Editing Dobsonfly

{{short description|Subfamily of insects}}
{{Redirect|Corydalinae|the subtribe in the poppy family|Papaveraceae#Fumarioideae}}
{{Automatic taxobox
| image = Corydalus cornutus MHNT.jpg
| image_caption = ''[[Corydalus cornutus]]''
| taxon = Corydalinae
| subdivision_ranks = Genera
| subdivision = See [[#Systematics|Systematics]]
}}

'''Dobsonflies''' are a [[subfamily]] of [[insect]]s, '''Corydalinae''', part of the [[Megaloptera]]n [[family (biology)|family]] [[Corydalidae]]. The [[larva]]e (commonly called '''hellgrammites''') are [[aquatic insect|aquatic]], living in [[stream]]s, and the adults are often found along streams as well. The nine [[genera]] of dobsonflies are distributed in the [[Americas]], [[Asia]], and [[South Africa]].

==Etymology==
The origin of the word "dobsonfly" is unclear. [[John Henry Comstock]] used the term in reference to these insects in his 1897 book ''Insect Life'',<ref name=jhc/> but did not explain it. He also mentioned that [[angling|anglers]] used the word "hellgrammite" for the aquatic larvae they used as bait, but the origin of this term is also unknown.<ref name="purdue"/>

==Description==
Adult dobsonflies are some of the largest non-[[Lepidopteran]] insects of [[temperate zones]] such as the United States and Canada, with a wingspan of up to {{cvt|18|cm|in}} in some species of ''[[Corydalus]]''.<ref name=sl>Stange, Lionel. "Alderflies and Dobsonflies." Encyclopedia of Entomology. 2nd ed. New York: Springer Publishing, 2008.</ref> The Asian ''[[Acanthacorydalis fruhstorferi]]'' can have a wingspan of up to {{cvt|21.6|cm|in}}, making it the largest dobsonfly and the [[List of largest insects|largest aquatic insect]] in the world by this measurement.<ref>{{cite web |url=http://www.guinnessworldrecords.com/world-records/428458-largest-aquatic-insect-pterygota |title=Largest aquatic insect (by wingspan) |website=Guinness World Records |access-date=4 April 2018 }}</ref><ref>{{cite web|url=http://www.cnn.com/2014/07/22/world/asia/giant-insect-china/|title=World's largest aquatic insect specimen found in China|first=Zoe|last=Li|date=July 22, 2014|access-date=June 3, 2016|publisher=CNN}}</ref> The [[insect wings|wings]] vary from a grayish to translucent shade, depending on the species, and the anal region of the hindwing is wide and folded at rest. Despite the large wings, adults are weak, fluttery fliers. The body is soft and coloration varies from yellow to dark shades of brown.<ref name=sl/> The body typically does not surpass {{cvt|7.5|cm|in}} in length,<ref name=sl/> although the largest Asian ''Acanthacorydalis'' may reach {{cvt|10.5|cm|in}}.<ref>{{cite journal| author1=Xing Yue Liu | author2=Ding Yang | author3=Si Qin Ge | author4=Xing Ke Yang | year=2005 | title=Phylogenetic review of the Chinese species of Acanthacorydalis (Megaloptera, Corydalidae) | journal=Zoologica Scripta | volume=34 | issue=4 | pages=373–387 | doi=10.1111/j.1463-6409.2005.00197.x | s2cid=84988447 }}</ref>

Adult males of many—but not all—species are easily recognized by their long, curving [[mandible (insect mouthpart)|mandibles]]. Examples of species with large-mandibled males include the genera ''Acanthacorydalis'', ''Corydalus'' and ''Platyneuromus'', while in ''Neoneuromus'', ''Nevromus'', ''Neurhermes'' and ''Protohermes'' the sexes are similar.<ref name=Liu2015>{{cite journal| author1=Xingyue Liu | author2=Fumio Hayashi | author3=Laura C. Lavine | author4=Ding Yang | year=2015 | title=Is diversification in male reproductive traits driven by evolutionary trade-offs between weapons and nuptial gifts? | journal=Proceedings of the Royal Society B: Biological Sciences | volume=282 | issue=1807 | pages=373–387 | doi=10.1098/rspb.2015.0247 | pmid=25925103 | pmc=4424648 }}</ref> In ''Corydalus cornutus'', a particularly long-mandibled species, these can reach up to {{cvt|4|cm|in}} in length and are used in competition for females.<ref name=sd>Simonsen, T. J., Dombroskie, J. J., and D. D. Lawrie (2008). Behavioral Observations on the Dobsonfly, ''Corydalus cornutus'' (Megaloptera: Corydalidae) with Photographic Evidence of the Use of the Elongate Mandibles in the Male. American Entomologist 64 167-169.</ref> It is possible that the mandibles may have been [[sexual selection|selected]] as [[secondary sex characteristics]] used by females to evaluate males during [[courtship display|courtship]]. Males cannot use these mandibles to bite because they are too long; on the other hand, females have short, heavily [[sclerite|sclerotized]] mandibles which enable them to deliver powerful bites when threatened.<ref name=sl/> Males of many species will also produce [[nuptial gift]]s in the form of packages of nutrient-rich  [[spermatophores]] that are eaten by the female partner after mating. This has been shown to be correlated to mandible size; in species where the males have large mandibles the "nuptial gift" is small or absent, while it is large in species where males lack the exaggerated mandibles. Two genera, ''[[Chloroniella]]'' and ''[[Chloronia]]'', are unusual in that the males lack large mandibles and do not produce "nuptial gifts".<ref name=Liu2015/> The [[antenna (biology)|antennae]] of males are also noticeably elongated, even longer than the mandibles.

[[Corydalinae]] is distinguished from closely related [[clades]] by the following [[synapomorphies]] (with exceptions in a few species): quadrate head with a postocular spine, ridge, and plane, non-pectinate antennae, four crossveins between the radius and the radial sector, and distinctive male terminalia with a well developed ninth [[gonostylus]].<ref name=sl/>

In regards to the larvae, [[entomologist]] [[John Henry Comstock]] wrote in his 1897 book Insect Life,<ref name=jhc>Comstock, John Henry (1897). Insect Life. Cornell University Library. Online.</ref> "In spite of its disagreeable appearance it is in some respects very interesting to students of Nature study." The larvae, commonly called hellgrammites, are perhaps better known than the adults due to their more readily findable nature. They are unusual in that although they are generally aquatic, taking in dissolved [[oxygen]] through abdominal lateral filaments and [[tracheal gills]], they also have [[Spiracle (arthropods)|spiracles]] that allow them to take in air directly when above water.<ref name=cr1997>Contreras-Ramos, Atilano. ''Corydalus''. Tree of Life Web Project. November 15, 1997.</ref>

Larvae of dobsonflies differ from those of their sister clade, the [[alderflies]], in that they bear eight pairs of lateral processes as well as anal [[prolegs]] with a pair of terminal hooks used to hold themselves to substrate, and also in that they lack a [[terminal filament]].<ref name=bt>Borror, Donald; Triplehorn, Charles; and Norman Johnson. An Introduction to the Study of Insects. 6th ed. New York: Harcourt Brace College Publishers, 1989. 358-363.</ref> At the end of the abdomen is a pair of claw-like structures. Body color is black or dark brown.

==Systematics==
There are about sixty species of dobsonflies.<ref name=sl/> Contreras-Ramos <ref name=cr1998>Contreras-Ramos, Atilano (1998). Systematics of the dobsonfly genus ''Corydalus'' Latreille (Megaloptera: Corydalidae). Thomas Say Monographs, Entomological Society of America. Lanham, MD. 360pp.</ref> suggests nine [[genera]] within Corydalinae, divided into four lineages. Working from "most basal" to "most derived" lineages, there are:

* The Protohermes lineage, containing the genera ''[[Neurhermes]]'' and ''[[Protohermes]]'', distributed from [[Northwest India]] to Indonesia, China, and Japan.
* The Chloroniella lineage, containing the monotypic genus ''[[Chloroniella]]'', found only in South Africa.
* The Nevromus lineage, containing the genera ''[[Acanthacorydalis]]'', ''[[Nevromus]]'', and ''[[Neoneuromus]]'', distributed from Northwest India to Southeast Asia.
* The Corydalus lineage, containing the genera ''[[Platyneuromus]]'', ''[[Chloronia]]'', and ''[[Corydalus]]'', distributed from southern Canada down to northern Argentina and south-east Brazil.

==Diet and behavior==
The larvae of dobsonflies live along the rocky bottoms of streams. Chiefly [[nocturnal|active during the night]], they ambush prey in the middle of [[riffles]] which supply plenty of oxygen and stir up prey.<ref name=hf1989>Hayashi, Fumio (1989). Radio Tracking and Activity Monitoring of the Dobsonfly Larva, ''Protohermes grandis'' (Megaloptera: Corydalidae). Oecologia 78 468-472.</ref> They are generalist predators; dissections have revealed that they primarily eat aquatic immatures of [[mayflies]], [[caddisflies]], [[stoneflies]], and [[chironomid]] midges.<ref name=hf1988>Hayashi, Fumio (1988). Prey selection by the dobsonfly larva, ''Protohermes grandis'' (Megaloptera: Corydalidae). Freshwater Biology 20 19-29.</ref> Although the larvae spend most of their lives under rocks below water, locals along Virginia and Pennsylvania rivers have reported emergences, known as "hellgrammite crawlings," during thunderstorms.<ref name=hd>Hall, Donald. Eastern Dobsonfly. Featured Creatures. University of Florida. April 2013.</ref>

The adults are also nocturnal, and are seldom seen as they hide under leaves in the [[canopy (biology)|canopy]] during the daytime. However, they do sometimes form aggregations under bridges or other structures along streams.<ref name=cr1997/> Since the adults live only about a week, they are not known to eat anything, although they have been reported to drink sweet solution in captivity.<ref name=cr1997/>

The dobsonfly may be attracted by [[mercaptan]], an indicator additive in natural gas and propane, and may behave as an [[animal sentinel]] in the presence of these gases.

==Life cycle==
The [[metamorphosis]] from larva to adult in dobsonflies is one of the simplest of the [[holometabolous]] orders,<ref name=sl/> yet the life cycle begins with an intriguing ritualized [[courtship display]] (most of the following comes from Simonsen et al. 2008 <ref name=sd/> and all pertains to ''Corydalus''; other courtship rituals remain unknown). Males compete with each other for females, aggressively fluttering the wings and trying to place their long mandibles underneath the body of the opponent in order to flip him into the air. Afterwards, the male approaches a female from the side and touches her with his antennae. At first the female reacts somewhat aggressively, moving the head from side to side with mandibles wide apart. However, she then allows the male to come closer and place his mandibles over her wings in a perpendicular position, a position he holds until the female signals reception to mating.

At least in ''[[Protohermes]]'', during copulation the male attaches to the female's [[insect genitalia|genitalia]] a large, globular [[spermatophore]] about {{cvt|4|mm|in|frac=8}} long and wide. The spermatophore consists of two parts: a large gelatinous mass, and a smaller seminal duct containing the sperm. After copulation, the female proceeds to spread her legs wide apart, curl the abdomen under the chest, and [[Nuptial gift|eat the gelatinous part of the spermatophore]].<ref name=hf1992>Hayashi, Fumio (1992). Large spermatophore production and consumption in dobsonflies ''Protohermes''. Japanese Journal of Entomology 60 59-66.</ref>

[[Oviposition]] occurs along rocky walls of streams at night, from May to September in ''Corydalus''.<ref name=cr1997/><ref name=mb>Mangan, Brian (1992). Oviposition of the Dobsonfly (''Corydalus cornutus'', Megaloptera) on a Large River. American Midland Naturalist 127 348-354.</ref> The females deposit coin-size [[egg]] masses containing on average one thousand grey, cylindrical eggs, each egg about {{cvt|1.5|mm|in|frac=16}} long and {{cvt|0.5|mm|in|frac=32}} wide. This mass is covered by a layer of a chalky, white substance, which probably protects the eggs from [[desiccation]] and overheating. Females tend to deposit egg masses at relatively few sites, resulting in grouped egg masses.<ref name=mb/>

One to two weeks after oviposition, the eggs hatch and the first [[instar]] larvae either fall directly into the stream or if not, immediately search for water. There the larvae live for up to five years, going through 10-12 instar [[moulting|molts]].<ref name=cr1997/> When they have finally reached maturity, the larvae leave the water and find a rock, log, or anthropogenic debris, typically close to the stream but sometimes up to 40 m away. There they construct a chamber for pupation and spend several days to several weeks as prepupae before shedding the exoskeleton and spending about a week to two weeks as [[pupae]].<ref name=cr1997/><ref>{{Cite journal|last1=Walsh|first1=B. D.|last2=Riley|first2=C. V.|date=2004|title=The Hellgrammite Fly (''Corydalis cornutus'', Linn.)|journal=American Entomologist|language=en|volume=50|issue=1|pages=50–51|doi=10.1093/ae/50.1.50|issn=2155-9902|doi-access=free}}</ref> The pupae are yellow-orange with dark spots on the dorsum of the abdomen, covered in minute setae, and exarate (i.e. the developing appendages and mouthparts are attached only at their proximal ends).<ref name=hd/> Although the males have a small tubercle on the [[prothoracic]] sternum and a slightly wider head than the females, the mandibles are not as noticeably divergent as in the adults.<ref name=mb1994>Mangan, Brian (1994). Pupation Ecology of the Dobsonfly ''Corydalus cornutus'' (Corydalidae: Megaloptera) along a Large River. Journal of Freshwater Ecology 9 57-62.</ref> Finally, the pupae emerge from the chamber, leaving behind the larval and pupal [[Exuviae|skin]]s.<ref name=cr1997/>

==Uses==
[[File:Hellgrammite in TN stream.JPG|thumb|Hellgrammite (larval form of the dobsonfly) found in a Tennessee stream]]
Hellgrammites are popular among [[angling|anglers]] as [[fishing bait|bait]] due to their large size, endurance, and ferocious temperament.<ref name=cr1997b>Contreras-Ramos, A. Corydalinae. Tree of Life Web Project. November 15, 1997.</ref> [[Smallmouth bass]], in particular, are very attracted to hellgrammites as bait, due to the insects' active movement in the water.<ref>https://www.nps.gov/neri/learn/nature/hellgrammites.htm</ref> [[John Henry Comstock]]<ref name=jhc/> suggested securing a net or wire screen to the rocky bottom of a creek and disturbing the rocks just upstream of the screen as a method to catch the larvae. They often run for relatively high prices at bait shops, leading to over-exploitation in some areas and regulation of sale in certain states.<ref name=mb/> Some anglers instead use [[fishing lure|lures]] in the shape of hellgrammites.<ref name=purdue>Turpin, T. [http://www.agriculture.purdue.edu/agcomm/newscolumns/archives/OSL/2013/August/130808OSL.html#.VC_SFZUtDrc Dobsonflies Look Vicious.] {{Webarchive|url=https://web.archive.org/web/20141006150630/http://www.agriculture.purdue.edu/agcomm/newscolumns/archives/OSL/2013/August/130808OSL.html#.VC_SFZUtDrc |date=2014-10-06 }} Purdue Agriculture News Columns. Purdue Extension. August 8, 2013.</ref>

Although not to the same extent as the larvae of mayflies and caddisflies, hellgrammites are intolerant of [[Water pollution|polluted waters]] and may have potential to be used as indicators of [[water quality]].<ref name=gc2004>Gullan, P. J., and P. S. Cranston. The Insects: An Outline of Entomology. Hoboken, NJ: Wiley Blackwell, 2004. 260.</ref>

As the adults are [[Phototaxis#Insects|strongly attracted to lights]], entomologists and collectors often use [[Blacklight|black light]] and [[mercury-vapor lamp|mercury-vapor light]] traps to capture them.<ref name=cr1997/>

==Gallery==
<gallery>
File: Protohermes grandis.jpg|''[[Protohermes grandis]]''
File:Corydalus cornutus MHNT.jpg|''[[Corydalus cornutus]]''
File:NevromusAustroindicus4.jpg|''[[Nevromus austroindicus]]''
Image:Dobsonfly Corydalus cornutus larvae.jpg|''[[Corydalus cornutus]]'' hellgrammite
</gallery>
{{Commons category-inline|Corydalinae}}

==References==
{{Reflist}}

==Further reading==
* Glorioso, M. (1981). [http://onlinelibrary.wiley.com/doi/10.1111/j.1365-3113.1981.tb00440.x/abstract "Systematics of the dobsonfly subfamily Corydalinae (Megaloptera: Corydalidae)"]. ''Systematic Entomology'' 6(3) 253–90. {{doi|10.1111/j.1365-3113.1981.tb00440.x}}.

==External links==
{{Wikispecies|Corydalinae}}

* [http://mbgnet.mobot.org/fresh/slide/dobson.htm Dobsonfly]. Aquatic Critters. Missouri Botanical Garden.
* [http://tolweb.org/tree?group=Corydalus&contgroup=Corydalinae Genus ''Corydalus'']. Tree of Life Web Project. 1997.
* [http://entomology.ifas.ufl.edu/creatures/misc/eastern_dobsonfly.htm Eastern dobsonfly, ''Corydalus cornutus'']. Featured Creatures. [[University of Florida|UF]]/[[Institute of Food and Agricultural Sciences|IFAS]]

{{Megaloptera}}
{{Taxonbar|from=Q2790845}}

[[Category:Aquatic insects]]
[[Category:Corydalidae]]