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Neuroptera

(Order)

Overview

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The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order contains some 6,010 species.1] Traditionally, the group that was once known as Planipennia, with the Neuroptera at that time also including alderflies, fishflies, dobsonflies and snakeflies, but these are now generally considered to be separate orders (the Megaloptera and Raphidioptera). Sometimes the name Neuropterida[Note 1] is used to refer to these three orders as a group. This is either placed at superorder rank, with the Endopterygota becoming an unranked clade above it, or the Endopterygota are maintained as a superorder, with an unranked Neuropterida being a part of them. Within the endopterygotes, the closest living relatives of the neuropteridan clade are the beetles. The common name lacewings is often used for the most widely known net-winged insects - the green lacewings (Chrysopidae) - but actually most members of the Neuroptera are referred to as some sort of "lacewing".

The adults of this order possess four membranous wings, with the forewings and hindwings about the same size, and with many veins. They have chewing mouthparts, and undergo complete metamorphosis.

Neuropterans first appeared during the Permian Period, and continued to diversify through the Mesozoic Era.[2] During this time several unusually large forms evolved, especially in the extinct family , often referred to as "the butterflies of the Jurassic" due to their large, patterned wings.[3]

Anatomy and biology

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Neuropterans are soft-bodied insects with relativel y few specialised features. They have large lateral compound eyes, and may or may not also have ocelli. Their mouthparts have strong mandibles suitable for chewing, and lack the various adaptations found in most other endopterygote insect groups.

They have four wings, which are usually similar in size and shape, have a generalised pattern of veins. Some Neuropterans have specialised sense organs in their wings, or have bristles or other structures to link their wings together during flight.[4]

The larvae are specialised predators, with elongated mandibles adapted for piercing and sucking. The larval body form varies between different families, depending on the nature of their prey. In general, however, they have three pairs of thoracic legs, each ending in two claws. The abdomen often has adhesive discs on the last two segments.[4]

Life cycle and ecology

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The larvae of most families are predators. Many chrysopids eat aphids and other pest insects, and have been used for biological control (either from commercial distributors but also abundant and widespread in nature). Larvae in various families cover themselves in debris (sometimes including dead prey insects) as camouflage, taken to an extreme in the ant lions, which bury themselves completely out of sight and ambush prey from "pits" in the soil. Larvae of some Ithonidae are root feeders, and larvae of Sisyridae are aquatic, and feed on freshwater sponges. A few mantispids are parasites of spider egg sacs.

As in other holometabolic orders, there is a pupal stage, generally enclosed in some form of cocoon composed of silk and soil or other debris. The pupa eventually cuts its way out of the cocoon with its mandibles, and may even move about for a short while before undergoing the moult to the adult form.[4]

Adults of many groups are also predatory, but some do not feed, or consume only nectar.

Taxonomy and systematics

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The understanding of neuropteran phylogeny has vastly improved since the mid-1990s, not the least courtesy of the ever-growing fossil record.[1] In 1995, for example, it was simply known that the Megaloptera and Raphidioptera were not part of the Neuroptera in the strict sense, and the Mantispoidea and part of the Myrmeleontoidea were the only groups that could be confirmed by cladistic analysis.[5] Though the relationships of some families remain to be fully understood, most major lineages of Neuropterida can nowadays be robustly placed in an evolutionary context.[Note 2]

A 49-million-year-old fossil wing of Palaeopsychops marringerae (Polystoechotidae), showing color pattern
Chrysoperla carnea (Chrysopidae)
Drepanepteryx phalaenoides (Hemerobiidae)
Ditaxis biseriata (Mantispidae)
Kalligramma haeckeli fossil (Kalligrammatidae)

Apart from a few groups that are quite basal or of uncertain position, the net-winged insects can be divided into two suborders, the Myrmeleontiformia and the Hemerobiiformia. The primitive Nevrorthidae, the most ancient group of living neuropterans, are sometimes considered a third suborder Nevrorthiformia or included in the Hemerobiiformia and more specifically in the Osmyloidea. But actually they are better considered a very basal lineage.[6]

Basal and unresolved forms

Suborder Hemerobiiformia

Suborder Myrmeleontiformia

Footnotes

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  1. ^ Also called "Neuropteroidea", though the ending "-oidea" is normally used for superfamilies.
  2. ^ See references in Haaramo (2008).[6]
  3. ^ "Neurorthidae" is a lapsus.
  4. ^ Not in Haaramo (2008), but this taxon, for some time considered obsolete following questions about the affiliations of the Osmylopsychopidae, warrants acceptance as the three families therein form a clade.
two claws. The abdo men often has adhesive discs on the last two segments.[4]

Life cycle and ecology

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The larvae of most families are predators. Many chrysopids eat aphids and other pest insects, and have been used for biological control (either from commercial distributors but also abundant and widespread in nature). Larvae in various families cover themselves in debris (sometimes including dead prey insects) as camouflage, taken to an extreme in the ant lions, which bury themselves completely out of sight and ambush prey from "pits" in the soil. Larvae of some Ithonidae are root feeders, and larvae of Sisyridae are aquatic, and feed on freshwater sponges. A few mantispids are parasites of spider egg sacs.

As in other holometabolic orders, there is a pupal stage, generally enclosed in some form of cocoon composed of silk and soil or other debris. The pupa eventually cuts its way out of the cocoon with its mandibles, and may even move about for a short while before undergoing the moult to the adult form.[4]

Adults of many groups are also predatory, but some do not feed, or consume only nectar.

Taxonomy and systematics

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The understanding of neuropteran phylogeny has vastly improved since the mid-1990s, not the least courtesy of the ever-growing fossil record.[1] In 1995, for example, it was simply known that the Megaloptera and Raphidioptera were not part of the Neuroptera in the strict sense, and the Mantispoidea and part of the Myrmeleontoidea were the only groups that could be confirmed by cladistic analysis.[5] Though the relationships of some families remain to be fully understood, most major lineages of Neuropteri da can nowadays be robustly placed in an evolutionary context.[Note 2]

A 49-million-year-old fossil wing of Palaeopsychops marringerae (Polystoechotidae), showing color pattern
Chrysoperla carnea (Chrysopidae)
Drepanepteryx phalaenoides (Hemerobiidae)
Ditaxis biseriata (Mantispidae)
Kalligramma haeckeli fossil (Kalligrammatidae)

Apart from a few groups that are quite basal or of uncertain position, the net-winged insects can be divided into two suborders, the Myrmeleontiformia and the Hemerobiiformia. The primitive Nevrorthidae, the most ancient group of living neuropterans, are sometimes considered a third suborder Nevrorthiformia or included in the Hemerobiiformia and more specifically in the Osmyloidea. But actually they are better considered a very basal lineage.[6]

Basal and unresolved forms

Suborder Hemerobiiformia

Suborder Myrmeleontiformia

Footnotes

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  1. ^ Also called "Neuropteroidea", though the ending "-oidea" is normally used for superfamilies.
  2. ^ See references in Haaramo (2008).[6]
  3. ^ "Neurorthidae" is a lapsus.
  4. ^ Not in Haaramo (2008), but this taxon, for some time considered obsolete following questions about the affiliations of the Osmylopsychopidae, warrants acceptance as the three families therein form a clade.

References

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  1. ^ a b David Grimaldi & Michael S. Engel (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 
  2. ^ A. G. Ponomarenko & D. E. Shcherbakov (2004). "New lacewings (Neuroptera) from the terminal Permian and basal Triassic of Siberia" (PDF). Paleontological Journal 38 (S2): S197?S203. http://www.palaeoentomolog.ru/Publ/PALS197.pdf. 
  3. ^ Michael S. Engel (2005). "A remarkable kalligrammatid lacewing from the Upper Jurassic of Kazakhstan (Neuroptera: Kalligrammatidae)". Transactions of the Kansas Academy of Science 108 (1): 59?62. doi:10.1660/0022-8443(2005)108[0059:ARKLFT]2.0.CO;2. 
  4. ^ a b c Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. pp. 447?450. ISBN 0-19-510033-6. 
  5. ^ John D. Oswald (1995). "Neuroptera. Lacewings, antlions, owlflies, etc.". Tree of Life Web Project. http://tolweb.org/Neuroptera/8220/1995.01.01. Retrieved September 2, 2010. 
  6. ^ a b Mikko Haaramo (March 11, 2008). "Neuroptera". Mikko's Phylogeny Archive. http://www.helsinki.fi/~mhaaramo/metazoa/protostoma/arthropoda/insecta/neuropterida/neuroptera.html. Retrieved April 27, 2008. 

External links

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Taxonomy

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The Order Neuroptera is further organized into finer groupings including:

Families

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Ascalaphidae

Owlflies are dragonfly-like insects with large bulging eyes and long knobbed antennae. They are neuropterans in the family Ascalaphidae; they are only distantly related to the true flies, and even more distant from the dragonflies and damselflies. They are diurnal or crepuscular predators of other flying insects, and are typically 5 cm (2.0 in) long. [more]

Babinskaiidae

[more]

Berothidae

The beaded lacewings, Berothidae, are a family of winged insects of the order Neuroptera. The family was first named by Anton Handlirsch in 1906. [more]

Brongniartiellidae

[more]

Chrysopidae

Green lacewings are insects in the large family Chrysopidae of the order Neuroptera. There are about 85 genera and (differing between sources) 1,300?2,000 species in this widespread group. Members of the genera Chrysopa and Chrysoperla are very common in North America and Europe; they are very similar and many of their species have been moved from one genus to the other time and again, and in the non-scientific literature assignment to Chrysopa and Chrysoperla can rarely be relied upon. Since they are the most familiar neuropterids to many people, they are often simply called "lacewings". But actually most of the diversity of Neuroptera are properly referred to as some sort of "lacewing", so common lacewings is preferable. [more]

Coniopterygidae

The dustywings, Coniopterygidae, are a family of Pterygota (winged insects) of the net-winged insect order (Neuroptera). About 460 living species are known. These tiny insects can usually be determined to genus with a hand lens according to their wing venation, but to distinguish species, examination of the genitals by microscope is usually necessary. [more]

Dilaridae

The pleasing lacewings, Dilaridae, are a family of winged insects of the order Neuroptera. [more]

Grammosmylidae

[more]

Hemerobiidae

Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings. These insects differ from the somewhat similar Chrysopidae (green lacewings) not only by the usual coloring but by the wing venation: hemerobiids having numerous long veins lacking in chrysopids. Some of the costal cross veins are forked, unlike in green lacewings. Imagines of subfamily mimic dead leaves. Hemerobiid larvae are usually less hairy than chrysopid larvae. [more]

Ithonidae

The Ithonidae or moth lacewings are a small family of winged insects of the insect order Neuroptera. This family contains a total of nine genera, seven living and two extinct. The modern moth lacewings have a notably disjunct distribution while the extinct genera had a more global range. The family is considered one of the most primitive living neuropteran families. Along with the giant lacewings, moth lacewings may be phytophagous. [more]

Kalligrammatidae

[more]

Mantispidae

Mantispidae is a family of small to moderate-sized net-winged insects, known as mantidflies, mantispids, mantid lacewings or mantis-flies. There are many genera with around 400 species worldwide, especially in the tropics and subtropics. Only 5 species of occur in Europe. [more]

Mesithonidae

[more]

Myrmeleonidae

[more]

Myrmeleontidae

Antlions are a group of insects in the family Myrmeleontidae (sometimes misspelled as "Myrmeleonidae"). The most well-known genus is . There are about 2,000 species. Strictly speaking, the term "antlion" applies to the larval form of the members of this family, but while several languages have their own terms for the adult, there is no widely used word for them in English. Very rarely, the adults are called "antlion lacewings". [more]

Nemopteridae

[more]

Nevrorthidae

[more]

Nymphidae

[more]

Nymphitidae

[more]

Osmylidae

Osmylidae are a small family of winged insects of the net-winged insect order Neuroptera. The osmylids are found all over the world except in North America. A common species through most of Europe is . [more]

Osmylitidae

[more]

Osmylopsychopidae

[more]

Permithonidae

[more]

Polystoechotidae

The Polystoechotidae or giant lacewings are a small family of winged insects of the insect order Neuroptera. This family contains three living and seven extinct genera. The modern giant lacewings have a notably disjunct distribution while the extinct genera had a more global range. The family is considered one of the most primitive living neuropteran families. Along with the moth lacewings, giant lacewings may be phytophagous. [more]

Psychopsidae

[more]

Rhachiberothidae

[more]

Sisyridae

The Sisyridae, commonly known as spongeflies or spongillaflies, is a family of winged insects of the order Neuroptera. Approximately 60 living species are known. [more]

Solenoptilidae

[more]

Stilbopterygidae

[more]

More info about the Family Stilbopterygidae may be found here.

References

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  1. ^ a b David Grimaldi & Michael S. Engel (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5. 
  2. ^ A. G. Ponomarenko & D. E. Shcherbakov (2004). "New lacewings (Neuroptera) from the terminal Permian and basal Triassic of Siberia" (PDF). Paleontological Journal 38 (S2): S197?S203. http://www.palaeoentomolog.ru/Publ/PALS197.pdf. 
  3. ^ Michael S . Engel (2005). "A remarkable kalligrammatid lacewing from the Upper Jurassic of Kazakhstan (Neuroptera: Kalligrammatidae)". Transactions of the Kansas Academy of Science 108 (1): 59?62. doi:10.1660/0022-8443(2005)108[0059:ARKLFT]2.0.CO;2. 
  4. ^ a b c Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. pp. 447?450. ISBN 0-19-510033-6. 
  5. ^ John D. Oswald (1995). "Neuroptera. Lacewings, antlions, owlflies, etc.". Tree of Life Web Project. http://tolweb.org/Neuroptera/8220/1995.01.01. Retrieved September 2, 2010. 
  6. ^ a b Mikko Haaramo (March 11, 2008). "Neuroptera". Mikko's Phylogeny Archive. http://www.helsinki.fi/~mhaaramo/metazoa/protostoma/arthropoda/insecta/neuropterida/neuroptera.html. Retrieved April 27, 2008. 

Footnotes

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  1. ^ Also called "Neuropteroidea", though the ending "-oidea" is normally used for superfamilies
  2. ^ Ponomarenko & Shcherbakov (2004)
  3. ^ Engel (2005)
  4. ^ a b c Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed.. Oxford University Press. pp. 447-450. ISBN 0-19-510033-6. 
  5. ^ Grimaldi & Engel (2005)
  6. ^ Oswald (1995)
  7. ^ a b See references in Haaramo (2008)
  8. ^ "Neurorthidae" is a lapsus
  9. ^ Not in Haaramo (2008), but this taxon, for some time considered obsolete following questions about the affiliations of the Osmylopsychopidae, warrants acceptance as the three families therein form a clade.

Sources

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Last Revised: August 24, 2012
2012/08/24 13:13:04