Molecular phylogeny of the green lacewings (Neuroptera: Chrysopidae)

Abstract  The first quantitative analysis of phylogenetic relationships of green lacewings (Chrysopidae) is presented based on DNA sequence data. A single nuclear and two mitochondrial genes are used in the analysis: carbomoylphosphate synthase (CPS) domain of carbamoyl-phosphate synthetase-aspartate transcarbamoylase-dihydroorotase (CAD) (i.e. rudimentary locus), large subunit ribosomal gene (16S) and cytochrome oxidase I (COI). This study represents the first use of the CAD gene to investigate phylogenetic relationships of the lacewings. DNA sequences for 33 chrysopid species from 18 genera, representing all subfamilies and tribes, were compared with outgroups sampled from families Hemerobiidae, Osmylidae and Polystoechotidae. Parsimony analyses of the combined data set recovered all of the previously established subfamilial and tribal groups as monophyletic clades (although relatively weakly supported) except Apochrysinae sensu lato . The enigmatic Nothancyla verreauxi Navás has historically been difficult to place in a subfamily group based on morphological characteristics; molecular data presented herein do not adequately resolve this problem.     

Behavioral responses of hybrid lacewings (Neuroptera: Chrysopidae) to courtship songs

The study of hybrid courtship songs and the behavioral responses of hybrids and parental individuals to hybrid songs can be useful in understanding the origin of reproductive isolation among species that differ mainly in their courtship songs. Here we test the hypotheses (a) that hybrid lacewings prefer hybrid songs to either of the parental songs from a cross betweenChrysoperla plorabunda andC. johnsoni, and (b) that parental individuals prefer their own song over those of hybrids. Analysis of songs showed that most features of hybrid songs are intermediate between the two parents. Hybrids organize their songs with a series of simple volleys like those ofC. plorabunda. Female hybrids from two reciprocal crosses and females and males of the parental species were presented with choices of hybrid and parental songs. Hybrids responded more to recordings of hybrid songs than to recordings ofC. plorabunda but did not differ in the responses given toC. johnsoni and hybrid songs. In contrast, males and females of both parental lines preferred to duet with recordings of their own song types and did not respond to hybrid songs. Our results demonstrate that hybrids would be at a disadvantage in nature, because neitherC. plorabunda norC. johnsoni will respond to their songs.     

The rare Chrysopidae (Neuroptera) of southwestern Europe

Quantitative surveys of the chrysopid fauna from southwestern Europe, namely the Iberian and Italian peninsulas, France south of 46° N, and the west-Mediterranean Islands, were analysed. A total of 56 species of Chrysopidae were reported, of which three species were abundant. These, Chrysoperla carnea (Stephens, 1836) sensu lato, Dichochrysa prasina (Burmeister, 1839) and D. flavifrons (Brauer, 1850), comprised a large percentage of the specimens. For the rarer species, comments are made on their distributions, the enhanced geographic range of exotic ones, and on levels of endemism and stenotopy.     

黄玛草蛉幼虫对草地贪夜蛾卵和低龄幼虫的捕食能力

【目的】明确本地优势天敌黄玛草蛉Mallada basalis对入侵我国的重大农业害虫草地贪夜蛾Spodoptera frugiperda卵及低龄幼虫的捕食能力和生防潜力。【方法】在实验室条件下采用功能反应模型评价了黄玛草蛉2和3龄幼虫对草地贪夜蛾卵及1和2龄幼虫的捕食能力。【结果】黄玛草蛉幼虫对草地贪夜蛾卵和低龄幼虫的捕食量均随猎物密度的升高而增加,最后趋于捕食饱和状态,而其捕食率随猎物密度的升高而逐步下降,对不同发育阶段的草地贪夜蛾均表现为Ⅱ型功能反应。黄玛草蛉2龄幼虫对草地贪夜蛾卵及1和2龄幼虫的瞬时攻击率a分别为0.150, 0.084和0.094,处理时间Th分别为0.282, 0.333和0.519 h,理论日最大捕食量T/Th分别为85106粒、72072头和46.242头;黄玛草蛉3龄幼虫对草地贪夜蛾卵及1和2龄幼虫的瞬时攻击率分别为2.018, 0.288和0.259,处理时间分别为0.102, 0.311和0.375 h,理论日最大捕食量分别为235294粒、77170头和64000头。【结论】黄玛草蛉2和3龄幼虫对草地贪夜蛾卵和低龄幼虫均具较强的捕食能力,其中黄玛草蛉3龄幼虫比2龄幼虫具有更强的捕食效率。     

不同虫态及虫龄大草蛉与七星瓢虫之间的集团内互作

【目的】为明确新疆棉田棉蚜 Aphis gossypii 捕食性天敌之间的集团内捕食效应及其对蚜虫数量的控制作用。【方法】本研究以优势天敌昆虫大草蛉 Chrysopa pallens 和七星瓢虫 Coccinella septempunctata 为对象,以棉蚜为猎物,在温室中利用盆栽棉花,首先观察了2种天敌昆虫之间各虫态及虫龄配对的19个处理在无蚜植株上共存24 h后的存活数,然后观察了2种天敌昆虫配对处理下棉苗上棉蚜数量随时间的变化趋势。【结果】在无蚜棉株上2种捕食性天敌昆虫共存24 h后的存活结果表明：（1）在发育阶段相同的配对组合中,若是成虫则均存活,若是１龄幼虫则大草蛉存活较多,若是末龄幼虫则七星瓢虫存活较多;（2）在有卵的组配中,除七星瓢虫卵不被大草蛉成虫所捕食外,其他5个组配处理中卵均被捕食;（3）在有蛹的配对组合中,除七星瓢虫蛹被大草蛉末龄幼虫捕食外,其他处理下蛹均不被捕食;（4）在成虫与幼虫的配对组合中,七星瓢虫成虫捕食较多的大草蛉1龄幼虫,但不捕食大草蛉末龄幼虫,而大草蛉成虫与七星瓢虫1龄或末龄幼虫之间不发生捕食;(5）在不同龄期幼虫的配对组合中,大草蛉末龄幼虫捕食七星瓢虫1龄幼虫,而七星瓢虫末龄幼虫捕食大草蛉1龄幼虫。在有蚜植株上2种捕食性天敌共存对棉蚜数量具有不同的控制作用：（1）2种捕食昆虫的幼虫各自单独存在（对照）下,蚜虫密度随时间而降低;（2）大草蛉幼虫与七星瓢虫幼虫或成虫配对处理下,棉蚜密度随时间而增大;（3）大草蛉成虫与七星瓢虫幼虫或成虫配对处理下,棉蚜密度随时间而减小。【结论】研究结果说明,大草蛉与七星瓢虫之间存在集团内捕食,但２种天敌共存对棉蚜的控制作用取决于大草蛉虫态, 若大草蛉为幼虫,可使蚜虫密度增大,若为成虫,则使蚜虫密度减小。     

Aggression and risk assessment during predatory interactions between first-instarSinea diadema (Fabricius) (Hemiptera: Reduviidae)

The behaviors exhibited by first-instarSinea diadema (F.) (Hemiptera: Reduviidae) during intraspecific interactions are described. Five distinct behaviors were characterized: posturing, striking, stalking, stridulating, and beaking. Distinct bouts of grappling were also noted. Interactions between unfed 2-day-old nymphs, unfed 3-day-old nymphs, and unfed 5-day-old nymphs were recorded and analyzed. The behavior of the nymphs changed with the duration of food deprivation. As the period of food deprivation increased from 2 to 5 days, the frequency of posturing decreased as the frequency of beaking increased. During noncannibalistic interactions, posturing or striking by one nymph was most frequently followed by posturing by the other nymph. In contrast, during cannibalistic interactions, posturing or striking was usually followed by beaking. The incidence of grappling increased with the duration of food deprivation. Moreover, grappling always preceded cannibalism. Although first instars were capable of capturing and killing conspecifics of the same age, cannibalism occurred only after 4 days of food deprivation. These results suggest that a decision-making process is involved. During interactions the nymphs can assess the vulnerability of the opponent. Whereas recently emerged nymphs usually forego the risk of attacking conspecifics, the attacks made by nymphs deprived of food are more persistent despite the danger of predation by their opponent.     

Aphidophagous predator diagnosis: Key to genera of European chrysopid larvae (Neur.: Chrysopidae)

Keys, based on the morphology of first- and third-instar larvae, are described for distinguishing the 13 genera of European Chrysopidae. The eggs, first-instar larvae and third-instar larvae of all genera are illustrated. Data on the biology and behaviour of the larvae are also presented.

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Résumé Dans ce travail nous présentons les caractères morphologiques des larves de Chrysopes qui sont utilisés dans la détermination des 13 genres qui habitent en Europe, ainsi que des clés pour l'identification des premiers et troisièmes stades larvaires (L-1 et L-3). Les stades L-1 et L-3 de tous les generes étudiés sont illustrés par des figures.

     

黄玛草蛉对木瓜粉蚧的捕食作用评价

为明确黄玛草蛉Mallada basalis(Walker)对木瓜粉蚧Paracoccus marginatus Williams and Granara de Willink的捕食作用,在实验室条件下观察了黄玛草蛉2龄、3龄幼虫对木瓜粉蚧2龄、3龄若虫及成虫的捕食选择偏好,同时采用捕食功能反应方法评价了黄玛草蛉2龄、3龄幼虫对木瓜粉蚧各虫态的捕食效能.结果表明,黄玛草蛉2龄、3龄幼虫偏好捕食木瓜粉蚧若虫,对木瓜粉蚧各虫态的捕食量均随着猎物密度的增加而增加符合HollingⅡ模型.其中,黄玛草蛉3龄幼虫具有较高的捕食能力,其对木瓜粉蚧2龄、3龄若虫及成虫的日最大捕食量分别为116头、81头和19头,a/Th值(a为瞬时攻击率,Th为处置单头猎物时间)分别为284.76、134.26和28.38.黄玛草蛉幼虫对猎物的寻找效应随猎物密度的增加而呈线性下降,且在相同猎物密度条件下,黄玛草蛉3龄幼虫对猎物的搜寻效应明显高于2龄幼虫.研究结果表明,黄玛草蛉对木瓜粉蚧具有较好的捕食能力.     

温度对中华通草蛉捕食麦长管蚜的影响

为明确温度对中华通草蛉 Chrysoperla sinica 捕食麦蚜能力的影响,充分发挥其控害潜能,本研究利用年龄-龄期两性生命表方法,研究了5个温度下(16℃、19℃、22℃、25℃、28℃)中华通草蛉幼虫对麦长管蚜 Sitobion avenae 的捕食能力。结果发现,1龄幼虫在19℃条件下捕食量最高,16℃条件下捕食量最低,且差异显著。2龄幼虫不同温度处理间捕食量均无显著性差异,3龄幼虫捕食麦长管蚜数量则随处理温度的提高而显著降低,而幼虫期总捕食量除28℃在不同温度间均无显著性差异。2龄幼虫平均捕食率高于1龄幼虫,但显著低于 3龄幼虫。年龄-龄期捕食率结果说明,温度升高缩短了幼虫发育时间,但提高了其捕食能力;虽然不同温度处理总捕食量不变,但高温缩短了幼虫捕食所消耗的时间。22℃条件下中华通草蛉雌成虫期长(96.63 d),产卵期长(55.36 d),繁殖力强(310.36),平均世代周期( T )短(48.99 d),容易产生世代重叠现象;内禀增长率( r )和周限增长率(λ)显示中华通草蛉种群呈正增长趋势。研究结果对于春季田间释放中华通草蛉防治麦长管蚜具有一定参考价值。     

Jerking in predaceous mites (Acari: Phytoseiidae) with emphasis on larvae

Jerking, a behavior involving a pronounced and often repeated lunging of the body, was studied in larvae among seven species of phytoseiid mites, Euseius finlandicus, Galendromus occidentalis, Neoseiulus californicus, Neoseiulus fallacis, Phytoseiulus macropilis, Phytoseiulus persimilis and Typhlodromus pyri, and in the nymphal stages of N. fallacis. This behavior was observed in larvae of six of the seven species and in all active immature stages of N. fallacis. Jerking was usually triggered by direct contact with a con- or heterospecific mite, although jerking occurred occasionally without direct contact in N. fallacis larvae. The larval jerking tendency (the mean fraction of contacts resulting in jerking) of a species was significantly correlated with a tendency to congregate with conspecifics. Congregated larvae often probed their immediate area and each other with the front legs, repeatedly jerking in response to contact with neighboring larvae. Species with larvae having higher jerking tendencies also jerked more repeatedly per jerking event. Among species, there was no consistent relationship between jerking tendency and larval age. Larvae of N. fallacis had a higher tendency to jerk than protonymphs and deutonymphs, and they had both the highest jerking and congregating tendencies among larvae of all species studied. An individual was more likely to jerk when approached by another mite than when it initiated contact, and when observed in N. fallacis larvae that initiated contact, jerking was sometimes aggressive. Jerking N. fallacis larvae experienced fewer and shorter periods of probing by the mouthparts and fewer attacks from cannibalistic adult females than anesthetized larvae that were unable to jerk.     

Chrysoperla rufilabris (Neuroptera: Chrysopidae) females do not avoid ovipositing in the presence of conspecific eggs

Previous experiments have demonstrated that green lacewing (Neuroptera: Chrysopidae) adults could be attracted to field crops using artificial honeydew. To be effective as a biological control method, such a technique would require that the increase in female abundance translate in an increase egg deposition. An experiment was conducted to evaluate whether the honeydew-feeding females of the green lacewing Chrysoperla rufilabris (Burmeister) avoid laying eggs in the presence of conspecific eggs. The potential risk associated with oviposition in a site already occupied by conspecific eggs was also studied. The preference of C. rufilabris larvae for kin and non-kin eggs and the susceptibility of C. rufilabris eggs to cannibalism relative to their age was determined. The results demonstrate that females are not reluctant to oviposit in the presence of conspecific eggs. Larvae show no preference for kin or non-kin eggs, and lacewing eggs become less susceptible to cannibalism as they age. This indicates that the risk of egg cannibalism by neonate in the field may be low. The results are discussed from ecological and biological control points of view.     

The inheritance of mating songs in two cryptic,sibling lacewing species (Neuroptera: Chrysopidae: Chrysoperla)

Speciation often involves incremental responses to natural selection and results in large scale genomic changes, but it may also occur abruptly and with little genetic imprint, as seen in some complexes of cryptic species. Recent attention has focused on sexual selection in rapid speciation, because it can disrupt premating signals that mediate reproductive isolation. Some models require that environmental adaptation assist sexual selection during speciation, while others show that populations can diverge through mate choice alone. We propose that speciation involving environmental adaptation in premating signals is likely to have a polygenic basis, while speciation due to arbitrary changes in premating signals could be accompanied by changes at just a few loci. The sibling species Chrysoperla plorabunda and Chrysoperla johnsoni belong to a large complex of cryptic species of green lacewings, and meet all criteria for speciation via sexual selection. We perform a genetic analysis of line crosses between them, focusing on their substrate-borne premating songs. Measurements of seven song features and their principal components are compared among parentals, F₁ and F₂ hybrids, and backcrosses. The distributions of phenotypes are consistent with a model of more than one, but not many, genes. Sex linkage and/or maternal effects are negligible. C. plorabunda is dominant for most traits affecting mate choice. Bayesian analyses of segregation variance show significant additive and epistatic effects on line means. A Bayesian Castle–Wright estimate suggests that relatively few effective factors are responsible for variation in volley period ( ), number of volleys per song ( ), and PC-1 ( ). Our results are consistent with simple genetic architecture of songs, supporting a role for major genes in premating isolation and strengthening the notion that mate choice alone, without significant environmental adaptation, has been responsible for generating new lacewing species.     

Trichome‐based host plant resistance of Lycopersicon species and the biocontrol agent Mallada signata: are they compatible?

Trichome‐based host plant resistance of Lycopersicon (Solanaceae) species offers the potential to reduce pesticide use in tomato production, but its compatibility with biocontrol agents is largely unknown. The effect of trichome‐based host plant resistance on the lacewing biocontrol agent, Mallada signata, was assessed for five accessions of L. cheesmanii, four accessions of Lycopersicon hirsutum, two accessions of Lycopersicon pennellii, and one Lycopersicon esculentum cultivar. An intact leaf was isolated from the whole plant using Tangletrap to coat the petiole and 20 green peach aphids [Myzus persicae (Sulzer) (Homoptera: Aphididae)] were placed on the leaf surface. After 24 h, 10 lacewings were placed on the leaf. The numbers of dead, trapped by exudates, untrapped and predated lacewings and aphids, and the numbers that had left the leaf were recorded a further 24 h later. Differences in insect designations between accessions were analysed using ANOVA. A General Linear Model (GLM), consisting of the densities of each trichome type and leaf area, was fitted to the data to determine the role of trichomes on the observed effects on lacewings and aphids. Lacewing mortality was greater on one accession of L. pennellii and one accession of L. hirsutum than on L. esculentum. The GLM indicated that type IV trichomes decreased the numbers of aphids predated, and increased cannibalism and, along with type III trichomes, increased entrapment‐related predator mortality. Although there were no differences in the numbers of predated aphids, with the majority predated for all accessions, the compatibility of trichome‐based host plant resistance of Lycopersicon species and the biocontrol of aphids by lacewings is questionable.     

Comparative biology,predation capacity and effect of an artificial diet on reproductive parameters of green lacewing Mallada boninensis (Neuroptera: Chrysopidae)

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Prey instar preference and functional responses of Mallada basalis (Walker) (Neuroptera: Chrysopidae) to different life stages of Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae)

Prey instar preference and functional responses of 2- and 3-instar Mallada basalis (Walker) (Neuroptera: Chrysopidae) larvae to 1- to 3-instar Phenacoccus solenopsis Tinsley (Hemiptera: Pseudococcidae) nymphs and adults were assessed in laboratory. Results indicated that both 2- and 3-instar M. basalis larvae preferred young over old P. solenopsis nymphs and adults were the least preferred. The 3-instar M. basalis larvae preyed more adult P. solenopsis than 2-instar larvae. Mallada basalis exhibited type II functional responses to prey densities: An increase in prey density leads to an increase in consumed preys. Regardless of P. solenopsis stages, the number of preys consumed by the 3-instar M. basalis larvae was greater than that by the 2-instar larvae. Attack rates and handling times differed depending on prey and predator stage combinations. The highest attack rate (1.1874) and lowest handling time (0.0040 h) were observed for the 3-instar M. basalis larvae fed on the 1-instar P. solenopsis nymphs. Regardless of P. solenopsis stages, the attack rate of 3-instar M. basalis was greater than 2-instar, whereas the reverse held regarding handling time. The findings collectively indicated that 3-instar M. basalis larvae have greater potential than 2-instar as efficient biological control agent of P. solenopsis. In developing real world biological control programs, however, the 2-instar M. basalis may be released if necessary since the final efficacy of the predator is the summation of the 2- and 3-instar M. basalis.     

Efficacy of the predator Mallada basalis (Neuroptera: Chrysopidae) on Tetranychus kanzawai and Panonychus citri (Acari: Tetranychidae) at different predator:prey release ratios

We compared population suppression of the phytophagous mites, Tetranychus kanzawai Kishida and Panonychus citri (McGregor), on papaya by second instar larvae of the green lacewing, Mallada basalis (Walker), at various predator:prey release ratios in the laboratory. Initially, we presented M. basalis with mixed age classes of each mite species separately at a density of approximately 30 mites per seedling. After 3 days, predator:prey ratios of 1:30, 1:15, and 1:10 resulted in reductions of T. kanzawai of 66.8%, 82.6%, and 83.3%, respectively, and reductions of P. citri of 41.8%, 75.5%, and 77.2%, respectively. Predation on individual age classes was approximately equal in both species, reinforcing previous findings that this predator does not show a preference among age classes. We next presented M. basalis with mixed populations of the two mite species in which there were equal numbers of each species and the density was as in the single species tests. Total mite reduction with both mite species present was 48.5%, 71.9%, and 74.5% at ratios of 1:30, 1:15, and 1:10, respectively; T. kanzawai was reduced by 50.5%, 77.4%, and 79.5%, respectively, and P. citri was reduced by 44.1%, 60.3%, and 63.2%, respectively. This study suggests that M. basalis has the potential for substantially suppressing populations of both T. kanzawai and P. citri on papaya at a predator:prey ratio of 1:15 or greater. However, evaluation under realistic agricultural settings is needed before specific recommendations about predator release rates can be made.     

Rearing of Chrysopa pallens (Rambur) (Neuroptera: Chrysopidae) on Artificial Diet

An artificial rearing method was designed for the generalist predator, Chrysopa pallens (Rambur). The rearing media were formulated based on, Orius strigicollis diets for feeding larvae, and two diets for adults were prepared with insect source and non‐insect as their main components. Development of the predators was successfully obtained with the diets for both larval and adult stages. The impact of these diets was recorded for growth during the larval stage and oviposition rates by the females. Total development period and mortality rate of, C. pallens were about 26.9 days and 11%, respectively. The females showed better reproduction with average of 2019 eggs over their life span of 89 days when the artificial diet with non‐insect source was provided. On overall basis the life expectancy and fecundity was better when compared with previous findings.     

Evolution of green lacewings (Neuroptera: Chrysopidae): a molecular supermatrix approach

We present a time‐calibrated phylogeny of the charismatic green lacewings (Neuroptera: Chrysopidae). Previous phylogenetic studies on the family using DNA sequences have suffered from sparse taxon sampling and/or limited amounts of data. Here we combine all available previously published DNA sequence data and add to it new DNA sequences generated for this study. We analysed these data in a supermatrix using Bayesian and maximum likelihood methods and provide a phylogenetic hypothesis for the family that recovers strong support for the monophyly of all subfamilies and resolves relationships among a large proportion of chrysopine genera. Chrysopinae tribes Leucochrysini and Belonopterygini were recovered as monophyletic sister clades, while the species‐rich tribe Chrysopini was rendered paraphyletic by Ankylopterygini. Relationships among the subfamilies were resolved, although with relatively low statistical support, and the topology varied based on the method of analysis. Greatest support was found for Apochrysinae as sister to Nothochrysinae and Chrysopinae, which is in contrast to traditional concepts that place Nothochrysinae as sister to the rest of the family. Divergence estimates suggest that the stem groups to the various subfamilies diverged during the Triassic‐Jurassic, and that stem groups of the chrysopine tribes diverged during the Cretaceous.