Hepialidae

Taxobox
name = Hepialidae

---

image_caption = "Gold Swift" male "calling"
regnum = Animalia
phylum = Arthropoda
classis = Insecta
ordo = Lepidoptera
subordo = Glossata
infraordo = Exoporia
superfamilia = Hepialoidea
familia = Hepialidae
familia_authority = Stephens, 1829
diversity = 60 genera and at least 587 species
diversity_link = Lepidopteran diversity
subdivision_ranks = Genera
subdivision =
*"Abantiades" Herrich-Schäffer, [1858]
*"Aenetus" Herrich-Schäffer, [1858]
*"Afrotheora" Nielsen and Scoble, 1986
*"Andeabatis" Nielsen and Robinson, 1983
*"Antihepialus" Janse, 1942
*"Aoraia" Dumbleton, 1966
*"Aplatissa" Viette, 1953
*"Bipectilis" Chus and Wang, 1985
*"Blanchardinella" Nielsen, Robinson & Wagner, 2000
*"Bordaia" Tindale, 1932
*"Calada" Nielsen and Robinson, 1983
*"Callipielus" Butler, 1882
*"Cibyra" Walker, 1856
*"Cladoxycanus" Dumbleton, 1966
*"Dalaca" Walker, 1856
*"Dalaca" auctt., nec Walker, 1856
*"Dioxycanus" Dumbleton, 1966
*"Druceiella" Viette, 1949
*"Dumbletonius"; auctt
*"Elhamma" Walker, 1856
*"Endoclita"; Felder, 1874
*"Eudalaca" Viette, 1950
*"Fraus" Walker, 1856
*"Gazoryctra" Hübner, [1820]
*"Gorgopis" Hübner, [1820]
*"Heloxycanus" Dugdale, 1994
*"Hepialiscus" Hampson, [1893]
*"Hepialus" Fabricius, 1775
*"Jeana" Tindale, 1935
*"Korscheltellus" Börner, 1920
*"Leto" Hübner, [1820]
*"Metahepialus" Janse, 1942
*"Napialus" Chu and Wang, 1985
*"Neohepialiscus" Viette, 1948
*"Oncopera"
*"Oxycanus" Walker, 1856
*"Palpifer" Hampson, [1893]
*"Parahepialiscus" Viette, 1950
*"Parapielus" Viette, 1949
*"Pfitzneriana" Viette, 1952
*"Pfitzneriella" Viette, 1951
*"Pharmacis" Hübner, [1820]
*"Phassodes" Bethune-Baker, 1905
*"Phassus" Walker, 1856
*"Phialuse" Viette, 1961
*"Phymatopus" Wallengren, 1869
*"Phymatopus" auctt. nec Wallengren, 1869
*"Puermytrans" Viette, 1951
*"Roseala" Viette, 1950
*"Schausiana" Viette, 1950
*"Sthenopis" auctt. nec Packard, [1865]
*"Thitarodes" Viette, 1968
*"Trichophassus" Le Cerf, 1919
*"Trictena" Meyrick, 1890
*"Triodia"
*"Wiseana" Viette, 1961
*"Xhoaphryx" Viette, 1953
*"Zelotypia" Scott, 1869
*"Zenophassus" Tindale, 1941

The Hepialidae is a family of insects in the lepidopteran order. Moths of this family are often referred to as swift moths or ghost moths.

Taxonomy and systematics

Hepialidae constitute by far the most diverse group of the infraorder Exoporia. There are 60 genera and at least 587 currently recognised species of these primitive moths recorded worldwide. The genera "Fraus" (endemic to Australia), "Gazoryctra" (Holarctic), "Afrotheora" (Southern African), and "Antihepialus" (African) are considered to be the most primitive, containing four genera and about 51 species with a mostly relictual southern Gondwanan distribution and are currently separated from the Hepialidae "sensu stricto" which might form a natural, derived groupNielsen, E.S., Robinson, G.S. and Wagner, D.L. 2000. Ghost-moths of the world: a global inventory and bibliography of the Exoporia (Mnesarchaeoidea and Hepialoidea) (Lepidoptera) "Journal of Natural History", 34(6): 823-878. [http://www.ingentaconnect.com/content/tandf/tnah/2000/00000034/00000006/art00003 Abstract] ] . The most diverse genera are "Oxycanus" with 73 species, "Endoclita" with 60 species, "Thitarodes" with 51 species and "Cibyra" with 50 species following a comprehensive catalogue of Exoporia. The relationships of the many genera are not yet well established; see below for an ordered synonymic generic checklist, and the Taxobox for navigation.

Morphology and identification

The family Hepialidae is considered to be very primitive with a number of structural differences to other moths including very short antennae and lack of a functional proboscis or frenulum (see Kristensen, 1999: 61-62 for details)Kristensen, N.P., (1999). The non-Glossatan Moths. Ch. 4, pp. 41-62 in Kristensen, N.P. (Ed.). "Lepidoptera, Moths and Butterflies". Volume 1: Evolution, Systematics, and Biogeography. Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom. Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.] . Like other Exoporia the sperm is transferred to the egg by an external channel between the "ostium" and the ovipore. Other non-ditrysian moths have a common cloaca. The moths are "homoneurous" with similar forewings and hindwings and are sometimes included as 'honorary' members of the Macrolepidoptera. Strictly speaking they are phylogenetically too basal and constitute Microlepidoptera, although hepialids range from very small moths to a wingspan record of 250 mm in "Zelotypia" . Because of their sometimes large size and striking colour patterns, they have received relatively more popular and taxonomic attention than most "micros". Many species display strong sexual dimorphism with males smaller but more boldly marked than females, or at high elevation, females of "Pharmacis" and "Aoraia" show "brachypterous" wing reduction [Sattler, K. (1991). A review of wing reduction in Lepidoptera. "Bulletin of the British Museum of Natural History (Entomology)", 60: 243-288.] .

Distribution

Hepialidae are distributed on ancient landmasses worldwide except Antarctica but with the surprising exceptions of Madagascar, the Caribbean islands and in Africa, Tropical West Africa. It remains to be borne out if these absences are real as "Aenetus cohici" was not long ago discovered in New Caledonia [http://www.sciencebuff.org/aenetus_cohici.php] . In the Oriental and Neotropical regions hepialids have diversified in rainforest environments but this not apparently the case in the Afrotropics. Hepialids mostly have low dispersive powers and do not occur on Oceanic islands with the exception of "Phassodes" on Fiji and Western Samoa and a few species in Japan and Kurile Islands. Whilst the type locality of "Eudalaca sanctahelena" is from the remote island of St Helena, this is thought to be an error for South Africa.

Behaviour

Swift moths are crepuscular and some species form leks, also though to have arisen independently in the hepialoid genus "Ogygioses" (Palaeosetidae). In most genera, males fly swiftly to virgin females that are calling with scent. In other genera, virgin females "assemble" upwind to displaying males [Mallet, J. 1984. Sex roles in the ghost moth "Hepialus humuli" (L.) with a review of mating in the Hepialidae (Lepidoptera). "Zoological Journal of the Linnean Society", 79: 67-82.] , which emit a musky pheromone from scales on the metathoracic tibiae. In such cases of sex role reversal, there may be visual cues also: males of the European Ghost Swift are possibly the most frequently noticed species, being white, ghostly and conspicuous when forming a lek at dusk [Andersson, S., Rydell, J., Svensson, M.G.E. (1998). Light, predation and the lekking behaviour of the ghost swift "Hepialus humuli" (L.) (Lepidoptera, Hepialidae). "Proceedings of the Royal Society B: Biological Sciences", 265: 1345-1351] . Sometimes they hover singly as if suspended from a thread or flying in a figure of eight motion. The chemical structure of some pheromones have been analysed [Schulz S., Francke W., König W.A., Schurig, V., Mori K., Kittmann R. and Schneider D. (1990). Male pheromone of swift moth, "Hepialus hecta" L. (Lepidoptera : Hepialidae). "Journal of chemical ecology", 16(12): 3511-3521.] .

Biology

The female does not lay its eggs in a specific location but scatters ("broadcasts") them while in flight, sometimes in huge numbers (29,000 were recorded from a single female "Trictena" [Tindale, N.B. (1932). Revision of the Australian ghost moths (Lepidoptera Homoneura, family Hepialidae). part 1, "Records of the South Australian Museum", 4: 497-536.] , which is presumably a world record for the Lepidoptera). The maggot-like larvae [http://ukmoths.org.uk/show.php?id=2692] feed in a variety of ways. Probably all Exoporia have concealed larvae, making silken tunnels in all manner of substrates. Some species feed on leaf litter, fungi [http://www.landcareresearch.co.nz/education/insects_spiders/pare/puriri_moth.asp] , mosses, decaying vegetation, ferns, gymnosperms and a wide span of monocot and dicot plants [Grehan, J.R. 1989. Larval feeding habits of the Hepialidae (Lepidoptera)"Journal of Natural History", 23(4): 803-824.] . There is very little evidence of hostplant specialisation; whilst the South African species "Leto venus" is restricted to the tree "Virgilia capensis" this may be a case of "ecological monophagy". A few feed on foliage (the .

Economic significance

Chinese medicine makes considerable use of the "mummies" collected of the caterpillar-attacking fungi "Cordyceps", and these can form an expensive ingredient. [Wu, Y. and Yuan, D. (1997). Biodiversity and conservation in China: a view from entomologists. "Entomologica Sinica", 4: 95-111.] [ [http://66.102.9.104/search?q=cache:FDIQNGSrnDoJ:www.ethnopharmacology.org/newsLetter/nl_2004/ISEnl4EE.pdf+Hepialidae+%2B+edible&hl=en&ct=clnk&cd=2&gl=uk 403 Forbidden ] ] The Witchetty grub (which are sometimes hepialid larvae) is a popular food sources especially among aboriginal Australians. In Central America and South America, hepialid larvae are also eaten [Ramos-Elorduy, J. (2002). Edible insects of Chiapas, Mexico. "Ecology of Food and Nutrition", 41(4): 271-299.] . However, some species of "Wiseana", "Oncopera", "Oxycanus", "Fraus" and "Dalaca" are considered pests of pastures in Australia, New Zealand and South America.

Faunas

Fauna of Europe

Source [http://www.faunaeur.org/full_results.php?id=65] and identification [Chinery, M. (1986). "Collins Guide to the Insects of Britain and Western Europe". (Reprinted 1991)] [Skinner, B. (1984). "Colour Identification Guide to Moths of the British Isles"]
*"Gazoryctra fuscoargentea" O. Bang-Haas 1927 - Northern Scandinavia
*"Gazoryctra ganna" (Hübner 1808)- Alps, northern Scandinavia, northern Russia
*"Hepialus humuli" Linnaeus 1758 Ghost Moth - Europe
*"Korscheltellus lupulinus" Linnaeus 1758 Common Swift - Europe
*"Pharmacis aemiliana" Costantini 1911 - Italy
*"Pharmacis anselminae" Teobaldelli 1977- Italy
*"Pharmacis bertrandi" Le Cerf 1936 - France
*"Pharmacis carna" Denis & Schiffermüller 1775 - Central and Eastern Europe
*"Pharmacis castillana" Oberthür 1883 - Spain
*"Pharmacis claudiae" Kristal & Hirneisen 1994 - Italy
*"Pharmacis fusconebulosa" De Geer 1778 Map-winged Swift - Europe
*"Pharmacis pyrenaica" Donzel 1838 - Pyrenees
*"Phymatopus hecta" Linnaeus 1758 Gold Swift - Central and northern Europe
*"Triodia adriaticus" Osthelder 1931 - Croatia, Macedonia, Greece, Crete
*"Triodia amasina" Herrich-Schäffer 1851 - Balkans
*"Triodia sylvina" Linnaeus 1761 Orange Swift - Europe

Cited literature

References

*Kristensen, N.P., (1999). The non-Glossatan Moths. Ch. 4, pp. 41-62 in Kristensen, N.P. (Ed.). "Lepidoptera, Moths and Butterflies". Volume 1: Evolution, Systematics, and Biogeography. Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom. Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
*Nielsen, E.S., Robinson, G.S. and Wagner, D.L. 2000. Ghost-moths of the world: a global inventory and bibliography of the Exoporia (Mnesarchaeoidea and Hepialoidea) (Lepidoptera) "Journal of Natural History", 34(6): 823-878.

External links

* [http://www.tolweb.org/Hepialidae Tree of Life]
* [http://www.ento.csiro.au/gallery/moths/Hepialidae Australian Moths Online]
* [http://www-staff.it.uts.edu.au/~don/larvae/hepi/hepialidae.html Hepialidae of Australia]
* [http://www.sciencebuff.org/genera_and_species.php Hepialidae of the World - List of Genera and Links to Species]
* [http://www.landcareresearch.co.nz/research/biodiversity/invertebratesprog/nzac/wiseana_spp_images.asp New Zealand Wiseana]
* [http://www.journals.royalsoc.ac.uk/(0dknfsv1d3dahdj040izqq45)/app/home/contribution.asp?referrer=parent&backto=issue,12,15;journal,215,318;linkingpublicationresults,1:102024,1 pdf Swift Moth Lekking]
* [http://internt.nhm.ac.uk/jdsml/research-curation/projects/lepindex/checklist.dsml?searchPageURL=index.dsml&UserID=&UserName=&sort=SCIENTIFIC_NAME_on_card&Current_superfamily=Hepialoidea&Current_family=Hepialidae&Current_subfamily=&Current_tribe=&recLimit=500 Lep index list of Hepialidae species]
* [http://www.pherobase.com/database/family/family-Hepialidae.php "Endoclita" and "Hepialus" pheromones]
* [http://www.ethnopharmacology.org/newsLetter/nl_2004/ISEnl4EE.pdf Abstract, counterfeit hepialid mummies]
* [http://www.landcareresearch.co.nz/education/insects_spiders/pare/puriri_moth.asp Puriri Moth]
* [http://www.landcareresearch.co.nz/research/biosystematics/invertebrates/nzac/wiseana_spp_images.asp "Wiseana"]
* [http://www.anu.edu.au/linguistics/nash/aust/nbt/obituary.html Obituary of Norman B. Tindale]

Generic checklist

*"Fraus" Walker, 1856
**="Hectomanes" Meyrick, 1980
**="Praus"; Pagenstacher, 1909
*"Gazoryctra" Hübner, [1820]
**="Garzorycta"; Hübner, [1826]
**="Gazoryctes"; Kirby, 1892
*"Afrotheora" Nielsen and Scoble, 1986
*"Antihepialus" Janse, 1942
**="Ptycholoma"; Felder, 1874
*"Bipectilis" Chus and Wang, 1985
*"Palpifer" Hampson, [1893]
**="Palpiphorus"; Quail, 1900
**="Palpiphora"; Pagenstacher, 1909
*"Eudalaca" Viette, 1950
**="Eudalacina" Paclt, 1953
*"Gorgopis" Hübner, [1820]
**="Gorcopis"; Walker, 1856
*"Metahepialus" Janse, 1942
*"Dalaca" Walker, 1856
**="Huapina" Bryk, 1945
**="Maculella" Viette, 1950
**="Toenga" Tindale, 1954
*"Callipielus" Butler, 1882
**="Stachyocera" Ureta, 1957
*"Blanchardinella" Nielsen, Robinson & Wagner, 2000
**="Blanchardina" Viette, 1950, nec Labbe, 1899
*"Calada" Nielsen and Robinson, 1983
*"Puermytrans" Viette, 1951
*"Parapielus" Viette, 1949
**="Lossbergiana" Viette, 1951
*"Andeabatis" Nielsen and Robinson, 1983
*"Druceiella" Viette, 1949
*"Trichophassus" Le Cerf, 1919
*"Phassus" Walker, 1856
*"Schausiana" Viette, 1950
*"Aplatissa" Viette, 1953
*"Pfitzneriana" Viette, 1952
*"Cibyra" Walker, 1856
*"Cibyra" ("Pseudodalaca" Viette, 1950)
*"Cibyra" ("Gymelloxes" Viette, 1952)
*"Cibyra" ("Alloaepytus" Viette, 1951)
*"Cibyra" ("Aeptus") Herrich-Schäffer, [1858]
*"Cibyra" ("Thiastyx" Viette, 1951)
*"Cibyra" ("Schaefferiana" Viette, 1950)
*"Cibyra" ("Paragorgopis" Viette, 1952)
*"Cibyra" ("Hepialyxodes" Viette, 1951)
*"Cibyra" ("Xytrops" Viette, 1951)
*"Cibyra" ("Cibyra" Walker, 1856)
*"Cibyra" ("Lamelliformia" Viette, 1952)
*"Cibyra" ("Tricladia" Felder, 1874)
**="Pseudophassus" Pfitzner, 1914
**="Parana" Viette, 1950
*"Cibyra" ("Pseudophilaenia" Viette, 1951)
*"Cibyra" ("Philoenia" Kirby, 1892)
**="Philaenia" auctt.
*"Cibyra" ("Yleuxas" Viette, 1951)
*"Phialuse" Viette, 1961
*"Roseala" Viette, 1950
*"Dalaca" auctt., nec Walker, 1856
*"Pfitzneriella" Viette, 1951
*"Aoraia" Dumbleton, 1966
**="Trioxycanus" Dumbleton, 1966
*"Triodia"
**="Alphus" Wallengren, 1869, nec Dejean, 1833
*"Korscheltellus" Börner, 1920
*"Pharmacis" Hübner, [1820]
*"Thitarodes" Viette, 1968
**="Forkalus" Chu and Wang, 1985
*"Phymatopus" Wallengren, 1869
**="Hepiolopsis" Börner, 1920
**="Phimatopus"; auctt.
*"Phymatopus" auctt. nec Wallengren, 1869
*"Hepialus" Fabricius, 1775
**="Hepiolus" Illiger, 1801
**="Epialus" Agassiz, 1847
**="Epiolus" Agassiz, 1847
**="Tephus" Wallengren, 1869
**="Trepialus"; Latreille, [1805]
*"Zenophassus" Tindale, 1941
*"Sthenopis" auctt. nec Packard, [1865]
*"Endoclita"; Felder, 1874
**="Endoclyta", Felder, 1875
**="Hypophassus", Le Cerf, 1919
**="Nevina", Tindale, 1941
**="Sahyadrassus", Tindale, 1941
**="Procharagia", Viette, 1949
*"Neohepialiscus" Viette, 1948
*"Elhamma" Walker, 1856
**="Perissectis" Meyrick, 1890
**="Pericentris"; Pagenstacher, 1909
**="Zauxieus" Viette, 1952
**="Theaxieus" Viette, 1952
*"Jeana" Tindale, 1935
*"Cladoxycanus" Dumbleton, 1966
*"Wiseana" Viette, 1961
**="Porina" Walker, 1956, nec d'Orbigny, 1852
**="Gorina"; Quail, 1899
**="Goryna"; Quail, 1899
**="Philpottia" Viette, 1950, nec Broun, 1915
*"Heloxycanus" Dugdale, 1994
*"Dumbletonius"; auctt
**="Trioxycanus" Dumbleton, 1966
*"Dioxycanus" Dumbleton, 1966
*"Napialus" Chu and Wang, 1985
*"Hepialiscus" Hampson, [1893]
*"Parahepialiscus" Viette, 1950
*"Xhoaphryx" Viette, 1953
*"Aenetus" Herrich-Schäffer, [1858]
**="Charagia" Walker, 1856
**="Phloiopsyche" Scott, 1864
**="Oenetus"; Kirby, 1892
**="Choragia"; Pagenstacher, 1909
**="Oenetes"; Oke, 1953
*"Leto" Hübner, [1820]
**="Ecto"; Pagenstacher, 1909
*"Zelotypia" Scott, 1869
**="Xylopsyche Swainson, 1851
**="Leto"; auctt
*"Oncopera"
**="Oncoptera" Walker, 1890
**="Paroncopera" Tindale, 1933
**="Onchopera"; Birket-Smith, 1974
**="Onchoptera"; Birket-Smith, 1974
*"Trictena" Meyrick, 1890
*"Bordaia" Tindale, 1932
**="Bordaja"; Chu and Wang, 1985
*"Abantiades" Herrich-Schäffer, [1858]
**="Pielus" Walker, 1856
**="Rhizopsyche" Scott, 1864
*"Oxycanus" Walker, 1856
**="Porina" Walker, 1856
**="Gorina"; Quail, 1899
**="Goryna"; Quail, 1899
**="Paraoxyxanus" Viette, 1950
*"Phassodes" Bethune-Baker, 1905