Snakefly

Snakeflies are a group of insects comprising the order Raphidioptera, which is divided into two families: Raphidiidae and Inocelliidae consisting of roughly 260 species.[1] Together with the Megaloptera they were formerly placed within the Neuroptera, but now these two are generally regarded as separate orders. Members of this order have been considered living fossils, as the phenotype of a species from the early Jurassic period (140 million years ago) closely resembles modern-day species.[2]

Snakefly
Temporal range: Lower Jurassic–Recent
Dichrostigma flavipes beentree
Female Dichrostigma flavipes
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
(unranked): Endopterygota
Order: Raphidioptera
Handlirsch, 1908
Suborders

Raphidiomorpha

Synonyms
  • Raphidiodea

Anatomy and life cycle

Adult snakeflies are characterized by having an elongate prothorax but no modification of the forelegs (as in Mantispidae). They have strong and relatively unspecialised mouthparts, and large compound eyes. Some species also have ocelli. The females typically have a long ovipositor, which they use to deposit their eggs into crevices in bark or rotting wood. The two pair of dragonfly-like wings are similar in size, with a primitive venation pattern, and a thickened costal margin (or "pterostigma").[3]

The larvae have large heads with projecting mandibles. The head and the first segment of the thorax are sclerotised, but the rest of the body is soft and fleshy. They have three pairs of true legs, but no prolegs. However, they do possess an adhesive organ on the abdomen, which they can use to fasten themselves to vertical surfaces.[3]

There is no set number of instars the larvae will go through, some species can have as many as 10-11. The larval stage usually takes around 2–3 years, but in some species can take as long as 6 years.[1] The final larval instar creates a cell in which the insect pupates. The pupa is fully capable of movement, and often leaves its cell for another location before the adult emerges. All snakeflies require a period of cool temperatures (probably around 0 °C) to induce pupation.[1] Depending on when the snakefly pupates determines the length of pupation, Most species pupate in the spring and can take a few days to 3 weeks. Some species seen in more tropical climates will pupate in the early summer and takes around 3 weeks before reaching adulthood. If the larvae begins pupation in the late summer or early fall it can take up to 10 months for pupation to complete.[1]

Snakeflies are extremely territorial and carnivorous organisms. They have been known to be an important predator to aphids and mites. Pollen has also been found in the guts of these organisms and it is unclear whether they require pollen for part of their lifecycle or if they prefer that as a food source as well. The larvae are also predacious and even though no current studies have been done on diet selection by larvae it is thought that they potentially feed on eggs and larvae of other insect species.[1]

Habitat and Species Distribution

Snakeflies are usually found in temperate coniferous forest. They are distributed widely around the globe: Europe and Asia, and can be found in certain regions of Africa, and western North America and Central America. In Africa they are only found in the mountains north of the sahara desert. In North America they are found west of the rocky mountains, and range from south west Canada all the way to the Mexican-Guatemalan Border which is the farthest south they have been found on the western hemisphere. In the eastern hemisphere, they can be found from Spain to Japan. Many species are found all throughout Europe and Asia with the southern edge of their range in northern Thailand and northern India.[1] Even though there is a large distribution of this insect order, individual species distribution is often very limited and some species are confined to a single mountain range.[2]

It was once thought that larvae were only found in the bark of trees, in part due to females' long ovipositor. Besides tree bark, larvae have also been found in soil, detritus, and around the roots of trees or smaller shrubs.[2] The eggs can absorb the nutrients through the soil or detritus before larvae hatch.

Predators and Parasitism

The main predators for snakeflies are wood foraging birds such as the tree creeper, great-spotted woodpecker, wood warbler, nuthatch, and Dunnock. The collared flycatcher which is a generalist forager, has also been known to feed on snakeflies. These are the only bird species that have been observed feeding on this species, but very little research has been done looking at the predators of snakeflies.[4]

Typically 5-15% of snakefly larvae are parasitized but rates as high as 50% have been observed in some species.[1] The insect order Hymenoptera is the largest group known to parasitize snakeflies; 90-95% of parasitized individuals are infected with Hymenoptera.

Pest Control

This order of insects has been considered a viable option in agriculture use. The main advantages to have this species as a pest control agent is that there are not many known predators to the species, and both adults and larvae are predacious. One of the disadvantages is that snakeflies undergo a long larval period, meaning it could take a long time to completely rid the crops of pests.[2]

Systematics

Distinguishing taxonomic features

The morphological characteristics which distinguish the order Raphidioptera from other insect orders are as follows :[5]

  • prothorax is lengthened giving long neck appearance
  • head has protruding eyes, long antennae and mandibles (chewing mouthparts).
  • has two pairs of identical wings.
  • ten-segmented abdomen without cerci.
  • lengthened ovipositors in females

Taxonomy

The Megaloptera, Neuroptera (in the modern sense) and Raphidioptera are very closely related, forming the group Neuropterida. This is either placed at superorder rank, with the Endopterygota - of which they are part - 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 Neuropterida are the beetles.

There are four extinct families known only from fossils.[6] Almost all known snakeflies belong to the suborder Raphidiomorpha.[6] The exception being the Jurassic family Priscaenigmatidae, placed in suborder Priscaenigmatomorpha.[6]

Extinct snakeflies are known from fossils dating from the Lower Jurassic to the Miocene.[6]

Raphidioptera as grouped according to Engel 2002 with updates according to Bechly and Wolf-Schwenninger, 2011 and Ricardo Pérez-de la Fuente et al (2012):[6][7][8]

Order Raphidioptera

  • Suborder Priscaenigmatomorpha Engel 2002
  • Suborder Raphidiomorpha
    • Family †Baissopteridae Martynova, 1961
      • Genus †Austroraphidia Willmann, 1994 (Lower Cretaceous; Brazil)
      • Genus †Baissoptera Martynova, 1961 (Upper Jurassic-Lower Cretaceous; Brazil, China, Russia)
      • Genus †Cretoraphidia Ponomarenko, 1993 (Upper Jurassic-Lower Cretaceous; Russia)
      • Genus †Cretoraphidiopsis Engel, 2002 (Lower Cretaceous; Mongolia)
      • Genus †Lugala Willmann, 1994 (Lower Cretaceous; Mongolia)
    • Family Inocelliidae Navás
      • Subfamily †Electrinocelliinae Engel, 1995
      • Subfamily Inocelliinae Engel, 1995
        • Genus Amurinocellia Aspöck & Aspöck, 1973 (Recent)
        • Genus Fibla Navás, 1915 (Eocene-Recent; Fossils: Baltic amber, Spain, USA)
        • Genus Indianoinocellia (Recent)
        • Genus Inocellia Schneider, 1843 (Recent)
        • Genus Negha Navas 1916 (Recent)
        • Genus Parainocellia (Recent)
        • Genus Sininocellia (Recent)
        • Genus †Succinofibla Aspöck & Aspöck, 2004 (Eocene; Baltic amber)
    • Family †Metaraphidiidae Bechly and Wolf-Schwenninger, 2011
      • Genus †Metaraphidia Whalley, 1985 (Lower Jurassic; England, Germany)
    • Family †Mesoraphidiidae Martynov, 1925 (= Alloraphidiidae, Huaxiaraphidiidae, Sinoraphidiidae, and Jilinoraphidiidae)
      • Subfamily Alloraphidiinae
        • Genus Alloraphidia Carpenter, 1967 (Upper Jurassic-Middle Cretaceous; Canada, China, Mongolia, Russia)
        • Genus Archeraphidia Ponomarenko, 1988 (Upper Jurassic-Lower Cretaceous; Russia, Mongolia)
        • Genus Pararaphidia Willmann, 1994 (Upper Jurassic-Lower Cretaceous; Russia, Mongolia)
      • Subfamily Mesoraphidiinae
        • Genus Baisoraphidia Ponomarenko, 1993 (Upper Jurassic-Lower Cretaceous; Russia)
        • Genus Cretinocellia Ponomarenko, 1988 (Lower Cretaceous; Mongolia)
        • Genus Huaxiaraphidia Hong, 1992 (Lower Cretaceous; China)
        • Genus Jilinoraphidia Hong and Chang, 1989 (Lower Cretaceous; China)
        • Genus Kezuoraphidia Willmann, 1994 (Lower Cretaceous; China)
        • Genus Mesoraphidia Martynov, 1925 (Upper Jurassic-Upper Cretaceous; China, Kazakhstan, Mongolia, USA)
        • Genus Proraphidia Martynova, 1947 (Upper Jurassic-Lower Cretaceous; England, Kazakhstan, Spain)
        • Genus Siboptera Ponomarenko, 1993 (Upper Jurassic-Lower Cretaceous; China, Russia)
        • Genus Sinoraphidia Hong, 1982 (Upper Jurassic; China)
        • Genus Xuraphidia Hong, 1992 (Lower Cretaceous; China)
        • Genus Yanoraphidia Ren, 1995 (Cretaceous)
        • Tribe Nanoraphidiini
          • Genus Cantabroraphidia Pérez-de la Fuente, Nel, Peñalver & Delclòs, 2010 (Albian; Spain)
          • Genus Grimaldiraphidia Bechly and Wolf-Schwenninger, 2011 (Turonian, New Jersey, USA)
          • Genus Nanoraphidia Engel, 2002 (Albian; Myanmar)
          • Genus Lebanoraphidia Bechly & Wolf-Schwenninger, 2011 (Neocomian, Lebanon)
      • Subfamily Ororaphidiinae
        • Genus Caloraphidia Ren, 1997 (Cretaceous, China)
        • Genus Necroraphidia Pérez-de la Fuente, Peñalver, Delclòs & Engel, 2012 (Albian, Spain)
        • Genus Ororaphidia Engel & Ren 2008 (Middle Jurassic; China)
        • Genus Styporaphidia Engel & Ren 2008 (Middle Jurassic; China)
      • Subfamily "incertae sedis"
        • Genus Alavaraphidia Pérez-de la Fuente, Peñalver, Delclòs & Engel, 2012 (Albian, Spain)
        • Genus Amarantoraphidia Pérez-de la Fuente, Peñalver, Delclòs & Engel, 2012 (Albian, Spain)
        • Genus Iberoraphidia Jepson, Ansorge & Jarzembowski, 2011 (Cretaceous)
    • Family Raphidiidae Latreille
      • Genus Africoraphidia (Recent)
      • Genus Agullla Navas 1914 (Recent)
      • Genus Alena Navas 1916 (Recent)
      • Genus Atlantoraphidia (Recent)
      • Genus Calabroraphidia (Recent)
      • Genus Dichrostigma (Recent)
      • Genus Harraphidia (Recent)
      • Genus Hispanoraphidia (Recent)
      • Genus Iranoraphidia (Recent)
      • Genus Italoraphidia (Recent)
      • Genus Mauroraphidia (Recent)
      • Genus Mongoloraphidia (Recent)
      • Genus Ohmella H. Aspöck & U. Aspöck (Oligocene-Recent)
        Ohmella coffini holotype MNHN part side direct lighting
        Ohmella coffini holotype female, Miocene, France
      • Genus Ornatoraphidia (Recent)
      • Genus Parvoraphidia (Recent)
      • Genus Phaeostigma (Recent)
      • Genus Puncha (Recent)
      • Genus Raphidia Linnaeus, 1758 (Eocene-Recent)
      • Genus Raphidilla (Recent)
      • Genus Subilla (Recent)
      • Genus Tadshikoraphidia (Recent)
      • Genus Tauroraphidia (Recent)
      • Genus Tjederiraphidia (Recent)
      • Genus Turcoraphidia (Recent)
      • Genus Ulrike (Recent)
      • Genus Venustoraphidia (Recent)
      • Genus Xanthostigma (Recent)
    • Family Incertae sedis
      • Genus †Archiinocellia Handlirsch, 1910 (Oligocene; Canada)
      • Genus †Arariperaphidia Martins-Neto & Vulcano, 1989 (Lower Cretaceous; Brazil)
Ohmella coffini holotype MNHN part side direct lighting
Ohmella coffini holotype female, Miocene, France

Footnotes

  1. ^ a b c d e f g Aspock, H (2002). "The Biology of Raphidioptera: A Review of Present Knowledge". Acta Zoologica Academiae Scientiarum Hungaricae.
  2. ^ a b c d Harring, E.; Aspock, H. (2002). "Molecular phylogeny of the Raphidiidae". Systematic Entomology. 36: 16–30. doi:10.1111/j.1365-3113.2010.00542.x.
  3. ^ a b Hoell, H.V., Doyen, J.T. & Purcell, A.H. (1998). Introduction to Insect Biology and Diversity, 2nd ed. Oxford University Press. pp. 445–446. ISBN 0-19-510033-6.CS1 maint: Multiple names: authors list (link)
  4. ^ Szentkiralyi, F.; Kristin, A. (2002). "Lacewings and Snakeflies as Prey for Bird Nestlings in Slovakian Forest Habitats". Acta Zoologica Academiae Scientiarum Hungaricae. 48.
  5. ^ Gillot, C. (1995). "Raphiodioptera". Entomology (2 ed.). pp. 293–295. ISBN 978-0-306-44967-3. Retrieved 14 November 2010.
  6. ^ a b c d e Engel, M.S. (2002). "The Smallest Snakefly(Raphidioptera: Mesoraphidiidae): A New Species in Cretaceous Amber from Myanmar, with a Catalog of Fossil Snakeflies". American Museum Novitates. 3363: 1–22. doi:10.1206/0003-0082(2002)363<0001:TSSRMA>2.0.CO;2. hdl:2246/2852.
  7. ^ Pérez-de la Fuente, R.; Peñalver, E.; Delclòs, X.; Engel, M.S. (2012). "Snakefly diversity in Early Cretaceous amber from Spain (Neuropterida, Raphidioptera)". ZooKeys. 204: 1–40. doi:10.3897/zookeys.204.2740. PMC 3391719. PMID 22787417.
  8. ^ Bechly, G.; Wolf-Schwenninger, K. (2011). "A new fossil genus and species of snakefly (Raphidioptera: Mesoraphidiidae) from Lower Cretaceous Lebanese amber, with a discussion of snakefly phylogeny and fossil history" (PDF). Insect Systematics and Evolution. 42 (2): 221–236. doi:10.1163/187631211X568164. Archived from the original (PDF) on 5 March 2014.

References

Agulla (snakefly)

Agulla is a genus of modern snakeflies in the family Raphidiidae.Agulla species are predatory, both as adults and larvae. They occur in North America in British Columbia and in the Western United States, namely in the Rocky Mountains and westward, including the southwestern deserts.

Agulla nixa

Agulla nixa is a species of snakefly that can be found in wooded areas in Texas and Mexico. Its common name is Texan snakefly. The species is 15-22 mm long and feeds on smaller insects. The female inserts its eggs in bark crevices which is also where the larvae eat.

Alavaraphidia

Alavaraphidia is an extinct genus of snakefly in the family Mesoraphidiidae. The genus is solely known from an Early Cretaceous, Albian age, fossil amber found in Spain. Currently, the genus comprises a single species, Alavaraphidia imperterrita.

Amarantoraphidia

Amarantoraphidia is an extinct genus of snakefly in the family Mesoraphidiidae. The genus is solely known from Early Cretaceous, Albian age, fossil amber found in Spain. Currently the genus comprises only a single species Amarantoraphidia ventolina.

Baissoptera

Baissoptera is an extinct genus of snakefly in the Baissopteridae family which was described by Martynova in 1961. Since 1961, it has been described three times; Carpenter in 1992, Ponomarenko in 1988 and Engel in 2002. According to J. Jepson et al. in 2011, the parent taxon is Baissopteridae. Fossils of the species have been found in Brazil, China, Spain and Russia.

Cantabroraphidia

Cantabroraphidia is an extinct genus of snakefly in the family Mesoraphidiidae. The genus is solely known from fossil amber found in Cantabria, northern Spain, dating to the Albian age of the Early Cretaceous Period. Currently the genus comprises a single species Cantabroraphidia marcanoi.

Fibla carpenteri

Fibla carpenteri is an extinct species of snakefly in the Inocelliidae genus Fibla. F. carpenteri is named in honor of the paleoentomologist Dr Frank Carpenter, for his vast knowledge and interest in Raphidioptera.The species is known from a single specimen, the holotype, deposited in the Harvard University, Museum of Comparative Zoology as specimen #9999. Dr. Michael S. Engel first studied and described the species after finding the specimen in the Harvard collections. He published his type description in the journal Psyche volume 102 published in 1995. Fairly well preserved in Eocene Baltic amber, the female individual has a torn forewing missing the distal portion, partial antennae, and the ovipositor is severed and missing the tip. There are also a number of small areas with "schimmel", a type of white mold sometimes present on arthropods in amber. With a total length, not including ovipositor or antennae, of just over 18 millimetres (0.71 in), Fibla carpenteri is the largest species of snakefly from amber and the largest species of the genus. As a whole the female shows no light color marking and was a fairly uniform dark brown to black coloration. The wings are hyaline with brown coloration of the vein structure and are slightly fuscous at the base. The pterostigma is also colored brown.

F. carpenteri is one of only four extinct Fibla which are known from the fossil record. Along with F. erigena F. carpenteri is one of two known from the baltic amber deposits, while F. cerdanica is from the Miocene of Spain and F. exusta is from the Eocene of the Florissant Formation, Colorado.

Florissantoraphidia funerata

Florissantoraphidia funerata is an extinct species of snakefly, originally assigned to the raphidiid genus Raphidia, but subsequently transferred to the genus Florissantoraphidia. The name F. funerata is derived from the Latin funeratus meaning to "bury" or "intern". The species is known from a single female specimen, the holotype, deposited in the Department of Palaeontology at the Natural History Museum in London as specimen number "In. 26922". Though they did not study the specimen, Horst Aspöck, Ulrike Aspöck and Hubert Rausch in the 1991 work Die Raphidiopteren der Erde noted and figured the specimen as an "unidentified raphidiid". Dr. Michael S. Engel first studied and described the species after finding the specimen in the Department of Palaeontology collections. He published his type description in the journal Transactions of the Kansas Academy of Science (Volume 106) in 2003.Florissantoraphidia funerata was found in the Florissant Formation, which has produced seven other species of snakefly, six in Raphidia and one in the Inocelliida genus Fibla. Out of the described snakefly specimens from the Florissant Formation, the F. funerata holotype is the most complete. Preserved as a compression fossil, the female individual is fossilized in a resting position giving a side view to the body and wings. Including the ovipositor, F. funerata is 16.5 millimetres (0.65 in) and has a forewing length of 10 millimetres (0.39 in). Of the known snakefly species, F. funerata is closest in appearance to the extinct F. mortua, which is also known from the Florissant Formation. The two cogeneric species can be separated by several features of the forewing, including a lack of terminal forks in veins along the posterior margin of the wing in F. funerata, and the radial cell bordering the pterostigma narrowing at the base in F. mortua. F. funerata was originally assigned to the genus Raphidia, but this assignment was made with hesitation by Dr. Engel as the characteristics used to separate living genera, the reproductive structures, are not preserved in most fossils and very rarely in compression fossils. Engel considered it is possible the Florrisant Formation species are members of one of the two nearctic genera Alena or Agulla. In a 2014 study Vladimir N. Makarkin and S. Bruce Archibald removed the species F. mortua and F. funerata from the genus Raphidia, and transferred them to a new, separate genus which the authors named Florissantoraphidia.

Iberoraphidia

Iberoraphidia is an extinct genus of snakefly in the family Mesoraphidiidae. The genus is solely known from a Cretaceous, Lower Barremian, fossil found in Spain. Currently the genus is composed of a single species, Iberoraphidia dividua.

Lebanoraphidia

Lebanoraphidia is an extinct genus of snakefly in the family Mesoraphidiidae. The genus is solely known from Cretaceous, Upper Neocomian, fossil amber found in Lebanon. Currently the genus is composed of a single species Lebanoraphidia nana.

Mesoraphidiidae

Mesoraphidiidae is an extinct family of snakeflies in the suborder Raphidiomorpha. The family lived from the Late Jurassic through the Late Cretaceous and is known from twenty-five genera. Mesoraphidiids have been found as both compression fossils and as inclusions in amber. The family was first proposed in 1925 by the Russian paleoentomologist Andrey Vasilyevich Martynov based on Upper Jurassic fossils recovered in Kazakhstan. The family was expanded in 2002 by the synonymizing of several other proposed snakefly families. The family was divided into three subfamilies and one tribe in a 2011 paper, further clarifying the relationships of the included genera.

Nanoraphidia

Nanoraphidia is an extinct genus of snakefly in the family Mesoraphidiidae containing the species Nanoraphidia electroburmica and Nanoraphidia lithographica.The genus name is derived from a combination of the Greek nanos (dwarf) and the snakefly genus Raphidia, and the species name from the Latin electrum (amber) and Burma, the former name of Myanmar. The genus is known from only the holotype, a single, partial adult, now deposited in the American Museum of Natural History as specimen number Bu-092. The amber specimen is from deposits in Tanai Village, Kachin State 105 kilometres (65 mi) northwest of Myitkyina, Myanmar.

The specimen has a badly disarticulated thorax and abdomen, which are present in the amber as integumental debris, however the head, wings, and anterior legs are in good condition. Nanoraphidia electroburmica is the smallest known Raphidioptera species, living or extinct, the fore wings being only 4.2 millimetres (0.17 in) long. The genus is also the first to be described from a specimen of old world amber. Nanoraphidia is separatable from other Mesoraphidiidae genera by the small size of specimens and distinct vein structure of the wings. The wings have an expanded costal area which is similar in appearance to the costal areas in Cretinocellia and Lugala. Both of these genera, however, belong to the Baissopteridae family and have many more crossveins and cells between Rs and M veins. The specimen shows N. electroburmica to have had a brown coloration to the body and a hyaline appearance to the wings.

Necroraphidia

Necroraphidia is an extinct genus of snakefly in the family Mesoraphidiidae. The genus is solely known from Early Cretaceous, Albian age, fossil amber found in Spain. Currently the genus comprises a single species, Necroraphidia arcuata.

Ororaphidia

Ororaphidia is an extinct genus of snakefly containing two species: the type species Ororaphidia megalocephala and Ororaphidia bifurcata.

Puncha

Puncha may refer to:

Puncha (community development block), an administrative division in Purulia Sadar East, West Bengal, India

Puncha, Purulia, a village, with a police station, in Purulia district, West Bengal, India.

Puncha (snakefly), a genus of snakeflies in the family Raphidiidae

Puncha ratzeburgi, a species of snakeflies in the family Raphidiidae

Puncha (snakefly)

Puncha is a genus of snakeflies in the family Raphidiidae. This monotypic genus contains one species:

Puncha ratzeburgiThese snakeflies are widely distributed in Central and Eastern Europe. They are predator, both as adults and larvae.

Raphidia

Raphidia is a genus of snakefly, mainly found in Europe.

Styporaphidia

Styporaphidia is a genus of snakefly, belonging to the extinct family Mesoraphidiidae, containing up to two species, the type species Styporaphidia magia and tentatively Styporaphidia? hispanica. The genus was named from the Greek stypos meaning "stem" or "stump" and Raphidia, the type genus for, and most often used as, a stem for generic names in the order Raphidioptera. The species name of S. magia is from the Greek word mageia meaning "magic" while the species name for S.? hispanica is from the Latin Hispania meaning "Spain" in reference to the type locality of the species.

Insect orders
Extant Raphidioptera families

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