Hepialidae

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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 moths worldwide.<ref>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</ref> The genera Fraus (endemic to Australia), Gazoryctra (Holarctic), Afrotheora (Southern African), and Antihepialus (African) are considered to be among the most basal, 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.<ref name="Nielsen00">Template:Cite journal</ref><ref>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</ref> The most diverse genera are Oxycanus with 78 species, Endoclita with 78 species, and Thitarodes with 80 species following a comprehensive catalogue of Exoporia.<ref name="Nielsen00"/> The relationships of the many genera are not yet well established; see below for an ordered synonymic generic checklist,<ref name="Nielsen00"/> and the Taxobox for navigation.

Morphology and identification

The Hepialidae represent one of the most basal families within the Lepidoptera, exhibiting numerous structural differences from other moths, such as very short antennae and the absence of a functional proboscis or frenulum (see Kristensen, 1999: 61–62 for details).<ref name="Kristensen99">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.</ref> Like other Exoporia, sperm is transferred to the egg via an external channel between the ostium and the ovipore. In contrast, other non-Ditrysian moths possess a common cloaca.<ref name="Nielsen00"/> Hepialids are homoneurous, with similar forewings and hindwings, and are occasionally regarded as 'honorary' members of the Macrolepidoptera despite their basal position. Phylogenetically, they are classified among the Microlepidoptera, though species range from very small moths to a record wingspan of 250 mm in Zelotypia.<ref name="Nielsen00"/> Due to their often large size and striking color patterns, they have attracted more popular and taxonomic interest than most "micros". Many species exhibit pronounced sexual dimorphism, with males typically smaller but more boldly marked than females; at high elevations, females of genera such as Pharmacis and Aoraia display "brachypterous" wing reduction.<ref>Template:Cite journal</ref>

Distribution

File:Abantiades latipennis.jpg
Abantiades latipennis, Tasmania, Australia

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.<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref><ref>This source is no longer available</ref> In the Oriental and Neotropical regions hepialids have diversified in rainforest environments, but this not apparently the case in the Afrotropics.<ref name ="Nielsen00"/> 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.<ref name="Nielsen00"/>

Behaviour

Swift moths are usually crepuscular and some species form leks, also thought to have arisen independently in the genus Ogygioses (Palaeosetidae).<ref name="Kristensen99"/> In most genera, males fly swiftly to virgin females that are calling with scent. In other genera, virgin females "assemble" upwind to displaying males,<ref>Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref> Sometimes they hover singly as if suspended from a thread or flying in a figure of eight motion.<ref name="Nielsen00"/> The chemical structures of some pheromones have been analysed.<ref>Template:Cite journal</ref>

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,<ref>Template:Cite journal</ref> which is presumably a world record for the Lepidoptera). The maggot-like larvae<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> 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,<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> mosses, decaying vegetation, ferns, gymnosperms and a wide span of monocot and dicot plants.<ref name="Nielsen00"/><ref>Template:Cite journal</ref> 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".<ref name="Nielsen00"/> 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.<ref>H. Buser, W.Huber and R. Joos 2000 Hepialidae – Wurzelbohrer. Pp. 61-96 in Schmetterlinge und ihre Lebensräume. Band 3. Pro Natura, Basel.</ref> The pupae can then climb up and down to adjust to changes in temperature and flooding.<ref>Template:Cite journal</ref> 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.<ref name="Kristensen99"/>

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.<ref name="Nielsen00"/><ref>Template:Cite journal</ref> 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.<ref>Template:Cite journal</ref> However, some species of Wiseana, Oncopera, Oxycanus, Fraus and Dalaca are considered pests of pastures in Australia, New Zealand, and South America.<ref name="Nielsen00"/>

Phylogeny

The Hepialidae were historically described as having "primitive" wing venation by John Henry Comstock (1893); in modern terms, this would rather be phrased and conceptualized as retaining ancestral wing venation. In his study Evolution of the Wings of Insects, he illustrated that the fore- and hindwings of Sthenopis (Hepialus) argenteomaculatus possess a five-branched radius, in contrast to other Lepidoptera where the hindwing radius is reduced to a single vein. This observation positioned the Hepialidae as a relict lineage preserving early-derived wing venation patterns.<ref>Template:Cite book</ref>

Faunas

Fauna of Europe

Source<ref>{{#invoke:citation/CS1|citation |CitationClass=web }}</ref> and identification<ref>Chinery, M. (1986). Collins Guide to the Insects of Britain and Western Europe. (Reprinted 1991)</ref><ref>Skinner, B. (1984). Colour Identification Guide to Moths of the British Isles</ref>

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

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References

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  • Comstock, J.H., (1893). Evolution of the Wings of Insects. The Wilder Quarter Century Book, Ithaca, NY.
  • 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.

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