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Ghost moth
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| Ghost moth | |
|---|---|
| Female | |
| Scientific classification | |
| Kingdom: | Animalia |
| Phylum: | Arthropoda |
| Class: | Insecta |
| Order: | Lepidoptera |
| Family: | Hepialidae |
| Genus: | Hepialus Fabricius, 1775 |
| Species: | H. humuli
|
| Binomial name | |
| Hepialus humuli | |
| Synonyms | |
|
List
| |
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 differences in size and wing color. The adults fly from June to August and are attracted to light. The species overwinters as a larva.[1] 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.[2]
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.[3]
Physiology and description
[edit]Female ghost moths have a wingspan of 50–70 mm. They have yellowish-buff forewings with darker linear markings and brown hindwings. Males are smaller, with a wingspan of 46–50 mm, and typically have white or silver wings.[1] However, in H. h. thulensis, found in Shetland and the Faroe Islands, there are buff-coloured individuals.
Mating
[edit]Male/male interactions
[edit]Lekking
[edit]The ghost swift aggregates in leks in order to attract female mates. Lekking occurs at dusk and typically lasts for 20–30 minutes.[3] 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.[4]
Displaying
[edit]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.[5] 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.[6] 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.[6]
Female/male interactions
[edit]Pheromones
[edit]Males perform a flight display and use both chemical and visual signals to attract females to their mating sites.[7] While aggregating in leks, male ghost swifts use pheromones.. These are emitted in order to attract a female, but they are not known to be used as an aphrodisiac. The main component of the male pheromone (in the distantly related species Phymatopus hecta) is (E,E)-α-Farnesene.[8]
The olfactory substances used to attract females are produced on the male's hind tibial brushes, and were said to be "goat-like", meaning the smell of the Goat Moth caterpillar.[7][9] Males may also be attracted to stationary females by olfactory stimuli.
Sexual dimorphism
[edit]
The ghost moth displays high levels of sexual dichromatism (see picture right). 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.[3] The upperside of males have unpigmented scales with elaborate morphology and meshwork that allow for light reflection and may aid in attracting females.[10] 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.[10] It is believed that there is behavioral dimorphism as well, with one study showing that females were more attracted to light than males.[7]
Predators
[edit]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.[4] The ghost moth lacks sophisticated predator defense systems and instead 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, especially at high latitudes where twilight is prolonged.
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. It is currently believed that the ghost moth's restricted flight patterns and low flight positions may be their main form of anti-predator defense.[4] 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, and most birds stop feeding well in advance of sunset.[3]
Life history
[edit]Egg
[edit]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.[2]

Larva
[edit]The ghost moth larvae grow up to 50 mm long and have a white opaque body with a red/brown head.[1] Their prothoracic plate is also red/brown, and their pinacula is dark brown. The young larvae feed on plant rootlets, while the older larvae feed on larger roots, stolons, and the lower regions of plant stems.[1] 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.[2] The larva cause damage to the plants they consume, with damage being the worst during the second summer of the larva's growth period.[1] The larvae typically feed in grasslands, lawns, and pastures and have been known to cause significant damage to the host species.[1] In the British Isles, the ghost moth larvae live in the soil and can commonly be found underneath the grass.[2]
Pupa
[edit]The ghost moth typically pupates during April or May after the two to three-year larval growth period has ended.[1]
Adult
[edit]Adults are most commonly found in June and July, and they are typically active around dusk. In particular, lekking and mating occur at dusk.[1]
Damage to crops
[edit]Ghost moth larvae are polyphagous—they are able to feed on multiple types of plants.[2] 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.[2]
Recorded food plants
[edit]- Arctium - burdock
- Asparagus
- Beta – beet
- Brassica
- Cannabis
- Chrysanthemum
- Dahlia
- Daucus – carrot
- Fagus – beech
- Fragaria – strawberry
- Fungi
- 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
Additional species which may be included in Hepialus
[edit]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
Hepialus humuli was first described by the Swedish taxonomist, Carl Linnaeus, in 1758. He named it Phalaena Noctua humuli — implying night moth of the hop vine. It was later allocated to the genus Hepialus (a fever), which was raised by Johan Christian Fabricius in 1775 and refers to its ″fitful, alternating flight″. The specific name humuli refers to the genus of hops (Humulus), on which Linnaeus wrongly thought the larvae fed on its roots.[11]
Subspecies
[edit]- Hepialus humuli humuli
- Hepialus humuli thulensis Newman, 1865 (Great Britain, Faroe Islands)
Ghost moth in folklore
[edit]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.[12] It is believed that the ghost moth is also referenced in the last passage of Wuthering Heights by Emily Brontë.[12]
Gallery
[edit]References
[edit]- ^ 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 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 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 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. S2CID 53167316.
- ^ a b c 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. Bibcode:1990JCEco..16.3511S. doi:10.1007/BF00982114. ISSN 0098-0331. PMID 24263445. S2CID 26903035.
- ^ R South 1909. The moths of the British Isles, volume 2. London, Frederick Warne.
- ^ a b Kaaber, Svend; Kristensen, Niels P.; Simonsen, Thomas J. (2009). "Sexual dimorphism and geographical male polymorphism in the ghost moth Hepialus humuli (Lepidoptera: Hepialidae): Scale ultrastructure and evolutionary aspects (PDF Download Available)". European Journal of Entomology. 106 (2): 303–313. doi:10.14411/eje.2009.036. Retrieved 2017-10-25.
- ^ Emmet, A Maitland (1991). The Scientific Names of the British Lepidoptera. Their history and meaning. Colchester: Harley Books. pp. 42, 43 & 197. ISBN 0-946589-35-6.
- ^ a b "Moth by Matthew Gandy from Reaktion Books". www.reaktionbooks.co.uk. Archived from the original on 2017-10-25. Retrieved 2017-10-25. Dead link
- Chinery, Michael (1986, reprinted 1991). Collins Guide to the Insects of Britain and Western Europe.
- Skinner, Bernard (1984). The Colour Identification Guide to Moths of the British Isles.
External links
[edit]- Kimber, Ian. "3.005 BF14 Ghost Moth Hepialus humuli (Linnaeus, 1758)". UKMoths. Retrieved 30 June 2019.
- Lepiforum e.V.
Ghost moth
View on GrokipediaTaxonomy and nomenclature
Classification and synonyms
The ghost moth is classified within the order Lepidoptera, superfamily Hepialoidea, family Hepialidae, genus Hepialus Fabricius, 1775, and species H. humuli (Linnaeus, 1758).[5] This placement positions it among the most basal lineages of extant moths, known collectively as ghost moths or swift moths.[6] The species was originally described by Carl Linnaeus in his Systema Naturae (1758) under the name Phalaena (Noctua) humuli, reflecting early taxonomic groupings of nocturnal Lepidoptera.[7] Historical synonyms include Phalaena humuli and Hepialus hethlandica Staudinger, 1871, though the latter is now considered a junior synonym based on morphological and distributional overlap.[8] Another early name, Hepialus lupulinus, was briefly associated but pertains more closely to related taxa like the common swift moth (Korscheltellus lupulina).[5] The 2023 revised world catalogue of Hepialidae by Grehan et al. confirms Hepialus as a monospecific genus limited to H. humuli, with no boundary shifts or major taxonomic revisions for this European species; it annotates the original Linnaean description and notes its distinction from Asian congeners previously misplaced in the genus.[9] This catalogue synthesizes nearly 400 years of literature, emphasizing stable classification amid broader family-wide rearrangements. Members of Hepialidae, including H. humuli, represent primitive moths evolutionarily, lacking the frenulum-retinaculum wing-coupling mechanism typical of derived Lepidoptera and instead employing a basal jugum structure for flight synchronization.[10] Adults also exhibit reduced mouthparts, often vestigial or absent, as they do not feed and rely on larval reserves for their brief lifespan.[6]Etymology
The common name "ghost moth" for Hepialus humuli originates from the strikingly pale, white wings of the adult males, which take on an ethereal, spirit-like appearance as they hover over grasslands at dusk in a display flight to attract females.[11] This hovering behavior, combined with the ghostly pallor, evokes images of apparitions, and the name may also draw from European folklore associating white moths with supernatural entities.[1][12] The scientific binomial Hepialus humuli traces its roots to Carl Linnaeus's 1758 description in Systema Naturae, where the species was originally named Phalaena (Noctua) humuli.[8] The specific epithet "humuli" refers to the hop plant (Humulus lupulus), as early observations linked the moth's larvae to feeding on hop roots, though subsequent studies revealed a broader range of host plants including grasses and herbs.[12] The genus Hepialus was established by Johan Christian Fabricius in 1775 to accommodate this and related species, deriving from the Ancient Greek "hēpialos," meaning "fever," in reference to the moths' erratic, feverish flight patterns.[13][14] Historically, the nomenclature of H. humuli evolved amid broader taxonomic shifts within the Hepialidae family, which encompasses both "ghost moths" and "swift moths" due to their rapid, darting flight. Early classifications placed the species under the catch-all genus Phalaena, leading to confusions with other swift-like moths in the Noctuidae, until Fabricius's reorganization highlighted distinguishing traits like the lack of a proboscis and primitive wing venation.[6] This dual common naming persists today, reflecting ongoing overlaps in popular and scientific identification of hepialid species across Europe.[15]Subspecies and related species
The ghost moth Hepialus humuli is divided into two primary subspecies: the nominate subspecies H. h. humuli, which occurs across mainland Europe, and H. h. thulensis, restricted to the Shetland Islands and Faroe Islands.[1][16] The subspecies H. h. thulensis (Newman, 1865) exhibits distinct morphological traits, including smaller size and males with buff or creamy white forewings marked by brown spots, contrasting with the uniformly white forewings of H. h. humuli males; females in both subspecies show yellow forewings with orange markings, though thulensis variants can display darker tones.[1][17][16] An additional variant, H. h. hethlandica (Staudinger, 1871), has been described from the Shetland Islands but is often subsumed under thulensis.[16] Genetic analyses, including mitochondrial COI and nuclear RpS5 markers, reveal cryptic homogeneity across European populations of H. humuli, with no unique genetic or genitalic differences supporting thulensis as distinct; this has prompted proposals to synonymize thulensis with the nominate form, though phenotypic variations warrant its continued recognition in recent catalogues.[18][16] Within the Hepialus genus, closely related species include H. sylvina (now classified as Triodia sylvina, the orange ghost or orange swift), which shares a broad European distribution with H. humuli and occupies similar grassy habitats, leading to potential distributional overlaps.[16] Other European Hepialidae, such as Phymatopus hecta (formerly H. hecta) and Korscheltellus lupulina (formerly H. lupulinus), also exhibit range overlaps with H. humuli, raising the possibility of hybridization in sympatric areas, though no verified interspecific hybrids have been documented.[16] Asian species formerly placed in Hepialus, particularly those in the genus Thitarodes (e.g., T. armoricanus and T. pui), have been subject to taxonomic debate regarding synonymy with Hepialus due to historical associations, but the 2023 global catalogue treats Thitarodes as a distinct genus with 80 species, primarily high-altitude forms from the Tibetan Plateau and Himalayas that show no distributional overlap with H. humuli.[16]Physical description
Adult morphology
The adult ghost moth, Hepialus humuli, exhibits a robust body structure typical of the Hepialidae family, with a hairy thorax covered in dense scales that provide insulation and camouflage.[19] The thorax is broad and sturdy, supporting powerful flight muscles suited for the moth's crepuscular hovering displays. Adults lack functional mouthparts, featuring a greatly reduced or absent proboscis, which renders them non-feeding and limits their adult lifespan to a few days focused on reproduction.[20] The antennae show sexual dimorphism, with males possessing bipectinate structures that enhance pheromone detection, while females have filiform antennae.[21] Wingspan varies by sex, measuring 40–50 mm in males and 50–70 mm in females, contributing to their medium-large size within the family.[1] Male forewings are typically cream-white and unpigmented, though geographical polymorphism occurs with variations such as yellow tones in certain populations (e.g., Faroe Islands), creating a ghostly appearance; whereas female forewings are yellowish-buff with darker linear markings for blending into vegetation; hindwings are brown in both sexes.[12][22] Wing venation in H. humuli follows the primitive pattern characteristic of Hepialidae, with similar fore- and hindwing configurations, including a present jugum for wing coupling and the absence of a distinct humeral angle at the forewing base, reflecting ancestral lepidopteran traits.[19][6] In males, the wing scales possess an elaborate internal cuticular meshwork that reflects light, producing a silvery sheen potentially aiding in mate attraction during low-light conditions.[23] This microstructure contrasts with the more standard ridged scales in females, underscoring adaptive differences in visual signaling.[23]Sexual dimorphism
The ghost moth, Hepialus humuli, exhibits pronounced sexual dimorphism, particularly in size, with females possessing a larger wingspan of 50–70 mm compared to males at 40–50 mm, a disparity that facilitates mate attraction during dusk lekking displays.[24][12] This size difference correlates with forewing lengths, where females' wings are approximately 2.4–2.9 mm longer than those of males (p < 0.01).[22] In coloration, males display silvery white wings that enhance visual signaling for attracting females from afar, while females have yellowish-buff forewings with darker linear markings and brown hindwings, providing cryptic camouflage against predators during egg-laying.[22][12] This dimorphism extends to wing scale ultrastructure: male scales feature an elaborate meshwork that reflects light for conspicuousness, whereas female scales are simpler, pigmented, and dentate for blending with vegetation.[22] Physiologically, males produce pheromones from bushy hairs on their hind legs, emitted during hovering to draw females to leks, contrasting with females' larger abdomens adapted for laying 200–1,600 eggs (mean ~600) over several days.[25][26]Distribution and habitat
Geographic range
The ghost moth (Hepialus humuli) is native to Europe, where it exhibits a widespread distribution spanning the Palaearctic region from the British Isles and Scandinavia in the north to northern Spain, southern Austria, and northern Italy in the south.[18][17] Its range is shaped by postglacial expansions from multiple refugia in the Alps, with genetic lineages indicating colonization of western and northern Europe from northwestern and northeastern groups.[18] The species is absent from the far south-eastern extremities of Europe and shows no established presence in Asia beyond potential minor overlaps in western boundary areas.[17] Northern limits extend to the Shetland Islands, Faroe Islands, and southern Scandinavia, reaching approximately 65°N in Fennoscandia, with subspecies such as H. h. thulensis in the northern islands (Shetland and Faroe).[22][17] In the British Isles, populations are well-distributed across England, Wales, Scotland, and Ireland, though absent from the Isles of Scilly.[27][28] Historical records confirm its presence in lowland areas throughout these regions, with phylogeographic evidence suggesting colonization of Britain via ancient land bridges like Doggerland.[18] Current extents remain stable, as indicated by citizen science observations on platforms like iNaturalist from 2020 to November 2025, which document ongoing localized populations without evidence of significant range shifts due to climate change.[29] In Britain specifically, abundance has declined by 73% between 1968 and 2002, making it a UK priority species for conservation, though globally it is Not Evaluated by the IUCN; distribution maps show persistence in core areas.[27][30]Habitat preferences
The ghost moth (Hepialus humuli) primarily inhabits damp grasslands, meadows, fens, and woodland edges featuring long grasses and weedy vegetation, where it can exploit suitable conditions for both larval development and adult lekking behaviors.[1][4] These environments provide the open, grassy swards essential for male hovering displays at dusk, often in uncut or rank growth areas that offer acoustic and visual cues for mating.[31] The species favors disturbed ground in both rural and urban settings, including field edges, rough meadows, and hedgerows adjacent to farmland with damp grassland.[32] Larval stages occupy moist soils within these habitats, where the subterranean caterpillars feed on grass and herbaceous roots, overwintering multiple times before pupation underground.[33][4] Such soils, often warm and damp after rainfall, support the typically two- to three-year larval period by maintaining humidity and nutrient availability from decaying organic matter.[34][1] Adult emergence and flight are concentrated in open fields near these larval sites, with leks forming over grassy patches to facilitate pheromone-based mate location.[35] The species occurs from lowlands up to approximately 1,000 meters in elevation, with populations in alpine refugia indicating adaptability to montane grasslands during postglacial expansions.[18][36] Recent research post-2020 underscores the vulnerability of these preferences to habitat fragmentation driven by agricultural intensification, which disrupts connected grassland networks essential for larval survival and adult dispersal in regions like southern Sweden.[37]Life cycle
Eggs
Female ghost moths deposit their eggs by scattering 200–1,600 of them loosely over vegetation or the soil surface while flying low over suitable habitats. This oviposition takes place over a four-day period following mating, with an average of approximately 600 eggs per female. The eggs are laid in loose clusters but remain unattached and not glued to the substrate, allowing them to fall among ground debris where they develop.[26][24] Under summer conditions, the eggs hatch after 10–14 days, though controlled laboratory experiments record an incubation period ranging from 11 to 24 days (mean of 18 days) at 20°C. In cooler climates, eggs exhibit diapause potential, remaining viable without hatching for at least six months at 5°C. Hatching success is high, with at least 80% viability observed even in relatively dry laboratory air.[26] Egg survival in the field is influenced by environmental factors, including vulnerability to desiccation—requiring near 100% humidity for optimal development, as is characteristic of Hepialidae eggs—and predation by ground-foraging invertebrates and birds that target the exposed clusters on the soil surface.[26][38]Larvae
The larvae of the ghost moth, Hepialus humuli, are whitish, maggot-like caterpillars with an orange-red head capsule that provides protection during feeding.[39] They undergo 12 to 14 instars, reaching full maturity at lengths of 30–50 mm.[26] This elongated body form is typical of hepialid larvae, adapted for subterranean life. These larvae are root-boring feeders, constructing silk-lined burrows and chambers in the soil from which they emerge to consume roots of grasses and herbaceous plants. They exhibit polyphagous habits, attacking a range of wild and cultivated species such as nettles (Urtica dioica), docks (Rumex spp.), and strawberries (Fragaria spp.), though they thrive in moist, grassy soils.[1] Feeding activity peaks in late autumn and spring, contributing to their slow growth over a biennial or sometimes triennial cycle.[26] The larvae overwinter twice, typically entering a state of reduced activity in soil chambers to endure cold temperatures, with development resuming in milder conditions.[1] This prolonged larval phase, spanning 2–3 years from hatching to pupation in spring, reflects their adaptation to temperate environments where optimal growth occurs at around 15°C.[26]Pupae
The pupal stage of the ghost moth (Hepialus humuli) is initiated in April or May, when fully grown larvae, after two years of root-feeding underground, construct silk-lined chambers in the soil for pupation.[39][40] The pupae are enclosed within a brown cocoon.[41] This non-feeding stage lasts 3–4 weeks, during which the pupa undergoes metamorphosis before adult emergence in late May to early June.[39] For emergence, the pupa actively wriggles upward through the soil to the surface, often leaving the empty pupal case protruding after the adult has eclosed.[41][42] Female pupae are larger than those of males, reflecting the sexual size dimorphism observed in adults.[1] Eclosion is triggered by rising soil temperatures in spring, with development proceeding optimally around 15°C.[43]Adults
Adult ghost moths (Hepialus humuli) emerge from pupae in late spring or early summer and possess no functional mouthparts, rendering them incapable of feeding as adults. They rely entirely on lipid reserves accumulated during the larval stage to fuel their brief lifespan, which typically lasts only a few days to a week.[1] This non-feeding physiology is characteristic of the Hepialidae family, prioritizing energy allocation toward reproduction over sustenance.[6] The flight period for adults spans from mid-June to late August in their native range across Europe, with peak activity occurring during the crepuscular hours around dusk. Males engage in short bursts of display flight, lasting 20–30 minutes beginning approximately 57 minutes after sunset, under low light intensities of 2.0–10.0 lux. This timing minimizes predation risk while facilitating mate location.[4][44] In terms of sensory adaptations, the compound eyes of adult ghost moths are suited for detecting movement and contrasts in low-light environments, supporting their dusk-active lifestyle. Notably, they lack ultrasonic hearing organs, providing no auditory defense against echolocating bats, which influences their restricted activity window to evade nocturnal predators.[44]Reproduction and behavior
Mating rituals
Males of the ghost moth (Hepialus humuli) form leks in open meadows, where groups of up to 10 individuals gather to perform synchronous hovering displays at dusk, typically lasting 20–30 minutes and starting about 57 minutes after sunset. These displays occur at light intensities between 10.0 and 2.0 lux, during which the white wings of males become highly conspicuous against the surrounding vegetation, aided by their sexual dimorphism in wing coloration.[35][45] During hovering, which involves vertical oscillations approximately 0.5 m above the ground, males emit pheromones from everted brush-like organs on their hind tibiae to attract females; the primary component of this male-produced scent is (E,E)-α-farnesene. Females respond by flying into the lek, landing near a selected male after assessing the group. Male-male competition manifests as aggressive chases and territorial hovering, with individuals eventually spacing 1–5 m apart to maintain display positions.[45][46] Once a female lands beside a male, courtship culminates in copulation, after which females typically mate only once per season while males engage in multiple matings, as evidenced by observations of distinct males pairing in successive encounters.[45]Predators and defenses
Adult ghost moths (Hepialus humuli) are primarily targeted by bats, particularly the northern bat (Eptesicus nilssonii), which exploits their lekking displays over fields at dusk.[47] Gleaning bats detect these moths acoustically in cluttered environments near vegetation.[31] Larvae face predation from birds, moles, and parasitic wasps, which attack the soil-dwelling stages and can significantly reduce populations in agricultural settings.[39] To counter these threats, ghost moths time their mating lekking to a brief dusk period, minimizing exposure to diurnal birds that cease foraging in low light and reducing encounters with certain bat species active earlier or later.[35] Lacking ultrasonic hearing organs common in many moths, adults rely on erratic, unpredictable flight patterns when bats approach, which disrupts predation attempts despite the absence of auditory evasion cues.[47] Larvae reside in soil burrows, where their pale bodies occur in the subterranean environment. Pupae develop underground in cocoons within larval burrows.https://www.bumblebee.org/invertebrates/LepidopteraHepialidae.htm[39]Ecology
Diet and food plants
The larvae of the ghost moth (Hepialus humuli) are polyphagous root-feeders, recorded on the roots of numerous herbaceous plants and grasses.[26] Representative host plants include grasses such as meadow grass (Poa spp.) and fescues (Festuca spp.), herbs like plantains (Plantago spp.) and docks (Rumex spp.), as well as cultivated species such as strawberry (Fragaria × ananassa) and lettuce (Lactuca sativa).[1][48] The larvae tunnel into the roots and basal stems of these hosts, consuming vascular tissue and disrupting nutrient and water transport within the plant.[39] This feeding strategy supports extended larval development, often spanning two years with multiple overwinterings.[33] Adult ghost moths possess vestigial mouthparts and do not feed during their short lifespan.[1] In northern regions of its range, such as Scotland and Scandinavia, larval hosts tend to favor plants characteristic of damp meadows, including moisture-loving grasses and forbs.[28]Role in ecosystems
The larvae of the ghost moth (Hepialus humuli) burrow extensively in soil while feeding on plant roots in grassland habitats.[26] As a common inhabitant of rough meadows and grasslands, the ghost moth serves as prey for higher trophic levels, including bats such as Eptesicus nilssonii, which detect and hunt adult males during their dusk lekking flights, thereby integrating the moth into food web dynamics.[49] Declines in its abundance signal broader reductions in farmland biodiversity and habitat quality.[50] Adult ghost moths play a minor role in pollination, limited by their short lifespan of a few days and absence of functional mouthparts, which prevents nectar consumption and sustained floral visitation; however, incidental contact with flowers during mating flights may support limited pollen transfer in grassland ecosystems.[1] The species occupies a key niche as a root-feeding herbivore in these habitats, linking underground processes to above-ground trophic interactions.Human interactions
Agricultural impact
The larvae of the ghost moth (Hepialus humuli) primarily damage agricultural crops by feeding on roots and stem bases, leading to wilting, stunted growth, and plant death in severe cases. In nurseries, this subterranean feeding affects herbaceous plants such as strawberries and chrysanthemums, where large populations can cause widespread loss of vigor or outright mortality.[51] In field settings, larvae target root crops like carrots and potatoes, as well as cereals and lettuce, by tunneling into roots and severing plants just below the soil surface, resulting in patchy establishment and reduced yields.[39][52] In modern contexts, particularly organic and regenerative farms, the pest poses a subtle but persistent threat to cereal crops following long-term grasslands, contributing to lodging and uneven development without widespread outbreaks.[53] Overall, it remains a minor, localized pest in the UK as of 2025, with no major national epidemics reported, though localized impacts can still affect crop marketability in vulnerable rotations.[39] Management strategies emphasize integrated pest management (IPM) approaches that avoid broad-spectrum pesticides to preserve beneficial organisms. Cultural practices, such as crop rotation to break the two-year larval life cycle and soil cultivation to expose larvae to predators like birds, moles, and ground beetles, provide effective non-chemical control.[39] Biological options include applying entomopathogenic nematodes (e.g., Steinernema or Heterorhabditis species) to moist soils in spring or autumn, targeting soil-dwelling larvae with high efficacy in garden and nursery settings when temperatures are between 12–20°C.[51] Post-2020 IPM guidelines prioritize these methods alongside monitoring for early damage signs, fostering natural enemies through habitat enhancements like flower-rich margins.[52]Cultural and folklore significance
In European folklore, the ghost moth (Hepialus humuli) has been associated with the souls of the departed due to the males' pale, white wings that create an ethereal appearance during their dusk flights.[54] This belief ties into broader traditions where white moths symbolize spirits or omens of death, particularly in British and Celtic contexts, where their fluttering was seen as restless souls seeking passage.[55] Historical records from 19th-century Britain further link such moths to death omens, with sightings interpreted as harbingers of misfortune or the presence of the deceased.[56] The ghost moth appears in literature as a symbol of the supernatural and the haunting past. A possible reference occurs in the final passage of Emily Brontë's Wuthering Heights (1847), where the narrator describes watching "moths fluttering among the heath and harebells," evoking the novel's themes of ghostly unrest; this imagery aligns with the ghost moth's habitat and appearance in the Yorkshire moors.[55] In modern culture, the ghost moth features in entomological art and conservation efforts to highlight its symbolic allure and ecological vulnerability. Organizations like Butterfly Conservation incorporate it into awareness campaigns, such as Halloween-themed posts that blend folklore with calls to protect moth habitats, emphasizing its role in biodiversity education.[54]References
- https://en.wiktionary.org/wiki/Hepialus
