The Hepialidae are a family of insects in the 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
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 (
Endemism to
Australia),
Gazoryctra (
Holarctic),
Afrotheora (
), and
Antihepialus (
) 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,
Synapomorphy 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.
[<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).[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 the sperm is transferred to the egg by an external channel between the ostium and the ovipore. Other non 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.[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,][ The chemical structures of some 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,[ 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 , at least in early 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.[
]
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.[
]
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[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]
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
-
Gorgopis Hübner, 1820
-
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)
-
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
-
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.
External links
