first developmental arrest and a decreased capacity for repeated diapause (Valencia, Spain; Marseille, France). Further experiments indicated that the photoperiodically controlled diapause reaction is also influenced by the number of photoperiodic cycles experienced during the period spent in each larval instar, which depends on temperature. The adaptive significance of obligatory and facultative repeated diapause, varying even among the offspring of a single female, may be to buffer the populations against the more extreme and, from year to year, unpredictable fluctuations in climatic conditions at the northern edge of the distribution.Abbreviations L₃ feeding 3rd larval instar - L₄D diapausing 4th larval instar - L₅D2 repeat-diapausing larval instar with second diapause at the 5th larval stage - LD light-dark cycle - KT shortday conditions (e.g. LD 8:16) - LT long-day conditions (e.g. LD 16:8)     

Molecular biogeography of the arctic-alpine disjunct burnet moth species Zygaena exulans (Zygaenidae, Lepidoptera) in the Pyrenees and Alps

Aim The phylogeography of ‘southern’ species is relatively well studied in Europe. However, there are few data about ‘northern’ species, and so we studied the population genetic structure of the arctic‐alpine distributed burnet moth Zygaena exulans as an exemplar. Location and methods The allozymes of 209 individuals from seven populations (two from the Pyrenees, five from the Alps) were studied by electrophoresis. Results All 15 analysed loci were polymorphic. The mean genetic diversities were moderately high (A: 1.99; H_e: 11.5; P: 65%). Mean genetic diversities were significantly higher in the Alps than in the Pyrenees in all cases. F_ST was 5.4% and F_IS was 10%. Genetic distances were generally low with a mean of 0.022 between large populations. About 62% of the variance between populations was between the Alps and the Pyrenees. The two samples from the Pyrenees had no significant differentiation, whereas significant differentiation was detected between the populations from the Alps (F_ST = 2.8%, P = 0.02). Main conclusion Zygaena exulans had a continuous distribution between the Alps and the Pyrenees during the last ice age. Most probably, the species was not present in Iberia, and the samples from the Pyrenees are derived from the southern edge of the glacial distribution area and thus became genetically impoverished. Post‐glacial isolation in Alps and Pyrenees has resulted in a weak genetic differentiation between these two disjunct high mountain systems.     

暗期干扰对环带锦斑蛾滞育诱导的影响

在短日照条件下,环带锦斑蛾Pseudopidorus fasciata以4龄幼虫进入冬季滞育,其滞育诱导的临界暗长为10.5 h。本文系统测试了暗期干扰实验［即在暗期采用单一的光（亦称光脉冲）干扰光周期反应的实验］对环带锦斑蛾幼虫滞育的抑制影响。幼虫孵化后在25℃,短日照L9∶D15(光9 h∶暗15 h),L10∶D14、L11∶D13及L12∶D12暗期的不同时段给予1 h或30 min光脉冲干扰。结果表明： 滞育是否被抑制主要取决于暗期的长度,当光脉冲干扰前的暗长（D₁）或光脉冲干扰后的暗长（D₂）超过临界暗长（10.5 h）时,100%的幼虫被诱导进入滞育;当D₁和D₂都短于（或等于）临界暗长时,滞育不同程度被抑制,抑制的效果则取决于所干扰的光周期、光脉冲的长度和光脉冲落入的位点。结果说明该虫滞育诱导的暗期干扰反应是基于临界暗长的时间测量。     

Genital Morphology and Copulatory Mechanism in Zygaena trifolii (Esper, 1783) (Insecta,Lepidoptera, Zygaenidae)

The coremata and male genital structures together with the associated muscles of Zygaena trifolii are described and illustrated in detail. Of the female genitalia only the external parts are dealt with. The closure of the male coremata pouch was found to be effected by a special muscle. Mating pairs were quickly fixed while in copula and dissected afterwards in order to examine the mechanism of copulation. It turns out that the female is held by the male at three points: first, the gonopods (= valvae) grip the female abdominal tip laterally. Secondly, the praephallus together with the everted endotheca (= vesica) are tightly locked inside the female ductus bursae. Thirdly, female postvaginal structures become evaginated and are then held between the spined anellus and the eighth abdominal tergum of the male.     

Little-known accessory glands in female Zygaena moths (Lepidoptera, Zygaenidae)

In the female genital system of Zygaena moths, an additional pair of accessory glands is present besides the Y-shaped sebaceous gland. The term 'Petersen's glands' is proposed for these organs. Anatomy, histology, histochemistry and cytology of Petersen's glands of Zygaena trifolii are described. The sac-like glands, situated in the extreme dorsocaudal part of the abdomen, can be divided into a purely secretory part consisting of acini with large pear-shaped gland cells and a reservoir part with combined secretory and storage function. The secretory cells of the acini are penetrated by long curved ductules or secretory end apparatuses having feltwork consisting of very fine filaments. The cytoplasm is characterized by abundance of smooth tubular endoplasmic reticulum (ATER) and the presence of peroxisomes. This cytoplasmic organization is in accordance with the chemical composition of the sticky secretion, which evidently consists completely of lipids. The ultrastructure of the epithelium lining the reservoir of the glands has both traits of secretory and of transporting epithelia. Besides contributing to the secretion, it may be involved in absorption of residual aqueous phase from the contents of the reservoir.     

Volatiles from the burnet moth Zygaena filipendulae (Lepidoptera) and associated flowers,and their involvement in mating communication

The burnet moth Zygaena filipendulae L. contains the cyanogenic glucosides linamarin and lotaustralin, which can be degraded to the volatiles hydrogen cyanide (HCN), acetone and 2‐butanone. Linamarin and lotaustralin are transferred from the male to female during mating and thus are considered to be involved in mating communication. Because volatile semiochemical cues play a major role in mating communication in many insect species, the emissions of HCN, acetone and 2‐butanone from Z. filipendulae are characterized in the present study, aiming to determine the interplay between the degradation products of cyanogenic glucosides and pheromones. The volatile emissions from Z. filipendulae and flowers inducing mating are measured using headspace solid‐phase micro‐extraction and gas chromatography‐mass spectrometry analysis. All Z. filipendulae life stages emit HCN, acetone and 2‐butanone. Virgin females show higher emissions than mated females, whereas mated males have higher emissions than virgin males. Hydrogen cyanide is only rarely detected in the course of male–female copulation. These observations indicate a role for the cyanogenic glucoside derived volatiles in female calling and male courtship behaviours, although not as a defence during copulation. Males rejected for mating by a female are accepted after injection of linamarin or lotaustralin, demonstrating that cyanogenic glucosides are also important for female assessment of the fitness of the male. Volatiles from flowers occupied during mate calling are also analyzed, and emissions from males and females result in the identification of novel putative pheromones for Z. filipendulae.     

Migration and Allee effects in the six-spot burnet moth Zygaena filipendulae

Abstract 1. Migration into local populations may increase the likelihood of persistence but emigration may decrease the persistence of small and isolated populations. The dispersal behaviour of a day-flying moth Zygaena filipendulae was examined to determine whether emigration is correlated positively or negatively with population size and host plant density.
2. A mark–release–recapture study showed that most moths moved small distances (< 40 m on average) and only 6% of movements were > 100 m.
3. Twenty-five individuals moved between populations, a measured exchange rate of 8%. Moths were more likely to move between patches that were close together and they moved to relatively large patches.
4. The fraction of residents increased with increasing population size in the patch and increasing host plant cover. Relatively high proportions of individuals left small patches with small moth populations.
5. Moths released in grassland lacking Lotus corniculatus (the host plant) tended to leave the area and biased their movement towards host plant areas, whereas those released within an area containing L. corniculatus tended to stay in that area.
6. Biased movement away from small populations and areas of low host plant density (normally with low population density) was found. This migration-mediated Allee effect is likely to decrease patch occupancy in metapopulations, the opposite of the rescue effect. The effects on metapopulation persistence are not known.     

Photoperiodic counter of diapause induction in Pseudopidorus fasciata (Lepidoptera: Zygaenidae)

Induction of larval diapause is a photoperiodically controlled event in the life history of the moth Pseudopidorus fasciata. In the present study, the photoperiodic counter of diapause induction has been systematically investigated. The required day number (RDN) for a 50% response was determined by transferring from a short night (LD 16:8) to a long night (LD 12:12) or vice versa at different times after hatching, The RND differed significantly between short- and long-night cycles at different temperatures. The RDN for long-night cycles at 20, 22, 25 and 28 degrees C was 11.5, 9.5, 7.5 and 8.5 days, respectively. The RDN for short-night cycles was 3 days at 22 degrees C and 5 days at 20 degrees C indicating that the effect of one short night was equivalent to the effect of 2-3 long nights effect. Night-interruption experiments of 24h photoperiods by a 1 h light pulse showed that the most crucial event for the photoperiodic time measurement in this moth was whether the length of pre-interruption (D(1)) or the post-interruption (D(2)) scotophases exceeded the critical night length (10.5 h). If D(1) or D(2) exceeded 10.5 h diapause was induced. The diapause-averting effect of a single short-night cycle (LD 16:8) against a background of long nights (LD 12:12) showed that the photoperiodic sensitivity was greatest during the first 7 days of the larval period and the highest sensitivity was on the fourth day. Both non-24 and 24 h light-dark cycle experiments revealed that the photoperiodic counter in P. fasciata is able to accumulate both long and short nights during the photosensitive period, but in different ways. The information from short-night cycles seems to be accumulated one by one in contrast to long-night cycles where six successive cycles were necessary for about 50% diapause induction and eight cycles for about 90% diapause. These results suggest the accumulation of long-night and short-night cycles may be based on different mechanisms.     

Photoperiodic clock of diapause induction in Pseudopidorus fasciata (Lepidoptera: Zygaenidae)

Pseudopidorus fasciata enters diapause as fourth instar larvae at short day lengths. Using 24-h light-dark cycles, the photoperiodic response curves in this species appeared to be similar with a critical night length of 10.5h at temperatures below 30 degrees C. At an average temperature of 30.5 degrees C, the critical night length had shifted to between 15 and 17h. In experiments using non-24-h light-dark cycles, it was clearly demonstrated that the dark period (scotophase) was the decisive phase for a diapause determination. In night interruption experiments using 24-h light-dark cycles, a 1-h light pulse at LD12:12 completely reversed the long night effect and averted diapause in all treatments. At LD 9:15 light pulses of 1-h, 30- or 15-min also averted diapause effectively when both the pre-interruption (D(1)) or the post-interruption scotophases (D(2)) did not exceed the critical night length. If D(1) or D(2) exceeded the critical night length diapause was induced. The most crucial event for the photoperiodic time measurement in this species is the length of the scotophase. A 10-min light pulse placed in the most photosensitive phase reversed diapause in over 50% of the individuals. Night interruption experiments under non-24-h light-dark cycles indicated that the photoperiodic clock measured only D(1) regardless of the length of D(2), suggesting that the most inductive cycles are often those in which L+D are close to 24h. In resonance experiments, this species showed a circadian periodicity at temperatures of 24.5 or 26 degrees C, but not at 30.5 and 23.3 degrees C. On the other hand, Bünsow and skeleton photoperiod experiments failed to reveal the involvement of a circadian system in this photoperiodic clock. These results suggest the photoperiodic clock in this species is a long-night measuring hourglass and the circadian effect found in the final expression of the photoperiodic response in the resonance experiments may be caused by a disturbing effect of the circadian system in unnatural regimes.     

Qualitative and quantitative studies on the compounds of the larval defensive secretion of Zygaena trifolii (Esper, 1783) (Insecta, Lepidoptera, Zygaenidae)

Beside the known existence of cyanoglucosides (linamarin and lotaustralin) and proteins the neurotoxin beta-cyanoalanine has been demonstrated for the first time in the defensive secretions of animals. It is proposed that beta-cyanoalanine is produced by metabolizing cyanide from the cyanoglucosides. The methanolic precipitated protein fraction contains high amounts of aspartic acid, glycine, alanine, leucine and serine, thus being similar to the composition of larval silks in Lepidoptera. The defensive secretion contains 85% water, 8% proteins, 7% cyanoglucosides, 0.3% beta-cyanoalanine and beta-glucosidase while beta-cyanoalanine-synthetase could only be detected in the haemolymph.     

Allozymic and morphometric analysis of populations in the Zygaena purpuralis complex (Lepidoptera, Zygaenidae)

In Central Europe and Asia Minor Zygaena purpuralis and Z. minos are two sibling species exhibiting a considerable geographic variation in several features. In this study we examine some populations from the Central and Southern Apennines (Italy) by comparing multivariate analyses of individual allozymic multilocus profiles and of individual morphometric profiles of male genitalia in order to assess their genetic and phenetic relationships.
Results indicate that populations of Z. purpuralis in the Apennines behave in rather different ways with respect to the occurrence of gene exchange with other gene pools. No gene flow appears to occur between two geographically close southern populations; however, extensive introgression, revealable at both the allozymic and morphometric level, seems to occur between some purpuralis and minos -like sympatric populations from Abruzzi. Evolutionary relationships among these peninsular Italy populations are discussed with reference to historical factors, that is, the occurrence of multiple Pleistocene refugia, which could have determined different degrees of isolation among gene pools. This situation is rather different from that previously outlined for Central Europe and present results suggest that the Z. purpuralis complex might include more than two biological species.     

Photoperiodism of diapause induction in Thyrassia penangae (Lepidoptera: Zygaenidae)

Thyrassia penangae enters winter diapause as a prepupa in a cocoon. Photoperiodism of diapause induction was systematically investigated in this moth. The photoperiodic response curves under 24-h light-dark cycles showed that this insect is a typical long-day species. The critical daylength was 13 h 30 min at 25 °C, 13 h at 30 °C and 12 h 20 min at 28 °C. Transferring experiments from a short day (LD 12:12) to a long day (LD 15:9) or vice versa indicated that photoperiodic sensitivity mainly occurs during the larval period. In experiments using non-24-h light-dark cycles, when the length of photophase exceeded the critical daylength (13.5 h), was diapause inhibited effectively, even when the length of scotophase exceeded the critical nightlength (10.5 h). Only when a long scotophase was combined with a short photophase, diapause was induced effectively. This result suggests that daylength measurement is more important than nightlength measurement in T. penangae. Night interruption experiments under 24-h light-dark cycles exhibited two points of apparent light sensitivity, but the photosensitive position was highly influenced by temperature and the length of scotophase. Nanda-Hamner experiments failed to reveal the involvement of a circadian system in this photoperiodic time measurement. All light-dark cycles from LD 12:12 to LD 12:72 resulted in a short day response, and all cycles from LD 14:4 to LD 14:72 resulted in a long day response, suggesting that photoperiodic time measurement in this moth is performed by a day-interval timer or an hourglass-like clock.     

Intimate roles for cyanogenic glucosides in the life cycle of Zygaena filipendulae (Lepidoptera, Zygaenidae)

Zygaena larvae sequester the cyanogenic glucosides (CNglcs) linamarin and lotaustralin from their food plants (Fabaceae) and also de novo biosynthesize these compounds. In Zygaenidae, CNglcs serve as defence compounds during the entire life cycle, and their content and ratio are tightly regulated. We demonstrate that Z. filipendulae males transfer a nuptial gift of CNglcs to females during mating, and that females prefer males with a higher content of CNglcs for mating. Average HCN emission from female imagines is 19 times higher than from males, suggesting that plumes of HCN emitted from the perching female may serve to attract flying males. Analysis of the linamarin and lotaustralin content and ratio within different tissues in Z. filipendulae larvae shows that integument and haemolymph constitute the main sites of CNglc deposition. The data suggest that CNglcs may serve an additional role as storage compounds of reduced nitrogen that is mobilized during the transition of the last instar larva to imago, most likely to provide nitrogen for chitin synthesis. At least one of the enzymes responsible for de novo biosynthesis of CNglcs in Z. filipendulae is located in the integument. In conclusion, CNglcs play many important and different roles during the entire life cycle of Z. filipendulae in addition to defence.     

Influences of daylength and temperature on the period of diapause and its ending process in dormant larvae of burnet moths (Lepidoptera,Zygaenidae)

The onset of larval diapause in the burnet moth Zygaena trifolii is clearly characterized by the larva molting into a specialized dormant morph. In a potentially bivoltine Mediterranean population (Marseille) two types of diapause can occur within 1 year: firstly, a facultative summer diapause of 3–10 weeks, and secondly, an obligate winter diapause, which can be lengthened by a period of thermal quiescence to several months in temperatures of 5°C. For the first time, three successive physiological periods have been experimentally distinguished within an insect dormancy (between onset of diapause and molting to the next non-diapause stage), using chilling periods of 30–180 days at 5°C, and varying conditions of photoperiod and temperature. These stages are: (1) a continuous Diapause-ending process (DEP); (2) thermal quiescence (Q); and finally, (3) a period of postdiapause development (PDD) before molting to the next larval instar. The result of transferring dormant larvae from chilling at 5°C to 20°C depended on the length of the chilling period. After chilling for 120–180 days, molting to the next instar occurred after 6–10 days, independent of daylength. This period corresponds with the duration of PDD. After shorter chilling periods (90, 60, 30 days and the control, 0 days) the period to eclosion increased exponentially, and included both the latter part of the previous diapause process and the 6–10 day period of PDD. However, photoperiod also influences the time to eclosion after chilling. Short daylength (8 h light / 16 h dark: LD 8/16) lengthened the diapause in comparison to long daylength (16 h light / 8 h dark: LD 16/8). Short daylength had a similar effect during chilling at 5°C, as measured by the longer time to eclosion after transfer. The shorter time to eclosion resulting from longer chilling periods (30–90 days) demonstrates that the state of diapause is continuously shortened at 5°C, and corresponds to the neuroendocrine controlled DEP. Presumably the DEP has already started after the onset of diapause. When chilling was continued after the end of the DEP, which ranged between 90 and 120 days, thermal quiescence (Q) followed (observed maximum 395 days). Different photoperiodic conditions during the pre-diapause inductive period modified diapause intensity (measured as the duration of diapause), in that a photoperiodic signal just below the critical photoperiod for diapause induction (LD 15/9) intensified diapause. Experiments simulating the summer diapause showed that PDD occurred in the range of 10–25°C. Higher temperatures (15 and 20°C) shortened the DEP at LD 16/8, so that at 20°C many individuals had already terminated diapause after 10–40 days and had molted after the 6–10 days of PDD. A temperature of 25°C unexpectedly lengthened the DEP to 110 days in several individuals. The ecological consequences and the adaptive significance of variation in the duration of the diapause are discussed in relation to the persistence of local populations predictably variable and rare climatic extremes throughout the year.     

Evolutionary history of the burnet moth genus Zygaena Fabricius, 1775 (Lepidoptera: Zygaenidae) inferred from nuclear and mitochondrial sequence data: phylogeny, host–plant association, wing pattern evolution and historical biogeography

Burnet moths of the genus Zygaena are a striking group of primarily diurnal Lepidoptera displaying an exceptional phenotypic plasticity. Previous attempts to elucidate the phylogenetic history of the group had been confounded by a perplexing pattern of characters or insufficient taxon sampling. In the present study, we infer a phylogeny of the genus Zygaena by analysing 5.4 kb of their nuclear and mitochondrial DNA. Eighty‐four of the 98 currently recognized species in this genus are considered, including representatives of all described species groups. RNA coding sequences are aligned with reference to zygaenoid moth specific secondary structure models of corresponding molecules. We conduct phylogenetic analyses within a Bayesian framework applying partition specific substitution parameters; covariation of paired sites in RNA gene sequences is accommodated by using doublet substitution models. The molecular data reveal that a considerable number of currently recognized species groups in Zygaena are not monophyletic. The traditional subgeneric classification proves to be artificial as well; Agrumenia and Zygaena (sensu stricto) are polyphyletic. Only the subgenus Mesembrynus can be confirmed as a monophyletic species cluster. Optimization of larval host–plant associations and forewing patterns on sampled trees of the Bayesian analyses suggest convergent evolution of similar wing pattern types in distantly related species clusters and a shift from cyanogenic to acyanogenic host‐plants. The phylogenetic results challenge the classic assumption that early species diversification in Zygaena took place in the Irano–Turkestanian region. Rather, the molecular data point to the western Mediterranean area as the geographical origin of the group and imply a subsequent colonization of the Middle East and Central Asia. We discuss the apparently convergent evolution of similar wing patterns in context with the chemical defence system of burnet moths and suggest a species group concept for the genus Zygaena that accounts for the recent findings. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society, 2007, 92 , 501–520.     

Induction and termination of prepupal summer diapause in Pseudopidorus fasciata (Lepidoptera: Zygaenidae)

The seasonal life cycle of the zygaenid moth, Pseudopidorus fasciata is complicated by two different developmental arrests: a winter diapause as a fourth larval instar and a summer diapause as a prepupa in a cocoon. Both larval diapause induction and termination are under photoperiodic control. Short days induce larval diapause with a critical daylength of 13.5h and long days terminate diapause with a critical daylength of 14h. In the present study photoperiodic control of summer diapause was investigated in Pseudopidorus fasciata. Under long photoperiods ranging from LD 14:10 to LD 18:6, only part of the population entered summer diapause, the rest continued to develop. The lowest number of prepupae entered diapause at LD 14:10, followed by LD 16:8 and LD 17:7. The highest incidence of diapause occurred with photoperiods of LD 15:9 and LD 18:6. By transferring the diapausing prepupae induced by various long photoperiods (LD 14:10, LD 15:9, LD 16:8, LD 17:7, LD 18:6) to LD 13:11, 25 degrees C, the duration of diapause induced by LD 14:10 was significantly shorter than those induced by longer photoperiods. By keeping aestivating prepupae induced by LD 15:9, 28 degrees C or by natural conditions at short photoperiods (LD 11:13 and LD 13:11) and at a long photoperiod (LD 15:9), the duration of diapause at LD 15:9 was more than twice as long as than those at LD 11:13 and LD 13:11. Moreover, adult emergence was highly dispersed with a high mortality at LD 15:9 but was synchronized with low mortality at LD 11:13 and LD 13:11. When the naturally induced aestivating prepupae were kept under natural conditions, the early aestivating prepupae formed in May exhibited a long duration of diapause (mean 126 days), whereas the later-aestivating prepupae formed in July exhibited a short duration of diapause (mean 69 days). These results indicate that aestivating prepupae require short or shortening photoperiod to terminate their diapause successfully. By transferring naturally induced aestivating prepupae to 25, 28 and 30 degrees C, the duration of diapause at the high temperature of 30 degrees C was significantly longer than those at 25 and 28 degrees C, suggesting that high temperature during summer also plays an important role in the maintenance of summer diapause in Pseudopidorus fasciata. All results reveal that summer diapause can serve as a "bet hedging" against unpredictable risks due to fluctuating environments or as a feedback mechanism to synchronize the period of autumn emergence.     

No evidence of quantitative signal honesty across species of aposematic burnet moths (Lepidoptera: Zygaenidae)

Many defended species use conspicuous visual warning signals to deter potential predators from attacking. Traditional theory holds that these signals should converge on similar forms, yet variation in visual traits and the levels of defensive chemicals is common, both within and between species. It is currently unclear how the strength of signals and potency of defences might be related: conflicting theories suggest that aposematic signals should be quantitatively honest, or, in contrast, that investment in one component should be prioritized over the other, while empirical tests have yielded contrasting results. Here, we advance this debate by examining the relationship between defensive chemicals and signal properties in a family of aposematic Lepidoptera, accounting for phylogenetic relationships and quantifying coloration from the perspective of relevant predators. We test for correlations between toxin levels and measures of wing colour across 14 species of day‐flying burnet and forester moths (Lepidoptera: Zygaenidae), protected by highly aversive cyanogenic glucosides, and find no clear evidence of quantitative signal honesty. Significant relationships between toxin levels and coloration vary between sexes and sampling years, and several trends run contrary to expectations for signal honesty. Although toxin concentration is positively correlated with increasing luminance contrast in forewing pattern in 1 year, higher toxin levels are also associated with paler and less chromatically salient markings, at least in females, in another year. Our study also serves to highlight important factors, including sex‐specific trends and seasonal variation, that should be accounted for in future work on signal honesty in aposematic species.     

Correlation Between the Mesodermal Male Genital Ducts and the Spermatophore Structure in a Ditrysian Moth,Zygaena trifolii (Esper, 1783) (Insecta,Lepidoptera, Zygaenidae)

Abstract The mesodermal male genital system and the spermatophore of Zygaena trifolii (Insecta, Lepidoptera, Zygaenidae) have been studied histologically. Whereas the long unpaired duct, the ductus mesospermaticus, has been found to consist of at least six distinctly different regions, the tissues and secretions of the seminal vesicles and the accessory glands were found to be quite uniform. Several of the different regions are separated from each other by epithelial septa (constrictions). The external structure and the internal organization of the spermatophore are described. Based on histological comparisons, the correlation between the contents of the spermatophore and the different parts of the male system is given.     

Geographic variation in diapause induction: the grape berry moth (Lepidoptera: Tortricidae)

Diapause in insects occurs in response to environmental cues, such as changes in photoperiod, and it is a major adaptation by which insects synchronize their activity with biotic resources and environmental constraints. For multivoltine agricultural insect pests, diapause initiation is an important consideration in management decisions, particularly toward the end of the growing season. The grape berry moth, Paralobesia viteana (Clemens), is the main insect pest affecting viticulture, and this insect responds to postsummer solstice photoperiods to initiate diapause. Because the range of grape berry moth extends from southern Canada to the southern United States, different populations are exposed to different photoperiodic regimes. We quantified the diapause response in grape berry moth populations from Arkansas, Michigan, New York, Pennsylvania, Texas, and Virginia, and observed latitudinal variation in diapause initiation. Populations from Michigan, New York, and Pennsylvania responded significantly different than those from Arkansas, Texas, and Virginia. We also observed, as a consequence of our experiments, that the timing of our laboratory studies influenced grape berry moth's response to photoperiod, ceteris paribus. Experiments that were conducted when grape berry moth would be naturally in diapause resulted in a significant higher proportion of diapausing pupae at photoperiods (i.e., >15 h) that generally do not induce diapause, suggesting that attention should be paid to the timing of behavioral and physiological experiments on insects. This relationship between photoperiod and diapause induction in grape berry moth across geographic regions will provide applicable knowledge to improve pest management decisions.     

高温对环带锦斑蛾幼虫滞育的抑制作用

本文报道了高温（31℃）对环带锦斑蛾幼虫滞育发生的抑制作用。当幼虫暴露于31℃ 时,所有个体都继续发育,与光周期无关。在诱导滞育的光周期条件(L12∶D12)下,光期的高温配合不同的暗期低温（15～28℃）,导致几乎所有个体滞育,但当暗期为5℃时,滞育率反而下降;相反,当光期的低温配合不同持续时间的暗期高温（31℃）时,则几乎所有的个体都继续发育,这说明高温在暗期发挥着重要的作用。在暗期给予不同时间长度（2、4、 6、8、10、12 h）的高温处理,结果表明一个2 h的高温处理就能有效地抑制滞育的发生。在暗期的不同时间给予4 h高温处理,显示了幼虫在暗期开始后的第一个4 h (18:00～22:00 )对高温最敏感,完全抑制了滞育的发生。最后讨论了高温调节滞育机制在该虫生活史上的适应意义。