The

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pa ...

order Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...

Neuroptera, or net-winged insects, includes the lacewings, mantidflies,

antlion The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the larvae ...

s, and their relatives. The order consists of some 6,000

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriat ...

. Neuroptera can be grouped together with the

Megaloptera Megaloptera is an order of insects. It contains the alderflies, dobsonflies and fishflies, and there are about 300 known species. The order's name comes from Ancient Greek, from ''mega-'' (μέγα-) "large" + ''pteryx'' (πτέρυξ) "win ...

and

Raphidioptera Snakeflies are a group of predatory insects comprising the order Raphidioptera with two extant families: Raphidiidae and Inocelliidae, consisting of roughly 260 species. In the past, the group had a much wider distribution than it does now; s ...

in the unranked taxon

Neuropterida The Neuropterida are a clade, sometimes placed at superorder level, of holometabolous insects with over 5,700 described species, containing the orders Neuroptera (lacewings, antlions), Megaloptera (alderflies, dobsonflies), and Raphidiopter ...

(once known as Planipennia) including: alderflies, fishflies, dobsonflies, and

snakeflies Snakeflies are a group of predatory insects comprising the order Raphidioptera with two extant families: Raphidiidae and Inocelliidae, consisting of roughly 260 species. In the past, the group had a much wider distribution than it does now; sn ...

. Adult Neuropterans have four membranous wings, all about the same size, with many

vein Veins are blood vessels in humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are the pulmonary and umbilical veins, both of which carry oxygenated ...

s. They have chewing mouthparts, and undergo complete metamorphosis. Neuropterans first appeared during the

Permian The Permian ( ) is a geologic period and stratigraphic system which spans 47 million years from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.9 Mya. It is the last period of the Paleo ...

period, and continued to diversify through the Mesozoic era. During this time, several unusually large forms evolved, especially in the extinct

family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...

Kalligrammatidae, often called "the butterflies of the

Jurassic The Jurassic ( ) is a geologic period and stratigraphic system that spanned from the end of the Triassic Period million years ago (Mya) to the beginning of the Cretaceous Period, approximately Mya. The Jurassic constitutes the middle period of ...

" for their large, patterned wings.

Anatomy and biology

Neuropterans are soft-bodied insects with relatively few specialized 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 Endopterygota (from Ancient Greek ''endon'' 'inner' + ''pterón'' 'wing' + New Latin ''-ota'' 'having'), also known as Holometabola, is a superorder of insects within the infraclass Neoptera that go through distinctive larval, pupal, and adult st ...

insect groups. They have four wings, usually similar in size and shape, and 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. 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 The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the to ...

often has adhesive discs on the last two segments.

Life cycle and ecology

The larvae of most families are

predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill t ...

s. Many chrysopids, hemerobids and coniopterygids eat aphids and other pest insects, and some 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 Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...

, 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 Ithonidae, commonly called moth lacewings and giant lacewings, is a small family of winged insects of the insect order Neuroptera. The family contains a total of ten living genera, and over a dozen extinct genera described from fossils. The mod ...

are root feeders, and larvae of Sisyridae are aquatic, and feed on freshwater

sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throu ...

s. A few mantispids are

parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson h ...

s of

spider Spiders (order Araneae) are air-breathing arthropods that have eight legs, chelicerae with fangs generally able to inject venom, and spinnerets that extrude silk. They are the largest order of arachnids and rank seventh in total species ...

egg sacs. As in other holometabolic orders, the

pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in thei ...

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

Beetle Beetles are insects that form the order Coleoptera (), in the superorder Endopterygota. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 400,000 describ ...

wasp A wasp is any insect of the narrow-waisted suborder Apocrita of the order Hymenoptera which is neither a bee nor an ant; this excludes the broad-waisted sawflies (Symphyta), which look somewhat like wasps, but are in a separate suborder ...

s, and some lake flies parasitize neuropteran larvae. Yang 2014 Sophogramma lii

Palaeopsychops marringerae Holotype SR 97-08-05 v2

Evolution

Neuropterans first appeared near the end of the

period, as shown by fossils of the Permithonidae from the Tunguska basin in Siberia and a similar fauna from Australia. The osmylids are of

or Early Cretaceous origin and may be the most ancient of the Neuropteran groups. The extinct osmylid '' Protosmylus'' is fossilized in middle Eocene

Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than ...

. The genus '' Burmaleon'' is described from two fossils of

Cenomanian The Cenomanian is, in the ICS' geological timescale, the oldest or earliest age of the Late Cretaceous Epoch or the lowest stage of the Upper Cretaceous Series. An age is a unit of geochronology; it is a unit of time; the stage is a unit in ...

age Burmese amber, implying crown group radiation in the Early Cretaceous or earlier. The family Kalligrammatidae lived from the Jurassic to Aptian (Lower Cretaceous) periods. Ithonidae are from the

to Recent, and the extinct lineages of the family were widespread geographically.

Phylogeny

Molecular analysis in 2018 using mitochondrial rRNA and mitogenomic data places the Megaloptera as sister to Neuroptera, and Raphidioptera as sister to this combined lineage, though these results were considered tentative. The

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...

record has contributed to the understanding of the group's phylogeny. Relationships within the Myrmeleontiformia are still in flux.

Taxonomy

Review of the Neropterid group orders by Engel, Winterton, and Breitkreuz (2018) included grouping of the Neuropteran families in a nested set of clades, an abandonment of the paraphyletic suborder "

Hemerobiiformia The Hemerobiiformia are a suborder of insects in the order Neuroptera. The phylogeny of the Neuroptera was explored in 2014 using mitochondrial DNA sequences. The results indicate that the traditional Hemerobiiformia are paraphyletic, meaning ...

" and redefinition of Myrmeleontiformia as a clade. Neuroptera *Superfamily

Coniopterygoidea Coniopterygoidea is a superfamily in the lacewing order Neuroptera which is considered the basal most linage in the order and sister to all other neuropteran clades. The superfamily includes the single living family Coniopterygidae (dustywings) ...

** family

Coniopterygidae The dustywings, Coniopterygidae, are a family of Pterygota (winged insects) of the net-winged insect order ( Neuroptera). About 460 living species are known.Engel & Grimaldi (2007) These tiny insects can usually be determined to genus with a ha ...

dustywings (Late Jurassic-Present) * Clade Euneuroptera **Superfamily

Osmyloidea Osmyloidea is a euneuropteran superfamily in the lacewing order Neuroptera sister to the superfamilies Dilaroidea, Mantispoidea, and the clade Neoneuroptera. The superfamily includes three living families and two extinct families described fr ...

*** Family

Osmylidae Osmylidae are a small family of winged insects of the net-winged insect order Neuroptera. The osmylids, also called lance lacewings, stream lacewings or giant lacewings, are found all over the world. There are around 225 extant species. Descrip ...

: osmylids (Early Jurassic-Present) *** Family Sisyridae: spongillaflies (Late Cretaceous-Present) *** Family Nevrorthidae (Late Cretaceous-Present) *** Family † Archeosmylidae (Permian-Triassic) *** Family † Saucrosmylidae (Middle Jurassic) **Superfamily

Dilaroidea Dilaridae is a family of Euneuropteran insects in the order Neuroptera, known as "pleasing lacewings". They were formerly placed in the paraphyletic superfamily Hemerobioidea, though the group is currently placed in the monophyletic superfamily ...

*** Family Dilaridae: pleasing lacewings (Late Cretaceous-Present) ** Superfamily

Mantispoidea Mantispoidea is a superfamily of euneuropteran insects in the order Neuroptera. The group was formerly placed in the paraphyletic suborder Hemerobiiformia, but is now considered sister to the superfamilies Dilaroidea and Osmyloidea. Mantisp ...

*** Family

Berothidae The Berothidae are a family of winged insects of the order Neuroptera. They are known commonly as the beaded lacewings. The family was first named by Anton Handlirsch in 1906. The family consists of 24 genera and 110 living species distributed ...

: beaded lacewings (Late Jurassic-Present) *** Family Mantispidae: mantidflies (including †

Dipteromantispidae Dipteromantispidae is an extinct family of neuropterans known from the Cretaceous period. Unlike other neuropterans, the family possesses only a single set of fully developed forewings, with the hindwings reduced to haltere like structures. The ...

) (Jurassic-Present) *** Family † Mesoberothidae (including † Mesithonidae) (Triassic) *** Family

Rhachiberothidae Rhachiberothidae, sometimes called thorny lacewings, are a family of winged insects in the order Neuroptera. The family has only 14 extant species in 4 genera found in Sub-Saharan Africa, but has a diverse fossil record extending back to the ...

: thorny lacewings (Early Cretaceous-Recent) ** Clade Neoneuroptera *** Superfamily Hemerobioidea (inc. Chrysopoidea) **** Family † Ascalochrysidae **** Family Chrysopidae: green lacewings (including † Mesochrysopidae) **** Family

Hemerobiidae Hemerobiidae is a family of Neuropteran insects commonly known as brown lacewings, comprising about 500 species in 28 genera. Most are yellow to dark brown, but some species are green. They are small; most have forewings 4–10 mm long (so ...

: brown lacewings **** Family † Osmylitidae **** Family † Solenoptilidae *** Clade Geoneuroptera **** Superfamily Ithonioidea ***** Family

: moth lacewings (includes Rapismatidae and Polystoechotidae) **** Clade Myrmeleontiformia ***** Superfamily Myrmeleontoidea (syn Nemopteroidea) ***** Family

Ascalaphidae Ascalaphidae is a family of insects in the order Neuroptera, commonly called owlflies; there are some 450 extant species. They are fast-flying crepuscular or diurnal predators of other flying insects, and have large bulging eyes and strongly kn ...

: owlflies (possibly in Myrmeleontoidea) ***** Family †

Babinskaiidae Babinskaiidae is an extinct family of neuropterans known from the Cretaceous period. They are part of the superfamily Myrmeleontoidea. Their distinguishing characters include: "long filiform antennae, narrowly elongated wings, with features su ...

***** Family

Myrmeleontidae The antlions are a group of about 2,000 species of insect in the neuropteran family Myrmeleontidae. They are known for the predatory habits of their larvae, which mostly dig pits to trap passing ants or other prey. In North America, the la ...

: antlions (includes Palaeoleontidae) ***** Family Nemopteridae: spoonwings etc (formerly in Myrmeleontoidea) ***** Family

Nymphidae Nymphidae, sometimes called split-footed lacewings, are a family of winged insects of the order Neuroptera. There are 35 extant species native to Australia and New Guinea. Nymphidae stand somewhat apart from other living Myrmeleontoidea. The a ...

: split-footed lacewings (includes Myiodactylidae) ***** Family † Rafaelianidae **** Superfamily Psychopsoidea ***** Family † Aetheogrammatidae ***** Family † Kalligrammatidae ***** Family † Osmylopsychopidae ( syn †Brongniartiellidae) ***** Family † Panfiloviidae (syn † Grammosmylidae) ***** Family † Prohemerobiidae ***** Family Psychopsidae: silky lacewings The fossil genus †'' Mesohemerobius'' from the Late Jurassic-Early Cretaceous of China has been treated as '' incertae sedis'' within Neuroptera, while the fossil families † Permoberothidae and † Permithonidae are treated as a sister group to clade Eidoneuroptera formed by Neuroptera + Megaloptera.

In human culture

The use of Neuroptera in biological control of insect pests has been investigated, showing that it is difficult to establish and maintain populations in fields of crops. Five species of Neuroptera are among 1681 insect species eaten by humans worldwide. The New Guinea Highland people claim to be able to maintain a muscular build and great stamina despite their low protein intake as a result of eating insects including Neuroptera.

Notes

References

External links

Illustrated database of Neuroptera (insects)

A database of Neuroptera related scientific literature

on the

University of Florida The University of Florida (Florida or UF) is a public land-grant research university in Gainesville, Florida. It is a senior member of the State University System of Florida, traces its origins to 1853, and has operated continuously on its ...

Institute of Food and Agricultural Sciences The University of Florida Institute of Food and Agricultural Sciences (UF/IFAS) is a teaching, research and Extension scientific organization focused on agriculture and natural resources. It is a partnership of federal, state, and county governmen ...

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