Relationships among the basal lineages of Noctuidae (Lepidoptera,Noctuoidea) based on eight gene regions

In this study, we clarify the relationships between the basal lineages in the moth family Noctuidae using DNA sequence data from eight independent gene regions. Data matrices (6.4 kbp) are analysed using parsimony and model‐based methods (maximum likelihood and Bayesian inference). Our results support the family Noctuidae as a monophyletic group in which most subfamilies have hindwing vein M2 reduced or absent. Our phylogenetic hypothesis suggests that in the Noctuidae, the plesiomorphic condition is that in which vein M2 arises about one‐third of the way up the discocellular vein between the origins of M1 and M3, mainly parallel to M3, and is of thickness similar to vein M3. Most Noctuidae lineages possess an apomorphic (derived) condition in which hindwing vein M2 is markedly reduced or totally absent, so that the cubital vein appears to be three‐branched and these lineages are hence referred to as ‘trifine’. However, Noctuidae also include a number of lineages in which vein M2 is unreduced, or only slightly reduced, and these are more problematic for morphological association with the family Noctuidae. Our results also show that the subfamily Acronictinae is not closely related to Pantheinae, but instead shows a closer association with Amphipyrinae. Among the major lineages of Noctuidae, we postulate a general trend, with numerous exceptions, in larval host plants from woody plants in the basal groups towards herb feeding in derived groups. Similarly, the major radiations of monocot‐feeding groups within the family Noctuidae are in the higher trifines. The following taxonomic changes are proposed: Thiacidinae, syn. nov., a junior synonym of Pantheinae, and Dyopsinae, stat. nov., are reinstated as a subfamily.     

Molecular phylogenetics of Erebidae (Lepidoptera,Noctuoidea)

As a step towards understanding the higher‐level phylogeny and evolutionary affinities of quadrifid noctuoid moths, we have undertaken the first large‐scale molecular phylogenetic analysis of the moth family Erebidae, including almost all subfamilies, as well as most tribes and subtribes. DNA sequence data for one mitochondrial gene (COI) and seven nuclear genes (EF‐1α, wingless, RpS5, IDH, MDH, GAPDH and CAD) were analysed for a total of 237 taxa, principally type genera of higher taxa. Data matrices (6407 bp in total) were analysed by parsimony with equal weighting and model‐based evolutionary methods (maximum likelihood), which revealed a well‐resolved skeleton phylogenetic hypothesis with 18 major lineages, which we treat here as subfamilies of Erebidae. We thus present a new phylogeny for Erebidae consisting of 18 moderate to strongly supported subfamilies: Scoliopteryginae, Rivulinae, Anobinae, Hypeninae, Lymantriinae, Pangraptinae, Herminiinae, Aganainae, Arctiinae, Calpinae, Hypocalinae, Eulepidotinae, Toxocampinae, Tinoliinae, Scolecocampinae, Hypenodinae, Boletobiinae and Erebinae. Where possible, each monophyletic lineage is diagnosed by autapomorphic morphological character states, and within each subfamily, monophyletic tribes and subtribes can be circumscribed, most of which can also be diagnosed by morphological apomorphies. All additional taxa sampled fell within one of the four previously recognized quadrifid families (mostly into Erebidae), which are now found to include two unusual monobasic taxa from New Guinea: Cocytiinae (now in Erebidae: Erebinae) and Eucocytiinae (now in Noctuidae: Pantheinae).     

The Noctuoidea (Lepidoptera) described by Linnaeus

Linnaeus described 166 nominal species of Noctuoidea, of which 144 (86.7%) are currently used as valid names, 18 are synonyms, one is a homonym and three taxa remain unknown. Seven taxa were described solely from illustrations in contemporary literature. Lectotypes are designated for 139 (87.4%) of the 159 remaining taxa, 122 of them from the Linnaean collection in London, 13 from the Queen Ludovica Ulrica's collection in Uppsala, Sweden, and four from other contemporary collections. Two Linnaean senior synonyms are introduced: Hada plebeja comb. nov. (Linnaeus, 1761) for the current H. nana (Hufnagel, 1766) and Oeonistis altica (Linnaeus, 1768) comb. nov. for Oe. entella (Cramer, 1779). Erebus occiduus (Linnaeus, 1758) syn. nov. is a junior subjective synonym of E. crepuscularis (Linnaeus, 1758); L. marmorides (Cramer, 1775) Stat. rev. should be reinstated for Letis occidua auct. In two species pairs the Linnaean names have been interpreted incorrectly. The lectotype designation of Phalaena prasinana Linnaeus, 1758, by Lempke (1947) is invalid because the specimen is not syntypic. According to the new lectotype, Pseudoips prasinanus (Linnaeus, 1758) comb. nov. replaces P. faganus (Fabricius, 1781), and Bena bicolorana (Fuessly, 1775) comb. nov. is available for the other species. Abrostola triplasia (Linnaeus, 1758) Stat. rev. replaces A. trigemina (Werneburg, 1864), and for the other species A. triparlila (Hufnagel, 1766) should be reinstated. One neotype designation [Eilema complanum (Linnaeus, 1758)], one lectotype designation [Xylena exsolela (Linnaeus, 1758)], and precedence of a junior synonym [Ophideres fullonia (Clerck, 1764)] over its unused senior synonym are to be referred to the ICZN; until their decision the current usage must be followed. The identity of Mesapamea secalis (Linnaeus, 1758) stat. rev. corresponds to current usage, but, under presence of syntypic material, the recent neotype designation by Lempke (1988) will be referred to the ICZN. The revision is mainly based on the discovery that the insect pins and the way specimens are prepared give evidence of the authenticity of the material. Contrary to the views of earlier authors, Linnaeus had his labels with the specific name pinned against the bottom of the drawers by the specimen pin. Because the labels have been moved twice during subsequent curations, their present position is less indicative of the authenticity than has been previously suggested. The origin of the so-called n-labels is discussed.     

Major lineages of Nolidae (Lepidoptera,Noctuoidea) elucidated by molecular phylogenetics

To elucidate the evolutionary relationships of the major lineages within the moth family Nolidae, we analysed a molecular dataset comprising eight independent gene regions (6.4 kbp), cytochrome c oxidase subunit I (COI) from the mitochondrial genome, and elongation factor‐1α (EF‐1α), ribosomal protein S5 (RpS5), carbamoylphosphate synthase domain protein (CAD), cytosolic malate dehydrogenase (MDH), glyceraldehyde‐3‐phosphate dehydrogenase (GAPDH), isocitrate dehydrogenase (IDH) and wingless genes from the nuclear genome, using parsimony and model‐based evolutionary methods (maximum likelihood and Bayesian inference). Our analyses revealed a well‐resolved phylogenetic hypothesis, again recovering the six previously recognized families within Noctuoidea (i.e. Oenosandridae, Notodontidae, Euteliidae, Erebidae, Nolidae and Noctuidae), and monophyly of the quadrifid Noctuoidea (i.e. Euteliidae, Erebidae, Nolidae and Noctuidae). The family Nolidae is diagnosed and characterized by two synapomorphies from morphology: construction of a ridged boat‐shaped cocoon that bears a vertical exit slit at one end; and two other morphological character states: elongation of the forewing retinaculum into a bar‐like or digitate condition and possession of a postpiracular counter‐tympanal hood. We present a new phylogenetic hypothesis for Nolidae consisting of eight strongly supported subfamilies, two of which are erected here: Diphtherinae, Risobinae, Collomeninae subfam. nov., Beaninae subfam. nov., Eligminae, Westermanniinae, Nolinae and Chloephorinae. Where we are able, each monophyletic lineage is diagnosed by morphological autapomorphies and within each subfamily, monophyletic tribes and subtribes are circumscribed, most of which are also diagnosable by morphological apomorphies. We also describe two new taxa: Gelastocerini trib. nov. and Etannina subtrib. nov. The Neotropical subfamily Diphtherinae, here newly circumscribed, is considered to be the plesiomorphic sister lineage to the rest of Nolidae. Diphtherinae are characterized by loss of the proximal pair of metatibial spurs in males and by the presence of a frontal tubercle, which is presumably associated with a derived strategy of emergence from the cocoon.     

Rapid antibody-based field test to distinguish between Helicoverpa armigera (Lepidoptera: Noctuidae) and Helicoverpa punctigera (Lepidoptera: Noctuidae)

Helicoverpa armigera (Hübner) and Helicoverpa punctigera (Wallengren) are the two most important insect pests of cotton production in Australia and require application of insecticides to control them. H. armigera has developed resistance to several insecticides but H. punctigera has not. Cost-effective management of insecticide resistance requires that growers be able to determine the proportion of H. armigera eggs or young larvae present on their crop before applying insecticides. This is impossible visually. We generated two monoclonal antibodies that reacted with the insect protein "lipophorin" and were capable of discriminating individuals of the two species at all life-stages. The antibodies were incorporated into a rapid test kit that was tested under field conditions over two growing seasons. Results obtained with the kit agreed closely with those obtained by rearing larvae through to second instar.     

The Lepidoptera of White Sands National Monument, Otero County, New Mexico, USA 3. A new species of Aleptina Dyar, 1902 (Lepidoptera, Noctuidae, Amphipyrinae, Psaphidini)

In 2006 the US National Park Service initiated a long-term study of the Lepidoptera at White Sands National Monument, Otero County, New Mexico. Aleptina arenariasp. n., described here, was discovered in 2008, the second year of the study. The adult moths and male and female genitalia are illustrated.     

Towards resolving the identities of the Graphium butterflies (Lepidoptera: Papilionidae) of Peninsular Malaysia

Graphium butterflies are famous in Peninsular Malaysia for their colourful wings, yet their taxonomy remains unresolved. The popular guides to Malaysian butterflies, place the species in one, two or three genera and identification to species using obscure morphological characters can be difficult, especially for the closely related species, G. bathycles bathycloides and G. chironides malayanum. We sequenced the COI mtDNA barcode for Graphium specimens in the Museum of Zoology, University of Malaya to test the utility of DNA barcoding for the identification of Graphium species. Additionally, we sequenced 28S rRNA to examine, in conjunction with COI, the phylogenetic relationships of these species and investigate the validity of Pathysa and Paranticopsis as distinct genera. We found that all species of Graphium possessed a distinctive cluster of DNA barcodes with the exception of specimens originally identified as G. bathycles bathycloides and G. chironides malayanum which shared DNA barcodes. On further examination we found that the morphological determinations were ambiguous as the specimens overlapped for diagnostic characters reported for each species. The COI and 28S rRNA phylogenetic trees showed a similar topology with Paranticopsis species forming a clade nested within a larger clade also comprising Pathysa species. Based on this topology, in order for Pathysa to be a valid genus, at least three other clades within Graphium s.l. would also have to be raised as genera.     

Larval hemocytes in Noctuidae (Insecta: Lepidoptera)

An attempt was made to relate certain physical characteristics of hemocytes and the taxonomy or ecophysiology of species in one family of insects. Composition of the hemocyte complex and features of the cell types varied widely among larvae of 85 species sampled from the family Noctuidae (Lepidoptera). The complex ranged from a basic one of 4 functioning types, the plasmatocytes, granulocytes, spherulocytes and oenocytoids, to ones that also included extreme variants of the plasmatocyte line. Except for the granolocytes, each type had features of size, form and structure that could vary also with species. Some species can be identified by these characters, but there is no apparent relationship of the characters to species biology and little to present taxonomic arrangements.     

New Records of Noctuoid Moths (Lepidoptera,Noctuoidea) in the Baikal Region

The results of long-term faunistic studies of the noctuoid moths in the Baikal region and the literature review are presented. An annotated check-list including 83 species of the families Notodontidae, Erebidae, Nolidae, and Noctuidae is given. Data on the previous records of the species from the study region are given. The Noctuoidea fauna of Irkutsk Province comprises 462 species, that of the Republic of Buryatia, 419 species, and that of the Baikal region as a whole, 510 species. The distribution limits are expanded for 17 species of the Noctuoidea complex: the new eastern range boundary is established for 11 species of Erebidae and Noctuidae, and the new western one is established for 5 representatives of Notodontidae, Erebidae and Noctuidae. Two species, Actebia confinis (Staudinger, 1881) and Actebia confusa (Alphéraky, 1882), are new to the Russian fauna. The trophic associations of Phidrimana amurensis (Staudinger, 1892) are established: the larvae of this Euro-Siberian species feed on the leaves of different species of elms (Ulmus japonica Rehder, Ulmus pumila L.). Three noctuid species, Calyptra lata (Butler, 1881), Bryophila granitalis (Butler, 1881) and Cosmia trapezinula (Filipjev, 1927), with large disjunctions of their ranges are reported; these moths are apparently relicts of the Atlantic Period of the Holocene.     

Molecular phylogeny of Lymantriinae (Lepidoptera,Noctuoidea, Erebidae) inferred from eight gene regions

To understand the evolutionary history of Lymantriinae and test the present higher‐level classification, we performed the first broad‐scale molecular phylogenetic analysis of the subfamily, based on 154 exemplars representing all recognized tribes and drawn from all major biogeographical regions. We used two mitochondrial genes (cytochrome c oxidase subunit I and 16S ribosomal RNA) and six nuclear genes (elongation factor‐1α, carbamoylphosphate synthase domain protein, ribosomal protein S5, cytosolic malate dehydrogenase, glyceraldehyde‐3‐phosphate dehydrogenase and wingless). Data matrices (in total 5424 bp) were analysed by parsimony and model‐based evolutionary methods (maximum likelihood and Bayesian inference). Based on the results of the analyses, we present a new phylogenetic classification for Lymantriinae composed of seven well‐supported tribes, two of which are proposed here as new: Arctornithini, Leucomini, Lymantriini, Orgyiini, Nygmiini, Daplasini trib. nov. and Locharnini trib. nov. We discuss the internal structure of each of these tribes and address some of the more complex problems with the genus‐level classification, particularly within Orgyiini and Nygmiini.     

Genetic similarity and distribution of cry genes of Bacillus thuringiensis pathogenic for Spodoptera exigua (Lepidoptera: Noctuidae) and Agrotis segetum (Lepidoptera: Noctuidae)

Crystals of Bacillus thuringiensis isolates, obtained from dead caterpillars, had potential toxicity against insects in the orders Diptera, Coleoptera, Hemiptera and Lepidoptera. Strain MPU B32b had the highest number and variety of cry genes and seemed to be the most interesting for further research on development as a novel biopesticide.     

Acoustic irradiation produced by flying moths (Lepidoptera,Noctuidae)

Characteristics of acoustic waves accompanying the flight of noctuid moths (Noctuidae) were measured. The low-frequency part of the spectrum is formed of a series of up to 17 harmonics of the wingbeat frequency (30–50 Hz) with a general tendency toward the decrease in the spectral density and the increase in the sound frequency. The root-mean-square level of the sound pressure from flapping wings was found to be 70–78 dB SPL. Besides low-frequency components, the flight of moths was accompanied by short ultrasonic pulses, which appeared with every wingbeat. Most of the spectral energy was concentrated within a range of 7–150 kHz with the main peaks at 60–110 kHz. The short-term pulses were divided into two or more subpulses with different spectra. The high-frequency pulses were produced at two phases of the wingbeat cycle: during the pronation of the wings at the highest point and at the beginning of their upward movement from the lowest point. In most of the specimens tested, the peak amplitude of sounds varied from 55 to 65 dB SPL at a distance of 6 cm from the insect body. However, in nine noctuid species, no high-frequency acoustic components were recorded. In these experiments, the acoustic flow from the flying moth within a frequency range of 2 to 20 kHz did not exceed the self-noise level of the microphone amplifier (RMS 18 dB SPL). Probable mechanisms of the high frequency acoustic emission during flight, the effect of these sounds on the auditory sensitivity of moths, and the possibility of their self-revealing to insectivorous bats are discussed. In addition, spectral characteristics of the moth echolocation clicks were more precisely determined within the higher frequency range (>100 kHz).     

The phylogeny of the Noctuidae (Lepidoptera)

A new phylogenetic system of the Noctuidae is proposed. The system should, however, be regarded as provisional because it is primarily based on only two character complexes: the male genitalia and the tympanal region. The presence of preabdominal brush-organs and the length of tibial spurs in the adult male, the presence of a ventral cervical gland and the lack of the SV 2 setae on the first abdominal segment of the larvae are considered. The results confirm the monophyly of the Noctuidae. The Herminiinae are considered to represent the sister-group of all other Noctuidae. Remaining noctuid subfamilies are grouped in a series of clades for which the corresponding autapomorphies are given. Some formerly accepted subfamilies, such as the Acontiinae, prove to be polyphyletic. Accordingly, these subfamilies are subdivided into separate lineages. A sister-group relationship between the Noctuidae and the Aganaidae is supported. These two families may be united with the Arctiidae to form a still broader clade.     

旋歧夜蛾形态和生活史观察

【目的】初步研究旋歧夜蛾Discestratrifolii(Hufnagel)的形态特征、生物学特性以及生活史。【方法】通过室内观察,数据测量与统计,总结与分析了旋歧夜蛾的形态、生物学特性等问题。【结果】在室内饲养条件下,旋歧夜蛾幼虫6龄,旋歧夜蛾完成1代的生命周期为(44.27±3.10)d,卵期(4.02±0.35)d,存活率96.97%±1.20%,幼虫期(17.87±1.40)d,存活率75.76%±6.94%,蛹期(10.48±0.80)d,存活率93.86%±0.46%,成虫期(12.62±2.82)d。【结论】基本明确了旋歧夜蛾的形态和生活史,有助于制定有效的旋歧夜蛾防治措施,同时为旋歧夜蛾其它研究奠定基础。     

Conventional resistance of experimental maize lines to corn earworm (Lepidoptera: Noctuidae), fall armyworm (Lepidoptera: Noctuidae), southwestern corn borer (Lepidoptera: Crambidae), and sugarcane borer (Lepidoptera: Crambidae)

Plant resistance is a useful component of integrated pest management for several insects that are economically damaging to maize, Zea mays L. In this study, 15 experimental lines of maize derived from a backcross breeding program were evaluated for resistance to corn earworm, Helicoverpa zea (Boddie); fall armyworm, Spodoptera frugiperda (J. E. Smith); southwestern corn borer, Diatraea grandiosella Dyar; and sugarcane borer, Diatraea saccharalis (F.). Experimental line 100-R-3 was resistant in the field to leaf feeding by fall armyworm and line 116-B-10 was resistant in the field to leaf feeding by fall armyworm and leaf and stalk feeding by southwestern corn borer. When corn earworm larvae were fed field harvested silks from experimental line 81-9-B in the laboratory, their pupal weights were significantly lower than the pupal weights of larvae that were fed silks from the resistant control, Zapalote Chico. Maysin levels lower than those commonly associated with corn earworm resistance were present in the resistant experimental line, 107-8-7, indicating a new basis confers resistance to corn earworm in this line. These resistant experimental lines will provide plant breeders with new sources of resistance to lepidopterous insects for the development of improved maize breeding populations.     

Cold hardiness of Helicoverpa zea (Lepidoptera: Noctuidae) pupae

An insect's cold hardiness affects its potential to overwinter and outbreak in different geographic regions. In this study, we characterized the response of Helicoverpa zea (Boddie) pupae to low temperatures by using controlled laboratory measurements of supercooling point (SCP), lower lethal temperature (LT(50)), and lower lethal time (LLTime). The impact of diapause, acclimation, and sex on the cold hardiness of the pupae also were evaluated. Sex did not significantly affect the SCP, LT(50), or LLTime. However, the mean SCP of diapausing pupae (-19.3°C) was significantly lower than nondiapausing pupae (-16.4°C). Acclimation of nondiapausing pupae to constant temperatures from 10 to 20°C before supercooling also produced a significantly lower SCP than nondiapausing pupae held at 25°C. The LT(50)s of nondiapausing and diapausing were not significantly different, but confirmed that H. zea pupae are chill-intolerant because these lethal temperatures are warmer than the corresponding mean SCPs. Diapausing pupae survived longer than nondiapausing pupae at the same, constant, cold temperatures, a finding consistent with the SCP results. Both of these results suggest enhanced cold hardiness in diapausing pupae. When laboratory results were compared with field temperatures and observed distributions of H. zea in the contiguous United States, the laboratory results corroborated what is currently perceived to be the northern overwintering limit of H. zea; approximately the 40(th) parallel. Moreover, our research showed that areas north of this limit are lethal to overwintering pupae not because of low temperature extremes, but rather the length of time spent at near-zero temperatures.     

Sex pheromone precursors in Spodoptera littoralis (Lepidoptera: Noctuidae)

Female pheromone gland extracts of Spodoptera littoralis were analyzed for pheromone precursors. Large amounts of fatty methyl esters were found and a positive relationship between the methyl esters and the pheromonal components was observed. The esters were identified on the basis of capillary gas chromatography, coupled gas chromatography with mass spectroscopy and dimethyl disulfide derivatization, and subsequent gas chromatography with mass spectrometry. The characteristic fatty esters of S. littoralis are methyl (Z)-9-tetradecenoate, (E)- and (Z)-11-tetradecenoate, (Z,E)-9,12-tetradecadienoate, (Z,E)-9,11-tetradecadienoate, and (Z)-11-hexadecenoate. The biosynthesis of the monosaturated acids involves probably the common E11 and Z11 desaturases and chain shortening. For the biosynthesis of the novel diene acids, we propose a second, specific desaturation of (Z)-9-tetradecenoate by an E11 desaturase to produce (Z,E)-9,11-tetradecadienoate or by an E12 desaturase to produce (Z,E)-9,12-tetradecadienoate.