Neuroptera

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Neuroptera
Temporal range: 299–0  Ma
O
S
D
C
P
T
J
K
Pg
N
Permian to recent
Crisopid July 2013-9a.jpg
Green lacewing
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Clade: Neuropterida
Order: Neuroptera
Linnaeus, 1758
clades

See Taxonomy

The insect order Neuroptera, or net-winged insects, includes the lacewings, mantisflies, antlions, and their relatives. The order consists of some 6,000 species. [1] Neuroptera is grouped together with the Megaloptera (alderflies, fishflies, and dobsonflies) and Raphidioptera (snakeflies) in the unranked taxon Neuropterida (once known as Planipennia).

Contents

Adult neuropterans have four membranous wings, all about the same size, 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 Kalligrammatidae, often called "the butterflies of the Jurassic" for their large, patterned wings. [3]

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 holometabolan insect groups.

They have four wings, usually similar in size and shape, and a generalised pattern of veins. [4] Some neuropterans have specialised sense organs in their wings, or have bristles or other structures to link their wings together during flight. [5]

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. [5]

Life cycle and ecology

Larva of Osmylus fulvicephalus, Osmylidae OsmylusLarva.jpg
Larva of Osmylus fulvicephalus , Osmylidae
Larva of Sisyra sp., Sisyridae SisyridaeLarva.jpg
Larva of Sisyra sp., Sisyridae

The larvae of most families are predators. Many chrysopids, hemerobiids 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). [6] [7]

Larvae in various families cover themselves in debris (including other insects, living and dead [8] ) 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, the pupal 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. [5]

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

Beetles, wasps, and some lake flies parasitize neuropteran larvae.

Lifecycle of lacewings LacewingsLyd.png
Lifecycle of lacewings

Evolution

One of the "butterflies of the Jurassic", Sophogramma lii (Kalligrammatidae) Yang 2014 Sophogramma lii.png
One of the "butterflies of the Jurassic", Sophogramma lii (Kalligrammatidae)

Neuropterans first appeared near the end of the Permian period, as shown by fossils of the Permithonidae from the Tunguska basin in Siberia and a similar fauna from Australia. [2]

Patterned wing of Paleogene (49 mya) fossil Palaeopsychops marringerae (Ithonidae) Palaeopsychops marringerae Holotype SR 97-08-05 v2.jpg
Patterned wing of Paleogene (49 mya) fossil Palaeopsychops marringerae (Ithonidae)

The osmylids are of Jurassic or Early Cretaceous origin and may be the most ancient of the Neuropteran groups. [9] The extinct osmylid Protosmylus is fossilized in middle Eocene Baltic amber. [10] The genus Burmaleon is described from two fossils of Cenomanian age Burmese amber, implying crown group radiation in the Early Cretaceous or earlier. [11] [12] The family Kalligrammatidae lived from the Jurassic to Aptian (Lower Cretaceous) periods. [13]

Ithonidae are from the Jurassic to Recent, and the extinct lineages of the family were widespread geographically. [14]

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. [15] [9] The fossil record has contributed to the understanding of the group's phylogeny. [1] [16] [17] [18] Relationships within the Myrmeleontiformia are still in flux. [19]


A phylogenomic analysis published in 2023 confirmed the topology of the neuropterid orders and found the relationships between the families of Neuropterida as shown in the following phylogenetic tree. [20]

Neuropterida

Raphidioptera (snakeflies) Snakefly R. confinis%3F (cropped).jpg

Megaloptera (alderflies and allies) Schlammfliege Sialis sp 5325.jpg

Neuroptera
Coniopterygoidea

Coniopterygidae (dustywings)

Euneuroptera
Osmyloidea

Osmylidae (giant lacewings) Oedosmylus sp crop.jpg

Nevrorthidae

Sisyridae (spongillaflies)

Dilaroidea

Dilaridae (pleasing lacewings)

Chrysopoidea

Chrysopidae (green lacewings) Chrysoperla carnea Guldoeje.jpg

Mantispoidea

Rhachiberothidae (thorny lacewings)

Mantispidae (mantidflies) Mantispidae fg1.jpg

Symphrasinae

Berothidae (beaded lacewings)

Neoneuroptera

Hemerobiidae (brown lacewings) Micromus variegatus01.jpg

Geoneuroptera
Ithonoidea

Ithonidae (moth lacewings) Rapisma sp- India crop.jpg

Myrmeleontiformia
Psychopsoidea

Psychopsidae (silky lacewings)

Silky Lacewing (6769953805).jpg

Myrmeleontoidea

Nymphidae (split-footed lacewings)

Nymphes myrmeleonoides (3155078680) crop.jpg

Nemopteridae (spoonwings)

Nemoptera sp. MHNT.ZOO.2004.0.736.jpg

Myrmeleontidae (antlions)

Distoleon tetragrammicus01.jpg

Ascalaphidae (owlflies)

Libelloides coccajus 210505.jpg


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" and redefinition of Myrmeleontiformia as a clade. [21]

Neuroptera

The fossil genus † Mesohemerobius Ping, 1928 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. [21]

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. [23]

Five species of Neuroptera are among 1681 insect species eaten by humans worldwide. [24]

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. [25]

Notes

  1. "Neurorthidae" is a lapsus .

Related Research Articles

<span class="mw-page-title-main">Chrysopidae</span> Family of insects

Green lacewings are insects in the large family Chrysopidae of the order Neuroptera. There are about 85 genera and 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 nonscientific literature assignment to Chrysopa and Chrysoperla can rarely be relied upon. Since they are the most familiar neuropterans to many people, they are often simply called "lacewings". Since most of the diversity of Neuroptera are properly referred to as some sort of "lacewing", common lacewings is preferable.

<span class="mw-page-title-main">Megaloptera</span> Order of insects

Megaloptera is an order of insects. It contains the alderflies, dobsonflies and fishflies, and there are about 300 known species.

<span class="mw-page-title-main">Snakefly</span> Order of insects

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; snakeflies are found in temperate regions worldwide but are absent from the tropics and the Southern Hemisphere. Recognisable representatives of the group first appeared during the Early Jurassic. They are a relict group, having reached their apex of diversity during the Cretaceous before undergoing substantial decline.

<span class="mw-page-title-main">Mantispidae</span> Family of insects

Mantispidae, known commonly as mantidflies, mantispids, mantid lacewings, mantisflies or mantis-flies, is a family of small to moderate-sized insects in the order Neuroptera. There are many genera with around 400 species worldwide, especially in the tropics and subtropics. Only five species of Mantispa occur in Europe. As their names suggest, members of the group possess raptorial forelimbs similar to those of the praying mantis, a case of convergent evolution.

<span class="mw-page-title-main">Owlfly</span> Family of insects

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 knobbed antennae. The larvae are ambush predators; some of them make use of self-decoration camouflage.

<span class="mw-page-title-main">Osmylidae</span> Family of insects

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 except North and Central America. There are around 225 extant species.

<span class="mw-page-title-main">Hemerobiidae</span> Family of insects

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. These insects differ from the somewhat similar Chrysopidae not only by the usual coloring but also by the wing venation: hemerobiids differ from chrysopids in having numerous long veins and forked costal cross veins. Some genera are widespread, but most are restricted to a single biogeographical realm. Some species have reduced wings to the degree that they are flightless. Imagines (adults) of subfamily Drepanepteryginae mimic dead leaves. Hemerobiid larvae are usually less hairy than chrysopid larvae.

<span class="mw-page-title-main">Neuropterida</span> Clade of insects

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

<span class="mw-page-title-main">Berothidae</span> Family of insects

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 discontinuously worldwide, mostly in tropical and subtropical regions. Numerous extinct species have also been described. Their ecology is poorly known, but in the species where larval stages have been documented, the larvae are predators of termites.

<span class="mw-page-title-main">Nevrorthidae</span> Family of insects

The Nevrorthidae are a small family of lacewings in the order Neuroptera. There are 19 extant species in four genera, with a geographically disjunct distribution: Nevrorthus, comprising 5 species with scattered distributions around the Mediterranean; Austroneurorthus, with two species known from southeastern Australia; Nipponeurorthus, comprising 11 species known from China and Japan; and Sinoneurorthus, known from a single species described from Yunnan Province, China. They are traditionally placed in the Osmyloidea, alongside Osmylidae and the spongillaflies (Sisyridae), but some research has considered them to be the sister group to the rest of Neuroptera. The larvae have unique straight jaws that are curved at the tips, and live as unspecialised predators in the sandy bottom sediments of clear, fast flowing mountain rivers and streams. They pupate underwater on the underside of stones. The adults are likely predators or feed on honeydew and other sugar-rich fluids.

<span class="mw-page-title-main">Psychopsidae</span> Family of insects

Psychopsidae is a family of winged insects of the order Neuroptera. They are commonly called silky lacewings.

<span class="mw-page-title-main">Sisyridae</span> Family of insects

Sisyridae, commonly known as spongeflies or spongillaflies, are a family of winged insects in the order Neuroptera. There are approximately 60 living species described, and several extinct species identified from the fossil record.

<span class="mw-page-title-main">Coniopterygidae</span> Family of insects

The dustywings, Coniopterygidae, are a family of Pterygota 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.

<span class="mw-page-title-main">Ithonidae</span> Family of insects

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 modern Ithonids 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. The family has been expanded twice, first to include the genus Rapisma, formerly placed in the monotypic family Rapismatidae, and then in 2010 to include the genera that had been placed into the family Polystoechotidae. Both Rapismatidae and Polystoechotidae have been shown to nest into Ithonidae sensu lato. The larvae of ithonids are grub-like, subterranean and likely phytophagous.

<i>Chrysopa</i>

Chrysopa is a genus of green lacewings in the neuropteran family Chrysopidae.

<span class="mw-page-title-main">Osmyloidea</span> Superfamily of insects

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 from the fossil record.

<span class="mw-page-title-main">Myrmeleontoidea</span> Superfamily of insects

Myrmeleontoidea is a neuropteran superfamily in the clade Myrmeleontiformia. Engel, Winteron, and Breitkreuz (2018) included the following families:

<span class="mw-page-title-main">Myrmeleontiformia</span> Suborder of insects

Myrmeleontiformia is an insect clade in the order Neuroptera, and which was historically treated as a suborder. The phylogeny of the Neuroptera has been explored using mitochondrial DNA sequences, and while issues remain for the order as a whole, such as "Hemerobiiformia" being paraphyletic, Myrmeleontiformia is generally agreed to be monophyletic, with one study giving the following cladogram:

<span class="mw-page-title-main">Dipteromantispidae</span> Extinct family of insects

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. They are generally small in size and possess raptorial forelegs. They are considered to belong to Mantispoidea, with an uncertain position within the clade. Some authors have suggested that they represent a subgroup of Mantispidae, and should instead be referred to as the subfamily Dipteromantispinae within that family.

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