Familial Clarification of Saucrosmylidae stat. nov. and New Saucrosmylids from Daohugou, China (Insecta, Neuroptera)

PubMed Central

Fang, Hui; Ren, Dong; Wang, Yongjie

2015-01-01

Backgound Saucrosmylids are characterized by the typically large body size, complicated venation and diverse wing markings, which were only discovered in Middle Jurassic of Daohugou, Ningcheng county, Inner Mongolia, China. Principal Findings Saucrosmylinae Ren, 2003, originally included as a subfamily in the Osmylidae, was transferred and elevated to family rank based on the definitive synapomorphic character. The updated definition of Saucrosmylidae stat. nov. was outlined in detail: presence of nygma and trichosors; diverse markings on membrane; complicated cross-veins; distal fusion of Sc and R1; expanded space between R1 and Rs having 2–7 rows of cells that should be a synapomorphic character of the family; proximal MP fork. And the previous misuses of Saucrosmylidae are also clarified. Furthermore, a new genus with a new species and an indeterminate species of Saucrosmylidae are described as Ulrikezza aspoeckae gen. et sp. nov. and Ulrikezza sp. from the Middle Jurassic of Daohugou, Inner Mongolia, China. A key to genera of Saucrosmylidae is provided. Conclusions/Significance The intriguing group represents a particular lineage of Neuroptera in the Mesozoic Era. The familial status of Saucrosmylidae was firstly advanced that clarified the former incorrect citation and use of the family name. As an extinct clade, many species of the saucrosmylids were erected just based on a single fore- or hindwing, and it should be realized that providing more stable characters is necessary when describing new lacewing taxa just based on an isolated hindwing. It is vital for the systematics of Saucrosmylidae. PMID:26485027

Flight mill performance of the lacewing Chrysoperla sinica (Neuroptera: Chrysopidae) as a function of age, temperature, and relative humidity.

PubMed

Liu, Zhongfang; McNeil, Jeremy N; Wu, Kongming

2011-02-01

The lacewing Chrysoperla sinica (Tjeder) (Neuroptera: Chrysopidae) is an important predator of several insect pests in China and has considerable potential as a biological control agent. An inoculative approach would be the releasing adults early in the season to ensure that populations are present before pest densities increase. However, an understanding of adult flight activity under different conditions is necessary to develop appropriate release strategies. Therefore, we used a 32-channel, computer-monitored flight mill system to determine the effect of age on the flight activity of unmated female and male adults. Both sexes had high total flight activity levels as well as the longest individual flight bouts 2 and 3 d after emergence. The effects of temperature (between 13 and 33 degrees C at 75% RH) and relative humidity (between 30 and 90% RH at 23 degrees C) on the flight activity of 3-d-old unmated adults also were determined. Flight activity declined at the lowest (13 degrees C) and highest (33 degrees C) temperatures tested, as well as at the lowest relative humidity (30% RH). These findings are discussed within the context of selecting the appropriate environmental conditions for releasing C. sinica.

First Transcriptome and Digital Gene Expression Analysis in Neuroptera with an Emphasis on Chemoreception Genes in Chrysopa pallens (Rambur)

PubMed Central

Li, Zhao-Qun; Zhang, Shuai; Ma, Yan; Luo, Jun-Yu; Wang, Chun-Yi; Lv, Li-Min; Dong, Shuang-Lin; Cui, Jin-Jie

2013-01-01

Background Chrysopa pallens (Rambur) are the most important natural enemies and predators of various agricultural pests. Understanding the sophisticated olfactory system in insect antennae is crucial for studying the physiological bases of olfaction and also could lead to effective applications of C. pallens in integrated pest management. However no transcriptome information is available for Neuroptera, and sequence data for C. pallens are scarce, so obtaining more sequence data is a priority for researchers on this species. Results To facilitate identifying sets of genes involved in olfaction, a normalized transcriptome of C. pallens was sequenced. A total of 104,603 contigs were obtained and assembled into 10,662 clusters and 39,734 singletons; 20,524 were annotated based on BLASTX analyses. A large number of candidate chemosensory genes were identified, including 14 odorant-binding proteins (OBPs), 22 chemosensory proteins (CSPs), 16 ionotropic receptors, 14 odorant receptors, and genes potentially involved in olfactory modulation. To better understand the OBPs, CSPs and cytochrome P450s, phylogenetic trees were constructed. In addition, 10 digital gene expression libraries of different tissues were constructed and gene expression profiles were compared among different tissues in males and females. Conclusions Our results provide a basis for exploring the mechanisms of chemoreception in C. pallens, as well as other insects. The evolutionary analyses in our study provide new insights into the differentiation and evolution of insect OBPs and CSPs. Our study provided large-scale sequence information for further studies in C. pallens. PMID:23826220

First Transcriptome and Digital Gene Expression Analysis in Neuroptera with an Emphasis on Chemoreception Genes in Chrysopa pallens (Rambur).

PubMed

Li, Zhao-Qun; Zhang, Shuai; Ma, Yan; Luo, Jun-Yu; Wang, Chun-Yi; Lv, Li-Min; Dong, Shuang-Lin; Cui, Jin-Jie

2013-01-01

Chrysopa pallens (Rambur) are the most important natural enemies and predators of various agricultural pests. Understanding the sophisticated olfactory system in insect antennae is crucial for studying the physiological bases of olfaction and also could lead to effective applications of C. pallens in integrated pest management. However no transcriptome information is available for Neuroptera, and sequence data for C. pallens are scarce, so obtaining more sequence data is a priority for researchers on this species. To facilitate identifying sets of genes involved in olfaction, a normalized transcriptome of C. pallens was sequenced. A total of 104,603 contigs were obtained and assembled into 10,662 clusters and 39,734 singletons; 20,524 were annotated based on BLASTX analyses. A large number of candidate chemosensory genes were identified, including 14 odorant-binding proteins (OBPs), 22 chemosensory proteins (CSPs), 16 ionotropic receptors, 14 odorant receptors, and genes potentially involved in olfactory modulation. To better understand the OBPs, CSPs and cytochrome P450s, phylogenetic trees were constructed. In addition, 10 digital gene expression libraries of different tissues were constructed and gene expression profiles were compared among different tissues in males and females. Our results provide a basis for exploring the mechanisms of chemoreception in C. pallens, as well as other insects. The evolutionary analyses in our study provide new insights into the differentiation and evolution of insect OBPs and CSPs. Our study provided large-scale sequence information for further studies in C. pallens.

Host Plant-Herbivore-Predator Interactions in Chrysoperla carnea (Neuroptera: Chrysopidae) and Myzus persicae (Homoptera: Aphididae) on Four Plant Species Under Laboratory Conditions.

PubMed

Farrokhi, Milad; Gharekhani, Gholamhossein; Iranipour, Shahzad; Hassanpour, Mahdi

2017-12-05

The common green lacewing, Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae), is a well-known biocontrol agent. The current study examined host plant-herbivore-predator interactions with C. carnea and Myzus persicae on four host plants (peach, almond, pepper, and potato). The experiments were carried out at 25 ± 1°C and 65 ± 5% RH at a photoperiod of 16:8 (L:D) h). Duration of the preadult growth period, adult longevity, fecundity, and population growth parameters were analyzed based on the age-stage, two-sex life table theory. The shortest and longest preadult developmental times of the predator were observed on the peach and potato, respectively. The highest and lowest predation rate, oviposition period, and male and female longevity of predator were also observed on the peach and potato, respectively. The lowest intrinsic rate of increase (r) and finite rate of increase (λ) were observed on the potato (0.1087 and 1.11 d-1, respectively) and the highest on the peach (0.1460 and 1.15 d-1, respectively). The maximum and minimum mean generation times (T) were 41.84 and 35.59 d in the potato and peach, respectively. Overall, peach was found to be a more appropriate host than the other host plants for development and predation fitness of C. carnea. These findings reveal that information on tritrophic interactions and subsequent life table evaluation of natural enemies improves integrated pest management programs. Published by Oxford University Press on behalf of Entomological Society of America 2017. This work is written by (a) US Government employee(s) and is in the public domain in the US.

The mitochondrial genome of the ascalaphid owlfly Libelloides macaronius and comparative evolutionary mitochondriomics of neuropterid insects

PubMed Central

2011-01-01

Background The insect order Neuroptera encompasses more than 5,700 described species. To date, only three neuropteran mitochondrial genomes have been fully and one partly sequenced. Current knowledge on neuropteran mitochondrial genomes is limited, and new data are strongly required. In the present work, the mitochondrial genome of the ascalaphid owlfly Libelloides macaronius is described and compared with the known neuropterid mitochondrial genomes: Megaloptera, Neuroptera and Raphidioptera. These analyses are further extended to other endopterygotan orders. Results The mitochondrial genome of L. macaronius is a circular molecule 15,890 bp long. It includes the entire set of 37 genes usually present in animal mitochondrial genomes. The gene order of this newly sequenced genome is unique among Neuroptera and differs from the ancestral type of insects in the translocation of trnC. The L. macaronius genome shows the lowest A+T content (74.50%) among known neuropterid genomes. Protein-coding genes possess the typical mitochondrial start codons, except for cox1, which has an unusual ACG. Comparisons among endopterygotan mitochondrial genomes showed that A+T content and AT/GC-skews exhibit a broad range of variation among 84 analyzed taxa. Comparative analyses showed that neuropterid mitochondrial protein-coding genes experienced complex evolutionary histories, involving features ranging from codon usage to rate of substitution, that make them potential markers for population genetics/phylogenetics studies at different taxonomic ranks. The 22 tRNAs show variable substitution patterns in Neuropterida, with higher sequence conservation in genes located on the α strand. Inferred secondary structures for neuropterid rrnS and rrnL genes largely agree with those known for other insects. For the first time, a model is provided for domain I of an insect rrnL. The control region in Neuropterida, as in other insects, is fast-evolving genomic region, characterized by AT

An annotated list of aquatic insects of Fort Sill, Oklahoma, excluding diptera with notes on several new state records

USGS Publications Warehouse

Zuellig, R.E.; Kondratieff, B.C.; Schmidt, J.P.; Durfee, R.S.; Ruiter, D.E.; Prather, I.E.

2006-01-01

Qualitative collections of aquatic insects were made at Fort Sill, Lawton, Oklahoma, between 2002 and 2004. Ephemeroptera, Plecoptera, Trichoptera, Odonata, Coleoptera, aquatic Heteroptera, Neuroptera, and Megaloptera were targeted. Additional records are included from a survey that took place in 1999. More than 11,000 specimens from more than 290 collections were examined. Based on the current understanding of aquatic insect systematics, 276 taxa distributed over 8 orders, 46 families, and 141 genera were identified. Twenty-three of the 276 taxa, Plauditus texanus Wiersema, Tricorythodes allectus (Needham), Palmacorixa nana walleyi Hungerford, Climacia chapini Partin and Gurney, Oxyethira forcipata Mosely, Oxyethira janella Denning, Triaenodes helo Milne, Ylodes frontalis (Banks), Acilius fraternus Harris, Coptotomus loticus Hilsenhoff, Coptotomus venustus (Say), Desmopachria dispersa Crotch, Graphoderus liberus (Say), Hydrovatus pustulatus (Melsheimer), Hygrotus acaroides (LeConte), Liodessus flavicollis (LeConte), Uvarus texanus (Sharp), Gyrinus woodruffi Fall, Haliplus fasciatus Aube, Haliplus lewisii Crotch, Haliplus tortilipenis Brigham & Sanderson, Chaetarthria bicolor Sharp, Epimetopus costatus complex, and Hydrochus simplex LeConte are reported from Oklahoma for the first time. The three most diverse orders included Coleoptera (86 species), Odonata (67 species) and Trichoptera (59 species), and the remaining taxa were distributed among Heteroptera, (30 species), Ephemeroptera (21 species), Plecoptera (6 species), Megaloptera (4 species), and Neuroptera (3 species). Based on previous published records, many of the species collected during this study were expected to be found at Fort Sill; however, 276 taxa of aquatic insects identified from such a small geographic area is noteworthy, especially when considering local climatic conditions and the relatively small size of Fort Sill (38,300 ha). Despite agricultural practices in Oklahoma, the dust bowl days

New findings of twisted-wing parasites (Strepsiptera) in Alaska

USGS Publications Warehouse

Mcdermott, Molly

2016-01-01

Strepsipterans are a group of insects with a gruesome life history and an enigmatic evolutionary past. Called ‘twisted-wing parasites’, they are minute parasitoids with a very distinct morphology (Figure 1). Alternatively thought to be related to ichneumon wasps, Diptera (flies), Coleoptera (beetles), and even Neuroptera (net-winged insects) (Pohl and Beutel, 2013); the latest genetic and morphological data support the sister order relationship of Strepsiptera and Coleoptera (Niehuis et al., 2012). Strepsipterans are highly modified, males having two hind wings and halteres instead of front wings or elytra. Unlike most parasitoids, they develop inside active, living insects who are sexually sterilized but not killed until or after emergence (Kathirithamby et al., 2015).

Rearing of Mallada basalis (Neuroptera: Chrysopidae) on modified artificial diets

PubMed Central

Li, Jun; Li, Zhigang; Han, Shichou

2017-01-01

Mallada basalis (Walker) has the potential to be a valuable biological control agent because of its predatory abilities, strong reproductive capacity, and broad prey range. This study aimed to improve on a previously used artificial diet for M. basalis, to achieve a longer oviposition period and greater survival rate and fecundity. We analyzed the development, survival, longevity, and reproduction of M. basalis (F1 and F2 generations) fed two artificial diets (AD1 and AD2). Both diets contained chicken egg yolk, beer yeast powder, honey, trehalose, seawater spirulina, and potassium sorbate. AD1 also contained sucrose and vitamin C. The duration of F1 1st larvae, F1 2nd larvae, F1 pupae, F2 egg, and F2 2nd larvae reared on AD1 were significantly shorter than those reared on AD2. F1 adult longevity and F2 oviposition period for AD1 (45.40 d and 31.00 d) were significantly longer than for AD2 (30.74 d and 20.80 d). All the following were significantly greater for AD1 compared with AD2: F1 female proportion, F1 daily oviposition, F1 female oviposition, F2 daily oviposition, F2 female oviposition, F1 emergence rate, F2 pupation rate, and F3 egg hatching rate. Moreover, for M. basalis fed AD2, the duration of F2 2nd and 3rd larvae (9.00 d and 8.64 d) were significantly longer than for F1 (4.70 d and 4.92 d). The F1 oviposition period (31.57 d) was significantly longer than F2 (20.80 d). The F2 female oviposition (189.20 egg/female) was significantly less than F1 (307.14 egg/female). We found that the oviposition period and female longevity of F1 reared on AD1 was longer than that reared on the artificial diet in a previous study. The daily oviposition and female oviposition of F1 from AD1 was larger, while the F2 egg hatching rate was greater compared with that from the previous diet. However, the offspring of M. basalis fed AD2 were less thrifty. We found diet AD1 supported development and reproduction better than AD2 and the diets in our previous study. These findings may contribute to the mass rearing of this economically important predatory green lacewing. PMID:28961246

Chromosome numbers in antlions (Myrmeleontidae) and owlflies (Ascalaphidae) (Insecta, Neuroptera).

PubMed

Kuznetsova, Valentina G; Khabiev, Gadzhimurad N; Krivokhatsky, Victor A

2015-01-01

A short review of main cytogenetic features of insects belonging to the sister neuropteran families Myrmeleontidae (antlions) and Ascalaphidae (owlflies) is presented, with a particular focus on their chromosome numbers and sex chromosome systems. Diploid male chromosome numbers are listed for 37 species, 21 genera from 9 subfamilies of the antlions as well as for seven species and five genera of the owlfly subfamily Ascalaphinae. The list includes data on five species whose karyotypes were studied in the present work. It is shown here that antlions and owlflies share a simple sex chromosome system XY/XX; a similar range of chromosome numbers, 2n = 14-26 and 2n = 18-22 respectively; and a peculiar distant pairing of sex chromosomes in male meiosis. Usually the karyotype is particularly stable within a genus but there are some exceptions in both families (in the genera Palpares and Libelloides respectively). The Myrmeleontidae and Ascalaphidae differ in their modal chromosome numbers. Most antlions exhibit 2n = 14 and 16, and Palparinae are the only subfamily characterized by higher numbers, 2n = 22, 24, and 26. The higher numbers, 2n = 20 and 22, are also found in owlflies. Since the Palparinae represent a basal phylogenetic lineage of the Myrmeleontidae, it is hypothesized that higher chromosome numbers are ancestral for antlions and were inherited from the common ancestor of Myrmeleontidae + Ascalaphidae. They were preserved in the Palparinae (Myrmeleontidae), but changed via chromosomal fusions toward lower numbers in other subfamilies.

Revision of the Neotropical green lacewing genus Ungla (Neuroptera, Chrysopidae)

PubMed Central

Tauber, Catherine A.; Sosa, Francisco; Albuquerque, Gilberto S.; Tauber, Maurice J.

2017-01-01

Abstract Here, Ungla Navás, 1914, a poorly known Neotropical genus is reviewed. Twenty-five valid species are recognized; seven of them are new to science: Ungla adamsi sp. n., U. elbergi sp. n., U. grandispiracula sp. n., U. mexicana sp. n., U. pennyi sp. n., U. quchapampa sp. n., U. stangei sp. n.; and five are transferred to Ungla from other genera: U. bolivari (Banks), U. chacranella (Banks), U. siderocephala (Navás), U. steinbachi (Navás), and U. banksi Tauber, new replacement name. In addition, ten new synonymies are identified. For each species, a full nomenclatural history, diagnosis, description or redescription with images, literature citations, and available information on the distribution and biology are provided. Name-bearing types were examined for each species, and images of most are included. Keys based on external features are provided for species identifications. As a result of this study, three generalizations appear: (1) The genital morphology of both males and females of Ungla species is very conserved. All species express a common structural pattern, the components of which vary only slightly among species. (2) Ungla species appear to fall into two geographically distinct groups: about one third (n=7) of the species are recorded from southern South America (specifically Argentina and Brazil) and the other approximately two thirds of the species (n=18) from more northern regions of Neotropical America [Andean and Caribbean regions, Central America, and southern Mexico (Chiapas)]. None of the species from either of the regions is known to overlap into the other region. (3) Available information on the immature stages and natural history of species in Ungla is meagre. PMID:28824280

Population Dynamics of Empoasca fabae (Hemiptera: Cicadellidae) in Central Iowa Alfalfa Fields

PubMed Central

Weiser Erlandson, L. A.; Obrycki, J. J.

2015-01-01

Adults and nymphs of Empoasca fabae Harris (Hemiptera: Cicadellidae) and adults of predatory species in the families Coccinellidae, Anthocoridae, Nabidae, Chrysopidae, and Hemerobiidae were sampled in Iowa alfalfa fields from June to September in 1999 and 2000. The relationship between each predatory taxa and E. fabae was examined using regression analysis. In 2000, all predators were found to be positively correlated with the presence of E. fabae during all periods sampled and most likely contributed to mortality. Orius insidiosus (Say) (Hemiptera: Anthoridae) was the most numerous insect predatory species; population numbers ranged from 0 to 1 and 0.1 to 3.7 adults per 0.25 m2 in 1999 and 2000, respectively. Partial life tables were constructed for E. fabae nymphs for two alfalfa-growing periods. Nymphs were grouped into three age intervals: first and second, third and fourth, and fifth instars. For the first alfalfa growing period examined, E. fabae nymphal mortality was 70% in 1999 and 49% in 2000. During the last growing period of each season (August–September), total nymphal mortality was relatively low (<25%). Adult E. fabae density ranged from 5.4 to 25.6 and 1.4–9.2 per 0.25 m2 in 1999 and 2000, respectively. E. fabae population peaks were similar for each age interval in all growing periods. This study provides further information on the population dynamics of E. fabae and its relationship with select predatory species in Iowa alfalfa fields. PMID:26320260

Genetic diversity analysis of Chrysopidae family (Insecta, Neuroptera) via molecular markers.

PubMed

Yari, Kheirollah; Mirmoayedi, Alinaghi; Marami, Marzieh; Kazemi, Elham; Kahrizi, Danial

2014-09-01

In entomology, improvement of molecular methods would be beneficial tools for accurate identification and detecting the genetic diversity of insect species to discover a corroborative evidence for the traditional classification based on morphology. The aim of this study was focused on RAPD-PCR method for distinguishing the genetic diversity between eight species of Chrysopidae family. In current research, many specimens were collected in different locations of Tehran province (Iran), between them 24 specimens were identified. The wing venation, male genitalia and other morphological characters were used for identification and also the sexing of species was recognized with study of external genitalia. Then, the DNA was extracted with CTAB method. The RAPD-PCR method was carried out with twenty random primers. The agarose gel electrophoresis was used for separation of the PCR products. Based on electrophoresis results, 133 bands were amplified and between them, 126 bands were poly-morph and others were mono-morph. Also, among the applied primers, the primers OPA02 with 19 bands and OPA03 with 8 bands were amplified the maximum and minimum of bands, respectively. The results showed that 80.35 and 73.21 % of genetic similarity existed between Chrysopa pallens-Chrysopa dubitans, and between the Chrysoperla kolthoffi and Chrysoperla carnea, respectively. The minimum (45.53 %) of genetic similarity was observed between C. kolthoffi and C. dubitans, and the maximum (0.80 %) was seen between C. pallens and C. dubitans.

New species and new distributional records of Neotropical Mantispidae (Insecta: Neuroptera).

PubMed

Ardila-Camacho, Adrian; Calle-tobÓn, Arley; Wolff, Marta; Stange, Lionel A

2018-04-23

The Neotropical fauna of Mantispidae is currently composed of 106 species. We provide new distributional records of Mantispidae from Colombia and Panama. Three new species are described, one in Symphrasinae from Colombia, and two in Mantispinae from Colombia and Panama. Haematomantispa nubeculosa (Navás, 1933) and Leptomantispa axillaris (Navás, 1908) are reported from Colombia for the first time, the former being the first record of the genus in the country. New locality records for other species previously known from Colombia are also given. For Panama, we report Anchieta fasciatella (Westwood, 1867) and Trichoscelia iridella (Westwood, 1867) for the first time, the former is herein newly transferred from Plega to Anchieta. Three names Mantispa confluens Navás, 1914, n. syn., Buyda apicata Navás, 1926, n. syn., and Mantispa neotropica Navás, 1933, n. syn., are here synonymized with Buyda phthisica (Gerstaecker, 1885). Updated keys for the genera of Mantispinae, and species of genera Trichoscelia, Buyda, and Climaciella from Colombia are included. With this new information, the known species richness of Mantispidae from Colombia increases from 21 to 26, and from 16 to 19 species in Panama.

A tool for developing an automatic insect identification system based on wing outlines

PubMed Central

Yang, He-Ping; Ma, Chun-Sen; Wen, Hui; Zhan, Qing-Bin; Wang, Xin-Li

2015-01-01

For some insect groups, wing outline is an important character for species identification. We have constructed a program as the integral part of an automated system to identify insects based on wing outlines (DAIIS). This program includes two main functions: (1) outline digitization and Elliptic Fourier transformation and (2) classifier model training by pattern recognition of support vector machines and model validation. To demonstrate the utility of this program, a sample of 120 owlflies (Neuroptera: Ascalaphidae) was split into training and validation sets. After training, the sample was sorted into seven species using this tool. In five repeated experiments, the mean accuracy for identification of each species ranged from 90% to 98%. The accuracy increased to 99% when the samples were first divided into two groups based on features of their compound eyes. DAIIS can therefore be a useful tool for developing a system of automated insect identification. PMID:26251292

Phylogeny and biogeography of southern African spoon-winged lacewings (Neuroptera: Nemopteridae: Nemopterinae).

PubMed

Sole, Catherine L; Scholtz, Clarke H; Ball, Jonathan B; Mansell, Mervyn W

2013-01-01

Nemopteridae are a charismatic family of lacewings characterised by uniquely extended hind wings. They are an ancient widespread group in the drier regions of the world. The family comprises two subfamilies, Crocinae (thread-wings) and Nemopterinae (spoon- and ribbon-wings). The present distribution of the family has been largely influenced by the vicariant events of plate tectonics, resulting in relict populations in some parts of the world and extensive evolutionary radiations in others, particularly southern Africa where the vast majority of the species are endemic to the Western and Northern Cape Provinces of South Africa. This study aimed to establish the validity of the 11 currently recognised genera and infer their biogeographic history using molecular sequence data from four gene regions. The hypothesis that the Cape nemopterines co-evolved with certain taxa in the Cape Floristic Region was also tested. Phylogenetic analysis supports seven of the 11 currently recognised genera. The crown age of the Nemopterinae is estimated to be at ca. 145.6 Mya, indicating that the group has been present since the late Jurassic. Most of the genera appear to have diversified during the middle Eocene and into the middle Miocene (ca. 44-11 Mya) with recent rapid radiation of several of the genera occurring during the late Miocene (ca. 6-4.5 Mya). While these data support an initial radiation with the Rushioideae (Aizoaceae) it is recommended that further study including observations and gut content be carried out. Copyright © 2012 Elsevier Inc. All rights reserved.

A new species of the genus Dilar Rambur (Neuroptera: Dilaridae) from Borneo

Treesearch

John D. Oswald; Nathan M. Schiff

2001-01-01

Dilar macleodi is described as a new species from lowland rainforest habitat in the Malaysian State of Sarawak on the island of Borneo. Diagnoses are provided to distinguish D. macleodi from the four other dilarid species that have been reported from the peninsula of Indochina or the Malay Archipelago.

[Spatio-temporal dynamic of green lacewings (Neuroptera: Chrysopidae) taxocenosis on natural ecosystems].

PubMed

Costa, Renildo I F; Souza, Brígida; Freitas, Sérgio De

2010-01-01

In order to study the interactions of green lacewings toxocenosis on natural ecossystems, samplings were carried out in the Parque Ecológico Quedas do Rio Bonito, located in Lavras, Alto Rio Grande region, South of Minas Gerais, Brazil. The species inventory was accomplished in two vegetation types: semi-evergreen forest and open field formations, including areas of montane grassland, rocky montane grassland and "cerrado". Insects were captured with a butterfly net during 2h, walking through each vegetation formation. Sampling resulted in 1,948 specimens belonging to 30 species, of which 14 were Chrysopini and 16 Leucochrysini. Representatives of these tribes were observed both in forest and in open field formations. Species of the genera Ceraeochrysa, Chrysoperla, Chrysopodes, Plesiochrysa and Leucochrysa were found in forests and in open field formations, except for Plesiochrysa. The highest richness and diversity of species were observed in the forest. The similarity index among the communities of green lacewings in the studied areas was 27%.

An Unlikely Silk: The Composite Material of Green Lacewing Cocoons

DOE Office of Scientific and Technical Information (OSTI.GOV)

Weisman, Sarah; Trueman, Holly E.; Mudie, Stephen T.

2009-01-15

Spiders routinely produce multiple types of silk; however, common wisdom has held that insect species produce one type of silk each. This work reports that the green lacewing (Mallada signata, Neuroptera) produces two distinct classes of silk. We identified and sequenced the gene that encodes the major protein component of the larval lacewing cocoon silk and demonstrated that it is unrelated to the adult lacewing egg-stalk silk. The cocoon silk protein is 49 kDa in size and is alanine rich (>40%), and it contains an {alpha}-helical secondary structure. The final instar lacewing larvae spin protein fibers of {approx}2 {mu}m diametermore » to construct a loosely woven cocoon. In a second stage of cocoon construction, the insects lay down an inner wall of lipids that uses the fibers as a scaffold. We propose that the silk protein fibers provide the mechanical strength of the composite lacewing cocoon whereas the lipid layer provides a barrier to water loss during pupation.« less

Ancient pinnate leaf mimesis among lacewings.

PubMed

Wang, Yongjie; Liu, Zhiqi; Wang, Xin; Shih, Chungkun; Zhao, Yunyun; Engel, Michael S; Ren, Dong

2010-09-14

Insects have evolved diverse methods of predator avoidance, many of which implicate complex adaptations of their wings (e.g., Phylliidae, Nymphalidae, Notodontidae). Among these, angiosperm leaf mimicry is one of the most dramatic, although the historical origins of such modifications are unclear owing to a dearth of paleontological records. Here, we report evidence of pinnate leaf mimesis in two lacewings (Neuroptera): Bellinympha filicifolia Y. Wang, Ren, Liu & Engel gen. et sp. nov. and Bellinympha dancei Y. Wang, Ren, Shih & Engel, sp. nov., from the Middle Jurassic, representing a 165-million-year-old specialization between insects and contemporaneous gymnosperms of the Cycadales or Bennettitales. Furthermore, such lacewings demonstrate a preangiosperm origin for leaf mimesis, revealing a lost evolutionary scenario of interactions between insects and gymnosperms. The current fossil record suggests that this enigmatic lineage became extinct during the Early Cretaceous, apparently closely correlated with the decline of Cycadales and Bennettitales at that time, and perhaps owing to the changing floral environment resulted from the rise of flowering plants.

Increased fitness and realized heritability in emamectin benzoate-resistant Chrysoperla carnea (Neuroptera: Chrysopidae).

PubMed

Mansoor, Muhammad Mudassir; Abbas, Naeem; Shad, Sarfraz Ali; Pathan, Attaullah Khan; Razaq, Muhammad

2013-10-01

The common green lacewing Chrysoperla carnea is a key biological control agent employed in integrated pest management (IPM) programs for managing various insect pests. A field collected population of C. carnea was selected for emamectin benzoate resistance in the laboratory and fitness costs and realized heritability were investigated. After five generations of selection with emamectin benzoate, C. carnea developed a 318-fold resistance to the insecticide. The resistant population had a relative fitness of 1.49, with substantially higher emergence rate of healthy adults, fecundity and hatchability and shorter larval duration, pupal duration, and development time compared to the susceptible population. Mean population growth rates; such as the intrinsic rate of natural population increase and biotic potential were higher for the emamectin benzoate selected population compared to the susceptible population. The realized heritability (h(2)) value of emamectin benzoate resistance was 0.34 in emamectin benzoate selected population of C. carnea. Chrysoperla species which show resistance to insecticides makes them compatible with those IPM systems where emamectin benzoate is employed.

Mesozoic lacewings from China provide phylogenetic insight into evolution of the Kalligrammatidae (Neuroptera)

PubMed Central

2014-01-01

Background The Kalligrammatidae are distinctive, large, conspicuous, lacewings found in Eurasia from the Middle Jurassic to mid Early Cretaceous. Because of incomplete and often inadequate fossil preservation, an absence of detailed morphology, unclear relationships, and unknown evolutionary trends, the Kalligrammatidae are poorly understood. Results We describe three new subfamilies, four new genera, twelve new species and four unassigned species from the late Middle Jurassic Jiulongshan and mid Early Cretaceous Yixian Formations of China. These kalligrammatid taxa exhibit diverse morphological characters, such as mandibulate mouthparts in one major clade and siphonate mouthparts in the remaining four major clades, the presence or absence of a variety of distinctive wing markings such as stripes, wing spots and eyespots, as well as multiple major wing shapes. Based on phylogenetic analyses, the Kalligrammatidae are divided into five principal clades: Kalligrammatinae Handlirsch, 1906, Kallihemerobiinae Ren & Engel, 2008, Meioneurinae subfam. nov., Oregrammatinae subfam. nov. and Sophogrammatinae subfam. nov., each of which is accorded subfamily-level status. Our results show significant morphological and evolutionary differentiation of the Kalligrammatidae family during a 40 million-year-interval of the mid Mesozoic. Conclusion A new phylogeny and classification of five subfamilies and their constituent genera is proposed for the Kalligrammatidae. These diverse, yet highly specialized taxa from northeastern China suggest that eastern Eurasia likely was an important diversification center for the Kalligrammatidae. Kalligrammatids possess an extraordinary morphological breadth and panoply of adaptations during the mid-Mesozoic that highlight our conclusion that their evolutionary biology is much more complex than heretofore realized. PMID:24912379

Chrysopa septempunctata (Neuroptera: Chrysopidae) Vitellogenin Functions Through Effects on Egg Production and Hatching.

PubMed

Liu, C; Mao, J; Zeng, F

2015-12-01

Vitellogenin (Vg) is a precursor of major egg storage protein, vitellin (Vt), and plays primary roles in reproduction of oviparous vertebrates and invertebrates. Chrysopa septempunctata Wesmael is an important and common predator of various insect pests. Here, we first cloned C. septempunctata Vg gene, CsVg. The complete CsVg cDNA was 5664 bp, which encodes an 1810-residues protein with a predicted molecular mass of 206.23 kDa. Expression profile revealed that CsVg mRNA first appeared on day 4 after emergence, maximally accumulated on day 10, and then declined gradually. RNAi mediated by injection of dsRNA depleted CsVg transcripts, significantly reduced egg-laying amount, and decreased egg hatching rate, suggesting that CsVg functions through effects on egg production and hatching in C. septempunctata. © The Authors 2015. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

Cryptochrysa Freitas Penny, a generic homonym, replaced by Titanochrysa Sosa Freitas (Neuroptera: Chrysopidae).

PubMed

Tauber, Catherine A; Sosa, Francisco; Contreras-Ramos, Atilano

2018-01-24

The green lacewing genus name Cryptochrysa Freitas Penny 2001 is identified as a junior homonym of Cryptochrysa Hampson 1926 (Lepidoptera: Noctuidae). Moreover, Titanochrysa Sosa Freitas 2012 is determined to be an available junior synonym of Cryptochrysa Freitas Penny. Thus, Titanochrysa becomes the substitute name for the preoccupied generic name. Here, we also provide (i) new information and images for Titanochrysa chloros (Freitas Penny) comb. nov., the only species ever included in the chrysopid genus Cryptochrysa, (ii) a clarified set of diagnostic features for the reconstituted genus Titanochrysa, and (iii) a key and images for identifying the six described Titanochrysa species.

Mesozoic lacewings from China provide phylogenetic insight into evolution of the Kalligrammatidae (Neuroptera).

PubMed

Yang, Qiang; Wang, Yongjie; Labandeira, Conrad C; Shih, Chungkun; Ren, Dong

2014-06-09

The Kalligrammatidae are distinctive, large, conspicuous, lacewings found in Eurasia from the Middle Jurassic to mid Early Cretaceous. Because of incomplete and often inadequate fossil preservation, an absence of detailed morphology, unclear relationships, and unknown evolutionary trends, the Kalligrammatidae are poorly understood. We describe three new subfamilies, four new genera, twelve new species and four unassigned species from the late Middle Jurassic Jiulongshan and mid Early Cretaceous Yixian Formations of China. These kalligrammatid taxa exhibit diverse morphological characters, such as mandibulate mouthparts in one major clade and siphonate mouthparts in the remaining four major clades, the presence or absence of a variety of distinctive wing markings such as stripes, wing spots and eyespots, as well as multiple major wing shapes. Based on phylogenetic analyses, the Kalligrammatidae are divided into five principal clades: Kalligrammatinae Handlirsch, 1906, Kallihemerobiinae Ren & Engel, 2008, Meioneurinae subfam. nov., Oregrammatinae subfam. nov. and Sophogrammatinae subfam. nov., each of which is accorded subfamily-level status. Our results show significant morphological and evolutionary differentiation of the Kalligrammatidae family during a 40 million-year-interval of the mid Mesozoic. A new phylogeny and classification of five subfamilies and their constituent genera is proposed for the Kalligrammatidae. These diverse, yet highly specialized taxa from northeastern China suggest that eastern Eurasia likely was an important diversification center for the Kalligrammatidae. Kalligrammatids possess an extraordinary morphological breadth and panoply of adaptations during the mid-Mesozoic that highlight our conclusion that their evolutionary biology is much more complex than heretofore realized.

Rates and patterns of molecular evolution in freshwater versus terrestrial insects.

PubMed

Mitterboeck, T Fatima; Fu, Jinzhong; Adamowicz, Sarah J

2016-11-01

Insect lineages have crossed between terrestrial and aquatic habitats many times, for both immature and adult life stages. We explore patterns in molecular evolutionary rates between 42 sister pairs of related terrestrial and freshwater insect clades using publicly available protein-coding DNA sequence data from the orders Coleoptera, Diptera, Lepidoptera, Hemiptera, Mecoptera, Trichoptera, and Neuroptera. We furthermore test for habitat-associated convergent molecular evolution in the cytochrome c oxidase subunit I (COI) gene in general and at a particular amino acid site previously reported to exhibit habitat-linked convergence within an aquatic beetle group. While ratios of nonsynonymous-to-synonymous substitutions across available loci were higher in terrestrial than freshwater-associated taxa in 26 of 42 lineage pairs, a stronger trend was observed (20 of 31, p binomial = 0.15, p Wilcoxon = 0.017) when examining only terrestrial-aquatic pairs including fully aquatic taxa. We did not observe any widespread changes at particular amino acid sites in COI associated with habitat shifts, although there may be general differences in selection regime linked to habitat.

Ancient pinnate leaf mimesis among lacewings

PubMed Central

Wang, Yongjie; Liu, Zhiqi; Wang, Xin; Shih, Chungkun; Zhao, Yunyun; Engel, Michael S.; Ren, Dong

2010-01-01

Insects have evolved diverse methods of predator avoidance, many of which implicate complex adaptations of their wings (e.g., Phylliidae, Nymphalidae, Notodontidae). Among these, angiosperm leaf mimicry is one of the most dramatic, although the historical origins of such modifications are unclear owing to a dearth of paleontological records. Here, we report evidence of pinnate leaf mimesis in two lacewings (Neuroptera): Bellinympha filicifolia Y. Wang, Ren, Liu & Engel gen. et sp. nov. and Bellinympha dancei Y. Wang, Ren, Shih & Engel, sp. nov., from the Middle Jurassic, representing a 165-million-year-old specialization between insects and contemporaneous gymnosperms of the Cycadales or Bennettitales. Furthermore, such lacewings demonstrate a preangiosperm origin for leaf mimesis, revealing a lost evolutionary scenario of interactions between insects and gymnosperms. The current fossil record suggests that this enigmatic lineage became extinct during the Early Cretaceous, apparently closely correlated with the decline of Cycadales and Bennettitales at that time, and perhaps owing to the changing floral environment resulted from the rise of flowering plants. PMID:20805491

Species abundance and potential biological control services in shade vs. sun coffee in Puerto Rico

USGS Publications Warehouse

Borkhataria, Rena R.; Collazo, Jaime A.; Groom, Martha J.

2012-01-01

Birds, lizards and insects were surveyed in three sun and three shade coffee plantations in Puerto Rico to provide a comprehensive comparison of biodiversity between plantations types and to identify potential interrelationships (e.g., biological or natural control services) between members of each taxon and coffee pests. Abundance of avian species, including insectivorous species, was significantly higher in shade coffee. Anolis cristatellus and A. stratulus were significantly more abundant in sun plantations whereas A. gundlachi and A. evermanni were detected more frequently in shade plantations. Insects in the orders Coleoptera, Diptera, Hemiptera, Hymenoptera, Lepidoptera, Neuroptera, and Psocoptera were significantly more abundant in shade coffee, while orthopterans were more abundant in sun. The coffee leaf miner (Leucoptera coffeela) and the flatid planthopper (Petrusa epilepsis) did not differ significantly between plantation types, nor did the abundance of the wasp complex that parasitizes the coffee leaf miner. These findings confirmed that shade plantations harbor a wide array of elements of biodiversity; but sun plantations may also harbor many elements of biodiversity, and in some cases, in higher abundance than in shade plantations.

The Green Lacewing, Chrysoperla carnea: Preference between Lettuce Aphids, Nasonovia ribisnigri, and Western Flower Thrips, Frankliniella occidentalis

PubMed Central

Shrestha, Govinda; Enkegaard, Annie

2013-01-01

This study investigated the prey preference of 3rd instar green lacewing, Chrysoperla carnea Stephens (Neuroptera: Chrysopidae), between western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and lettuce aphids, Nasonovia ribisnigri (Mosley) (Hemiptera: Aphididae) in laboratory experiments at 25 ± 1° C and 70 ± 5% RH with five prey ratios (10 aphids:80 thrips, 25 aphids:65 thrips, 45 aphids:45 thrips, 65 aphids:25 thrips, and 80 aphids:10 thrips). Third instar C. carnea larvae readily preyed upon both thrips and aphids, with thrips mortality varying between 40 and 90%, and aphid mortality between 52 and 98%. Chrysoperla carnea had a significant preference for N. ribisnigri at two ratios (10 aphids:80 thrips, 65 aphids:25 thrips), but no preference for either prey at the other ratios. There was no significant linear relationship between preference index and prey ratio, but a significant intercept of the linear regression indicated an overall preference of C. carnea for aphids with a value of 0.651 ± 0.054. The possible implications of these findings for control of N. ribisnigri and F. occidentalis by C. carnea are discussed. PMID:24205864

The green lacewing, Chrysoperla carnea: preference between lettuce aphids, Nasonovia ribisnigri, and Western flower thrips, Frankliniella occidentalis.

PubMed

Shrestha, Govinda; Enkegaard, Annie

2013-01-01

This study investigated the prey preference of 3(rd) instar green lacewing, Chrysoperla carnea Stephens (Neuroptera: Chrysopidae), between western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), and lettuce aphids, Nasonovia ribisnigri (Mosley) (Hemiptera: Aphididae) in laboratory experiments at 25 ± 1° C and 70 ± 5% RH with five prey ratios (10 aphids:80 thrips, 25 aphids:65 thrips, 45 aphids:45 thrips, 65 aphids:25 thrips, and 80 aphids:10 thrips). Third instar C. carnea larvae readily preyed upon both thrips and aphids, with thrips mortality varying between 40 and 90%, and aphid mortality between 52 and 98%. Chrysoperla carnea had a significant preference for N. ribisnigri at two ratios (10 aphids:80 thrips, 65 aphids:25 thrips), but no preference for either prey at the other ratios. There was no significant linear relationship between preference index and prey ratio, but a significant intercept of the linear regression indicated an overall preference of C. carnea for aphids with a value of 0.651 ± 0.054. The possible implications of these findings for control of N. ribisnigri and F. occidentalis by C. carnea are discussed.

Nomenclatorial changes and redescriptions of three of Navás’ Leucochrysa (Nodita) species (Neuroptera, Chrysopidae)

PubMed Central

Catherine A., Tauber; Gilberto S., Albuquerque; Maurice J., Tauber

2011-01-01

Abstract Three species that Navás described – Leucochrysa (Nodita) azevedoi Navás, 1913, Leucochrysa (Nodita) camposi (Navás, 1933) and Leucochrysa (Nodita) morenoi (Navás, 1934) – have received recent taxonomic attention. All three have many similar external features; indeed Navás himself, as well as subsequent authors, have confused the species with each other. Here, (a) misidentifications are corrected; (b) a neotype of Leucochrysa azevedoi is designated; (c) Leucochrysa (Nodita) morenoi, previously synonymized with Leucochrysa (Nodita) camposi, is recognized as a valid species [Reinstated status] All three species are redescribed and illustrated, with special emphasis on the types. Leucochrysa (Nodita) azevedoi was found to be relatively common in agricultural areas along Brazil’s Atlantic coast. The two other species are known only from their type localities: Leucochrysa (Nodita) camposi – coastal Ecuador, and Leucochrysa (Nodita) morenoi – Quito, Ecuador. PMID:21594110

Escaping compound eye ancestry: the evolution of single-chamber eyes in holometabolous larvae.

PubMed

Buschbeck, Elke K

2014-08-15

Stemmata, the eyes of holometabolous insect larvae, have gained little attention, even though they exhibit remarkably different optical solutions, ranging from compound eyes with upright images, to sophisticated single-chamber eyes with inverted images. Such optical differences raise the question of how major transitions may have occurred. Stemmata evolved from compound eye ancestry, and optical differences are apparent even in some of the simplest systems that share strong cellular homology with adult ommatidia. The transition to sophisticated single-chamber eyes occurred many times independently, and in at least two different ways: through the fusion of many ommatidia [as in the sawfly (Hymenoptera)], and through the expansion of single ommatidia [as in tiger beetles (Coleoptera), antlions (Neuroptera) and dobsonflies (Megaloptera)]. Although ommatidia-like units frequently have multiple photoreceptor layers (tiers), sophisticated image-forming stemmata tend to only have one photoreceptor tier, presumably a consequence of the lens only being able to efficiently focus light on to one photoreceptor layer. An interesting exception is found in some diving beetles [Dytiscidae (Coleoptera)], in which two retinas receive sharp images from a bifocal lens. Taken together, stemmata represent a great model system to study an impressive set of optical solutions that evolved from a relatively simple ancestral organization. © 2014. Published by The Company of Biologists Ltd.

Action of neem oil (Azadirachta indica A. Juss) on cocoon spinning in Ceraeochrysa claveri (Neuroptera: Chrysopidae).

PubMed

Scudeler, Elton Luiz; Garcia, Ana Silvia Gimenes; Padovani, Carlos Roberto; Santos, Daniela Carvalho

2013-11-01

Neem oil is a biopesticide that disturbs the endocrine and neuroendocrine systems of pests and may interfere with molting, metamorphosis and cocoon spinning. The cocoon serves protective functions for the pupa during metamorphosis, and these functions are dependent on cocoon structure. To assess the changes in cocoon spinning caused by neem oil ingestion, Ceraeochrysa claveri larvae, a common polyphagous predator, were fed with neem oil throughout the larval period. When treated with neem oil, changes were observed on the outer and inner surfaces of the C. claveri cocoon, such as decreased wall thickness and impaired ability to attach to a substrate. These negative effects may reduce the effectiveness of the mechanical and protective functions of cocoons during pupation, which makes the specimen more vulnerable to natural enemies and environmental factors. © 2013 Elsevier Inc. All rights reserved.

Suitability of Microtheca ochroloma (Coleoptera: Chrysomelidae) for the Development of the Predator Chrysoperla rufilabris (Neuroptera: Chrysopidae).

PubMed

Niño, Angie A; Cave, Ronald D

2015-08-01

Microtheca ochroloma Stål, the yellowmargined leaf beetle, is one of the most destructive pests of crucifer vegetables on organic farms. Larvae of the green lacewing Chrysoperla rufilabris Burmeister have been observed preying on M. ochroloma, but no studies have evaluated the suitability of M. ochroloma as prey for C. rufilabris or the efficacy of this predator as a biological control agent of the pest. This study quantified the killing rate, developmental time, and survivorship of C. rufilabris when offered eggs and larvae of M. ochroloma at 10, 15, 20, or 25°C. Mean number of prey killed daily increased from 8.4 eggs and 4.0 larvae at 15°C to 18.6 eggs and 10.2 larvae at 25°C. However, predator larvae killed 78% fewer total eggs at 25°C than at 15°C; total number of first-instar prey killed did not vary significantly with temperature. Mean developmental time of predator larvae decreased from 75.5 d at 15°C to 26.6 d at 25°C when they were fed eggs, whereas it decreased from 54.0 d at 15°C to 21.4 d at 25°C when they were fed larvae. Predator survivorship was reduced by 80% at 15°C and no larvae survived at 10°C. We conclude that C. rufilabris can complete development on a diet of eggs of M. ochroloma, but its effectiveness to control M. ochroloma populations will be lessened during cool months, from November to April, when crucifers are produced in Florida and the beetle is actively developing, reproducing, and causing crop damage. © The Authors 2015. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For Permissions, please email: journals.permissions@oup.com.

Larvae of five horticulturally important species of Chrysopodes (Neuroptera, Chrysopidae): shared generic features, descriptions and keys

PubMed Central

Silva, Patrícia S.; Tauber, Catherine A.; Albuquerque, Gilberto S.; Tauber, Maurice J.

2013-01-01

Abstract An expanded list of generic level larval characteristics is presented for Chrysopodes; it includes a reinterpretation of the mesothoracic and metathoracic structure and setation. Keys, descriptions and images of Semaphoront A (first instar) and Semaphoront B (second and third instars) are offered for identifying five species of Chrysopodes (Chrysopodes) that are commonly reported from horticultural habitats in the Neotropical region. PMID:23653514

Bt Proteins Have No Detrimental Effects on Larvae of the Green Lacewing, Chrysopa pallens (Rambur) (Neuroptera: Chrysopidae).

PubMed

Ali, I; Zhang, S; Muhammad, M S; Iqbal, M; Cui, J-J J-J

2018-06-01

Biosafety of a genetically modified crop is required to be assessed prior to its commercialization. For this, a suitable artificial diet was developed and used to establish a dietary exposure test for assessing the toxicity of midgut-active Bt insecticidal proteins on Chrysopa pallens (Rambur). Subsequently, this dietary exposure test was used to evaluate the toxicity of the proteins Cry1Ab, Cry1Ac, Cry1Ah, Cry1Ca, Cry1F, Cry2Aa, Cry2Ab, and Vip3Aa on C. pallens larvae. Temporal stability, bioactivity, and the intake of the insecticidal proteins were confirmed by enzyme-linked immunosorbent assay and a sensitive-insect bioassay. The life history characteristics, such as survival, pupation, adult emergence, 7-day larval weight, larval developmental time, and emerged male and female fresh weights remained unaffected, when C. pallens were fed the pure artificial diet (negative control) and the artificial diets containing 200 μg/g of each purified protein: Cry1Ab, Cry1Ac, Cry1Ah, Cry1Ca, Cry1F, Cry2Aa, Cry2Ab, or Vip3Aa. On the contrary, all of the life history characteristics of C. pallens larvae were adversely affected when fed artificial diet containing boric acid (positive control). The results demonstrate that diets containing the tested concentrations of Cry1Ab, Cry1Ac, Cry1Ah, Cry1Ca, Cry1F, Cry2Aa, Cry2Ab, and Vip3Aa have null effects on C. pallens larvae. The outcome indicates that genetically modified crops expressing the tested Bt proteins are safe for the lacewing, C. pallens.

Predation and behavioral changes in the neotropical lacewing Chrysoperla externa (Hagen) (Neuroptera: Chrysopidae) exposed to lambda-cyhalothrin.

PubMed

Luna, René Fernando; Bestete, Luziani Rezende; Torres, Jorge Braz; da Silva-Torres, Christian Sherley Araújo

2018-05-24

Pyrethroid insecticides are widely recommended against various defoliating pest species, but usually lack efficacy against sucking pests such as aphids, which are preferred prey of lacewing larvae. Interaction of pyrethroids and lacewings are likely to occur in fields infested by both defoliating and sap-sucking pests and should provide a complementary control. Therefore, our study aimed to estimate dose-mortality curves and behavioral changes by Chrysoperla externa exposed to lambda-cyhalothrin. We tested the susceptibility of two populations from different locations and insecticide history exposure through topical application. Based on the LD 50 -calculated, the population exhibiting the greater LD 50 was exposed to resistance enhancement (Sel) by treating larvae once for seven successive generations. The population with lower LD 50 was kept without selection (Nsel). Subsequently, walking, predation and oviposition behavioral after exposure to dried insecticide residues were investigated. After seven generations with insecticide selection, the resistance rations between Sel and Nsel populations were 5.85- and 9.37-fold higher for larvae and 3.38- and 2.75-fold higher for adults, respectively. Selected females caged in partially treated arenas laid similar eggs number on both treated and untreated surfaces, while Nsel females laid fewer eggs on treated surfaces. Insecticide repellency was not observed in either population, although irritability was observed for Nsel larvae. Selected larvae confined on fully and partially treated surfaces walked further, for a longer time, and with greater speeds compared to Nsel larvae. Furthermore, Sel and Nsel larvae had reduced predation rates when confined on treated surfaces, and Nsel larvae consumed less prey than Sel larvae. Results indicate changes in susceptibility, behavior, and predation rate of C. externa following exposure to lambda-cyhalothrin.

A morphological and life history comparison between desert populations of a sit-and-pursue antlion, in reference to a co-occurring pit-building antlion.

PubMed

Scharf, Inon; Filin, Ido; Subach, Aziz; Ovadia, Ofer

2009-10-01

Although most antlion species do not construct pits, the vast majority of studies on antlions focused on pit-building species. We report here on a transplant experiment aiming to test for morphological and life history differences between two desert populations of a sit-and-pursue antlion species, Lopezus fedtschenkoi (Neuroptera: Myrmeleontidae), originating from habitats, which mainly differ in plant cover and productivity. We raised the antlion larvae in environmental chambers simulating either hyper-arid or Mediterranean climate. We found significant differences in the morphology and life history of L. fedtschenkoi larvae between the two populations. For example, the larvae originating from the more productive habitat pupated faster and had a higher growth rate. In agreement with the temperature-size rule, antlions reached higher final mass in the colder Mediterranean climate and exhibited a higher growth rate, but there was no difference in their developmental time. Observed differences in morphology between populations as well as those triggered by climate growing conditions could be explained by differences in size allometry. We also provide a quantitative description of the allometric growth axis, based on 12 morphological traits. Comparing the responses of L. fedtschenkoi with those observed in a co-occurring pit-building antlion indicated that there were neither shape differences that are independent of size nor was there a difference in the plasticity level between the two species.

Suitability of leguminous cover crop pollens as food source for the green lacewing Chrysoperla externa (Hagen) (Neuroptera: Chrysopidae).

PubMed

Venzon, Madelaine; Rosado, Maria C; Euzébio, Denise E; Souza, Brígida; Schoereder, José H

2006-01-01

Diversification of crops with species that provide suitable pollen for predators may reduce pest population on crops by enhancing predator effectiveness. In this paper we evaluated the suitability of leguminous cover crop pollens to the predatory green lacewing Chrysoperla externa (Hagen). The predator is commonly found in coffee agroecosystems and the plant species tested were pigeon pea and sunn hemp, which are used in organic coffee systems. Newly emerged females and males of C. externa were reared on diets containing pollen of pigeon pea, sunn hemp, or castor bean, used as a control. The reproductive success of C. externa was evaluated when females fed the pollen species and when honey was added to the diets, to verify the predator need for an extra carbohydrate source. Similar intrinsic growth rates were found for females fed on pigeon pea pollen and on sunn hemp pollen but these rates increased significantly when honey was added to the diets. Females fed with pigeon pea pollen plus honey and with sunn hemp pollen plus honey had higher intrinsic growth rates than those fed with castor bean pollen plus honey. Females fed on castor bean pollen only or on honey only, did not oviposit. Leguminous pollen species were equally suitable for C. externa especially when they were complemented with honey. The results suggest that to successfully enhance predator effectiveness, organic coffee plantation should be diversified with plant providing pollen in combination with plant providing nectar.

Effects of the chitin synthesis inhibitor buprofezin on survival and development of immatures of Chrysoperla rufilabris (Neuroptera: Chrysopidae).

PubMed

Liu, T X; Chen, T Y

2000-04-01

Effects of buprofezin (Applaud), a chitin synthesis inhibitor, on survival and development of eggs, three instars, and pupae of Chrysoperla rufilabris (Burmeister) were determined in the laboratory. Buprofezin at three tested concentrations (100, 500, and 1,000 mg [AI]/liter) did not affect the viability and development of eggs when the eggs were treated, or third instars and pupae when those stages were treated. Although the degree of effects by buprofezin on larvae varied with instar, buprofezin at the higher concentrations (500 and 1,000 mg [AI]/liter) reduced survival rates 17-47% and prolonged the overall development from first instars to adult emergence by 2 or 3 d when first instars were treated, indicating that the first instar is the most vulnerable stage. When second instars were treated, the survival of C. rufilabris from second instars to pupae was not significantly affected. However, the developmental time from second instar to adult emergence was longer in the treatments with the highest concentration (1,000 mg [AI]/liter) than that with the lowest concentration (100 mg [AI]/liter). The compatibility of buprofezin with natural enemies in integrated pest management programs is discussed.

Taxonomic notes on Babinskaiidae from the Cretaceous Burmese amber, with the description of a new species (Insecta, Neuroptera)

PubMed Central

Hu, Jiahui; Lu, Xiumei; Wang, Bo; Liu, Xingyue

2018-01-01

Abstract Babinskaiidae is an extinct lacewing family of the superfamily Myrmeleontoidea. Hitherto, nine species of seven genera are described from the Lower and mid-Cretaceous. Here a new species of Babinskaiidae is described from Cretaceous Burmese amber, namely Parababinskaia makarkini sp. n. The new species possesses an A2 vein in the hind wing, suggesting that the loss of this vein might not be an autapomorphy of Babinskaiidae. The female of Electrobabinskaia burmana Lu, Zhang & Liu, 2017 is also described for the first time based on two specimens with their abdomens perfectly preserved, exhibiting a specialised sternum VI with paired elongate projections. A brief discussion of female genital characters is provided, which may increase our understanding of the morphology and phylogenetic position of Babinskaiidae. PMID:29674912

Taxonomic notes on Babinskaiidae from the Cretaceous Burmese amber, with the description of a new species (Insecta, Neuroptera).

PubMed

Hu, Jiahui; Lu, Xiumei; Wang, Bo; Liu, Xingyue

2018-01-01

Babinskaiidae is an extinct lacewing family of the superfamily Myrmeleontoidea. Hitherto, nine species of seven genera are described from the Lower and mid-Cretaceous. Here a new species of Babinskaiidae is described from Cretaceous Burmese amber, namely Parababinskaia makarkini sp. n. The new species possesses an A2 vein in the hind wing, suggesting that the loss of this vein might not be an autapomorphy of Babinskaiidae. The female of Electrobabinskaia burmana Lu, Zhang & Liu, 2017 is also described for the first time based on two specimens with their abdomens perfectly preserved, exhibiting a specialised sternum VI with paired elongate projections. A brief discussion of female genital characters is provided, which may increase our understanding of the morphology and phylogenetic position of Babinskaiidae.

A cute and highly contrast-sensitive superposition eye - the diurnal owlfly Libelloides macaronius.

PubMed

Belušič, Gregor; Pirih, Primož; Stavenga, Doekele G

2013-06-01

The owlfly Libelloides macaronius (Insecta: Neuroptera) has large bipartite eyes of the superposition type. The spatial resolution and sensitivity of the photoreceptor array in the dorsofrontal eye part was studied with optical and electrophysiological methods. Using structured illumination microscopy, the interommatidial angle in the central part of the dorsofrontal eye was determined to be Δϕ=1.1 deg. Eye shine measurements with an epi-illumination microscope yielded an effective superposition pupil size of about 300 facets. Intracellular recordings confirmed that all photoreceptors were UV-receptors (λmax=350 nm). The average photoreceptor acceptance angle was 1.8 deg, with a minimum of 1.4 deg. The receptor dynamic range was two log units, and the Hill coefficient of the intensity-response function was n=1.2. The signal-to-noise ratio of the receptor potential was remarkably high and constant across the whole dynamic range (root mean square r.m.s. noise=0.5% Vmax). Quantum bumps could not be observed at any light intensity, indicating low voltage gain. Presumably, the combination of large aperture superposition optics feeding an achromatic array of relatively insensitive receptors with a steep intensity-response function creates a low-noise, high spatial acuity instrument. The sensitivity shift to the UV range reduces the clutter created by clouds within the sky image. These properties of the visual system are optimal for detecting small insect prey as contrasting spots against both clear and cloudy skies.

Comparative Toxicities of Newer and Conventional Insecticides: Against Four Generalist Predator Species.

PubMed

Prabhaker, Nilima; Naranjo, Steven; Perring, Thomas; Castle, Steven

2017-12-05

Generalist insect predators play an essential role in regulating the populations of Bemisia tabaci and other pests in agricultural systems, but may be affected negatively by insecticides applied for pest management. Evaluation of insecticide compatibility with specific predator species can provide a basis for making treatment decisions with the aim of conserving natural enemies. Eleven insecticides representing six modes of action groups were evaluated for toxicity against four predator species and at different developmental stages. Full-concentration series bioassays were conducted on laboratory-reared or insectary-supplied predators using Petri dish and systemic uptake bioassay techniques. Highest toxicities were observed with imidacloprid and clothianidin against first and second instar nymphs of Geocoris punctipes (Say) (Hemiptera: Geocoridae). Later instar nymphs were less susceptible to neonicotinoid treatments based on higher LC50s observed with imidacloprid, thiamethoxam, and dinotefuran against third or fourth instar nymphs. The pyrethroid insecticide bifenthrin was highly toxic against adults of G. punctipes and Orius insidiosus (Say) (Hemiptera: Anthocoridae). Standard concentration/mortality evaluation of nonacute toxicity insecticides, including buprofezin, pyriproxyfen, spirotetramat, and spiromesifen, was inconclusive in terms of generating probit statistics. However, low mortality levels of insects exposed for up to 120 h suggested minimal lethality with the exception of pyriproxyfen that was mildly toxic to Chrysoperla rufilabris (Burmeister) (Neuroptera: Chrysopidae). Published by Oxford University Press on behalf of Entomological Society of America 2017. This work is written by (a) US Government employee(s) and is in the public domain in the US.

Seasonal variation in the populations of Polyphagotarsonemus latus and Tetranychus bastosi in physic nut (Jatropha curcas) plantations.

PubMed

Rosado, Jander F; Picanço, Marcelo C; Sarmento, Renato A; da Silva, Ricardo Siqueira; Pedro-Neto, Marçal; Carvalho, Marcos Alberto; Erasmo, Eduardo A L; Silva, Laila Cristina Rezende

2015-07-01

Studies on the seasonal variation of agricultural pest species are important for the establishment of integrated pest control programs. The seasonality of pest attacks on crops is affected by biotic and abiotic factors, for example, climate and natural enemies. Besides that, characteristics of the host plant, crop management, location and the pests' bioecology also affect this seasonality. The mites Polyphagotarsonemus latus (Prostigmata: Tarsonemidae) and Tetranychus bastosi (Prostigmata: Tetranychidae) are the most important pests in the cultivation of physic nut, Jatropha curcas (Euphorbiaceae). All parts of J. curcas can be used for a wide range of purposes. In addition many researchers have studied its potential for use as neat oil, as transesterified oil (biodiesel), or as a blend with diesel. However studies about physic nut pests have been little known. The objective of this study was to assess the seasonal variation of P. latus and T. bastosi in physic nut. This study was conducted at three sites in the state of Tocantins, Brazil. We monitored climatic elements and the densities of the two mite species and of their natural enemies for a period of 2 years. Attack by P. latus occurred during rainy seasons, when the photoperiod was short and the physic nut had new leaves. In contrast, attack by T. bastosi occurred during warmer seasons with longer photoperiods and stronger winds. Populations of both mites and their natural enemies were greater in sites with greater plant diversity adjacent to the plantations. The predators found in association with P. latus and T. bastosi were Euseius concordis (Acari: Phytoseiidae), spiders, Stethorus sp. (Coleoptera: Coccinellidae) and Chrysoperla sp. (Neuroptera: Chrysopidae).

Resistance of green lacewing, Chrysoperla carnea Stephens to nitenpyram: Cross-resistance patterns, mechanism, stability, and realized heritability.

PubMed

Mansoor, Muhammad Mudassir; Raza, Abu Bakar Muhammad; Abbas, Naeem; Aqueel, Muhammad Anjum; Afzal, Muhammad

2017-01-01

The green lacewing, Chrysoperla carnea Stephens (Neuroptera: Chrysopidae) is a major generalist predator employed in integrated pest management (IPM) plans for pest control on many crops. Nitenpyram, a neonicotinoid insecticide has widely been used against the sucking pests of cotton in Pakistan. Therefore, a field green lacewing strain was exposed to nitenpyram for five generations to investigate resistance evolution, cross-resistance pattern, stability, realized heritability, and mechanisms of resistance. Before starting the selection with nitenpyram, a field collected strain showed 22.08-, 23.09-, 484.69- and 602.90-fold resistance to nitenpyram, buprofezin, spinosad and acetamiprid, respectively compared with the Susceptible strain. After continuous selection for five generations (G1-G5) with nitenpyram in the laboratory, the Field strain (Niten-SEL) developed a resistance ratio of 423.95 at G6. The Niten-SEL strain at G6 showed no cross-resistance to buprofezin and acetamiprid and negative cross-resistance to spinosad compared with the Field strain (G1). For resistance stability, the Niten-SEL strain was left unexposed to any insecticide for four generations (G6-G9) and bioassay results at G10 showed that resistance to nitenpyram, buprofezin and spinosad was stable, while resistance to acetamiprid was unstable. The realized heritability values were 0.97, 0.16, 0.03, and -0.16 to nitenpyram, buprofezin, acetamiprid and spinosad, respectively, after five generations of selection. Moreover, the enzyme inhibitors (PBO or DEF) significantly decreased the nitenpyram resistance in the resistant strain, suggesting that resistance was due to microsomal oxidases and esterases. These results are very helpful for integration of green lacewings in IPM programs. Copyright © 2016 Elsevier B.V. All rights reserved.

Two common and problematic leucochrysine species – Leucochrysa (Leucochrysa) varia (Schneider) and L. (L.) pretiosa (Banks) (Neuroptera, Chrysopidae): redescriptions and synonymies

PubMed Central

Tauber, Catherine A.; Sosa, Francisco; Albuquerque, Gilberto S.

2013-01-01

Abstract We dedicate this article to the memory of Sergio de Freitas, FCAV-UNESP, Jaboticabal, São Paulo, Brazil (deceased, 2012). He was an active and enthusiastic Neuropterist and the cherished mentor and friend of Francisco Sosa. Leucochrysa McLachlan is the largest genus in the Chrysopidae, yet it has received relatively little taxonomic attention. We treat two problematic and common Leucochrysa species – Leucochrysa (Leucochrysa) varia (Schneider, 1851) and Leucochrysa (Leucochrysa) pretiosa (Banks, 1910). Both are highly variable in coloration and were described before the systematic importance of chrysopid genitalia was recognized. Recent studies show that these species occur within a large complex of cryptic species and that they have accumulated a number of taxonomic problems. We identify new synonymies for each of the species–for Leucochrysa (Leucochrysa) varia: Leucochrysa (Leucochrysa) ampla (Walker, 1853), Leucochrysa internata (Walker, 1853), and Leucochrysa (Leucochrysa) walkerina Navás, 1913; for Leucochrysa (Leucochrysa) pretiosa: Leucochrysa (Leucochrysa) erminea Banks, 1946. The synonymy of Leucochrysa delicata Navás, 1925 with Leucochrysa (Leucochrysa) pretiosa is stabilized by the designation of a neotype. The following species, which were previously synonymized with Leucochrysa (Leucochrysa) varia or Leucochrysa (Leucochrysa) pretiosa, are reinstated as valid: Leucochrysa (Leucochrysa) phaeocephala Navás, 1929, Leucochrysa (Leucochrysa) angrandi (Navás, 1911), and Leucochrysa (Leucochrysa) variata (Navás, 1913). To help stabilize Leucochrysa taxonomy, lectotypes are designated for Allochrysa pretiosa and Allochrysa variata. Finally, Leucochrysa vegana Navás, 1917 is considered a nomen dubium. PMID:23805050

Side Effects of Neem Oil on the Midgut Endocrine Cells of the Green Lacewing Ceraeochrysa claveri (Navás) (Neuroptera: Chrysopidae).

PubMed

Scudeler, E L; Santos, D C

2014-04-01

We described the ultrastructure of Ceraeochrysa claveri (Navás) midgut endocrine cells in larva, pupa, and adult, and evaluated the side effects of ingested neem oil, a botanical insecticide obtained from the seeds of the neem tree (Azadirachta indica), on these cells. During the larval period, C. claveri were fed (ad libitum) Diatraea saccharalis (F.) eggs treated with neem oil at concentrations of 0.5%, 1%, or 2%. Transmission electron microscopy showed that two subtypes of endocrine cells, namely granular and vesicular, occurred in the midgut epithelium during the three stages of the life cycle. Both cell types did not reach the midgut lumen and were positioned basally in the epithelium. The endocrine cells did not show extensive infoldings of the basal plasma membrane, and there were numerous secretory granules in the basal region of the cytoplasm. In the granular endocrine cells, the granules were completely filled with a dense matrix. In the vesicular endocrine cells, the main secretory products consisted of haloed vesicles. Ultrastructural examination indicated that only the granular endocrine cells exhibited signs of morphologic changes of cell injury present in all life cycle stages after the larvae were chronically exposed to neem oil by ingestion. The major cellular damage consisted of dilatation and vesiculation of the rough endoplasmic reticulum and the development of smooth endoplasmic reticulum and mitochondrial swelling. Our data suggest that cytotoxic effects on midgut endocrine cells can contribute to a generalized disruption of the physiological processes in this organ due to a general alteration of endocrine function.

Post-exposure temperature influence on the toxicity of conventional and new chemistry insecticides to green lacewing Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae)

PubMed Central

Mansoor, Muhammad Mudassir; Afzal, Muhammad; Raza, Abu Bakar M.; Akram, Zeeshan; Waqar, Adil; Afzal, Muhammad Babar Shahzad

2014-01-01

Chrysoperla carnea (Stephens) is an important biological control agent currently being used in many integrated pest management (IPM) programs to control insect pests. The effect of post-treatment temperature on insecticide toxicity of a spinosyn (spinosad), pyrethroid (lambda cyhalothrin), organophosphate (chlorpyrifos) and new chemistry (acetamiprid) to C. carnea larvae was investigated under laboratory conditions. Temperature coefficients of each insecticide tested were evaluated. From 20 to 40 °C, toxicity of lambda cyhalothrin and spinosad decreased by 2.15- and 1.87-fold while toxicity of acetamiprid and chlorpyrifos increased by 2.00 and 1.79-fold, respectively. The study demonstrates that pesticide effectiveness may vary according to environmental conditions. In cropping systems where multiple insecticide products are used, attention should be given to temperature variation as a key factor in making pest management strategies safer for biological control agents. Insecticides with a negative temperature coefficient may play a constructive role to conserve C. carnea populations. PMID:25972753

Tri-trophic interactions between Bt cotton, the herbivore Aphis gossypii Glover (Homoptera: Aphididae), and the predator Chrysopa pallens (Rambur) (Neuroptera: Chrysopidae).

PubMed

Guo, Jian-Ying; Wan, Fang-Hao; Dong, Liang; Lövei, Gábor L; Han, Zhao-Jun

2008-02-01

Tri-trophic impacts of transgenic Bacillus thuringiensis (Bt) cotton GK12 and NuCOTN 99B were studied using a predator, the great lacewing Chrysopa pallens (Rambur), and its prey, the cotton aphid Aphis gossypii Glover, in laboratory feeding experiments. The parental nontransgenic cotton cultivar of GK12 was used as control. The predator was fed with uniform (aphids from a single cultivar) or mixed prey (aphids from the three cotton cultivars provided on alternate days). Mortality and development of the immature stages, pupal body mass, adult sex ratio, fecundity, and egg viability of C. pallens were measured. When fed GK12-originated aphid prey, pupal body mass of C. pallens was significantly higher than that of the control, more females emerged, and these females laid significantly more eggs. Other parameters were not impacted. Females emerging from larvae maintained on NuCOTN 99B-originated prey laid fewer eggs than those maintained on GK12. Other measurements did not differ significantly between the two Bt cotton cultivars. Compared with the control, mixed feeding significantly prolonged pupal development time and increased pupal body mass and percentage of females but did not affect other parameters. These results indicate that C. pallens is sensitive to aphid prey from different cotton cultivars. Transgenic Bt cotton GK12-originated aphid prey has no adverse impact on survival, development, and fecundity of C. pallens. Between the two Bt cotton cultivars, NuCOTN 99B-originated aphid prey provided to C. pallens in the larval stage may lower female fecundity. Mixed feeding of C. pallens with the two Bt cotton-originated prey and non-Bt prey may have some adverse impacts on pupal development.

Demographic analysis, a comparison of the jackknife and bootstrap methods, and predation projection: a case study of Chrysopa pallens (Neuroptera: Chrysopidae).

PubMed

Yu, Ling-Yuan; Chen, Zhen-Zhen; Zheng, Fang-Qiang; Shi, Ai-Ju; Guo, Ting-Ting; Yeh, Bao-Hua; Chi, Hsin; Xu, Yong-Yu

2013-02-01

The life table of the green lacewing, Chrysopa pallens (Rambur), was studied at 22 degrees C, a photoperiod of 15:9 (L:D) h, and 80% relative humidity in the laboratory. The raw data were analyzed using the age-stage, two-sex life table. The intrinsic rate of increase (r), the finite rate of increase (lambda), the net reproduction rate (R0), and the mean generation time (T) of Ch. pallens were 0.1258 d(-1), 1.1340 d(-1), 241.4 offspring and 43.6 d, respectively. For the estimation of the means, variances, and SEs of the population parameters, we compared the jackknife and bootstrap techniques. Although similar values of the means and SEs were obtained with both techniques, significant differences were observed in the frequency distribution and variances of all parameters. The jackknife technique will result in a zero net reproductive rate upon the omission of a male, an immature death, or a nonreproductive female. This result represents, however, a contradiction because an intrinsic rate of increase exists in this situation. Therefore, we suggest that the jackknife technique should not be used for the estimation of population parameters. In predator-prey interactions, the nonpredatory egg and pupal stages of the predator are time refuges for the prey, and the pest population can grow during these times. In this study, a population projection based on the age-stage, two-sex life table is used to determine the optimal interval between releases to fill the predation gaps and maintain the predatory capacity of the control agent.

Post-exposure temperature influence on the toxicity of conventional and new chemistry insecticides to green lacewing Chrysoperla carnea (Stephens) (Neuroptera: Chrysopidae).

PubMed

Mansoor, Muhammad Mudassir; Afzal, Muhammad; Raza, Abu Bakar M; Akram, Zeeshan; Waqar, Adil; Afzal, Muhammad Babar Shahzad

2015-05-01

Chrysoperla carnea (Stephens) is an important biological control agent currently being used in many integrated pest management (IPM) programs to control insect pests. The effect of post-treatment temperature on insecticide toxicity of a spinosyn (spinosad), pyrethroid (lambda cyhalothrin), organophosphate (chlorpyrifos) and new chemistry (acetamiprid) to C. carnea larvae was investigated under laboratory conditions. Temperature coefficients of each insecticide tested were evaluated. From 20 to 40 °C, toxicity of lambda cyhalothrin and spinosad decreased by 2.15- and 1.87-fold while toxicity of acetamiprid and chlorpyrifos increased by 2.00 and 1.79-fold, respectively. The study demonstrates that pesticide effectiveness may vary according to environmental conditions. In cropping systems where multiple insecticide products are used, attention should be given to temperature variation as a key factor in making pest management strategies safer for biological control agents. Insecticides with a negative temperature coefficient may play a constructive role to conserve C. carnea populations.

Effects of neem oil (Azadirachta indica A. Juss) on midgut cells of predatory larvae Ceraeochrysa claveri (Navás, 1911) (Neuroptera: Chrysopidae).

PubMed

Scudeler, Elton Luiz; dos Santos, Daniela Carvalho

2013-01-01

The effects of ingested neem oil, a botanical insecticide obtained from the seeds of the neem tree, Azadirachta indica, on the midgut cells of predatory larvae Ceraeochrysa claveri were analyzed. C. claveri were fed on eggs of Diatraea saccharalis treated with neem oil at a concentration of 0.5%, 1% and 2% during throughout the larval period. Light and electron microscopy showed severe damages in columnar cells, which had many cytoplasmic protrusions, clustering and ruptured of the microvilli, swollen cells, ruptured cells, dilatation and vesiculation of rough endoplasmic reticulum, development of smooth endoplasmic reticulum, enlargement of extracellular spaces of the basal labyrinth, intercellular spaces and necrosis. The indirect ingestion of neem oil with prey can result in severe alterations showing direct cytotoxic effects of neem oil on midgut cells of C. claveri larvae. Therefore, the safety of neem oil to non-target species as larvae of C. claveri was refuted, thus the notion that plants derived are safer to non-target species must be questioned in future ecotoxicological studies. Copyright © 2012 Elsevier Ltd. All rights reserved.

Neem oil (Azadirachta indica A. Juss) affects the ultrastructure of the midgut muscle of Ceraeochrysa claveri (Navás, 1911) (Neuroptera: Chrysopidae).

PubMed

Scudeler, Elton Luiz; Garcia, Ana Silvia Gimenes; Pinheiro, Patricia Fernanda Felipe; Santos, Daniela Carvalho Dos

2017-01-01

Cytomorphological changes, by means of ultrastructural analyses, have been used to determine the effects of the biopesticide neem oil on the muscle fibers of the midgut of the predator Ceraeochrysa claveri. Insects, throughout the larval period, were fed eggs of Diatraea saccharalis treated with neem oil at a concentration of 0.5%, 1% or 2%. In the adult stage, the midgut was collected from female insects at two stages of adulthood (newly emerged and at the start of oviposition) and processed for ultrastructural analyses. In the newly emerged insects obtained from neem oil treatments, muscle fibers showed a reduction of myofilaments as well as swollen mitochondria and an accumulation of membranous structures. Muscular fibers responded to those cellular injuries with the initiation of detoxification mechanisms, in which acid phosphatase activity was observed in large vesicles located at the periphery of the muscle fiber. At the start of oviposition in the neem oil treated insects, muscle fibers exhibited signs of degeneration, containing vacant areas in which contractile myofilaments were reduced or completely absent, and an accumulation of myelin structures, a dilatation of cisternae of sarcoplasmic reticulum, and mitochondrial swelling and cristolysis were observed. Enzymatic activity for acid phosphatase was present in large vesicles, indicating that mechanisms of lytic activity during the cell injury were utilized but insufficient for recovery from all the cellular damage. The results indicate that the visceral muscle layer is also the target of action of neem oil, and the cytotoxic effects observed may compromise the function of that organ. Copyright © 2016 Elsevier GmbH. All rights reserved.

The movement of proteins across the insect and tick digestive system.

PubMed

Jeffers, Laura A; Michael Roe, R

2008-02-01

The movement of intact proteins across the digestive system was shown in a number of different blood-feeding and non-blood-feeding insects in the orders Blattaria, Coleoptera, Diptera, Hemiptera, Lepidoptera, Orthoptera, Neuroptera and Siphonaptera, as well as in two tick families Ixodidae and Argasidae. Protein movement was observed for both normal dietary and xenobiotic proteins, which suggest that the mechanism for transfer is not substrate specific. The number of studies on the mechanism of movement is limited. The research so far suggests that movement can occur by either a transcellular or an intercellular pathway in the ventriculus with most of the research describing the former. Transfer is by continuous diffusion with no evidence of pinocytosis or vesicular transport common in mammalian systems. Proteins can move across the digestive system without modification of their primary or multimeric structure and with retention of their functional characteristics. Accumulation in the hemolymph is the result of the protein degradation rate in the gut and hemolymph and transfer rate across the digestive system and can be highly variable depending on species. Research on the development of delivery systems to enhance protein movement across the insect digestive system is in its infancy. The approaches so far considered with some success include the use of lipophilic-polyethylene glycol (PEG) polymers, the development of fusion proteins with lectins, reduced gut protease activity and the development of amphiphilic peptidic analogs. Additional research on understanding the basic mechanisms of protein delivery across the insect digestive system, the importance of structure activity in this transfer and the development of technology to improve movement across the gut could be highly significant to the future of protein and nucleic acid-based insecticide development as well as traditional chemical insecticidal technologies.

Side-effects of glyphosate on the life parameters of Eriopis connexa (Coleoptera: Coccinelidae) in Argentina.

PubMed

Mirande, L; Haramboure, M; Smagghe, G; Piñeda, S; Schneider, M I

2010-01-01

In Argentina, transgenic soybean crop (Roundup Ready, RR) has undergone a major expansion over the last 15 years, with the consequent increase of glyphosate applications, a broad-spectrum and post emergence herbicide. Soybean crops are inhabited by several arthropods. Eriopis connexa Germar (Coleoptera: Coccinelidae) is a predator associated to soybean soft-bodies pest and have a Neotropical distribution. Nowadays, it is being considered a potentially biological control agent in South America. The objectives of this work were to evaluate the side-effects of glyphosate on larvae (third instar) and adults of this predator. Commercial compound and the maximum registered concentrations for field use were employed: GlifoGlex 48 (48% glyphosate, 192 mg a.i./litre, Gleba Argentina S.A.). The exposure was by ingestion through the treated prey (Rophalosiphum padi) or by drinking treated water during 48 h for treatment of the adult. The herbicide solutions were prepared using distilled water as solvent. The bioassays were carried out in the laboratory under controlled conditions: 23 +/- 0.5 degrees C, 75 +/- 5% RH and 16:8 (L:D) of photoperiod. Development time, weight of pupae, adult emergence, pre-oviposition period, fecundity and fertility were evaluated as endpoints. Larvae from glyphosate treatment molted earlier than controls. In addition, the weight of pupae, longevity, fecundity and fertility were drastically reduced in treated organisms. The reductions were more drastic when the treatments were performed at the third larval stage than as adult. The reproduction capacity of the predator was the most affected parameter and could be related to a hormonal disruption by glyphosate in the treated organisms. This work can confirm the deleterious effects of this herbicide on beneficial organisms. Also, it agrees with prior studies carried out on other predators associated to soybean pest, such as Chrysoperla externa (Neuroptera: Chrysopidae) and Alpaida veniliae (Araneae

The morphology and evolution of the female postabdomen of Holometabola (Insecta).

PubMed

Hünefeld, Frank; Missbach, Christine; Beutel, Rolf Georg

2012-07-01

In the present article homology issues, character evolution and phylogenetic implications related to the female postabdomen of the holometabolan insects are discussed, based on an earlier analysis of a comprehensive morphological data set. Hymenoptera, the sistergroup of the remaining Holometabola, are the only group where the females have retained a fully developed primary ovipositor of the lepismatid type. There are no characters of the female abdomen supporting a clade Coleopterida + Neuropterida. The invagination of the terminal segments is an autapomorphy of Coleoptera. The ovipositor is substantially modified in Raphidioptera and distinctly reduced in Megaloptera and Neuroptera. The entire female abdomen is extremely simplified in Strepsiptera. The postabdomen is tapering posteriorly in Mecopterida and retractile in a telescopic manner (oviscapt). The paired ventral sclerites of segments VIII and IX are preserved, but valvifers and valvulae are not distinguishable. In Amphiesmenoptera sclerotizations derived from the ventral appendages VIII are fused ventromedially, forming a solid plate, and the appendages IX are reduced. The terminal segments are fused and form a terminal unit which bears the genital opening subapically. The presence of two pairs of apophyses and the related protraction of the terminal unit by muscle force are additional autapomorphies, as is the fusion of the rectum with the posterior part of the genital chamber (cloaca). Antliophora are supported by the presence of a transverse muscle between the ventral sclerites of segment VIII. Secondary egg laying tubes have evolved independently within Boreidae (absent in Caurinus) and in Tipulomorpha. The loss of two muscle associated with the genital chamber are likely autapomorphies of Diptera. The secondary loss of the telescopic retractability of the postabdomen is one of many autapomorphies of Siphonaptera. Copyright © 2012 Elsevier Ltd. All rights reserved.

[Effects of Beauveria bassiana on Myzus persicae and its two predaceous natural enemies].

PubMed

Zhu, Hong; Luo, Xu-mei; Song, Jin-xin; Fan, Mei-zhen; Li, Zeng-zhi

2011-09-01

A Beauveria bassiana strain Bb21 was isolated from naturally infected green peach aphid Myzus persicae (Hemiptera: Aphididae). The effects of the strain on M. persicae and its two predaceous natural enemies Chrysoperla carnea (Neuroptera: Chrysopidae) and Harmonia axyridis (Coleoptera: Coccinellidae) were investigated under laboratory conditions. Bb21 had strong pathogenicity to M. persicae, with the LD50 of 97 conidia x mm(-2) (45-191, 95% confidence interval), but was less pathogenic to the second instar nymph of C. carnea, with the LD50 of 1089 conidia x mm(-2). The LD50 for C. carnea was 10.2 times higher than that for M. persicae. The pathogenicity of Bb21 to H. axyridis was very weak, with a low infection rate of 13% even at a high concentration 5 x 10(8) conidia x mL(-1). The Bb21 at low conidia concentration had less effect on the developmental period and fecundity of the two predaceous natural enemies. However, when applied at the high concentration 5 x 10(8) spores x mL(-1), Bb21 shortened the larval stage of H. axyridis averagely by 1.4 d and decreased the adult emergence rate and fecundity by 33% and 14%, respectively, and shortened the larval stage of C. carnea averagely by 0.7 d and decreased the adult emergence rate and fecundity by 24% and 11%, respectively. Since the LD50 for green peach aphid was much lower than that for the two predaceous natural enemies, and had very low effect on the adult emergence rate and fecundity of the two predators at the concentration recommended for field spray, Bb21 could be applied as a biocontrol agent of M. persicae in the integrated management of pernicious organisms.

IPM-compatibility of foliar insecticides for citrus: Indices derived from toxicity to beneficial insects from four orders

PubMed Central

Michaud, J.P.; Grant, A.K.

2003-01-01

A series of compounds representing four major pesticide groups were tested for toxicity to beneficial insects representing four different insect orders: Coleoptera (Coccinellidae), Hemiptera (Anthocoridae), Hymenoptera (Aphelinidae), and Neuroptera (Chrysopidae). These materials included organophosphates (methidathion, esfenvalerate and phosmet), carbamates (carbofuran, methomyl and carbaryl), pyrethroids (bifenthrin, fenpropathrin, zeta-cypermethrin, cyfluthrin and permethrin) and the oxadiazine indoxacarb. Toxicity to coccinellid and lacewing species was assessed by treating 1st instar larvae with the recommended field rate of commercial products, and two 10 fold dilutions of these materials, in topical spray applications. Adult Aphytis melinus Debach and 2nd instar Orius insidiosus (Say) were exposed to leaf residues of the same concentrations for 24 h. ANOVA performed on composite survival indices derived from these data resolved significant differences among materials with respect to their overall toxicity to beneficial insects. Cyfluthrin, fenpropathrin and zeta-cypermethrin all increased the developmental time of the lacewing and one or more coccinellid species for larvae that survived topical applications. Bifenthrin increased developmental time for two coccinellid species and decreased it in a third. Indoxacarb (Avaunt® WG, DuPont Corp.) ranked highest overall for safety to beneficial insects, largely because of its low dermal toxicity to all species tested. Zeta-cypermethrin (Super Fury®, FMC Corporation) received the second best safety rating, largely because of its low toxicity as a leaf residue to A. melinus and O. insidiosus. Phosmet (Imidan® 70W, Gowan Co.) and methidathion (Supracide® 25W, Gowan Co.) ranked high for safety to coccinellid species, but compounds currently recommended for use in citrus such as fenpropathrin (Danitol® 2.4EC, Sumimoto Chem. Co.) and carbaryl (Sevin® XLR EC, Rhone Poulenc Ag. Co.) ranked very low for IPM

Lethal and sublethal effects of pesticides on Chrysoperla carnea larvae (Neuroptera: Chrysopidae) and the influence of rainfastness in their degradation pattern over time.

PubMed

Maia, Jader Braga; Carvalho, Geraldo Andrade; Medina, Pilar; Garzón, Agustín; Gontijo, Pablo da Costa; Viñuela, Elisa

2016-07-01

The predator Chrysoperla carnea is a model species for the study of non-target effects of pesticides under different scenarios: registration of plant protection products under the European Union and effects of the Bt toxin. Laboratory and persistence studies were carried out with six pesticides currently used in corn crops in Spain that were applied at their maximum field recommended concentrations. The assessed end-points were larval mortality, survivorship until adult stage, duration of the larval and pupal periods, fecundity, fertility and sex ratio of the emerged adults. Based on the total effect (lethal and sublethal) caused to L3 larvae in contact with fresh residues in the laboratory, pendimethalin was harmless (IOBC 1), lambda-cyhalothrin, abamectin, and hexythiazox were slightly harmful (IOBC 2), deltamethrin was moderately harmful (IOBC 3) and chlorpyrifos was harmful (IOBC 4). Afterwards, the residues of the two most toxic pesticides in the lab (deltamethrin and chlorpyrifos) were aged under greenhouse conditions (22 ± 2 °C, 40 ± 10 % R.H., 16.9 μmol m(-2) s(-1) UV radiation) in the presence and absence of artificial rainfall (10 l m(-2) h(-1), applied 24 h after pesticide application). Deltamethrin was classified as short lived (IOBC A) in both cases. However, degradation of chlorpyrifos residues was accelerated in the presence of rainfall, leading to the classification as slightly persistent (IOBC B), while in absence of rainfall it behaved as persistent (IOBC D). Every pesticide can be recommended for inclusion in corn IPM programs where the predator is present except chlorpyrifos that exhibited high direct toxicity in the lab and prolonged residual action even in the presence of rainfall.

Foraging on and consumption of two species of papaya pest mites, Tetranychus kanzawai and Panonychus citri (Acari: tetranychidae) by Mallada basalis (Neuroptera: Chrysopidae)

USDA-ARS?s Scientific Manuscript database

Tetranychus kanzawai Kishida and Panonychus citri (McGregor) are two major acarine pests of the principal papaya variety in Taiwan, and they often co-occur in the same papaya screenhouses. This study measured prey acceptability, foraging schedule, short-term consumption rate, and handling time of la...

Attraction of Chrysoperla carnea complex and Chrysopa spp. lacewings (Neuroptera: Chrysopidae) to aphid sex pheromone components and a synthetic blend of floral compounds in Hungary.

PubMed

Koczor, Sándor; Szentkirályi, Ferenc; Birkett, Michael A; Pickett, John A; Voigt, Erzsébet; Tóth, Miklós

2010-12-01

The deployment of synthetic attractants for the manipulation of lacewing populations as aphid predators is currently used in integrated pest management. This study investigates a synthetic bait comprising floral compounds previously found to attract the Chrysoperla carnea complex, and, for the first time, the aphid sex pheromone components (1R,4aS,7S,7aR)-nepetalactol and (4aS,7S,7aR)-nepetalactone, in field experiments in Hungary, for their ability to manipulate lacewing populations. The synthetic floral bait attracted both sexes of the Chrysoperla carnea complex, and Chrysopa formosa Brauer showed minimal attraction. The aphid sex pheromone compounds alone attracted males of C. formosa and C. pallens (Rambur). When the two baits were combined, Chrysopa catches were similar to those with aphid sex pheromone baits alone, but carnea complex catches decreased significantly (by 85-88%). As the floral bait alone attracted both sexes of the carnea complex, it showed potential to manipulate the location of larval density via altering the site of oviposition. Aphid sex pheromone compounds alone attracted predatory males of Chrysopa spp. and can potentially be used to enhance biological control of aphids. For the carnea complex, however, a combination of both baits is not advantageous because of the decrease in adults attracted. Assumptions of intraguild avoidance underlying this phenomenon are discussed. Copyright © 2010 Society of Chemical Industry.

Comparative mitogenomics of Braconidae (Insecta: Hymenoptera) and the phylogenetic utility of mitochondrial genomes with special reference to Holometabolous insects

PubMed Central

2010-01-01

sister-group of Diptera. Neuropterida (Neuroptera + Megaloptera), and a sister-group relationship with (Diptera + Mecoptera) were supported across all analyses. Conclusions Our comparative studies indicate that mitochondrial genomes are a useful phylogenetic tool at the ordinal level within Holometabola, at the superfamily within Hymenoptera and at the subfamily level within Braconidae. Variation at all of these hierarchical levels suggests that the utility of mitochondrial genomes is likely to be a valuable tool for systematics in other groups of arthropods. PMID:20537196

Comparative mitogenomics of Braconidae (Insecta: Hymenoptera) and the phylogenetic utility of mitochondrial genomes with special reference to Holometabolous insects.

PubMed

Wei, Shu-jun; Shi, Min; Sharkey, Michael J; van Achterberg, Cornelis; Chen, Xue-xin

2010-06-11

Diptera. Neuropterida (Neuroptera + Megaloptera), and a sister-group relationship with (Diptera + Mecoptera) were supported across all analyses. Our comparative studies indicate that mitochondrial genomes are a useful phylogenetic tool at the ordinal level within Holometabola, at the superfamily within Hymenoptera and at the subfamily level within Braconidae. Variation at all of these hierarchical levels suggests that the utility of mitochondrial genomes is likely to be a valuable tool for systematics in other groups of arthropods.

Olfactory responses of Plutella xylostella natural enemies to host pheromone, larval frass, and green leaf cabbage volatiles.

PubMed

Reddy, G V P; Holopainen, J K; Guerrero, A

2002-01-01

The parasitoids Trichogramma chilonis (Hymenoptera: Trichogrammatidae) and Cotesia plutellae (Hymenoptera: Braconidae), and the predator Chrysoperla carnea (Neuroptera: Chrysopidae), are potential biological control agents for the diamondback moth, Plutella xylostella (Lepidoptera: Yponomeutidae). We present studies on the interactions between these bioagents and various host-associated volatiles using a Y olfactometer. T chilonis was attracted to a synthetic pheromone blend (Z11-16:Ald, Z11-16:Ac, and Z11-16:OH in a 1:1:0.01 ratio), to Z11-16:Ac alone, and to a 1:1 blend of Z11-16:Ac and Z11-16:Ald. C. plutellae responded to the blend and to Z11-16:Ac and Z11-16:Ald. Male and female C. carnea responded to the blend and to a 1:1 blend of the major components of the pheromone, although no response was elicited by single compounds. Among the four host larval frass volatiles tested (dipropyl disulfide, dimethyl disulfide, allyl isothiocyanate, and dimethyl trisulfide), only allyl isothiocyanate elicited significant responses in the parasitoids and predator, but C. plutellae and both sexes of C. carnea did respond to all four volatiles. Among the green leaf volatiles of cabbage (Brassica oleracea subsp. capitata), only Z3-6:Ac elicited significant responses from T. chilonis, C. plutellae, and C. carnea, but C. plutellae also responded to E2-6:Ald and Z3-6:OH. When these volatiles were blended with the pheromone, the responses were similar to those elicited by the pheromone alone, except for C. carnea males, which had an increased response. The effect of temperature on the response of the biological agents to a mixture of the pheromone blend and Z3-6:Ac was also studied. T. chilonis was attracted at temperatures of 25-35 degrees C, while C. plutellae and C. carnea responded optimally at 30-35 degrees C and 20-25 degrees C, respectively. These results indicate that the sex pheromone and larval frass volatiles from the diamondback moth, as well as volatile compounds from

Shallot Aphids, Myzus ascalonicus, in Strawberry: Biocontrol Potential of Three Predators and Three Parasitoids

PubMed Central

Enkegaard, Annie; Sigsgaard, Lene; Kristensen, Kristian

2013-01-01

The parasitization capacity of 3 parasitoids and the predation capacity of 3 predators towards the shallot aphid, Myzus ascalonicus Doncaster (Homoptera: Aphididae), on strawberry, Fragaria x ananassa Duchesne (Rosales: Rosaceae) cv. Honeoye, were examined in laboratory experiments. In Petri dish assays, both Aphidius colemani Viereck (Hymenoptera: Aphidiidae) and A. ervi Haliday readily stung shallot aphids, with no significant difference in stinging frequency between the two species. A. ervi induced a significantly higher mortality (79.0 ± 7.2%) in terms of stung aphids compared with A. colemani (55.3 ± 4.1%); however, only a minor fraction (2.7 ± 1.8% and 7.1 ± 3.1%, respectively) of the killed aphids resulted in formation of mummies, presumably due to a physiological response to parasitism. The low percentage of mummification precludes the use of either Aphidius species in anything but inundative biocontrol. In similar set-ups, Aphelinus abdominalis (Dalman) (Hymenoptera: Aphelinidae) killed almost half (49.6 ± 5.3%) of the exposed aphids through host feeding. In addition, 23.2 ± 7.3% of non-host-fed aphids developed into mummified aphids, and 38.1 ± 13.2% of non-host-fed aphids died from other parasitoid-induced causes. However, the host feeding rate was reduced to only 1.2 ± 0.8%, and no significant parasitization mortality was observed on strawberry plants, suggesting that host plants interfered with A. abdominalis activity. This parasitoid does not, therefore, seem to be suited to either inoculative or inundative biocontrol of shallot aphids in strawberry. The three predators studied were the green lacewing, Chrysoperla carnea Steph. (Neuroptera: Chrysopidae), the two-spotted lady beetle, Adalia bipunctata L. (Coleoptera: Coccinellidae), and the gall midge Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae). Third instars of all 3 predators readily preyed upon the shallot aphid in Petri dish set-ups with significant differences in daily

An assessment of arthropod prey resources at Nakula Natural Area Reserve, a potential site of reintroduction for Kiwikiu (Pseudonestor xanthophrys) and Maui `Alauahio (Parareomyza montana).

USGS Publications Warehouse

Banko, Paul C.; Peck, Robert W.; Cappadonna, Justin; Steele, Claire; Leonard, David L.; Mounce, Hanna L.; Becker, Dusti; Swinnerton, Kirsty

2015-01-01

), which comprised 90% of all prey items for 50 adult birds and 98% of all prey for two nestlings. Caterpillars were also the most important prey for Maui ‘alauahio (43% for 104 adult birds) although spiders (Araneae, 16%), beetles (12%) and true bugs, planthoppers and psyllids (Hemiptera; 12%) were also important. Caterpillars were generally the most abundant type of arthropod in the foliage of koa and ‘ōhi‘a, although spiders, beetles and hemipterans were also common. Total arthropod biomass and caterpillar biomass at Nakula was as great, or greater, than that observed at Hanawi and Waikamoi per unit of foliage of both koa and ‘ōhi‘a. Spiders generally dominated the bark fauna on both koa and ‘ōhi‘a at all sites although isopods (Isopoda), millipedes (Myriapoda: Millipeda) and lacewings (Neuroptera) were also abundant at Waikamoi and Hanawi. Total arthropod biomass on bark, as well as the biomass of several individual taxa, was significantly lower at Nakula than the other sites. Our measurement of the density of beetle exit holes in dead koa branches found no difference between Nakula and Waikamoi. Finally, no difference existed in the abundance of arthropods (primarily caterpillars and moth pupae) within ‘ākala stems among sites. With the exception of bark surfaces, our results suggest that the arthropod prey base for birds on primary foraging substrates at Nakula is similar to that found at two sites within the current range of kiwikiu and Maui ‘alauahio. However, our results should be viewed with caution because they are limited to the scale of individual branch, tree, or ‘ākala stem. To complete the assessment, our results should be scaled up to the landscape level by determining the density of each substrate within each site. Key arthropod prey of kiwikiu and Maui ‘alauahio are available at Nakula and, as habitat restoration continues, food abundance should increase to the point at which populations of these birds can be supported.