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.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 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.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 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).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, 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.This source is no longer available 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.H. Buser, W.Huber and R. Joos 2000 Hepialidae – Wurzelbohrer. Pp. 61-96 in Schmetterlinge und ihre Lebensräume. Band 3. Pro Natura, Basel. 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 identificationChinery, 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 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, =Garzorycta; Hübner, =Gazoryctes; Kirby, 1892 Afrotheora Nielsen and Scoble, 1986 Antihepialus Janse, 1942 =Ptycholoma; Felder, 1874 Bipectilis Chus and Wang, 1985 Palpifer Hampson, =Palpiphorus; Quail, 1900 =Palpiphora; Pagenstacher, 1909 Eudalaca Viette, 1950 =Eudalacina Paclt, 1953 Gorgopis Hübner, =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, 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, 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, Zenophassus Tindale, 1941 Sthenopis auctt. nec Packard, 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, Parahepialiscus Viette, 1950 Xhoaphryx Viette, 1953 Aenetus Herrich-Schäffer, =Charagia Walker, 1856 =Phloiopsyche Scott, 1864 =Oenetus; Kirby, 1892 =Choragia; Pagenstacher, 1909 =Oenetes; Oke, 1953 Leto Hübner, =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, =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