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

Contents

1 Taxonomy and systematics 2 Morphology and identification 3 Distribution 4 Behaviour 5 Biology 6 Economic significance 7 Faunas

7.1 Fauna of Europe

8 Generic checklist 9 Cited literature 10 References 11 External links

Taxonomy and systematics[edit] The Hepialidae
Hepialidae
constitute by far the most diverse group of the infraorder Exoporia. The 60 genera contain at least 587 currently recognised species of these primitive moths worldwide. The genera Fraus (endemic to Australia), Gazoryctra
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
Gondwanan
distribution and are currently separated from the Hepialidae
Hepialidae
sensu stricto which might form a natural, derived group.[2] The most diverse genera are Oxycanus
Oxycanus
with 73 species, Endoclita
Endoclita
with 60 species, Thitarodes with 51 species and Cibyra with 50 species following a comprehensive catalogue of Exoporia.[2] The relationships of the many genera are not yet well established; see below for an ordered synonymic generic checklist,[2] and the Taxobox for navigation. Morphology and identification[edit] The family Hepialidae
Hepialidae
is considered to be very primitive, with a number of structural differences to other moths including very short antennae and the lack of a functional proboscis or frenulum (see Kristensen, 1999: 61-62 for details).[3] Like other Exoporia the sperm is transferred to the egg by an external channel between the ostium and the ovipore. Other nonditrysian moths have a common cloaca.[2] The moths are homoneurous with similar forewings and hindwings, and are sometimes included as 'honorary' members of the Macrolepidoptera, though archaic they are. 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.[2] Because of their sometimes large size and striking colour patterns, they have received 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.[4] Distribution[edit]

Abantiades
Abantiades
latipennis, Tasmania, Australia

Hepialidae
Hepialidae
are distributed on ancient landmasses worldwide except Antarctica
Antarctica
but with the surprising exceptions of Madagascar, the Caribbean
Caribbean
islands and in Africa, tropical West Africa. It remains to be borne out if these absences are real as Aenetus
Aenetus
cohici was not long ago discovered in New Caledonia.[5] In the Oriental
Oriental
and Neotropical regions hepialids have diversified in rainforest environments, but this not apparently the case in the Afrotropics.[2] Hepialids mostly have low dispersive powers and do not occur on oceanic islands with the exception of Phassodes
Phassodes
on Fiji
Fiji
and Western Samoa
Western Samoa
and a few species in Japan
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.[2] Behaviour[edit] Swift moths are crepuscular and some species form leks, also thought to have arisen independently in the genus Ogygioses (Palaeosetidae).[3] In most genera, males fly swiftly to virgin females that are calling with scent. In other genera, virgin females "assemble" upwind to displaying males,[6] 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.[7] Sometimes they hover singly as if suspended from a thread or flying in a figure of eight motion.[2] The chemical structures of some pheromones have been analysed.[8] Biology[edit] 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,[9] which is presumably a world record for the Lepidoptera). The maggot-like larvae [1] 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[2], mosses, decaying vegetation, ferns, gymnosperms and a wide span of monocot and dicot plants.[2][10] There is very little evidence of hostplant specialisation; whilst the South African species Leto venus
Leto venus
is restricted to the tree Virgilia capensis
Virgilia capensis
this may be a case of "ecological monophagy".[2] A few feed on foliage (the austral 'oxyacanine' genera which may drag foliage into their feeding tunnel: Nielsen et al., 2000: 825). Most feed underground on fine roots, at least in early instars and some then feed internally in tunnels in the stem or trunk of their hostplants. The pupa has rows of dorsal spines on the abdominal segments as in other lower members of the Heteroneura.[3] Economic significance[edit] Chinese medicine makes considerable use of the "mummies" collected of the caterpillar-attacking fungi Ophiocordyceps sinensis, and these can form an expensive ingredient.[2][11] [12] The Witchetty grub
Witchetty grub
(which are sometimes hepialid larvae) is a popular food source especially among aboriginal Australians. In Central America
Central America
and South America, hepialid larvae are also eaten.[13] However, some species of Wiseana, Oncopera, Oxycanus, Fraus and Dalaca are considered pests of pastures in Australia, New Zealand, and South America.[2] Faunas[edit] Fauna of Europe[edit] Source [3] and identification[14] [15]

Gazoryctra
Gazoryctra
fuscoargenteus O. Bang-Haas 1927 - Northern Scandinavia Gazoryctra
Gazoryctra
ganna (Hübner 1808) - Alps, northern Scandinavia, northern Russia Hepialus
Hepialus
humuli Linnaeus 1758 (ghost moth) - Europe Korscheltellus
Korscheltellus
lupulina 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 pyrenaicus Donzel 1838 - Pyrenees Phymatopus
Phymatopus
hecta Linnaeus 1758 (gold swift) - Central and northern Europe Triodia adriaticus
Triodia adriaticus
Osthelder 1931 - Croatia, Macedonia, Greece, Crete Triodia amasinus
Triodia amasinus
Herrich-Schäffer 1851 - Balkans Triodia sylvina
Triodia sylvina
Linnaeus 1761 (orange swift) - Europe

Generic checklist[edit]

Fraus Walker, 1856

=Hectomanes Meyrick, 1980 =Praus; Pagenstacher, 1909

Gazoryctra
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
Palpifer
Hampson, [1893]

=Palpiphorus; Quail, 1900 =Palpiphora; Pagenstacher, 1909

Eudalaca
Eudalaca
Viette, 1950

=Eudalacina Paclt, 1953

Gorgopis
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
Korscheltellus
Börner, 1920 Pharmacis Hübner, [1820] Thitarodes Viette, 1968

=Forkalus Chu and Wang, 1985

Phymatopus
Phymatopus
Wallengren, 1869

=Hepiolopsis Börner, 1920 =Phimatopus; auctt.

Phymatopus
Phymatopus
auctt. nec Wallengren, 1869 Hepialus
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
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
Heloxycanus
Dugdale, 1994 Dumbletonius; auctt

=Trioxycanus Dumbleton, 1966

Dioxycanus Dumbleton, 1966 Napialus Chu and Wang, 1985 Hepialiscus
Hepialiscus
Hampson, [1893] Parahepialiscus Viette, 1950 Xhoaphryx Viette, 1953 Aenetus
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
Zelotypia
Scott, 1869

=Xylopsyche Swainson, 1851 =Leto; auctt

Oncopera

=Oncoptera Walker, 1890 =Paroncopera Tindale, 1933 =Onchopera; Birket-Smith, 1974 =Onchoptera; Birket-Smith, 1974

Trictena
Trictena
Meyrick, 1890 Bordaia Tindale, 1932

=Bordaja; Chu and Wang, 1985

Abantiades
Abantiades
Herrich-Schäffer, [1858]

=Pielus Walker, 1856 =Rhizopsyche Scott, 1864

Oxycanus
Oxycanus
Walker, 1856

=Porina Walker, 1856 =Gorina; Quail, 1899 =Goryna; Quail, 1899 =Paraoxyxanus Viette, 1950

Phassodes
Phassodes
Bethune-Baker, 1905

Cited literature[edit]

^ " Animal
Animal
biodiversity: An outline of higher-level classification and survey of taxonomic richness - Lepidoptera" (PDF). mapress.com. Retrieved 5 April 2018.  ^ a b c d e f g h i j k l 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.Abstract ^ a b c 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
Animal
Kingdom. Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York. ^ Sattler, K. (1991). A review of wing reduction in Lepidoptera. Bulletin of the British Museum of Natural History (Entomology), 60: 243-288. ^ "Buffalo Museum of Science - Home". www.sciencebuff.org. Retrieved 5 April 2018.  ^ Mallet, J. 1984. Sex roles in the ghost moth Hepialus
Hepialus
humuli (L.) with a review of mating in the Hepialidae
Hepialidae
(Lepidoptera). Zoological Journal of the Linnean Society, 79: 67-82. ^ Andersson, S., Rydell, J., Svensson, M.G.E. (1998). Light, predation and the lekking behaviour of the ghost swift Hepialus
Hepialus
humuli (L.) (Lepidoptera, Hepialidae). Proceedings of the Royal Society B: Biological Sciences, 265: 1345-1351 ^ 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. ^ Tindale, N.B. (1932). Revision of the Australian ghost moths ( Lepidoptera
Lepidoptera
Homoneura, family Hepialidae). part 1, Records of the South Australian Museum, 4: 497-536. ^ Grehan, J.R. 1989. Larval feeding habits of the Hepialidae (Lepidoptera) Journal of Natural History, 23(4): 803-824. ^ Wu, Y. and Yuan, D. (1997). Biodiversity
Biodiversity
and conservation in China: a view from entomologists. Entomologica Sinica, 4: 95-111. ^ "Wayback Machine" (PDF). archive.org. 8 August 2007. Retrieved 5 April 2018.  ^ Ramos-Elorduy, J. (2002). Edible insects of Chiapas, Mexico. Ecology of Food and Nutrition, 41(4): 271-299. ^ Chinery, M. (1986). Collins Guide to the Insects
Insects
of Britain and Western Europe. (Reprinted 1991) ^ Skinner, B. (1984). Colour Identification Guide to Moths of the British Isles

References[edit]

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[edit]

Data related to Hepialidae
Hepialidae
at Wikispecies

Wikimedia Commons has media related to Lepidoptera.

Tree of Life Australian Moths Online Hepialidae
Hepialidae
of Australia Hepialidae
Hepialidae
of the World - List of Genera and Links to Species Lep index list of Hepialidae
Hepialidae
species Endoclita
Endoclita
and Hepialus
Hepialus
pheromones Abstract, counterfeit hepialid mummies Images of Hepialidae
Hepialidae
species in New Zealand Obituary of Norman B. Tindale

v t e

Extant Lepidoptera
Lepidoptera
families

Kingdom: Animalia Phylum: Arthropoda Class: Insecta Subclass: Pterygota Infraclass: Neoptera Superorder: Endopterygota

Suborder Zeugloptera

Micropterigoidea

Micropterigidae
Micropterigidae
(mandibulate archaic moths)

Suborder Aglossata

Agathiphagoidea

Agathiphagidae (kauri moths)

Suborder Heterobathmiina

Heterobathmioidea

Heterobathmiidae

Suborder Glossata

Dacnonypha

Eriocranioidea

Eriocraniidae

Acanthoctesia

Acanthopteroctetoidea

Acanthopteroctetidae (archaic sun moths)

Lophocoronina

Lophocoronoidea

Lophocoronidae

Neopseustina

Neopseustoidea

Neopseustidae (archaic bell moths)

Exoporia

Hepialoidea

Anomosetidae Hepialidae
Hepialidae
(swift moths, ghost moths) Neotheoridae (Amazonian primitive ghost moths) Palaeosetidae (miniature ghost moths) Prototheoridae (African primitive ghost moths)

Mnesarchaeoidea

Mnesarchaeidae (New Zealand primitive moths)

H e t e r o n e u r a

M o n o t r y s i a

Incurvarioidea

Adelidae
Adelidae
(fairy longhorn moths) Cecidosidae Crinopterygidae Heliozelidae Incurvariidae Prodoxidae
Prodoxidae
(yucca moths)

Andesianoidea

Andesianidae (Andean endemic moths)

Nepticuloidea

Nepticulidae
Nepticulidae
(pigmy, or midget moths) Opostegidae
Opostegidae
(white eyecap moths)

Palaephatoidea

Palaephatidae (Gondwanaland moths)

Tischerioidea

Tischeriidae (trumpet leaf miner moths)

D i t r y s i a

Simaethistoidea

Simaethistidae

Tineoidea

Acrolophidae
Acrolophidae
(burrowing webworm moths) Arrhenophanidae Eriocottidae (Old World spiny-winged moths) Lypusidae Psychidae (bagworm moths) Tineidae
Tineidae
(fungus moths)

Gracillarioidea

Bucculatricidae
Bucculatricidae
(ribbed cocoon makers) Douglasiidae (Douglas moths) Gracillariidae Roeslerstammiidae

Yponomeutoidea

Acrolepiidae
Acrolepiidae
(false diamondback moths) Bedelliidae Glyphipterigidae
Glyphipterigidae
(sedge moths) Heliodinidae Lyonetiidae Plutellidae Yponomeutidae (ermine moths) Ypsolophidae

Gelechioidea

Autostichidae Batrachedridae Blastobasidae Coleophoridae
Coleophoridae
(case-bearers, case moths) Cosmopterigidae
Cosmopterigidae
(cosmet moths) Elachistidae
Elachistidae
(grass-miner moths) Gelechiidae
Gelechiidae
(twirler moths) Lecithoceridae
Lecithoceridae
(long-horned moths) Metachandidae Momphidae
Momphidae
(mompha moths) Oecophoridae
Oecophoridae
(concealer moths) Pterolonchidae Scythrididae
Scythrididae
(flower moths) Xyloryctidae
Xyloryctidae
(timber moths)

Galacticoidea

Galacticidae

Zygaenoidea

Heterogynidae Zygaenidae
Zygaenidae
(burnet, forester, or smoky moths) Himantopteridae Lacturidae Somabrachyidae Megalopygidae (flannel moths) Aididae Anomoeotidae Cyclotornidae Epipyropidae
Epipyropidae
(planthopper parasite moths) Dalceridae
Dalceridae
(slug caterpillars) Limacodidae
Limacodidae
(slug, or cup moths)

Cossoidea

Cossidae
Cossidae
(carpenter millers, or goat moths) Dudgeoneidae (dudgeon carpenter moths)

Sesioidea

Brachodidae (little bear moths) Castniidae
Castniidae
(castniid moths: giant butterfly-moths, sun moths) Sesiidae
Sesiidae
(clearwing moths)

Choreutoidea

Choreutidae
Choreutidae
(metalmark moths)

Tortricoidea

Tortricidae
Tortricidae
(tortrix moths)

Urodoidea

Urodidae
Urodidae
(false burnet moths)

Schreckensteinioidea

Schreckensteiniidae
Schreckensteiniidae
(bristle-legged moths)

Epermenioidea

Epermeniidae
Epermeniidae
(fringe-tufted moths)

Alucitoidea

Alucitidae (many-plumed moths) Tineodidae (false plume moths)

Pterophoroidea

Pterophoridae
Pterophoridae
(plume moths)

Whalleyanoidea

Whalleyanidae

Immoidea

Immidae

Copromorphoidea

Copromorphidae (tropical fruitworm moths) Carposinidae
Carposinidae
(fruitworm moths)

Hyblaeoidea

Hyblaeidae
Hyblaeidae
(teak moths)

Pyraloidea

Pyralidae
Pyralidae
(snout moths) Crambidae
Crambidae
(grass moth)

Thyridoidea

Thyrididae
Thyrididae
(picture-winged leaf moths)

Mimallonoidea

Mimallonidae (sack bearer moths)

Lasiocampoidea

Lasiocampidae
Lasiocampidae
(eggars, snout moths, or lappet moths)

Bombycoidea

Anthelidae
Anthelidae
(Australian lappet moth) Bombycidae
Bombycidae
(silk moths) Brahmaeidae
Brahmaeidae
(Brahmin moths) Carthaeidae (Dryandra moth) Endromidae
Endromidae
(Kentish glory and relatives) Eupterotidae Lemoniidae Saturniidae
Saturniidae
(saturniids) Sphingidae
Sphingidae
(hawk moths, sphinx moths and hornworms) Phiditiidae

Noctuoidea

Doidae Erebidae
Erebidae
(underwing, tiger, tussock, litter, snout, owlet moths) Euteliidae Noctuidae
Noctuidae
(daggers, sallows, owlet moths, quakers, cutworms, darts) Nolidae
Nolidae
(tuft moths) Notodontidae
Notodontidae
(prominents, kittens) Oenosandridae

Drepanoidea

Epicopeiidae
Epicopeiidae
(oriental swallowtail moths) Drepanidae
Drepanidae
(hook-tips)

Geometroidea

Sematuridae Uraniidae Geometridae (geometer moths)

Cimelioidea

Cimeliidae (gold moths)

Calliduloidea

Callidulidae
Callidulidae
(Old World butterfly-moths)

Superfamily unassigned

Millieriidae

Rhopalocera (butterflies)

Hedyloidea

Hedylidae
Hedylidae
(American moth-butterflies)

Hesperioidea

Hesperiidae (skippers)

Papilionoidea (true butterflies)

Lycaenidae
Lycaenidae
(gossamer-winged butterflies: blues, coppers and relatives) Nymphalidae
Nymphalidae
(brush-footed, or four-footed butterflies) Papilionidae (swallowtail butterflies) Pieridae
Pieridae
(whites, yellows, orangetips, sulphurs) Riodinidae
Riodinidae
(metalmarks)

Note: division Monotrysia
Monotrysia
is not a clade.

Taxonomy of the Lepidoptera Lists by region

Taxon identifiers

Wd: Q580250 BAMONA: Hepialidae BugGuide: 19714 EoL: 834 EPPO: 1HEPIF Fauna Europaea: 65 Fossilworks: 71427 GBIF: 3556 ITIS:

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