Hepiolus Illiger, 1801 Epialus Agassiz, 1847 Epiolus Agassiz, 1847 Tephus Wallengren, 1869
Noctua humuli Linnaeus, 1758 Hepialus thulensis Newman, 1865 Hepialus humulator Haworth, 1802 Hepialus hethlandica Staudinger, 1871 Hepialus rosea Petersen, 1902 Hepialus albida Spuler, 1910 Hepialus azuga Pfitzner, 1912 Hepialus grandis Pfitzner, 1912 Hepialus dannenbergi Stephan, 1923 Hepialus pusillus Stephan, 1923 Hepialus rufomaculata Lempke, 1938 Hepialus albida Bytinski-Salz, 1939 Hepialus roseoornata Bytinski-Salz, 1939 Hepialus uniformis Bytinski-Salz, 1939 Hepialus faeroensis Dahl, 1954 Hepialus fumosa Cockayne, 1955 Hepialus radiata Cockayne, 1955 Hepialus postnigrescens Lempke, 1961 Hepialus postrufescens Lempke, 1961 Hepialus griseomaculata van Wisselingh, 1965 Hepialus thuleus
The ghost moth (Hepialus humuli), also known as the ghost swift, 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 differences in size and wing color. The adults fly from June to August and are attracted to light. The species overwinters as a larva. The term ghost moth is sometimes used as a general term for all Hepialids. The ghost moth gets its name from the hovering display flight of the male, sometimes slowly rising and falling, over open ground to attract females. In a suitable location several males may display together in a lek. The larva is whitish and maggot-like and feeds underground on the roots of a variety of wild and cultivated plants (see list below). The species can be an economically significant pest in forest nurseries.
1 Physiology and description 2 Mating
2.1 Male/male interactions
2.1.1 Lekking 2.1.2 Displaying
2.2 Female/male interactions
3 Sexual Dimorphism 4 Predators 5 Life history
5.1 Egg 5.2 Larva 5.3 Pupa 5.4 Adult
6 Damage to crops 7 Ghost moth in folklore 8 Subspecies 9 Recorded food plants 10 Additional species which may be included in Hepialus 11 Gallery 12 References 13 External links
Physiology and description Female ghost moths have a wingspan of 50-70mm. They have yellowish-buff forewings with darker linear markings and brown hindwings. Males are smaller, with a wingspan of 46-50mm, and typically have white or silver wings. However, in H. h. thulensis, found in Shetland and the Faroe Islands, there are buff-coloured individuals. Mating Male/male interactions Lekking The ghost swift aggregates in leks in order to attract female mates. Lekking occurs at dusk and typically lasts for 20–30 minutes. During the lekking period, incident light intensities between 10.0 and 2.0 lux have been found to increase the brightness contrast between the background (grass/plants) and male moths' silver/white wings. It is thus believed that the male wing color may have evolved as a secondary adaptation to aid in the moth's visibility. Displaying The male ghost swifts display by hovering directly above vegetation, while occasionally shifting slowly horizontally. The displaying male only occasionally made vertical movements to shift display positions. Females are attracted to the displaying males in leks, and once a female chooses a male she will pass within a few centimeters of him. The male will follow the female, who will land and beat her wings, signaling that the male may approach her. The two moths will then copulate, and the male moth may return to the lek and display again afterwards. Female/male interactions Pheromones Males perform a flight display and use both chemical and visual signals to attract females to their mating sites. In addition to aggregating in leks, male ghost swifts also use pheromones to attract a mate. The pheromones are emitted in order to attract a female, but they are not used as an aphrodisiac. The main component of the male pheromone is (E,E)-a-Famesene.
The olfactory substances used to attract females are produced on the male's hind-tibial brushes. Males may also be attracted to stationary females by olfactory stimuli. In addition, the male ghost moths produce a goat-like smell from their hind tibiae while in flight
Female ghost moth (left) and male ghost moth (right)
The ghost moth displays high levels of sexual dichromatism (see picture below). Female ghost moths are a yellow brownish color, while males have silver/white wings. It has been suggested that the difference in wing color between males and females is used for visual epidemic signaling. The upper side of males have un-pigmented scales with elaborate morphology and meshwork that allow for light reflection and may aid in attracting females. The females lack the intricate morphology of the males. The underside of both the male and female ghost moth is a uniform grey/brown color. It is believed that there is behavioral dimorphism as well, with one study showing that females were more attracted to light than males. Predators Common predators of ghost moths include several species of bats and birds. These predators are attracted to the moths during the male flight displays. Eptesicus nilssonii, the northern bat, has often been documented preying on lekking ghost moths. The ghost moth is a member of the family Hepialidae, an early branch of Lepidoptera. Species in the Hepialidae lack several predator defense systems, including ultrasonic hearing. The ghost moth lacks sophisticated predator defense systems, and instead the ghost moth restricts its sexual behavior to a short period during dusk to reduce its predation risk. Despite these precautions, the moth is still at a large predation risk. It is believed that the deaf moths, such as the family Hepialidae, predate the predatory bats that may have driven the evolution of ultrasonic hearing. Members of the Hepialidae also lack adaptations that allow for erratic and maneuverable flight as well as color mimicking, two traits that aid in predator defense. It is currently believed that the ghost moth’s restricted flight patterns and low flight positions may be the ghost swift’s main form of predatory defense. The ghost moth displays for only 20–30 minutes at dusk, which aids in predator avoidance, as most bats typically do not start feeding until after dusk, when it is darker outside. Life history Egg On average, most female ghost moths will lay around 600 eggs over four days, but a female can lay anywhere from 200 to 1,600 eggs. Larva The ghost moth larvae grow up to 50mm long and have a white opaque body with a red/brown head. Their prothoracic plate is also red/brown, and their pinacula is dark brown. The young larvae feed on plant rootlets, while the older large feed on larger roots, stolons, and the lower regions of plant stems. The larval growth is very slow, and the developmental period can last for two to three years. The larva have at least 12 instars, but further research is needed to see if there may be more instars during higher temperatures. The larva cause damage to the plants they consume, with damage being the worst during the second summer of the larva’s growth period. The larvae typically feed in grasslands, lawns, and pastures and have been known to cause significant damage to the host species. In the British Isles, the ghost moth larvae live in the soil and can commonly be found underneath the grass. Pupa The ghost moth typically pupates during the April or May after the two to three year larval growth period has ended. Adult Adults are most commonly found in June and July, and they are typically active around dusk. In particular, lekking and mating occurs at dusk. Damage to crops Ghost moth larvae are polyphagous—they are able to feed on multiple types of plants. Larvae born on crops will typically feed on the roots and can cause significant damage to the crops. Ghost moths frequently cause damage to strawberries, lettuce, and chrysanthemum plants. Ghost moth in folklore It is believed that the common name of “ghost moth” for Hepialus humuli may have originated from European folklore, as there are numerous references to white moths being the souls of the departed. It is believed that the ghost moth is also referenced in the last passage of Wuthering Heights by Emily Brontë. Subspecies
Hepialus humuli humuli Hepialus humuli thulensis Newman, 1865 (Great Britain, Faroe Islands)
Recorded food plants
Arctium - burdock Asparagus Beta - beet Brassica Cannabis Chrysanthemum
Dahlia Daucus - carrot Fagus - beech Fragaria - strawberry Helianthus - Jerusalem artichoke Humulus - hop
Lactuca - lettuce Pastinaca - parsnip Phaseolus - bean Pisum - pea Poaceae - grasses Quercus - oak
Rumex - dock/sorrel Solanum - potato Taraxacum officinale - Dandelion Urtica - nettle Fungi
Additional species which may be included in Hepialus Chinese authors retain a number of species in Hepialus. Most of these are placed in the genus Thitarodes by others. Species retained in Hepialus include:
Hepialus bibelteus F.R. Shen & Y.S. Zhou, 1997 Hepialus biruensis S.Q. Fu, 2002 Hepialus deqinensis X.C. Liang, 1988 Hepialus haimaensis X.C. Liang, 1988
Hepialus latitegumenus F.R. Shen & Y.S. Zhou, 1997 Hepialus pui G.R. Zhang, D.X. Gu & X. Liu, 2007 Hepialus xiaojinensis Y.Q. Tu, K.S. Ma & D.L. Zhang, 2009
Ghost moth on a Faroese stamp
^ a b c d e f g h Alford, David V. (2012-05-30). Pests of Ornamental Trees, Shrubs and Flowers: A Colour Handbook, Second Edition. CRC Press. ISBN 9781840766288. ^ a b c d Andersson, S.; Rydell, J.; Svensson, M. G. E. (1998-07-22). "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. ISSN 0962-8452. PMC 1689211 . ^ a b c d e f Edwards, C. A. (May 1964). "The bionomics of swift moths. I.—The ghost swift moth, Hepialus humuli (L.)". Bulletin of Entomological Research. 55 (1): 147–160. doi:10.1017/S000748530004935X. ISSN 1475-2670. ^ a b c Rydell, Jens (1998-08-07). "Bat defence in lekking ghost swifts (Hepialus humuli), a moth without ultrasonic hearing". Proceedings of the Royal Society of London B: Biological Sciences. 265 (1404): 1373–1376. doi:10.1098/rspb.1998.0444. ISSN 0962-8452. PMC 1689220 . PMID 9721686. ^ Mallet, James (1984). "Sex roles in the ghost moth Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera)". Zoological Journal of the Linnean Society. 80 (1): 67–82. doi:10.1111/j.1096-3642.1984.tb02320.x. ISSN 0024-4082. ^ a b Turner, John R. G. (1976-02-01). "Sexual behaviour: female swift moth is not the aggressive partner". Animal Behaviour. 24 (1): 188–190. doi:10.1016/S0003-3472(76)80113-3. ^ a b c d Mallet, James (1984-01-01). "Sex roles in the ghost moth Hepialus humuli (L.) and a review of mating in the Hepialidae (Lepidoptera)". Zoological Journal of the Linnean Society. 80 (1): 67–82. doi:10.1111/j.1096-3642.1984.tb02320.x. ISSN 1096-3642. ^ Schulz, S.; Francke, W.; König, W. A.; Schurig, V.; Mori, K.; Kittmann, R.; Schneider, D. (December 1990). "Male pheromone of swift moth, Hepialus hecta L. (Lepidoptera: Hepialidae)". Journal of Chemical Ecology. 16 (12): 3511–3521. doi:10.1007/BF00982114. ISSN 0098-0331. PMID 24263445. ^ a b "Sexual dimorphism and geographical male polymorphism in the ghost moth Hepialus humuli (Lepidoptera: Hepialidae): Scale ultrastructure and evolutionary aspects (PDF Download Available)". ResearchGate. Retrieved 2017-10-25. ^ a b "Moth by Matthew Gandy from Reaktion Books". www.reaktionbooks.co.uk. Retrieved 2017-10-25.
Chinery, Michael: Collins Guide to the Insects of Britain and Western Europe, 1986 (Reprinted 1991) Skinner, Bernard: Colour Identification Guide to Moths of the British Isles, 1984
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Ghost moth up UKmoths Fauna Europaea Lepiforum.de
Wd: Q476878 EoL: 956497 EPPO: HEPIHU Fauna Europaea: 431770 GBIF: 1828637 iNaturalist: 205224 LepIndex: 137124.0