The Hepialidae are a

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

insects Insects (from Latin ') are pancrustacean 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 j ...

in the

lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...

n 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 Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...

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 (Mne ...

. The 60

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

contain at least 587 currently recognised

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

of these primitive

moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...

s worldwide. The genera ''

Fraus In Roman mythology, Fraus was the goddess or personification of treachery and fraud. She was daughter of Orcus Orcus ( la, Orcus) was a god of the underworld, punisher of broken oaths in Etruscan and Roman mythology. As with Hades, the nam ...

'' (

endemic Endemism is the state of a species being found 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 found elsew ...

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

), ''

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 *''Gazo ...

'' (

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 subregion of the African continent, south of the Congo and Tanzania. The physical location is the large part of Africa to the south of the extensive Congo River basin. Southern Africa is home to a number of ...

n), and ''

Antihepialus ''Antihepialus'' is a genus of moths of the family Hepialidae. There are four described species, found in southern and eastern Africa. Species *'' Antihepialus antarcticus'' – South Africa *'' Antihepialus capeneri'' – South Africa *'' An ...

'' (

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

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, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stage ...

distribution and are currently separated from the Hepialidae ''sensu stricto'' which might form a natural,

derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...

group.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): 823–87
Abstract
/ref> The most diverse genera are '' Oxycanus'' with 73 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 60 species, '' Thitarodes'' with 51 species and ''Cibyra'' with 50 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 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 mouthparts used for feeding and sucking. In vertebrates, a proboscis is an elong ...

frenulum A frenulum (or frenum, plural: 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 h ...

(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 is the male reproductive 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 with a tail known as a flagellum, whi ...

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 inseminated by the spermatophores ejected by the aedeagus of a male insect during copulation. The spermatophores that pass through the ovipore are stored in most insect species i ...

. Other non

ditrysia The Ditrysia are a natural group or clade of insects in the lepidopteran order containing both butterflies and moths. They are so named because the female has two distinct sexual openings: one for mating, and the other for laying eggs (in contra ...

n moths have a common

cloaca In animal anatomy, a cloaca ( ), plural cloacae ( or ), is the posterior orifice that serves as the only opening for the digestive, reproductive, and urinary tracts (if present) of many vertebrate animals. All amphibians, reptiles and birds, a ...

. 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 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 about, a monophy ...

, though archaic they are. Strictly speaking, they are phylogenetically 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, and are thus harder to ...

, although hepialids range from very small moths to a wingspan record of 250 mm in ''

Zelotypia ''Zelotypia'' is a monotypic moth genus of the family Hepialidae. The only described species is ''Z. stacyi'', the bentwing ghost moth, which is only found in Queensland and New South Wales, Australia. This is a very large species with a wings ...

''. 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 the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...

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

elevation The elevation of a geographic 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 surface (see Geodetic datum § Vert ...

, females of ''

Pharmacis ''Pharmacis'' is a genus of moths of the family Hepialidae. There are eight described species found in Eurasia. Species *'' Pharmacis aemilianus'' - Italy *'' Pharmacis anselminae'' - Italy *'' Pharmacis bertrandi'' - France *'' Pharmacis carna' ...

'' and '' Aoraia'' 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, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...

but with the surprising exceptions of

Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...

, the

Caribbean The Caribbean (, ) ( es, El Caribe; french: la Caraïbe; ht, Karayib; nl, De Caraïben) is a region of the Americas that consists of the Caribbean Sea, its islands (some surrounded by the Caribbean Sea and some bordering both the Caribbean Se ...

islands and in Africa, tropical

West Africa West Africa or Western Africa is the westernmost region of Africa. The United Nations defines Western Africa as the 16 countries of Benin, Burkina Faso, Cape Verde, The Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Liberia, Mali, Maurit ...

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

New Caledonia ) , anthem = "" , image_map = New Caledonia on the globe (small islands magnified) (Polynesia centered).svg , map_alt = Location of New Caledonia , map_caption = Location of New Caledonia , mapsize = 290px , subdivision_type = Sovereign st ...

. In the

Oriental The Orient is a term for the East in relation to Europe, traditionally comprising anything belonging to the Eastern world. It is the antonym of ''Occident'', the Western World. In English, it is largely a metonym for, and coterminous with, the ...

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 bioge ...

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 Fiji ( , ,; fj, Viti, ; Fiji Hindi: फ़िजी, ''Fijī''), 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 ...

and

Western Samoa Samoa, officially the Independent State of Samoa; sm, Sāmoa, and until 1997 known as Western Samoa, is a Polynesian island country consisting of two main islands ( Savai'i and Upolu); two smaller, inhabited islands ( Manono and Apolima); ...

and a few species in

Japan Japan ( ja, 日本, or , and formally , ''Nihonkoku'') is an island country in East Asia. It is situated in the northwest Pacific Ocean, and is bordered on the west by the Sea of Japan, while extending from the Sea of Okhotsk in the north ...

and

Kurile Islands The Kuril Islands or Kurile Islands (; rus, Кури́льские острова́, r=Kuril'skiye ostrova, p=kʊˈrʲilʲskʲɪjə ɐstrɐˈva; Japanese: or ) are a volcanic archipelago currently administered as part of Sakhalin Oblast in the ...

. Whilst the type locality of ''

Eudalaca sanctahelena ''Eudalaca sanctahelena'' is a species of moth of the family Hepialidae. It was described from Saint Helena Saint Helena () is a British overseas territory located in the South Atlantic Ocean. It is a remote volcanic tropical island west ...

'' is from the remote island of St Helena, 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, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...

and some species form

leks A lek is an aggregation of male animals gathered to engage in competitive displays and courtship rituals, known as lekking, to entice visiting females which are surveying prospective partners with which to mate. A lek can also indicate an avail ...

, also thought to have arisen independently in the genus ''

Ogygioses ''Ogygioses'' is a genus of moths of the family Palaeosetidae. ''Ogygioses luangensis'' is found in Thailand and the other ''Ogygioses'' species are found in Taiwan.Ghost-moths of the world: a global inventory and bibliography of the Exoporia (Mne ...

'' (

Palaeosetidae The Palaeosetidae or miniature ghost moths are a family of insects in the order Lepidoptera contained within the superfamily Hepialoidea. Taxonomy and systematics The Palaeosetidae are a primitive family of Hepialoidea with four currently reco ...

). 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 musky

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 w ...

on the metathoracic

tibiae The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...

. 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 Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...

in a specific location but scatters ("broadcasts") them while in flight, sometimes in huge numbers (29,000 were recorded from a single female ''

Trictena ''Abantiades'' is a genus of moths of the family Hepialidae. There are 14 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 ...

'', which is presumably a world record for the Lepidoptera). The maggot-like

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

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 funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...

, mosses, decaying vegetation,

fern A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...

gymnosperms The gymnosperms ( lit. revealed seeds) are a group of seed-producing plants that includes conifers, cycads, ''Ginkgo'', and gnetophytes, forming the clade Gymnospermae. The term ''gymnosperm'' comes from the composite word in el, γυμνό� ...

and a wide span of

monocot Monocotyledons (), commonly referred to as monocots, (Lilianae ''sensu'' Chase & Reveal) are grass and grass-like flowering plants (angiosperms), the seeds of which typically contain only one embryonic leaf, or cotyledon. They constitute one of ...

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, t ...

plants. There is very little evidence of hostplant specialisation; whilst the South African species ''

Leto venus ''Leto'' is a monotypic moth genus of the family Hepialidae described by Jacob Hübner in 1820. The only described species is ''Leto venus'', described by Pieter Cramer in 1780, which is endemic to South Africa. The larval food plant is ''Virgi ...

'' 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 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 below the sur ...

s, at least in early

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

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 * Dorsal c ...

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 Order ( la, ordo) is one of the eight major hierarchical taxonomic ran ...

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 Sym ...

-attacking fungi ''

Ophiocordyceps sinensis ''Ophiocordyceps sinensis'' (formerly known as ''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 in the mead ...

'', 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 cossid moth ''Endoxyla leucomochla'', which fee ...

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

Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...

and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

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

'' and '' Dalaca'' 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 (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 ''Gazoryctra fuscoargenteus'' is a moth of the family Hepialidae. It is found from northern Scandinavia to Siberia and the Kamchatka Peninsula. The wingspan is 37–42 mm for males and 39–50 mm for females. Adults are on wing from th ...

'' O. Bang-Haas 1927 – Northern

Scandinavia Scandinavia; Sámi languages: /. ( ) is a subregion#Europe, subregion in Northern Europe, with strong historical, cultural, and linguistic ties between its constituent peoples. In English usage, ''Scandinavia'' most commonly refers to Denmark, ...

*''

Gazoryctra ganna ''Gazoryctra ganna'' is a moth of the family Hepialidae. It is found in Sweden, Finland, Russia (Siberia), France, Switzerland, Austria, Italy and Kazakhstan. The wingspan The wingspan (or just span) of a bird or an airplane is the distan ...

'' (Hübner 1808) –

Alps The Alps () ; german: Alpen ; it, Alpi ; rm, Alps ; sl, Alpe . are the highest and most extensive mountain range system that lies entirely in Europe, stretching approximately across seven Alpine countries (from west to east): France, Sw ...

, northern Scandinavia, northern

Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...

*''

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

'' 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 in the far south-east. Female ghost moths are larger than males, and exhibit sexual dimorphism with their differ ...

) – Europe *''

Korscheltellus lupulina The common swift (''Korscheltellus lupulina'') is a moth of the family Hepialidae. It was previously placed in the genus '' Hepialus''. It is a common, often abundant European species. The species was described by Carl Linnaeus in his 1758 10t ...

'' Linnaeus 1758 (

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

) – Europe *''

aemiliana'' Costantini 1911 –

Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...

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

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of Overseas France, overseas regions and territories in the Americas and the Atlantic Ocean, Atlantic, Pacific Ocean, Pac ...

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

castillana'' Oberthür 1883 –

Spain , image_flag = Bandera de España.svg , image_coat = Escudo de España (mazonado).svg , national_motto = ''Plus ultra'' (Latin)(English: "Further Beyond") , national_anthem = (English: "Royal March") , i ...

*'' Pharmacis claudiae'' Kristal & Hirneisen 1994 – Italy *'' Pharmacis fusconebulosa'' De Geer 1778 (

map-winged swift The map-winged swift (''Pharmacis fusconebulosa'') is a moth belonging to the family Hepialidae and has a patchy distribution throughout Eurasia. The species was first described by Charles De Geer in 1778. It was previously placed in the genus ' ...

) – Europe *'' Pharmacis pyrenaicus'' Donzel 1838 –

Pyrenees The Pyrenees (; es, Pirineos ; french: Pyrénées ; ca, Pirineu ; eu, Pirinioak ; oc, Pirenèus ; an, Pirineus) is a mountain range straddling the border of France and Spain. It extends nearly from its union with the Cantabrian Mountains to C ...

*''

Phymatopus hecta The gold swift (''Phymatopus hecta'') is a moth belonging to the family Hepialidae. Until recently it was placed in the genus ''Hepialus''. The species was Species description, first described by Carl Linnaeus in his 1758 10th edition of Systema ...

'' Linnaeus 1758 (

gold swift The gold swift (''Phymatopus hecta'') is a moth belonging to the family Hepialidae. Until recently it was placed in the genus '' Hepialus''. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. Moths ...

) – Central and northern Europe *'' Triodia adriaticus'' Osthelder 1931 –

Croatia , image_flag = Flag of Croatia.svg , image_coat = Coat of arms of Croatia.svg , anthem = "Lijepa naša domovino"("Our Beautiful Homeland") , image_map = , map_caption = , capit ...

North Macedonia North Macedonia, ; sq, Maqedonia e Veriut, (Macedonia before February 2019), officially the Republic of North Macedonia,, is a country in Southeast Europe. It gained independence in 1991 as one of the successor states of Socialist Feder ...

Greece Greece,, or , romanized: ', officially the Hellenic Republic, is a country in Southeast Europe. It is situated on the southern tip of the Balkans, and is located at the crossroads of Europe, Asia, and Africa. Greece shares land borders with ...

Crete Crete ( el, Κρήτη, translit=, Modern: , Ancient: ) is the largest and most populous of the Greek islands, the 88th largest island in the world and the fifth largest island in the Mediterranean Sea, after Sicily, Sardinia, Cyprus, and ...

*''

Triodia amasinus ''Triodia amasinus'' is a species of moth belonging to the family Hepialidae. It was described by Gottlieb August Wilhelm Herrich-Schäffer in 1851, and it is known from Turkey, Hungary, Romania, Bulgaria, the Republic of Macedonia, Albania and ...

'' Herrich-Schäffer 1851 –

Balkans The Balkans ( ), also known as 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 throughout the who ...

*'' 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 Euro ...

) – Europe

Generic checklist

*''

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

'' Hübner,

820 __NOTOC__ Year 820 ( DCCCXX) was a leap year starting on Sunday (link will display the full calendar) of the Julian calendar. Events By place Abbasid Caliphate *Abbasid caliph Al-Ma'mun appointed Isa ibn Yazid al-Juludi as Abbasid gove ...

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

'' Nielsen and Scoble, 1986 *''

'' Janse, 1942 **=''Ptycholoma''; Felder, 1874 *'' Bipectilis'' Chus and Wang, 1985 *''

Palpifer ''Palpifer'' is a genus of moths of the family Hepialidae described by George Hampson in 1893. There are 10 described species found in south and east Asia and parts of Mexico. Species of the genus possess large rounded and ascending palpi. Anten ...

'' Hampson,

893 __FORCETOC__ Year 893 ( DCCCXCIII) was a common year starting on Monday (link will display the full calendar) of the Julian calendar. Events By place Europe * Vladimir, ruler (''khan'') of the Bulgarian Empire, is dethroned by his fat ...

**=''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 *'' Gorg ...

'' Hübner,

**=''Gorcopis''; Walker, 1856 *''

Metahepialus ''Metahepialus'' is a genus of moths of the family Hepialidae. There are two described species, all endemic to South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Afric ...

'' Janse, 1942 *'' Dalaca'' 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 Chile, officially the Republic of Chile, is a country in the western part of South A ...

'' 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 ''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 Hemis ...

'' Nielsen and Robinson, 1983 *'' Druceiella'' Viette, 1949 *'' Trichophassus'' Le Cerf, 1919 *''

Phassus ''Phassus'' is a genus of moths of the family Hepialidae. There are 21 described species distributed from Mexico south to Brazil. Species *''Phassus absyrtus'' - Brazil *''Phassus agrionides'' - Brazil *''Phassus aurigenus'' - Costa Rica *''Phas ...

'' Walker, 1856 *''

Schausiana ''Schausiana'' is a monotypic moth genus of the family Hepialidae described by Pierre Viette in 1950. The only described species is ''Schausiana trojesa'', described by William Schaus in 1901, which is endemic to Mexico. The wingspan The wi ...

'' Viette, 1950 *'' Aplatissa'' Viette, 1953 *'' Pfitzneriana'' Viette, 1952 *''

Cibyra Cibyra or Kibyra (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 the chief city of a ...

'' 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 (link will display the full calendar) of the Julian calendar. Events By place Europe * Summer – King Louis the German, summoned by the disaffected Frankish ...

*''Cibyra'' ('' Thiastyx'' Viette, 1951) *''Cibyra'' ('' Schaefferiana'' Viette, 1950) *''Cibyra'' ('' Paragorgopis'' 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'' Viette, 1950 *'' Dalaca'' auctt., nec Walker, 1856 *'' Pfitzneriella'' Viette, 1951 *'' Aoraia'' Dumbleton, 1966 **=''Trioxycanus'' Dumbleton, 1966 *'' Triodia'' **=''Alphus'' Wallengren, 1869, nec Dejean, 1833 *''

Korscheltellus ''Korscheltellus'' is a genus of moths of the family Hepialidae. It consists of two widespread species, the familiar common swift (''K. lupulina'') of Europe and the conifer swift (''K. gracilis'') of Canada and the eastern United States. ''K. g ...

'' Börner, 1920 *''

'' Hübner,

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

Phymatopus ''Phymatopus'' (originally: ''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 500 species and 30 genera. The genus was erected by Hans Daniel Jo ...

'' 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 in the far south-east. Female ghost moths are larger than males, and exhibit sexual dimorphism with their differ ...

'' 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 (link will display the full calendar) of the Julian calendar. Events By place Byzantine Empire * Siege of Patras: Local Slavic tribes of the Peloponnese lay siege t ...

*'' Zenophassus'' Tindale, 1941 *''

Sthenopis ''Sthenopis'' is a genus of moths of the family Hepialidae. There are eight described species found in North America and China. Species

*''Sthenopis argenteomaculatus'' (silver-spotted ghost moth) – Canada/United States :*Recorded food p ...

'' auctt. nec Packard,

865 __NOTOC__ Year 865 ( DCCCLXV) was a common year starting on Monday (link will display the full calendar) of the Julian calendar. Events By place Europe * King Louis the German divides the East Frankish Kingdom among his three sons. C ...

*''

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

Neohepialiscus ''Neohepialiscus'' is a Monotypic taxon, monotypic moth genus of the family Hepialidae. The only described species is ''N. algeriensis'' of Algeria and Tunisia. References External linksHepialidae genera
Hepialidae Monotypic moth genera ...

'' 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''; id, Papua, or , historically ) is the world's second-largest island wit ...

'' 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 *'' 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 ''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 () or Gramineae () is a large and nearly ubiquitous family of monocotyledonous flower ...

'' Hampson,

*'' 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 (; grc-gre, Λητώ , ''Lētṓ'', or , ''Lātṓ'' in Ancient Greek dialects#Provenance, Doric Greek) is a goddess and the mother of Apollo, the god of music, and Artemis ...

'' Hübner,

**=''Ecto''; Pagenstacher, 1909 *''

'' Scott, 1869 **=''Xylopsyche'' Swainson, 1851 **=''Leto''; auctt *'' Oncopera'' **=''Oncoptera'' Walker, 1890 **=''Paroncopera'' Tindale, 1933 **=''Onchopera''; Birket-Smith, 1974 **=''Onchoptera''; Birket-Smith, 1974 *''

'' Meyrick, 1890 *''

Bordaia ''Bordaia'' is a genus of moths of the family Hepialidae. There are five described species, all endemic to Australia Australia, officially the Commonwealth of Australia, is a sovereign country comprising the mainland of the Australian ...

'' Tindale, 1932 **=''Bordaja''; Chu and Wang, 1985 *''

Abantiades ''Abantiades'' is a genus of moths of the family Hepialidae. There are 14 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 ''Euca ...

'' 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

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 LifeAustralian 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

{{Taxonbar, from=Q580250 Moth families Taxa named by James Francis Stephens