The Hepialidae are a

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

insects Insects (from Latin ') are hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of jointed ...

in the

lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

. 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 Order () is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between Family_(biology), family and Class_(biology), class. In biological classification, the order is a taxonomic rank used in the classific ...

Exoporia The Exoporia are a group of primitive Lepidoptera comprising the superfamilies Mnesarchaeoidea and Hepialoidea.Nielsen, E.S., Robinson, G.S. and Wagner, D.L. 2000. Ghost-moths of the world: a global inventory and bibliography of the Exoporia (M ...

. The 82

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

contain at least 700 currently recognised

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

of these primitive

moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...

s worldwide. The genera ''

Fraus In Roman mythology, Fraus was the goddess of personification of treachery and fraud. She was daughter of Orcus and Night ( Nyx). She was depicted with a woman's face, the body of a snake, and on her tail the sting of a scorpion.Johann Joachim Esc ...

'' (

endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

), ''

Gazoryctra ''Gazoryctra'' is a genus of moths of the family Hepialidae. There are 14 described species found in Eurasia, Canada and the United States. Species *'' Gazoryctra chishimana'' – Kuril Islands *'' Gazoryctra confusus'' – United States *'' G ...

'' (

Holarctic The Holarctic realm is a biogeographic realm that comprises the majority of habitats found throughout the continents in the Northern Hemisphere. It corresponds to the floristic Boreal Kingdom. It includes both the Nearctic zoogeographical reg ...

), ''

Afrotheora ''Afrotheora'' is a genus of moths of the family Hepialidae. There are 7 described species, all found in southern Africa. They are considered to be one of the more primitive genera of the Hepialidae, with short antennae and lacking a functional p ...

'' (

Southern Africa Southern Africa is the southernmost region of Africa. No definition is agreed upon, but some groupings include the United Nations geoscheme for Africa, United Nations geoscheme, the intergovernmental Southern African Development Community, and ...

n), and '' Antihepialus'' (

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

n) are considered to be the most primitive, containing four genera and about 51 species with a mostly relictual southern

Gondwanan Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia, Zealandia, Arabia, and the ...

distribution and are currently separated from the Hepialidae ''sensu stricto'' which might form a natural, derived group. The most diverse genera are ''

Oxycanus ''Oxycanus'' is a genus of moths in the family Hepialidae. There are 71 described species found in Australia and New Guinea. Species *'' Oxycanus aedisima'' – Australia *'' Oxycanus aegrus'' – New Guinea *'' Oxycanus albostrigata'' – Austr ...

'' with 78 species, ''

Endoclita ''Endoclita'' is a genus of moths of the family Hepialidae. There are 60 described species found in eastern and southeast Asia and the Indian subcontinent. Species *'' Endoclita aboe'' – India *'' Endoclita absurdus'' – China *'' Endoclita ...

'' with 78 species, and''

Thitarodes ''Thitarodes'' is a genus of moths of the family Hepialidae. In English ''Thitarodes'' is known as "ghost moth". They are found in eastern Asia. The majority are restricted to the Tibetan Plateau. Often in Chinese entomological nomenclature ''Thi ...

'' 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 synonym (taxonomy), 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 A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...

frenulum A frenulum or frenum (: frenula or frena, from the Latin ''frēnulum'', "little bridle", the diminutive of ''frēnum'') is a small fold of tissue that secures the motion of a mobile organ in the body. In human anatomy Frenula on the human ...

(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 Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...

is transferred to the egg by an external channel between the ostium and the

ovipore An ovipore is a pore-like sexual organ of a female insect that is insemination, inseminated by the spermatophores ejected by the aedeagus of a male insect during copulation (zoology), copulation. The spermatophores that pass through the ovipore ar ...

. Other non

ditrysia Ditrysia is a clade of lepidopterans that contains both butterflies and a majority of moth species. They are named for the fact that the female has two distinct sexual openings: one for mating, and the other for laying eggs. About 98% of know ...

n moths have a common

cloaca A cloaca ( ), : cloacae ( or ), or vent, is the rear orifice that serves as the only opening for the digestive (rectum), reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles, birds, cartilagin ...

. The moths are homoneurous with similar forewings and hindwings, and are sometimes included as 'honorary' members of the

Macrolepidoptera Macrolepidoptera is a group within the insect Order (biology), order Lepidoptera. Traditionally used for the larger butterflies and moths as opposed to the "microlepidoptera", this group is artificial. However, it seems that by moving some taxa ...

, though archaic they are. Strictly speaking, they are

phylogenetically In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical data ...

too basal and constitute

Microlepidoptera Microlepidoptera (micromoths) is an artificial (i.e., unranked and not monophyletic) grouping of moth families, commonly known as the "smaller moths" ( micro, Lepidoptera). These generally have wingspans of under 20 mm, so are harder to iden ...

, 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 Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...

, with males smaller but more boldly marked than females, or at high

elevation The elevation of a geographic location (geography), ''location'' is its height above or below a fixed reference point, most commonly a reference geoid, a mathematical model of the Earth's sea level as an equipotential gravitational equipotenti ...

, females of '' Pharmacis'' and ''

Aoraia ''Aoraia'' is a genus of moths of the family Hepialidae. There are 13 described species, all endemic to New Zealand. The type species of this genus is ''Porina dinodes'' Meyrick, 1890. This genus contains some large species with a wingspan of up ...

'' show "brachypterous" wing reduction.

Distribution

Hepialidae are distributed on ancient landmasses worldwide except

Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean (also known as the Antarctic Ocean), it contains the geographic South Pole. ...

but with the surprising exceptions of

Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...

, the

Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...

islands and in Africa, tropical

West Africa West Africa, also known as Western Africa, is the westernmost region of Africa. The United Nations geoscheme for Africa#Western Africa, United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Gha ...

. It remains to be borne out if these absences are real as '' Aenetus cohici'' was not long ago discovered in

New Caledonia New Caledonia ( ; ) is a group of islands in the southwest Pacific Ocean, southwest of Vanuatu and east of Australia. Located from Metropolitan France, it forms a Overseas France#Sui generis collectivity, ''sui generis'' collectivity of t ...

. In the

Oriental The Orient is a term referring to the East in relation to Europe, traditionally comprising anything belonging to the Eastern world. It is the antonym of the term ''Occident'', which refers to the Western world. In English, it is largely a meto ...

and

Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...

regions hepialids have diversified in rainforest environments, but this not apparently the case in the

Afrotropics The Afrotropical realm is one of the Earth's eight biogeographic realms. It includes Sub-Saharan Africa, the southern Arabian Peninsula, the island of Madagascar, and the islands of the western Indian Ocean. It was formerly known as the Ethiopi ...

. Hepialids mostly have low dispersive powers and do not occur on oceanic islands with the exception of ''Phassodes'' on

Fiji Fiji, officially the Republic of Fiji, is an island country in Melanesia, part of Oceania in the South Pacific Ocean. It lies about north-northeast of New Zealand. Fiji consists of an archipelago of more than 330 islands—of which about ...

and

Western Samoa Samoa, officially the Independent State of Samoa and known until 1997 as Western Samoa, is an island country in Polynesia, part of Oceania, in the South Pacific Ocean. It consists of two main islands ( Savai'i and Upolu), two smaller, inhabit ...

and a few species in

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

and

Kurile Islands The Kuril Islands or Kurile Islands are a volcanic archipelago administered as part of Sakhalin Oblast in the Russian Far East. The islands stretch approximately northeast from Hokkaido in Japan to Kamchatka Peninsula in Russia, separating the ...

. Whilst the type locality of '' Eudalaca sanctahelena'' is from the remote island of

St Helena Saint Helena (, ) is one of the three constituent parts of Saint Helena, Ascension and Tristan da Cunha, a remote British overseas territory. Saint Helena is a volcanic and tropical island, located in the South Atlantic Ocean, some 1,874 km ...

, this is thought to be an error for South Africa.

Behaviour

Swift moths are usually

crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...

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 A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...

from

scales Scale or scales may refer to: Mathematics * Scale (descriptive set theory), an object defined on a set of points * Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original * Scale factor, a number ...

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

s have been analysed.

Biology

The female does not lay its

eggs An egg is an organic vessel in which an embryo begins to develop. Egg, EGG or eggs may also refer to: Biology * Egg cell, the female reproductive cell (gamete) in oogamous organisms Food * Eggs as food Places * Egg, Austria * Egg, Switzerland ...

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

larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...

e feed in a variety of ways. Probably all

have concealed larvae, making silken tunnels in all manner of substrates. Some species feed on leaf litter,

fungi A fungus (: fungi , , , or ; or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified as one ...

mosses Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta ('' sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and ho ...

, decaying vegetation,

fern The ferns (Polypodiopsida or Polypodiophyta) are a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. They differ from mosses by being vascular, i.e., having specialized tissue ...

gymnosperms The gymnosperms ( ; ) are a group of woody, perennial Seed plant, seed-producing plants, typically lacking the protective outer covering which surrounds the seeds in flowering plants, that include Pinophyta, conifers, cycads, Ginkgo, and gnetoph ...

and a wide span of

monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are flowering plants whose seeds contain only one Embryo#Plant embryos, embryonic leaf, or cotyledon. A monocot taxon has been in use for several decades, but ...

and

dicot The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, ...

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 Feeding is the process by which organisms, typically animals, obtain food. Terminology often uses either the suffixes -vore, -vory, or -vorous from Latin ''vorare'', meaning "to devour", or -phage, -phagy, or -phagous from Greek φαγε ...

". 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

root In vascular plants, the roots are the plant organ, organs of a plant that are modified to provide anchorage for the plant and take in water and nutrients into the plant body, which allows plants to grow taller and faster. They are most often bel ...

s, at least in early

instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...

s 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. Th
pupa
has rows of

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage The fus ...

spines on the abdominal segments as in other lower members of the

Heteroneura Heteroneura is a natural group (or clade) in the insect order Lepidoptera that comprises over 99% of all butterflies and moths. This is the sister group of the infraorder Exoporia The Exoporia are a group of primitive Lepidoptera comprising ...

Economic significance

Chinese medicine makes considerable use of the "mummies" collected of the

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...

-attacking fungi ''

Ophiocordyceps sinensis ''Ophiocordyceps sinensis'' (synonym (taxonomy), synonym ''Cordyceps sinensis''), known colloquially as caterpillar fungus, is an entomopathogenic fungus (a fungus that grows on insects) in the family Ophiocordycipitaceae. It is mainly found ...

'', and these can form an expensive ingredient. The

witchetty grub The witchetty grub (also spelled witchety grub or witjuti grub) is a term used in Australia for the large, white, wood-eating larvae of several moths. In particular, it applies to the larvae of the Cossidae, cossid moth ''Endoxyla leucomochla'' ...

(which are sometimes hepialid larvae) is a popular food source especially among aboriginal Australians. In

Central America Central America is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

, hepialid larvae are also eaten. However, some species of ''

Wiseana ''Wiseana'' is a genus of ghost moths, collectively known as porina, of the family (biology), family Hepialidae. There are seven described species, all endemic (ecology), endemic to New Zealand. Some species of this genus are a major pest in New ...

'', '' Oncopera'', ''

'', ''

'' and ''

Dalaca ''Dalaca'' is a genus of moths of the family Hepialidae. There are 23 described species found throughout South America as far north as Panama. The larvae feed on grasses. Species *''Dalaca chilensis'' - Chile *''Dalaca chiriquensis'' - Panama *' ...

'' 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 John Henry Comstock (February 24, 1849 – March 20, 1931) was an eminent researcher in entomology and arachnology and a leading educator. His work provided the basis for classification of butterflies, moths, and scale insects. Career Comstock w ...

(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 identificationSkinner, B. (1984). ''Colour Identification Guide to Moths of the British Isles'' *'' Gazoryctra fuscoargenteus'' O. Bang-Haas 1927 – Northern

Scandinavia Scandinavia is a subregion#Europe, subregion of northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. ''Scandinavia'' most commonly refers to Denmark, Norway, and Sweden. It can sometimes also ...

*'' Gazoryctra ganna'' (Hübner 1808) –

Alps The Alps () are some of the highest and most extensive mountain ranges in Europe, stretching approximately across eight Alpine countries (from west to east): Monaco, France, Switzerland, Italy, Liechtenstein, Germany, Austria and Slovenia. ...

, northern Scandinavia, northern

Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...

*'' Hepialus humuli'' Linnaeus 1758 (

ghost moth The ghost moth or ghost swift (''Hepialus humuli'') is a moth of the family Hepialidae. It is common throughout Europe, except for the far south-east. Female ghost moths are larger than males, and exhibit sexual dimorphism with their differenc ...

) – Europe *'' Korscheltellus lupulina'' Linnaeus 1758 (

common swift The common swift (''Apus apus'') is a medium-sized bird, superficially similar to the barn swallow or Common house martin, house martin but somewhat larger, though not stemming from those passerine species, being in the order Apodiformes. The re ...

) – Europe *'' Pharmacis aemiliana'' Costantini 1911 –

Italy Italy, officially the Italian Republic, is a country in Southern Europe, Southern and Western Europe, Western Europe. It consists of Italian Peninsula, a peninsula that extends into the Mediterranean Sea, with the Alps on its northern land b ...

*'' Pharmacis anselminae'' Teobaldelli 1977 – Italy *'' Pharmacis bertrandi'' Le Cerf 1936 –

France France, officially the French Republic, is a country located primarily in Western Europe. Overseas France, Its overseas regions and territories include French Guiana in South America, Saint Pierre and Miquelon in the Atlantic Ocean#North Atlan ...

*'' Pharmacis carna'' Denis & Schiffermüller 1775 – Central and Eastern Europe *'' Pharmacis castillana'' Oberthür 1883 –

Spain Spain, or the Kingdom of Spain, is a country in Southern Europe, Southern and Western Europe with territories in North Africa. Featuring the Punta de Tarifa, southernmost point of continental Europe, it is the largest country in Southern Eur ...

*'' Pharmacis claudiae'' Kristal & Hirneisen 1994 – Italy *'' Pharmacis fusconebulosa'' De Geer 1778 ( map-winged swift) – Europe *'' Pharmacis pyrenaicus'' Donzel 1838 –

Pyrenees The Pyrenees are a mountain range straddling the border of France and Spain. They extend nearly from their union with the Cantabrian Mountains to Cap de Creus on the Mediterranean coast, reaching a maximum elevation of at the peak of Aneto. ...

*'' Phymatopus hecta'' Linnaeus 1758 ( gold swift) – Central and northern Europe *'' Triodia adriaticus'' Osthelder 1931 –

Croatia Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...

North Macedonia North Macedonia, officially the Republic of North Macedonia, is a landlocked country in Southeast Europe. It shares land borders with Greece to the south, Albania to the west, Bulgaria to the east, Kosovo to the northwest and Serbia to the n ...

Greece Greece, officially the Hellenic Republic, is a country in Southeast Europe. Located on the southern tip of the Balkan peninsula, it shares land borders with Albania to the northwest, North Macedonia and Bulgaria to the north, and Turkey to th ...

Crete Crete ( ; , Modern Greek, Modern: , Ancient Greek, Ancient: ) is the largest and most populous of the Greek islands, the List of islands by area, 88th largest island in the world and the List of islands in the Mediterranean#By area, fifth la ...

*'' Triodia amasinus'' Herrich-Schäffer 1851 –

Balkans The Balkans ( , ), corresponding partially with the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throug ...

*'' Triodia sylvina'' Linnaeus 1761 (

orange swift The orange swift or orange moth (''Triodia sylvina'') is a moth belonging to the family Hepialidae. The species was first described by Carl Linnaeus in 1761 and was previously placed in the genus ''Hepialus''. It is distributed throughout Europ ...

) – Europe

Generic checklist

*''

'' Walker, 1856 **=''Hectomanes'' Meyrick, 1980 **=''Praus''; Pagenstacher, 1909 *''

'' Hübner, 820**=''Garzorycta''; Hübner, 826**=''Gazoryctes''; Kirby, 1892 *''

'' Nielsen and Scoble, 1986 *'' Antihepialus'' Janse, 1942 **=''Ptycholoma''; Felder, 1874 *'' Bipectilis'' Chus and Wang, 1985 *'' Palpifer'' Hampson, 893**=''Palpiphorus''; Quail, 1900 **=''Palpiphora''; Pagenstacher, 1909 *''

Eudalaca ''Eudalaca'' is a genus of moths of the family Hepialidae. There are 35 described species, most restricted to South Africa but a few found further north in the continent. Species *'' Eudalaca aequifascia'' - East Africa *'' Eudalaca albiplumis'' ...

'' Viette, 1950 **=''Eudalacina'' Paclt, 1953 *''

Gorgopis ''Gorgopis'' is a genus of moths of the family Hepialidae. There are 28 described species found in southern and eastern Africa. Species *'' Gorgopis alticola'' - Tanzania *'' Gorgopis angustiptera'' *'' Gorgopis annulosa'' - South Africa *'' Gor ...

'' Hübner, 820**=''Gorcopis''; Walker, 1856 *'' Metahepialus'' Janse, 1942 *''

'' Walker, 1856 **=''Huapina'' Bryk, 1945 **=''Maculella'' Viette, 1950 **=''Toenga'' Tindale, 1954 *'' Callipielus'' Butler, 1882 **=''Stachyocera'' Ureta, 1957 *''

Blanchardinella ''Blanchardinella'' is a genus of moths of the family Hepialidae. It consists of only one species, ''Blanchardinella venosus'', which is found in Chile. The genus was named in honour of Émile Blanchard Charles Émile Blanchard (6 March 1819 ...

'' Nielsen, Robinson & Wagner, 2000 **=''Blanchardina'' Viette, 1950, nec Labbe, 1899 *''

Calada ''Calada'' is a genus of moths of the family Hepialidae. There are two described species, both endemic to Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , ...

'' Nielsen and Robinson, 1983 *'' Puermytrans'' Viette, 1951 *'' Parapielus'' Viette, 1949 **=''Lossbergiana'' Viette, 1951 *''

Andeabatis ''Andeabatis'' is a monotypic moth genus of the family Hepialidae. The only species is ''Andeabatis chilensis'' of southern South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisph ...

'' Nielsen and Robinson, 1983 *'' Druceiella'' Viette, 1949 *'' Trichophassus'' Le Cerf, 1919 *'' Phassus'' Walker, 1856 *'' Schausiana'' Viette, 1950 *'' Aplatissa'' Viette, 1953 *'' Pfitzneriana'' Viette, 1952 *''

Cibyra Cibyra or Kibyra (Greek language, Greek: ), also referred to as Cibyra Magna, was an Ancient Greek city near the modern town of Gölhisar, in Burdur Province. It lay outside the north-western limits of the ancient province of Lycia and was th ...

'' Walker, 1856 *''Cibyra'' ('' Pseudodalaca'' Viette, 1950) *''Cibyra'' ('' Gymelloxes'' Viette, 1952) *''Cibyra'' ('' Alloaepytus'' Viette, 1951) *''Cibyra'' ('' Aeptus'') Herrich-Schäffer,

858 __NOTOC__ Year 858 ( DCCCLVIII) was a common year starting on Saturday of the Julian calendar. Events By place Europe * Summer – King Louis the German, summoned by the disaffected Frankish nobles, invades the West Frankish Kin ...

*''Cibyra'' ('' Thiastyx'' Viette, 1951) *''Cibyra'' ('' Schaefferiana'' Viette, 1950) *''Cibyra'' (''

Paragorgopis ''Paragorgopis'' is a genus of picture-winged flies in the family Ulidiidae. Species * ''Paragorgopis argyrata'' * '' Paragorgopis cancellata'' * '' Paragorgopis clathrata'' * '' Paragorgopis discrepans'' * '' Paragorgopis euryale'' * ''Paragorg ...

'' Viette, 1952) *''Cibyra'' ('' Hepialyxodes'' Viette, 1951) *''Cibyra'' ('' Xytrops'' Viette, 1951) *''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 ''Roseala'' is a monotypic moth genus of the family Hepialidae. The only described species is ''R. bourgognei'' which is endemic to Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is ...

'' Viette, 1950 *''

'' auctt., nec Walker, 1856 *'' Pfitzneriella'' Viette, 1951 *''

'' Dumbleton, 1966 **=''Trioxycanus'' Dumbleton, 1966 *'' Triodia'' **=''Alphus'' Wallengren, 1869, nec Dejean, 1833 *'' Korscheltellus'' Börner, 1920 *'' Pharmacis'' Hübner, 820*''

'' Viette, 1968 **=''Forkalus'' Chu and Wang, 1985 *''

Phymatopus ''Phymatopus'' (originally: ''Phalaena Noctua'' Linnaeus, 1758) is a genus of moths of the family Hepialidae (commonly referred to as swift moths or ghost moths), which consists of around 700 species and 82 genera. The genus was erected by Hans ...

'' Wallengren, 1869 **=''Hepiolopsis'' Börner, 1920 **=''Phimatopus''; auctt. *''

'' auctt. nec Wallengren, 1869 *''

Hepialus The ghost moth or ghost swift (''Hepialus humuli'') is a moth of the family Hepialidae. It is common throughout Europe, except for the far south-east. Female ghost moths are larger than males, and exhibit sexual dimorphism with their differenc ...

'' Fabricius, 1775 **=''Hepiolus'' Illiger, 1801 **=''Epialus'' Agassiz, 1847 **=''Epiolus'' Agassiz, 1847 **=''Tephus'' Wallengren, 1869 **=''Trepialus''; Latreille,

805 __NOTOC__ Year 805 ( DCCCV) was a common year starting on Wednesday of the Julian calendar, the 805th year of the Common Era (CE) and Anno Domini (AD) designations, the 805th year of the 1st millennium, the 5th year of the 9th century, and the ...

*'' Zenophassus'' Tindale, 1941 *'' Sthenopis'' auctt. nec Packard,

865 __NOTOC__ Year 865 ( DCCCLXV) was a common year starting on Monday of the Julian calendar. Events By place Europe * King Louis the German divides the East Frankish Kingdom among his three sons. Carloman receives Bavaria (with more l ...

*''

''; Felder, 1874 **=''Endoclyta'', Felder, 1875 **=''Hypophassus'', Le Cerf, 1919 **=''Nevina'', Tindale, 1941 **=''Sahyadrassus'', Tindale, 1941 **=''Procharagia'', Viette, 1949 *'' Neohepialiscus'' Viette, 1948 *''

Elhamma ''Elhamma'' is a genus of moths of the family Hepialidae. There are four described species found in Australia and New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands ...

'' Walker, 1856 **=''Perissectis'' Meyrick, 1890 **=''Pericentris''; Pagenstacher, 1909 **=''Zauxieus'' Viette, 1952 **=''Theaxieus'' Viette, 1952 *'' Jeana'' Tindale, 1935 *''

Cladoxycanus ''Cladoxycanus'' is a monotypic genus of moths belonging to the family Hepialidae. It consists of only one species, ''Cladoxycanus minos'', which is endemic to New Zealand. ''C. minos'' was first described by George Hudson George Hudson (pr ...

'' Dumbleton, 1966 *''

'' Viette, 1961 **=''Porina'' Walker, 1956, nec d'Orbigny, 1852 **=''Gorina''; Quail, 1899 **=''Goryna''; Quail, 1899 **=''Philpottia'' Viette, 1950, nec Broun, 1915 *''

Heloxycanus ''Heloxycanus patricki'', also known as the sphagnum porina moth, is a species of moth of the family Hepialidae, the ghost moths. It is the only member of the genus ''Heloxycanus''. This species is endemic to New Zealand. It has been classified ...

'' Dugdale, 1994 *''

Dumbletonius ''Dumbletonius'' is a genus of moths of the family Hepialidae. There are two described species, both endemic to New Zealand. The genus was named in honour of Lionel Jack Dumbleton Lionel Jack Dumbleton (1905 – 25 September 1976) was a ...

''; auctt **=''Trioxycanus'' Dumbleton, 1966 *'' Dioxycanus'' Dumbleton, 1966 *''

Napialus ''Napialus'' is a genus of moths of the family Hepialidae. There are three described species, all endemic to China. Species *'' Napialus hunanensis'' - Hunan *'' Napialus kulingi'' *'' Napialus chongquingensis'' - Chongqing ChongqingPost ...

'' Chu and Wang, 1985 *''

Hepialiscus ''Hepialiscus'' is a genus of moths of the family Hepialidae. There are four described species found in Taiwan and Nepal. The larvae feed on grass Poaceae ( ), also called Gramineae ( ), is a large and nearly ubiquitous family (biology), fa ...

'' Hampson, 893*'' Parahepialiscus'' Viette, 1950 *'' Xhoaphryx'' Viette, 1953 *''

Aenetus ''Aenetus'' is a genus of moths of the family Hepialidae. There are 24 described species found in Indonesia, New Guinea, New Caledonia, Australia and New Zealand. Most species have green or blue forewings and reddish hindwings, but some are predomi ...

'' Herrich-Schäffer,

**=''Charagia'' Walker, 1856 **=''Phloiopsyche'' Scott, 1864 **=''Oenetus''; Kirby, 1892 **=''Choragia''; Pagenstacher, 1909 **=''Oenetes''; Oke, 1953 *''

Leto In ancient Greek mythology and Ancient Greek religion, religion, Leto (; ) is a childhood goddess, the daughter of the Titans Coeus and Phoebe (Titaness), Phoebe, the sister of Asteria, and the mother of Apollo and Artemis.Hesiod, ''Theogony' ...

'' Hübner, 820**=''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 ''Abantiades'' is a genus of moths of the family Hepialidae. There are 37 described species, all found exclusively in Australia. The group includes some large species with a wingspan of up to 160 mm. The larvae feed on the roots of ''Eucal ...

'' Herrich-Schäffer,

**=''Pielus'' Walker, 1856 **=''Rhizopsyche'' Scott, 1864 *''

'' 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.

External links

*
Tree of Life

Australian Moths Online

Hepialidae of the World – List of Genera and Links to SpeciesLepIndex list of Hepialidae genera and species''Endoclita'' and ''Hepialus'' pheromonesAbstract, counterfeit hepialid mummiesImages of Hepialidae species in New Zealand

{{Authority control Moth families Taxa named by James Francis Stephens