Entomopathogenic Nematodes for Biological Control of Pryeria sinica Moore (Lepidoptera: Zygaenidae) and Persistence on Euonymus japonica Thunberg Foliage

Korean entomopathogenic nematodes (EPN), Steinernema carpocapsae Pocheon strain, Steinernema sp. GSNUS-4 strain, Steinernema sp. GSNUS-14 strain, Steinernema sp. GSNUS-16 strain, Heterorhabditis sp. GSNUH-2 strain, and Heterorhabditis sp. GSNUH-3 strain were evaluated for the biological control of pellucid zygaenid, Pryeria sinica Moore (Lepidoptera: Zygaenidae), a leaf-feeding insect pest of Euonymus japonica Thunberg. In addition, persistence of S. carpocapsae Pocheon strain on E. japonica foliage was checked.Nematode species, strain, and concentration influenced infectivity of Korean EPNs against 3rd instar of P. sinica larvae. S. carpocapsae Pocheon strain was the most effective nematode.Larval mortality by S. carpocapsae Pocheon strain in the exposure experiments was low in both light and dark conditions representing 36.7% at 100 infective juveniles/larva in each dark and light condition. Mortality by fenitrothion, however, was 100% 3 days later. Infectivity of S. carpocapsae Pocheon strain was not different between non-exposed foliage and exposed foliage to sunlight for 30 min.Mortality of 4th instar of P. sinica larvae was very low for both S. carpocapsae Pocheon alone and S carpocapsae Pocheon with spreading agent. Application time of nematodes, shading, and leaf position on the tree influenced persistence of S. carpocapsae Pocheon strain on E. japonica foliage. Nematode survival on foliage was positively correlated with shading level and higher at the lowest level than the middle or upper level of the tree.     

Some beneficial effects of aggregation in young larvae ofPryeria sinica moore (Lepidoptera: Zygaenidae)

The effects of group size on the survival and development of young larvae of Pryeria sinicaMoore were investigated by laboratory and field experiments. Under laboratory conditions, about 20% of isolated larvae died of unsuccessful feeding in the first instar, however, larvae survived successfully in aggregations of four or more individuals. In the field, larvae emerge in early spring and wait for new leaves to open before feeding. In this period, the larger the group size of hatchlings the survival rate became higher. The nest-web spun by hatchlings was considered to play an important role in protecting them from desiccation. In the period that larvae began to feed on leaves, more than 36 larvae are necessary to aggregate for the successful establishment of feeding groups. The nest-web played an important role also in the establishment of feeding group. However, the natural group size of the first instar larvae was larger than the minimum group size to spin a sound nest-web in the field experiment. On the other hand, in later stage, larvae in a large group did not have an excess advantage in survival or developmental rate over larvae in a small group. It was found that the experiments on survival and developmental rates could not explain the reason that this species maintain large compact groups in the most part of larval period.     

The phylogenetic relationships of Chalcosiinae (Lepidoptera, Zygaenoidea, Zygaenidae)

The chalcosiine zygaenid moths constitute one of the most striking groups within the lower-ditrysian Lepidoptera, with highly diverse mimetic patterns, chemical defence systems, scent organs, copulatory mechanisms, hostplant utilization and diapause biology, plus a very disjunctive biogeographical pattern. In this paper we focus on the genus-level phylogenetics of this subfamily. A cladistic study was performed using 414 morphological and biochemical characters obtained from 411 species belonging to 186 species-groups of 73 genera plus 21 outgroups. Phylogenetic analysis using maximum parsimony leads to the following conclusions: (1) neither the current concept of Zygaenidae nor that of Chalcosiinae is monophyletic; (2) the previously proposed sister-group relationship of Zygaeninae + Chalcosiinae is rejected in favour of the relationship (Zygaeninae + ((Callizygaeninae + Cleoda ) + ( Heteropan + Chalcosiinae))); (3) except for the monobasic Aglaopini, none of the tribes sensu Alberti (1954 ) is monophyletic; (4) chalcosiine synapomorphies include structures of the chemical defence system, scent organs of adults and of the apodemal system of the male genitalia. A paired metathoracic androconial organ and a series of abdominal tergal corematal organs have been discovered, both being new to Lepidoptera. Due to highly homoplastic patterns in copulatory structures and wings that demonstrate significant sexual dimorphism, polymorphism and mimicry, 17 of the 69 'true' chalcosiine genera ( c . 25%) are shown to be either paraphyletic or polyphyletic. The present classification is therefore very misleading. Reductions of various parts of the male genitalia in some groups are accompanied by morphological and functional replacement involving the 8th abdominal segment. A prominent but convergent lock and key mechanism is revealed.  © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society , 2005, 143 , 161–341.     

Zygaenidae (Lepidoptera) of Omsk Province

The fauna of the family Zygaenidae of Omsk Province is reviewed. 12 species are listed, among which 7 are recorded from this territory for the first time.     

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.     

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.     

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。     

The complete mitochondrial genome of Eterusia aedea (Lepidoptera,Zygaenidae) and comparison with other zygaenid moths

Zygaenidae comprises >1036 species, including many folivorous pests in agriculture. In the present study, the complete mitochondrial genome (mitogenome) of a major pest of tea trees, Eterusia aedea was determined. The 15,196-bp circular genome contained the common set of 37 mitochondrial genes (including 13 protein-coding genes, two rRNA genes, and 22 tRNA genes) and exhibited the similar genomic features to reported Zygaenidae mitogenome. Comparative analyses of Zygaenidae mitogenomes showed a typical evolutionary trend of lepidopteran mitogenomes. In addition, we also investigated the gene order of lepidopteran mitogenomes and proposed that the novel gene order trnA-trnR-trnN-trnE-trnS-trnF from Zygaenidae and Gelechiidae and most other gene rearrangements of this tRNA cluster evolved independently. Finally, the mitogenomic phylogeny of Lepidoptera was reconstructed based on multiple mitochondrial datasets. And all the phylogenetic results revealed the sister relationships of Cossoidea and Zygaenoidea with both BI and ML methods, which is the first stable mitogenomic evidence for this clade.     

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.     

Characterization of microsatellite loci for Reissita simonyi (Rebel, 1899) (Lepidoptera,Zygaenidae)

The newly developed microsatellite set for Reissita simonyi ( Rebel 1899 ) (Lepidoptera, Zyganidae) is presented. In total, 14 loci turned out to be highly polymorphic. The microsatellite set will be used for a population genetic analysis of this endemic species in Southern Arabia. In the focus of this study stands the genetic differentiation and variability of isolated populations, which are assumed to be based on habitat fragmentation due to natural processes and human impact. Moreover, this new microsatellite set showed accurate band patterns in the Palaearctic genus Zygaena which seems to be promising for further studies.     

Phylogeny, systematics and evolution of mimetic wing patterns of Eterusia moths (Lepidoptera, Zygaenidae, Chalcosiinae)

Abstract. The aim of the present study was to investigate the phylogeny, systematics and evolution of the mimetic wing patterns of Eterusia, a day-flying moth genus that exhibits great morphological diversity, as well as the highest insular differentiation in eastern Asia and which has the most chaotic taxonomic history in the family Zygaenidae. We examined the wing patterns of the insects involved using visible and ultraviolet light (both reflectance and fluorescence). The phylogeny of thirty-four taxa, including all the recognized species of Eterusia plus two species of Soritia as outgroups, was reconstructed based on eighty adult morphological characters, including forty-one derived from colour patterns. Phylogenetic relationships based on the whole dataset revealed that (1) the most current concept of Eterusia is monophyletic, and (2) different types of mimetic pattern show different levels of phylogenetic conservation. To investigate the evolution of their colour patterns we inactivated all the relevant characters and reconstructed another phylogeny, which was found to differ significantly from the one based on the whole character set in the position of the E. risa species group. We used these phylogenetic hypotheses to test evolutionary predictions based on conventional Müllerian mimicry and quasi-Batesian mimicry dynamics. The results of permutation–tail–probability tests showed that the coloration characters are phylogenetically conserved, thus justifying a Müllerian interpretation. However, when comparing the observed topologies with hypothetical trees constrained to fit perfect Müllerian or quasi-Batesian scenarios using the Kishino–Hasegawa test, the observed phylogenies were more consistent with the phylogenetic prediction of quasi-Batesian mimicry. Therefore, we consider that applying these two phylogenetic methods to justify mimicry models may not always be practical. Finally, the taxonomy of Eterusia is revised. In total, two new species (E. austrochinensis, E. guanxiana), one new subspecies (E. risa palawanica) and four new synonyms (E. lativitta and E. fasciata of E. sublutea, E. coelestina of E. subcyanea, E. angustipennis gaedei of E. angustipennis angustipennis) are established.     

Unusual location and structure of female pheromone glands in Theresimima (= Ino) ampelophaga Bayle-Berelle (Lepidoptera : Zygaenidae)

Morphological and behavioural studies strongly suggest that the sex pheromone glands of female Theresimima (= Ino) ampelophaga (Lepidoptera : Zygaenidae) are situated on the anterior part of the 3rd–5th abdominal tergites. The glandular epithelium consists of 2 cell types: gland cells and wrapping cells. The gland cells have a central microvilli-lined cavity which is in contact with the lumen of the hair. These hairs are exposed during the calling behaviour, and the pheromone is probably given off via pores in the scale wall.     

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.     

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.     

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.     

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.     

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。     

Lemyra flammeola (Moore) (Lepidoptera,Arctiidae), new to Korea

An arctiine species Lemyra flammeola (Moore) is newly recorded from Korea. Three specimens of L. flammeola were collected on the island of Jejudo in September of 2009. Here we provide a diagnosis and systematic account of L. flammeola. Up to now 79 species of Arctiidae have been recorded from the Korean peninsula.     

韩国斑蛾科三种幼虫的形态记述(鳞翅目)

在韩国己记述的斑蛾科昆虫有17种,这些斑蛾的幼虫主要食叶或卷叶,多数为害果树和林木等。幼虫身体短粗,头缩于前胸内,体上常有毛瘤或毛簇,腹足趾钩为单序中带。本文详细地记述了韩国斑蛾科Illiberis tenuis(Butler),Chalcosia remota(Walker)和Pryeria sinica Moore老熟幼虫的形态特征,并提供了形态特征图。