The Hepialidae constitute by far the most diverse group of the infraorderExoporia. The 60 genera contain at least 587 currently recognised species of these primitive moths worldwide. The genera Fraus (endemic to Australia), Gazoryctra (Holarctic), Afrotheora (Southern African), and Antihepialus (African) 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, derived group.[2] The most diverse genera are Oxycanus with 73 species, Endoclita with 60 species, Thitarodes with 51 species and Cibyra with 50 species following a comprehensive catalogue of Exoporia.[2] The relationships of the many genera are not yet well established; see below for an ordered synonymicgeneric checklist,[2] and the Taxobox for navigation.
Morphology and identificationEdit
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).[3] Like other Exoporia the sperm is transferred to the egg by an external channel between the ostium and the ovipore. Other nonditrysian moths have a common cloaca.[2] 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.[2] 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.[4]
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.[5] In the Oriental and Neotropical regions hepialids have diversified in rainforest environments, but this not apparently the case in the Afrotropics.[2] 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.[2]
BehaviourEdit
Swift moths are usually crepuscular and some species form leks, also thought to have arisen independently in the genus Ogygioses (Palaeosetidae).[3] In most genera, males fly swiftly to virgin females that are calling with scent. In other genera, virgin females "assemble" upwind to displaying males,[6] which emit a musky pheromone from scales on the metathoracictibiae. 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.[7] Sometimes they hover singly as if suspended from a thread or flying in a figure of eight motion.[2] The chemical structures of some pheromones have been analysed.[8]
BiologyEdit
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,[9] which is presumably a world record for the Lepidoptera). The maggot-like larvae[10] feed in a variety of ways. Probably all Exoporia have concealed larvae, making silken tunnels in all manner of substrates. Some species feed on leaf litter, fungi,[11]mosses, decaying vegetation, ferns, gymnosperms and a wide span of monocot and dicot plants.[2][12] 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".[2] A few feed on foliage (the austral 'oxyacanine' genera which may drag foliage into their feeding tunnel: Nielsen et al., 2000: 825). Most feed underground on fine roots, at least in early instars and some then feed internally in tunnels in the stem or trunk of their hostplants. Root-feeding larvae travelling through soil make silk-lined tunnels. Before pupating they make a vertical tunnel, which can be up to 10 cm deep, with an exit close to the ground surface.[13] The pupae can then climb up and down to adjust to changes in temperature and flooding.[14] Before the adult moth emerges, the pupa protrudes half way out at the ground surface. The pupa has rows of dorsal spines on the abdominal segments as in other lower members of the Heteroneura.[3]
Economic significanceEdit
Chinese medicine makes considerable use of the "mummies" collected of the caterpillar-attacking fungi Ophiocordyceps sinensis, and these can form an expensive ingredient.[2][15] The witchetty grub (which are sometimes hepialid larvae) is a popular food source especially among aboriginal Australians. In Central America and South America, hepialid larvae are also eaten.[16] However, some species of Wiseana, Oncopera, Oxycanus, Fraus and Dalaca are considered pests of pastures in Australia, New Zealand, and South America.[2]
PhylogenyEdit
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.[17]
^van Nieukerken; et al. (2011). "Order Lepidoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness" (PDF). Zootaxa. 3148: 212–221. doi:10.11646/zootaxa.3148.1.41.
^ abcdefghijklNielsen, Ebbe S.; Robinson, Gaden S.; Wagner, David 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. doi:10.1080/002229300299282. S2CID 86004391.
^ abcKristensen, 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.
^Sattler, Klaus (1991). "A review of wing reduction in Lepidoptera". Bulletin of the British Museum (Natural History), Entomology. 60 (2): 243–288.
^. www.sciencebuff.org. Archived from the original on 27 September 2007. Retrieved 5 April 2018.
^Mallet, James (1984). "Sex roles in the ghost moth Hepialus humuli (L.) with a review of mating in the Hepialidae (Lepidoptera)". Zoological Journal of the Linnean Society. 79: 67–82. doi:10.1111/j.1096-3642.1984.tb02320.x.
^Andersson, S.; Rydell, J.; Svensson, M. G. E. (1998). "Light, predation and the lekking behaviour of the ghost swift Hepialus humuli (L.) (Lepidoptera, Hepialidae)". Proceedings of the Royal Society B: Biological Sciences. 265 (1403): 1345–1351. doi:10.1098/rspb.1998.0440. PMC1689211.
^Schulz, Stefan; Francke, Wittko; König, Wilfried A.; Schurig, Volker; Mori, Kenji; Kittmann, Rolf; Schneider, Dietrich (1990). "Male pheromone of swift moth, Hepialus hecta L. (Lepidoptera: Hepialidae)". Journal of Chemical Ecology. 16 (12): 3511–3521. doi:10.1007/BF00982114. PMID 24263445. S2CID 26903035.
^Tindale, Norman Barnett (1932). "Revision of the Australian ghost moths (Lepidoptera Homoneura, family Hepialidae). part 1". Records of the South Australian Museum. 4: 497–536.
^. www.landcareresearch.co.nz. Archived from the original on 2004-04-23.
^Grehan, J. R. (1989). "Larval feeding habits of the Hepialidae (Lepidoptera)". Journal of Natural History. 23 (4): 803–824. doi:10.1080/00222938900770421.
^H. Buser, W.Huber and R. Joos 2000 Hepialidae – Wurzelbohrer. Pp. 61-96 in Schmetterlinge und ihre Lebensräume. Band 3. Pro Natura, Basel.
^Atijegbe, S. R.; Mansfield, S.; Rostás, M.; Ferguson, C. M.; Worner, S. (2020). "The remarkable locomotory ability of Wiseana (Lepidoptera: Hepialidae) pupae: an adaptation to predation and environmental conditions?". The Wētā. 54: 19–31.
^Wu, Yanru; Yuan, Decheng (1997). "Biodiversity and conservation in China: a view from entomologists". Entomologica Sinica. 4 (2): 95–111. doi:10.1111/j.1744-7917.1997.tb00078.x. S2CID 86294063.
^Ramos-Elorduy, Julieta (2002). "Edible insects of Chiapas, Mexico". Ecology of Food and Nutrition. 41 (4): 271–299. doi:10.1080/03670240214081. S2CID 84107193.
^J. H., Comstock (1893). Evolution of the Wings of Insects. Ithaca, NY: The Wilder Quarter Century Book.
^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
ReferencesEdit
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 linksEdit
Wikimedia Commons has media related to Hepialidae.
Images of Hepialidae species in New Zealand 2015-11-07 at the Wayback Machine
Obituary of Norman B. Tindale
October 23, 2023
hepialidae, family, insects, lepidopteran, order, moths, this, family, often, referred, swift, moths, ghost, moths, gold, swift, male, calling, scientific, classificationdomain, eukaryotakingdom, animaliaphylum, arthropodaclass, insectaorder, lepidopterasuperf. The Hepialidae are a family of insects in the lepidopteran order Moths of this family are often referred to as swift moths or ghost moths HepialidaeGold swift male calling Scientific classificationDomain EukaryotaKingdom AnimaliaPhylum ArthropodaClass InsectaOrder LepidopteraSuperfamily HepialoideaFamily HepialidaeStephens 1829GeneraAbantiades Herrich Schaffer 1858 Aenetus Herrich Schaffer 1858 Afrotheora Nielsen and Scoble 1986 Ahamus Z W Zou amp G R Zhang 2010 Andeabatis Nielsen and Robinson 1983 Antihepialus Janse 1942 Aoraia Dumbleton 1966 Aplatissa Viette 1953 Bipectilus Chus and Wang 1985 Blanchardinella Nielsen Robinson amp Wagner 2000 Bordaia Tindale 1932 Calada Nielsen and Robinson 1983 Callipielus Butler 1882 Cibyra Walker 1856 Cladoxycanus Dumbleton 1966 Dalaca Walker 1856 Dalaca auctt nec Walker 1856 Dioxycanus Dumbleton 1966 Druceiella Viette 1949 Dumbletonius auctt Elhamma Walker 1856 Endoclita Felder 1874 Eudalaca Viette 1950 Fraus Walker 1856 Gazoryctra Hubner 1820 Gorgopis Hubner 1820 Heloxycanus Dugdale 1994 Hepialiscus Hampson 1893 Hepialus Fabricius 1775 Jeana Tindale 1935 Korscheltellus Borner 1920 Leto Hubner 1820 Metahepialus Janse 1942 Napialus Chu and Wang 1985 Neohepialiscus Viette 1948 Oncopera Walker 1856 Oxycanus Walker 1856 Palpifer Hampson 1893 Parahepialus Z W Zou amp D R Zhang 2010 Parahepialiscus Viette 1950 Parapielus Viette 1949 Parathitarodes Ueda 1999 Pfitzneriana Viette 1952 Pfitzneriella Viette 1951 Pharmacis Hubner 1820 Phassodes Bethune Baker 1905 Phassus Walker 1856 Phialuse Viette 1961 Phymatopus Wallengren 1869 Phymatopus auctt nec Wallengren 1869 Puermytrans Viette 1951 Roseala Viette 1950 Schausiana Viette 1950 Sthenopis auctt nec Packard 1865 Thitarodes Viette 1968 Trichophassus Le Cerf 1919 Trictena Meyrick 1890 Triodia Hubner 1820 Wiseana Viette 1961 Xhoaphryx Viette 1953 Zelotypia Scott 1869 Zenophassus Tindale 1941 Oiophassus J F Zhang 1989 Prohepialus Piton 1940 Protohepialus Pierce 1945Diversity 1 62 genera and at least 606 species Contents 1 Taxonomy and systematics 2 Morphology and identification 3 Distribution 4 Behaviour 5 Biology 6 Economic significance 7 Phylogeny 8 Faunas 8 1 Fauna of Europe 9 Generic checklist 10 Cited literature 11 References 12 External linksTaxonomy and systematics EditThe Hepialidae constitute by far the most diverse group of the infraorder Exoporia The 60 genera contain at least 587 currently recognised species of these primitive moths worldwide The genera Fraus endemic to Australia Gazoryctra Holarctic Afrotheora Southern African and Antihepialus African 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 derived group 2 The most diverse genera are Oxycanus with 73 species Endoclita with 60 species Thitarodes with 51 species and Cibyra with 50 species following a comprehensive catalogue of Exoporia 2 The relationships of the many genera are not yet well established see below for an ordered synonymic generic checklist 2 and the Taxobox for navigation Morphology and identification EditThe 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 3 Like other Exoporia the sperm is transferred to the egg by an external channel between the ostium and the ovipore Other nonditrysian moths have a common cloaca 2 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 2 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 4 Distribution Edit nbsp Abantiades latipennis Tasmania AustraliaHepialidae 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 5 In the Oriental and Neotropical regions hepialids have diversified in rainforest environments but this not apparently the case in the Afrotropics 2 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 2 Behaviour EditSwift moths are usually crepuscular and some species form leks also thought to have arisen independently in the genus Ogygioses Palaeosetidae 3 In most genera males fly swiftly to virgin females that are calling with scent In other genera virgin females assemble upwind to displaying males 6 which emit a musky pheromone from scales on the metathoracic tibiae In such cases of sex role reversal there may be visual cues also males of the European ghost swift are possibly the most frequently noticed species being white ghostly and conspicuous when forming a lek at dusk 7 Sometimes they hover singly as if suspended from a thread or flying in a figure of eight motion 2 The chemical structures of some pheromones have been analysed 8 Biology EditThe 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 9 which is presumably a world record for the Lepidoptera The maggot like larvae 10 feed in a variety of ways Probably all Exoporia have concealed larvae making silken tunnels in all manner of substrates Some species feed on leaf litter fungi 11 mosses decaying vegetation ferns gymnosperms and a wide span of monocot and dicot plants 2 12 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 2 A few feed on foliage the austral oxyacanine genera which may drag foliage into their feeding tunnel Nielsen et al 2000 825 Most feed underground on fine roots at least in early instars and some then feed internally in tunnels in the stem or trunk of their hostplants Root feeding larvae travelling through soil make silk lined tunnels Before pupating they make a vertical tunnel which can be up to 10 cm deep with an exit close to the ground surface 13 The pupae can then climb up and down to adjust to changes in temperature and flooding 14 Before the adult moth emerges the pupa protrudes half way out at the ground surface The pupa has rows of dorsal spines on the abdominal segments as in other lower members of the Heteroneura 3 Economic significance EditChinese medicine makes considerable use of the mummies collected of the caterpillar attacking fungi Ophiocordyceps sinensis and these can form an expensive ingredient 2 15 The witchetty grub which are sometimes hepialid larvae is a popular food source especially among aboriginal Australians In Central America and South America hepialid larvae are also eaten 16 However some species of Wiseana Oncopera Oxycanus Fraus and Dalaca are considered pests of pastures in Australia New Zealand and South America 2 Phylogeny EditThe 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 17 Faunas EditFauna of Europe Edit Source 18 and identification 19 20 Gazoryctra fuscoargenteus O Bang Haas 1927 Northern Scandinavia Gazoryctra ganna Hubner 1808 Alps northern Scandinavia northern Russia Hepialus humuli Linnaeus 1758 ghost moth Europe Korscheltellus lupulina Linnaeus 1758 common swift Europe Pharmacis aemiliana Costantini 1911 Italy Pharmacis anselminae Teobaldelli 1977 Italy Pharmacis bertrandi Le Cerf 1936 France Pharmacis carna Denis amp Schiffermuller 1775 Central and Eastern Europe Pharmacis castillana Oberthur 1883 Spain Pharmacis claudiae Kristal amp Hirneisen 1994 Italy Pharmacis fusconebulosa De Geer 1778 map winged swift Europe Pharmacis pyrenaicus Donzel 1838 Pyrenees Phymatopus hecta Linnaeus 1758 gold swift Central and northern Europe Triodia adriaticus Osthelder 1931 Croatia North Macedonia Greece Crete Triodia amasinus Herrich Schaffer 1851 Balkans Triodia sylvina Linnaeus 1761 orange swift EuropeGeneric checklist EditFraus Walker 1856 Hectomanes Meyrick 1980 Praus Pagenstacher 1909 Gazoryctra Hubner 1820 Garzorycta Hubner 1826 Gazoryctes Kirby 1892 Afrotheora Nielsen and Scoble 1986 Antihepialus Janse 1942 Ptycholoma Felder 1874 Bipectilis Chus and Wang 1985 Palpifer Hampson 1893 Palpiphorus Quail 1900 Palpiphora Pagenstacher 1909 Eudalaca Viette 1950 Eudalacina Paclt 1953 Gorgopis Hubner 1820 Gorcopis Walker 1856 Metahepialus Janse 1942 Dalaca Walker 1856 Huapina Bryk 1945 Maculella Viette 1950 Toenga Tindale 1954 Callipielus Butler 1882 Stachyocera Ureta 1957 Blanchardinella Nielsen Robinson amp 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 Schaffer 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 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 Borner 1920 Pharmacis Hubner 1820 Thitarodes Viette 1968 Forkalus Chu and Wang 1985 Phymatopus Wallengren 1869 Hepiolopsis Borner 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 Schaffer 1858 Charagia Walker 1856 Phloiopsyche Scott 1864 Oenetus Kirby 1892 Choragia Pagenstacher 1909 Oenetes Oke 1953 Leto Hubner 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 Schaffer 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 1905Cited literature Edit van Nieukerken et al 2011 Order Lepidoptera Linnaeus 1758 In Zhang Z Q Ed Animal biodiversity An outline of higher level classification and survey of taxonomic richness PDF Zootaxa 3148 212 221 doi 10 11646 zootaxa 3148 1 41 a b c d e f g h i j k l Nielsen Ebbe S Robinson Gaden S Wagner David 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 doi 10 1080 002229300299282 S2CID 86004391 a b c 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 Sattler Klaus 1991 A review of wing reduction in Lepidoptera Bulletin of the British Museum Natural History Entomology 60 2 243 288 Buffalo Museum of Science Home www sciencebuff org Archived from the original on 27 September 2007 Retrieved 5 April 2018 Mallet James 1984 Sex roles in the ghost moth Hepialus humuli L with a review of mating in the Hepialidae Lepidoptera Zoological Journal of the Linnean Society 79 67 82 doi 10 1111 j 1096 3642 1984 tb02320 x Andersson S Rydell J Svensson M G E 1998 Light predation and the lekking behaviour of the ghost swift Hepialus humuli L Lepidoptera Hepialidae Proceedings of the Royal Society B Biological Sciences 265 1403 1345 1351 doi 10 1098 rspb 1998 0440 PMC 1689211 Schulz Stefan Francke Wittko Konig Wilfried A Schurig Volker Mori Kenji Kittmann Rolf Schneider Dietrich 1990 Male pheromone of swift moth Hepialus hecta L Lepidoptera Hepialidae Journal of Chemical Ecology 16 12 3511 3521 doi 10 1007 BF00982114 PMID 24263445 S2CID 26903035 Tindale Norman Barnett 1932 Revision of the Australian ghost moths Lepidoptera Homoneura family Hepialidae part 1 Records of the South Australian Museum 4 497 536 Ghost Moth Larva UKmoths Puriri moth NZAC Pare www landcareresearch co nz Archived from the original on 2004 04 23 Grehan J R 1989 Larval feeding habits of the Hepialidae Lepidoptera Journal of Natural History 23 4 803 824 doi 10 1080 00222938900770421 H Buser W Huber and R Joos 2000 Hepialidae Wurzelbohrer Pp 61 96 in Schmetterlinge und ihre Lebensraume Band 3 Pro Natura Basel Atijegbe S R Mansfield S Rostas M Ferguson C M Worner S 2020 The remarkable locomotory ability of Wiseana Lepidoptera Hepialidae pupae an adaptation to predation and environmental conditions The Weta 54 19 31 Wu Yanru Yuan Decheng 1997 Biodiversity and conservation in China a view from entomologists Entomologica Sinica 4 2 95 111 doi 10 1111 j 1744 7917 1997 tb00078 x S2CID 86294063 Ramos Elorduy Julieta 2002 Edible insects of Chiapas Mexico Ecology of Food and Nutrition 41 4 271 299 doi 10 1080 03670240214081 S2CID 84107193 J H Comstock 1893 Evolution of the Wings of Insects Ithaca NY The Wilder Quarter Century Book Fauna Europaea Archived from the original on October 1 2007 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 IslesReferences EditComstock 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 Edit nbsp Wikimedia Commons has media related to Hepialidae nbsp Data related to Hepialidae at Wikispecies Tree of Life Australian Moths Online Hepialidae of Australia Archived 2006 09 27 at the Wayback Machine Hepialidae of the World List of Genera and Links to Species LepIndex list of Hepialidae genera and species Endoclita and Hepialus pheromones Abstract counterfeit hepialid mummies Images of Hepialidae species in New Zealand Archived 2015 11 07 at the Wayback Machine Obituary of Norman B Tindale Retrieved from https en wikipedia org w index php title Hepialidae amp oldid 1180609827, wikipedia, wiki, book, books, library,
