# Hepialidae

The Hepialidae are 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

The Hepialidae constitute by far the most diverse group of the infraorder Exoporia. The 82 genera contain at least 700 currently recognised species of these primitive moths 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 group. The most diverse genera are Oxycanus with 78 species, Endoclita with 78 species, andThitarodes with 80 species following a comprehensive catalogue of Exoporia.<1>] 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 the lack of a functional proboscis or frenulum (see Kristensen, 1999: 61–62 for details). 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. 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. 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.

## 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. 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 usually crepuscular and some species form leks, also thought to have arisen independently in the 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, which emit a 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. Sometimes they hover singly as if suspended from a thread or flying in a figure of eight motion. The chemical structures of some pheromones have been analysed.

## 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, which is presumably a world record for the Lepidoptera). The maggot-like larvae 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, mosses, decaying vegetation, ferns, gymnosperms and a wide span of monocot and dicot plants. 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 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. Root-feeding larvae travelling through soil make silk-lined tunnels. Before pupating they make a vertical tunnel, which can be up to 10 cm deep, with an exit close to the ground surface. The pupae can then climb up and down to adjust to changes in temperature and flooding. Before the adult moth emerges, the pupa protrudes half way out at the ground surface. The pupa has rows of dorsal spines on the abdominal segments as in other lower members of the Heteroneura.

## Economic significance

Chinese medicine makes considerable use of the "mummies" collected of the caterpillar-attacking fungi Ophiocordyceps sinensis, and these can form an expensive ingredient. The witchetty grub (which are sometimes hepialid larvae) is a popular food source especially among aboriginal Australians. In Central America and South America, hepialid larvae are also eaten. However, some species of Wiseana, Oncopera, Oxycanus, Fraus and Dalaca are considered pests of pastures in Australia, New Zealand, and South America.

## Phylogeny

The Hepialidae were identified as having primitive wing venation by John Henry Comstock (1893). In his study of Evolution of the Wings of Insects he shows that the fore and hind wings of Sthenopis (Hepialus) argenteomaculatus maintain a five branched radius while in the remainder of the Lepidoptera the hind wing radius is merged into one vein. This identifies the Hepialidae as a primitive relict of primitive wing venation.

## Faunas
### Fauna of Europe

Source and identification
* Gazoryctra fuscoargenteus O. Bang-Haas 1927 – Northern Scandinavia
* Gazoryctra ganna (Hübner 1808) – Alps, northern Scandinavia, northern Russia
* Hepialus humuli Linnaeus 1758 (ghost moth) – Europe
* 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 hecta Linnaeus 1758 (gold swift) – Central and northern Europe
* Triodia adriaticus Osthelder 1931 – Croatia, North Macedonia, Greece, Crete
* Triodia amasinus Herrich-Schäffer 1851 – Balkans
* Triodia sylvina Linnaeus 1761 (orange swift) – Europe

## 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

## Cited literature

* ↑ John R. Grehan, Carlos G.C. Mielke, John R.G. Turner, and John E. Nielsen. 2023. A revised world catalogue of Ghost Moths (Lepidoptera: Hepialidae) with taxonomic and biological annotations. ZooNova 28: 1-313
* ↑ van Nieukerken; et al. (2011). "Order Lepidoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness" (PDF). Zootaxa. 3148: 212–221. doi:10.11646/zootaxa.3148.1.41. Archived (PDF) from the original on 2019-10-13. Retrieved 2023-02-22.
* ↑ John R. Grehan, Carlos G.C. Mielke, John R.G. Turner, and John E. Nielsen. 2023. A revised world catalogue of Ghost Moths (Lepidoptera: Hepialidae) with taxonomic and biological annotations. ZooNova 28: 1-313
* 1 2 3 4 5 6 7 8 9 10 11 12 Nielsen, Ebbe S.; Robinson, Gaden S.; Wagner, David 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. Bibcode:2000JNatH..34..823N. doi:10.1080/002229300299282. S2CID 86004391.
* ↑ John R. Grehan, Carlos G.C. Mielke, John R.G. Turner, and John E. Nielsen. 2023. A revised world catalogue of Ghost Moths (Lepidoptera: Hepialidae) with taxonomic and biological annotations. ZooNova 28: 1-313
* 1 2 3 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.
* ↑ Sattler, Klaus (1991). "A review of wing reduction in Lepidoptera". Bulletin of the British Museum (Natural History), Entomology. 60 (2): 243–288. Archived from the original on 2022-10-13. Retrieved 2023-02-22.
* ↑ "Buffalo Museum of Science – Home". www.sciencebuff.org. Archived from the original on 27 September 2007. Retrieved 5 April 2018.
* ↑ This source is no longer available
* ↑ Mallet, James (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. doi:10.1111/j.1096-3642.1984.tb02320.x.
* ↑ 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 (1403): 1345–1351. doi:10.1098/rspb.1998.0440. PMC 1689211.
* ↑ Schulz, Stefan; Francke, Wittko; König, Wilfried A.; Schurig, Volker; Mori, Kenji; Kittmann, Rolf; Schneider, Dietrich (1990). "Male pheromone of swift moth, Hepialus hecta L. (Lepidoptera: Hepialidae)". Journal of Chemical Ecology. 16 (12): 3511–3521. Bibcode:1990JCEco..16.3511S. doi:10.1007/BF00982114. PMID 24263445. S2CID 26903035.
* ↑ Tindale, Norman Barnett (1932). "Revision of the Australian ghost moths (Lepidoptera Homoneura, family Hepialidae). Part 1". Records of the South Australian Museum. 4: 497–536. Archived from the original on 2023-11-08. Retrieved 2024-03-17.
* ↑ "Ghost Moth Larva | UKmoths".
* ↑ "Puriri moth: NZAC Pare". www.landcareresearch.co.nz. Archived from the original on 2004-04-23.
* ↑ Grehan, J. R. (1989). "Larval feeding habits of the Hepialidae (Lepidoptera)". Journal of Natural History. 23 (4): 803–824. Bibcode:1989JNatH..23..803G. doi:10.1080/00222938900770421.
* ↑ H. Buser, W.Huber and R. Joos 2000 Hepialidae – Wurzelbohrer. Pp. 61-96 in Schmetterlinge und ihre Lebensräume. Band 3. Pro Natura, Basel.
* ↑ Atijegbe, S. R.; Mansfield, S.; Rostás, M.; Ferguson, C. M.; Worner, S. (2020). "The remarkable locomotory ability of Wiseana (Lepidoptera: Hepialidae) pupae: an adaptation to predation and environmental conditions?". The Wētā. 54: 19–31.
* ↑ Wu, Yanru; Yuan, Decheng (1997). "Biodiversity and conservation in China: a view from entomologists". Entomologica Sinica. 4 (2): 95–111. Bibcode:1997InsSc...4...95W. doi:10.1111/j.1744-7917.1997.tb00078.x. S2CID 86294063. Archived from the original on 2024-05-17. Retrieved 2023-02-22.
* ↑ Ramos-Elorduy, Julieta (2002). "Edible insects of Chiapas, Mexico". Ecology of Food and Nutrition. 41 (4): 271–299. Bibcode:2002EcoFN..41..271R. doi:10.1080/03670240214081. S2CID 84107193.
* ↑ J. H., Comstock (1893). Evolution of the Wings of Insects. Ithaca, NY: The Wilder Quarter Century Book.
* ↑ "Fauna Europaea". Archived from the original on October 1, 2007.
* ↑ 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