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            Introduction                                                                                                                                        Contents

 

Entomology:  NEUROPTERA 1

Kingdom:  Animalia, Phylum: Arthropoda

Subphylum: Hexapoda: Class: Insecta: Order: Neuroptera

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Pteragota:  Holometabola

  Order:  Neuroptera (28 Families)

    General Summary

      Raphidiodea

      Planipennia

    Morphology & Habits

    Families of Neuroptera

    Sample Examinations

    References      Citations

 

General Summary of Neuroptera

 

          There are two pairs of large, broad wings, the posterior ones having a large anal field. The longitudinal veins branch freely and cross-veins are common, especially behind the anterior border.  A pterostigma or pigmented area located laterally on the anterior border is either missing or weakly defined. At rest the wings are held over the back in a roof-like manner.

 


          Their larvae are carnivorous and always aquatic.  They bear on the abdomen a series of pairs of gills that are jointed and which are moved by intrinsic muscles.  Sialis lutaria, the alder fly alderfly occurs in Northern Europe. Corydalus of North and South America and Archichauliodes of Australia and New Zealand are other examples (
Borradaile & Potts, 1958).

 

          Suborder Raphidiodea:  are insects of small size. They have two pairs of similar hyaline wings with freely branching longitudinal veins and a well-marked pterostigma are characteristic. The head projects forwards and is flattened above and narrowed toward the prothoracic junction. This, together with the elongated sub-cylindrical prothorax and the elongated 10th segment of the abdomen, gives them the name "Snakeflies."  The larvae are terrestrial.  Raphidia is Northern European.

 

         Suborder Planipennia (Lacewings and Ant lions):  There is a wide range of form and size in this group, both very large and very small insects included. There is also much variation in the wings because in some forms, e.g. Ithone, the two pairs of wings are identical while in others, e.g. Nemoptera, the posterior ones are elongated to narrow strap-like structures several times larger than the body.  However, all except the small Coniopterygidae agree in having an abundantly branching venation with many cross-veins. Metamorphosis is complete.


          Larvae of the majority of Planipennia are terrestrial as in Myrmeleon, the ant lion.  However, some like Sisyra are aquatic and parasitic in he freshwater sponge, Spongilla.   General features of the larvae of Planipennia are the forward extended, curved and pointed mandibles and maxillae, which are so arranged as to form between them at each side a food tube up which the blood of their prey is drawn.  Larvae are mostly predaceous.  Hemerobius, Sisyra, Chrysopa, etc. are found in Northern Europe (
Borradaile & Potts, 1958).

 

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Morphology & Habits

 

          The name Neuroptera means, "nerve-winged."  They have complete metamorphosis and their wings are developed internally.  The larvae are either aquatic or terrestrial, e.g., the Dobsonfly has aquatic larvae while ant and aphis lions have terrestrial larvae.

 

          The larvae have chewing mouthparts but feed by sucking.  They possess a pincher-like mandible with opposing maxillae.  The mandible is grooved, the maxilla acting as a cover.  Together they form a food channel that connects with the mouth cavity.

 

 

          Green lacewing larvae are important aphid predators. 

 

          Pupation occurs on land and the pupae are active.  The adult possesses large mandibles also, while the wing venation is similar to the damselfly (Odonata).  All Neuroptera have knobbed antennae.

 

          Ant lions may live several years, especially if food is not too plentiful.  The adults are either predators or nonfeeding.  Many species are highly beneficial as predators of destructive insects. The eggs are laid on a stalk as a protection from predation by other members of the same species.  At first the eggs are laid in the prone position, but they later spring up.

 

 

          Aphis lions are called "green lacewings" in the adult stage.

 

          There are some modifications within the order.  Some adults possess raptorial legs.  Sialidae larvae have gills on the abdomen, which are located laterally and are segmented.

 

          Neuroptera show some affinities with other groups such as Coleoptera to which they are closely related.

 

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Families of Neuroptera

 

          Some of the common families of Neuroptera are distinguished according to their shapes and habits as follows (see Borror et al. 1989 for details):

 

             Apochrysidae. -- <Habits>; <Adults>

 

 

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          Ascalaphidae. -- <Habits>; <Adults> & <Juveniles> -- Owl flies resemble dragonflies with long antennae.  Ululodes hyalina Latr. of the southern United States and Central America was studied by McClendon (1902).  Eggs are laid in groups of 57-75 in a double row at the end of a twig, and the mass is fenced off slightly below the base by several circles of "repagula" placed on end.  These repagula are thought to be aborted eggs, produced by certain ovarian tubules at the same time that others form normal eggs.  This formation is thought to protect the egg mass from natural enemies.  The incubation period is 9-10 days.

 

 

Larvae hide in depression in soil or under the edges of stones and cover their bodies with sand or dust.  While awaiting prey, the huge mandibles are held widely separated.  The closing of the jaws is seemingly triggered by contact, and the prey is usually paralyzed within seconds by the bite.  The body fluids of the prey are absorbed through a duct formed by the fitting together of curved mandible and maxilla.  There are 3 larval instars, and development takes ca. 62 days.

 

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          Berothidae. -- <Habits>; <Adults>   Berothidae or beaded lacewings, are a family of winged insects of the order Neuroptera. The family was first named by Anton Handlirsch in 1906.

 

 

 

 

          Berothidae is very closely related to the thorny lacewings which are often included in the Berothidae as subfamily "Rhachiberothinae" but occasionally are considered the distinct family Rhachiberothidae or included as a subfamily in the mantidflies.  Mantidflies, family Mantispidae, are another group of rather close relatives, and at least some of the probably paraphyletic group of fossil forms collectively called "Mesithonidae" also seem to be quite close.  The family consists of 22 genera and 100 living species distributed discontinuously world wide with most being subtropical to tropical.[1] Eleven extinct genera with a total of thirteen species have been described from the fossil record.

 

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          Chrysopidae. -- <Habits>; <Adults> & <Juveniles> -- Common lacewings green & 2nd largest family in order.  Chrysopids are considered wholly beneficial and have been used in augmentation release programs against homopterous pests throughout the world.  A number of species of Chrysopa were introduced to New Zealand for use in the control of aphid and mealybug pests and also against Chermidae attacking pines (Clausen 1940/62).

 

 

 

 

Chrysopidae feed on a variety of soft-bodied insects, but mostly on aphids and mealybugs.  Leafhoppers, thrips, lecaniine Coccidae, mites, etc., may also be attacked.  Extensive feeding occurs at intervals on eggs of Lepidoptera.  Larvae of Chrysopa rufilabris Burm. have been found to pierce leaf tissue with the mandibles in order to feed on larvae of Agromyza jucunda v.d.W. in their mines.  This species is a valuable predator of red mites on cotton, the larvae consuming an average of 80 per day during the entire developmental period (Clausen 1940/62).  Generally, adults feed on the same insects that serve as prey for larvae, although their activities in this respect are less.  Extended early accounts of the biology and behavior of Chrysopidae are by Wildermuth (1916), Smith (1921, 1922b) and Withycombe (1923).

 

Adults usually live 4-6 weeks.  Oviposition occurs the day following emergence from the cocoon and mating, but occasional species pass the winter as adults and oviposit the following spring and summer.  Killington (1935) referred to a spermatophore being produced at mating by Nathanica fulviceps Steph., although Withycombe (1923) did not find one.  The number of eggs laid by the different species varies, the maximum being recorded by Smith of 617 by a female of Chrysopa occulata Fitch in 42 days.  The general average is thought to be 100-200.  Killington (1936) cited oviposition records, among which are those by Okamoto of 550 eggs from C. nipponensis Okam. and by Withycombe of 480 eggs from a female of C. phyllochroma Wesm.

 

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          Coniopterygidae. -- <Habits>; <Adults> & <Juveniles> -- Dusty wings are tiny insects of only 3 mm in length.  Both adults and immatures feed on small, relatively inactive prey such as coccids, mites and aphids.  They are mainly found on shrubs and trees, although some species seem to be confined to low vegetation.  Adults are active fliers especially at sunset, when both sexes are attracted to lights.  Fontenellea maroccana Carp. & Lest. attacks Orthesia in North Africa and an undetermined species was observed feeding on Cryptoparlatoria leucaspis Lind. on cryptomeria in Japan (T. Ishii cited by Clausen, 1940).  Withycombe (1923, 1924a) found several species to be predaceous on Phylloxera, his observations indicating that the feeding range of the various species is wide.  Conwentzia psociformis Curt. is associated with oak in England where it feeds on Phylloxera but also on diaspine Coccidae, red mites, etc (Arrow 1917). 

 

 

Eggs are laid singly on infested foliage.  They are oval in outline, flattened dorsoventrally, and slightly pointed at the micropylar end.  The chorion surface bears reticulate markings.  Eggs of C. hageni Banks are yellowish-pink, although some may have an orange tint (Quayle 1913).  Conwentzia psociformis lays a total of ca. 200 eggs.

 

The number of larval instars was noted as 4 for C. hageni (Quayle 1913).  Larvae of this species feed on all stages of red mites, the body contents being entirely sucked out from a single puncture.  One larva consumed 226 red mites during its feeding period.  The oval, flattened cocoons of Conwentzia are usually found on the underside of leaves or on bark.  They consist of a double layer of silk with loosely woven margins.  Cocoons of Semidalis aleyrodiformis Steph. do not have a clear double layer of silk.  Withycombe (1923, 1924a) found that the pupal skin is often left within the cocoon rather than discarded after adult emergence.

 

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          Corydalidae. -- <Habits>; <Adults>   Dobson flies & fish flies are usually longer than 25 mm.  damselflies with long, narrow, many-veined wings and a long slender abdomen.   The family Corydalidae contains megalopterous insects known as dobsonflies and fishflies. There are about 12 species that occur mainly throughout the Northern Hemisphere, both temperate and tropical, and South America.

 

 

          They are large Megaloptera, with a body usually larger than 25 mm (1 inch). They may have long filamentous antennae, though in male fishflies they are characteristically feathered. Ocelli are present; the fourth tarsal segment is cylinder-shaped. The four large wings are translucent, smoky grey, or mixed, and the anterior pair is slightly longer than the posterior one.

 

          The eastern dobsonfly, is the most well-known North American species among the dobsonflies. These genera have distinctive elongated mandibles in males and form the subfamily Corydalinae. The genera in which the males have normal mandibles, called fishflies, form the subfamily Chauliodinae. The summer fishfly, Chauliodes pectinicornis, is perhaps the best-known of these in North America; its immense mating swarms in the Upper Mississippi River region fill the air on a few summer nights each year much like mayflies in certain regions of Europe, leaving millions of carcasses to be cleaned up the next day.

 

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          Dilaridae. -- <Habits>; <Adults>   Pleasing lacewings.  Adults resemble small brown moths: wings rounded, hairy, (and in Nallachius americanus, transparent except for many light brown spots, a number of which coalesce to form irregular bands across the wing); costal cross veins of wings not forked; male antennae pectinate; female ovipositor long, recurved over abdomen. Wing venation:

 

 

 

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          Hemerobiidae. -- <Habits>; <Adults> & <Juveniles> -- Brown lacewings.  Hemerobiidae are known as  "brown lacewings" and are distributed worldwide.  There are around 60 species known in North America.  Diagnostic characters include forked costal cross-veins, filiform or moniliform antennae and the forewing having two or more R-s veins. 

 

Hemerobiids resemble chrysopids as far as their predaceous habits are concerned, but they may be distinguished on the basis of several subtle characteristics as follows:  Hemerobiid larvae never carry trash as do some chrysopid larvae; the eggs are not stalked and are attached by their sides to leaves and bark (Withycombe 1922, 1923).  Eggs are whitish, grayish or even pink and elongate-oval, with the chorion often pitted.  They bear a knob-like or disk-like micropyle, often of considerable size (Clausen 1940/62).  The chorion is densely studded with glossy papillate projections.  The egg breaker is saw-shaped.  Eggs of H. pini are pale cream colored, that changes to brown before hatching (Clausen 1940/62).

 

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            Inocelliidae. -- <Habits>; <Adults>   Small insects with front wings of 11-17 mm. 

 

 

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          Ithonidae. -- <Habits>; <Adults> & <Juveniles> -- A rare group with a wingspread of 35-40 mm.  The Australian Ithone fusca Newm. is a predator on various soft-bodied insects, especially scarab larvae that occur in sandy soil (Tillyard 1922).  Clausen (1940) noted that other research has shown that the larvae are unable to feed on other insects and most likely obtain food from plant roots.  In the United States the family is rare, being represented by one species, Oliarces clara Banks from southern California.  The wingspread of this species is 35-40 mm and it resembles Scialis with bleached wings.  Larvae are scarabaeiform and phytophagous.

 

 

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          Mantispidae. -- <Habits>; <Adults> & <Juveniles> (= old Mantidae) -- Mantid flies resemble mantids (Orthoptera).  Adult mantispids or "mantidflies" have large raptorial forelegs, similar to those of Mantidae, which are for capturing other insects.  The larvae are predaceous in the egg sacs of spiders (Mantispa & Climaciella) or on larvae of bees and wasps (Plega).  Brauer (1869b) gave the first and most complete account of one of these species, Mantispa styriaca Poda, that develops in the egg sacs of Lycosidae and related spiders.  Bristowe (1932) supplemented this account.  The eggs are stalked like those of Chrysopidae and are laid in autumn in clusters on tree bark, etc., seemingly without relation to the host.  Hatching occurs ca. 3 weeks later, and young larvae hibernate without feeding.  The following spring or early summer, when the spider egg masses have been laid, they search them out, tearing a hole in the covering to enter.  They then wait until the spider eggs hatch, after which they feed rapidly and extensively.  There is a molt immediately after hatching, and a second one (the last) after some feeding.  The 3rd instar scarabaeiform larvae differ from the active campodeiform 2nd instar (see Clausen 1940 for diagram)..  Its head is small, abdomen very large and the legs are rudimentary.  it is not able to move in an orderly fashion.  The larva completes feeding and spins its yellow oblong cocoon in which the pupa is formed within the unbroken larval skin.  After a while the pupa forces its way out of the old larval skin, cocoon, and the host egg sac, wanders about for a while and eventually casts its skin to release the adult.

 

 

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          Myiodactylidae. -- <Habits>; Adults> & <Juveniles> -- Split-footed lacewings often included with the  Nymphidae.    Clausen (1940) noted this as a separate family of Neuroptera, with only a few genera and species that were restricted to the Australian region and some South Pacific islands.

 

 

Larvae of Osmylops pallidus Banks are rather circular in outline, and bear conspicuous mandibles.  They inhabit the underside of eucalyptus leaves where they prey on other insects (see Clausen 1940 for diagram).

 

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          Myrmeleontidae. -- <Habits>; <Adults> & <Juveniles> -- Ant lions are the largest family in this order.  This family includes the well-known ant lions, named thus because of the habit of larvae feeding mainly on ants and their ferocious appearance and manner of attack.  Larvae are also sometimes called "doodlebugs."  The genus Myrmeleon is highly specialized, and because of its cosmopolitan distribution and abundance, is the dominant group in the Neuroptera with the possible exception of Chrysopidae.  There are ca. 89 species in North America, being most abundant in the South and West.  Adults resemble damselflies, with long slender abdomens.  However, they differ in being softer-bodied and having rather long clubbed antennae and in possessing a different wing venation.  They are feeble fliers that are frequently attracted to lights.  The wings are either clear or irregularly spotted.

 

Reaumur (1742) published an early detailed account of the behavior of Myrmeleon carius L., the accuracy of which was verified by subsequent researchers.  Wheeler (1930) provided a review of the biology and behavior of this species.  The eggs are small, oval in outline, and are laid during autumn in small groups, adhering end to end, in the sand.  Hatching occurs shortly thereafter, and a feeding period occurs before hibernation.  The young ant lion is slender, with conspicuous mandibles and a tough integument, which is essential not only for defense but for protection against desiccation.  There are several types of setae on the dorsum which serve to transmit the stimuli from falling sand particles to indicate the presence of prey in the pit.

 

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          Nemopteridae. -- <Habits>; <Adults> & <Juveniles> -- Generally included in the Nymphidae.    Clausen (1940) noted this as a separate family of Neuroptera, with just a few species that are closely related morphologically to Myrmeleontidae.  Very long and narrow hind wings distinguish adults.  Larvae show a decided constriction between the head and thorax, which reaches its greatest development in the grotesque Necrophylus arenarius Roux, a species occurring in the tombs of Egypt and under ledges along the Nile.  The neck is very slender and as long as the rest of the body.  Larvae live in the dust on the floors of caves, in neglected buildings and in other sheltered spots.

 

 

 

 

 

          Croce filipennis Westw. (Ghosh 1910, Imms 1911) lives in abandoned buildings where the larvae prey on passing insects, especially Dermestidae and Psocidae, the latter of which is probably preferred.  The eggs are oval and the cocoons spherical, both of which are also found in the dust on the floor.  The life cycle is one year, of which the egg and pupal stages take 10-12 and 18-22 days, respectively.

 

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          Nymphidae. -- <Habits>; <Adults> & <Juveniles> -- Split-footed lacewings.  Clausen (1940) noted this as a separate family of Neuroptera.  Nymphus myrmeleonides Leach is found in Australia and certain islands of the South Pacific.  Larvae are found under debris, each covered so that only the front of the head and the mandibles are exposed.  They are believed to prey on wood ants, which are common in their environment.  They form a spherical, parchment-like cocoon (Clausen 1940/62).

 

 

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          Osmylidae. -- <Habits>; <Adults> & <Juveniles> -- A small family of Giant lacewings.  Clausen (1940) noted this as a separate family of Neuroptera, with only a few species, the adults of which are relatively small.  Withycombe (1923) and Killington (1936) gave detailed accounts of Osmylus fulvicephalus Scop.  The larvae are amphibious and probe in mud with their long, slender mandibles for Chironomus spp. larvae and other Diptera inhabiting the wet, mossy borders of streams, etc.  Eggs are elongate-oval, somewhat flattened, with the chorion bearing reticulate markings.  The micropyle is knob-like.  They are laid in rows on some object near the water's edge.  hatching occurs in ca. 22 days, and larvae mature the following spring.  The cocoon is formed in damp moss, and adults emerge 10-12 days after.

 

 

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          Polystoechotidae. -- <Habits>; <Adults>   Giant lacewings with a wing spread of 40-75 mm.  Generally included in the Ithonidae.

 

 

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          Psychopsidae. -- <Habits>; <Adults> & <Juveniles> -- Silky lacewings.  Clausen (1940) noted this as a separate family of Neuroptera with little known habits.  Psychopsis elegans Guer. was studied by Tillyard (1918).  The larvae are predaceous on various insects occurring on trunks and branches of Eucalyptus and related trees.  Oval eggs are laid in small groups on bark during midsummer.  Larvae feed for long intervals, and thus the period of development extends over two seasons.  Egg incubation requires 12 days, and hibernation occurs in the 1st larval instar.  The second winter is passed in the 3rd instar after feeding is completed.  Cocoons resemble those of Chrysopa, and are found in bark crevices.  They spend ca. 3 weeks in the cocoon stage.

 

 

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          Raphidiidae. -- <Habits>; <Adults> & <Juveniles> -- The front wing length varies from 6-17 mm.  Carpenter (1936) gave an early review regarding the behavior of raphidids, most of which were European species.  Adults of Raphidia notata F. were found to feed readily on aphids in the laboratory.  Larger insects such as flies and beetle larvae were attacked only after they had been injured and were thus unable to put up a defense.  Adult Agulla unicolor Carp. fed on freshly killed Chironomidae.  They showed a definite cannibalistic tendency also. 

 

 

 

          Raphidiidae is a small family with less than 100 described species.  There are 17 species of Raphidia known in North America.  They are mainly palearctic in distribution, but are found on all continents except Australia.  Important characters of these "snake flies" include an elongated prothorax with forelegs at the posterior end; front legs not raptorial; forewing R-s with more than two branches; ocelli are absent; ovipositor is exserted and antennae are setiform.  The wings of adults vary from 6-17 mm long, and females are somewhat larger than males.

 

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          Sialidae. -- <Habits>; <Adults> & <Juveniles> -- Alder flies, which are dark colored and about 25 mm. long.  The alderflies are a small family and the larvae of Sialis and several other genera are entirely aquatic feeding partly on other aquatic insects and smaller animals in their environment.  Adults are dark, ca. 25 mm. max. long.  The eggs are laid in masses on vegetation over or near water.  The larvae differ from the Corydalidae by having a terminal filament, seven pairs of lateral filaments and no hooked anal prolegs.

 

 

 

         Sialids have not been used in biological pest control, but are certainly important in the natural control of insects in aquatic habitats. 

 

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          Sisyridae. -- <Habits>; <Adults> & <Juveniles> -- Spongilla flies appear as brownish lacewings.  This is a small family of "spongillaflies" with just a few genera and species.  There are 6 species in North America, three each in genera Climacia and Sisyra.  The Rs vein in the former has a single fork while the latter has 2 forks located proximad of the pterostigma.  Larvae of Sisyra fuscata F. are entirely aquatic, sucking the body fluids of fresh-water sponges.  The larvae are distinguished by having paired, jointed gills on the venter of the first 7 abdominal segments.  Eggs are laid in clusters that are covered with a web of white silk, on foliage or other objects hanging over the water (Clausen 1940/62). 

 

 

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             Stilbopterygidae. -- <Habits>; <Adults>   Indecisive group related to the Myrmeleontidae.

 

 

 

 

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             Sympherobiidae. -- <Habits>; <Adults>   Indecisive group of Neuroptera.

 

 

 

 

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             Trichomatidae. -- <Habits>; <Adults>   Generally considered a subfamily of Berothidae

 

 

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Details of Insect Taxonomic Groups

 

          Examples of beneficial species occur in almost every insect order, and considerable information on morphology and habits has been assembled.  Therefore, the principal groups of insect parasitoids and predators provide details that refer to the entire class Insecta.  These details are available at <taxnames.htm>.

 

 

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References

 

 

 

 

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