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Wikipedia / 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 moths worldwide. 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. The most diverse genera are Oxycanus with 78 species, Endoclita with 78 species, and Thitarodes with 80 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 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). 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. 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. 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.

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 10cm 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 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.

Faunas

Fauna of Europe

Source and identification

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]

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

Cibyra (Tricladia Felder, 1874)

  • =Pseudophassus Pfitzner, 1914

  • =Parana Viette, 1950

  • Cibyra (Pseudophilaenia Viette, 1951)

Cibyra (Philoenia Kirby, 1892)

Aoraia Dumbleton, 1966

  • =Trioxycanus Dumbleton, 1966

Triodia

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

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

References

  • 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.