Editing Hummingbird hawk-moth

{{Short description|Species of moth}}
{{Speciesbox
| image = Hummingbird hawk moth (Macroglossum stellatarum) in flight.jpg
| image_caption = In flight
| image2 = Macroglossum stellatarum au repos summer 2006.jpg
| image2_caption = At rest
| taxon = Macroglossum stellatarum
| authority = ([[Carl Linnaeus|Linnaeus]], [[Lepidoptera in the 10th edition of Systema Naturae|1758]])
| synonyms = *''Sphinx stellatarum'' <small>Linnaeus, 1758</small>
}}

[[File:Distribution map Macroglossum stellatarum.svg|thumb|300px|Distribution map: yellow, winter in the Northern Hemisphere; green, year round; blue, summer in the Northern Hemisphere]]
The '''hummingbird hawk-moth''' ('''''Macroglossum stellatarum''''') is a species of [[hawk moth]] found across temperate regions of [[Eurasia]]. The species is named for its similarity to [[hummingbird]]s, as they feed on the nectar of tube-shaped flowers using their long [[proboscis]] while hovering in the air; this resemblance is an example of [[convergent evolution]]. 

The hummingbird hawk-moth was [[Species description|first described]] by [[Carl Linnaeus]] in his 1758 [[10th edition of Systema Naturae|10th edition of ''Systema Naturae'']]. As of 2018, its entire genome and mitogenome have been sequenced.<ref>{{Cite journal |last1=Li |first1=Jun |last2=Zhang |first2=Yaoyao |last3=Hu |first3=Kunjie |last4=Zhao |first4=Yaqi |last5=Lin |first5=Ruirui |last6=Li |first6=Yan |last7=Huang |first7=Zhuoran |last8=Zhang |first8=Xu |last9=Geng |first9=Xuexia |date=2018-07-01 |title=Mitochondrial genome characteristics of two Sphingidae insects (''Psilogramma increta'' and ''Macroglossum stellatarum'') and implications for their phylogeny |journal=International Journal of Biological Macromolecules |volume=113 |pages=592–600 |doi=10.1016/j.ijbiomac.2018.02.159 |pmid=29501752 |s2cid=3983266 |issn=0141-8130}}</ref>

==Distribution==

The hummingbird hawk-moth is distributed throughout the northern [[Old World]] from Portugal to Japan, but it breeds mainly in warmer climates (southern Europe, North Africa, and points east). Three generations are produced in a year in Spain.<ref>{{cite journal |title=The year-round phenology of ''Macroglossum stellatarum'' (Linnaeus, 1758) at a Mediterranean area of South of Spain (Lepidoptera: Sphingidae). |journal= SHILAP Revista de Lepidopterología|year=2017 |volume=45 |issue=180 |pages=625–633 |author=Cuadrado, M.|doi=10.57065/shilap.885 |url=https://www.redalyc.org/articulo.oa?id=45553890013 }}</ref> There is evidence that the population in the [[British Isles]] is actively expanding its range, as numbers have been consistently increasing. In addition, it is believed that this population is becoming resident instead of migratory, as warmer temperatures due to [[climate change]] are allowing individuals to [[Overwintering|overwinter]].<ref>{{Cite web |last=Ornithology |first=British Trust for |date=2022-08-10 |title=Record summer for 'hummingbirds' in UK gardens |url=https://www.bto.org/about-bto/press-releases/record-summer-%E2%80%98hummingbirds%E2%80%99-uk-gardens |access-date=2024-05-31 |website=BTO - British Trust for Ornithology |language=en}}</ref>

It is a strong flier, dispersing widely in the summer.<ref>{{Cite journal |last=Herrera |first=Carlos M. |date=1992-02-01 |title=Activity pattern and thermal biology of a day-flying hawkmoth (''Macroglossum stellatarum'') under Mediterranean summer conditions |journal=Ecological Entomology |volume=17 |issue=1 |pages=52–56 |doi=10.1111/j.1365-2311.1992.tb01038.x |bibcode=1992EcoEn..17...52H |issn=1365-2311 |hdl=10261/44693 |s2cid=85320151 |hdl-access=free}}</ref> However it rarely survives the winter in northern latitudes (e.g. north of the [[Alps]] in Europe, north of the [[Caucasus]] in Russia).

Moths in the genus ''[[Hemaris]]'', also of the family Sphingidae, are known as "hummingbird moths" in the US, and "bee moths" in Europe.{{Citation needed|date=September 2020}}<ref>P. Waring  and M. Townsend. 2003, and later editions ''Field Guide to the Moths of Britain and Ireland''</ref> This sometimes causes confusion between this species and the North American genus.

==Life cycle==
Two or more broods are produced each year. The adult may be encountered at any time of the year, especially in the south of the range, where there may be three or four broods. It overwinters as an adult in a crevice among rocks, trees, and buildings.<ref name="pittaway"/> On very warm days it may emerge to feed in mid-winter. Unlike other moths, they have no sexual dimorphism in the size of their antennal lobes.<ref name=":1">{{Cite journal |last1=Kelber |first1=Almut |last2=Warrant |first2=Eric |last3=Basil el Jundi |last4=Charalabidis |first4=Alice |last5=Heinze |first5=Stanley |last6=Stöckl |first6=Anna |date=2016-05-17 |title=Differential investment in visual and olfactory brain areas reflects behavioural choices in hawk moths |journal=Scientific Reports |volume=6 |pages=26041 |doi=10.1038/srep26041 |pmid=27185464 |pmc=4869021 |issn=2045-2322 |bibcode=2016NatSR...626041S}}</ref>

===Ova===
The glossy pale green ova (eggs) are spherical with a {{convert|1|mm|in|2|adj=on}} diameter. They are said to look like the flower buds of the host plant ''[[Galium]]'', and that is where the female lays them. They hatch 6 to 8 days after laying.<ref name="pittaway"/> Up to 200 eggs may be laid by one female, each on a separate plant.

===Larvae===
Newly hatched [[larva]]e are clear yellow, and in the second [[instar]] assume their green coloration. The larva is green with two grey stripes bordered in cream along the sides and with a horn at the rear end typical of sphingids. The horn is purplish red, changing to blue with an orange tip in the last instar.<ref name="pittaway"/> They feed fully exposed on the top of the host plant and rest in among a tangle of stems. Although dependent on warmth and sun, the larval stage can be as rapid as 20 days.
<gallery mode=packed>
File:Macroglossum.stellatarum.caterpillar.3088.Liosi.jpg|Larva of ''Macroglossum stellatarum''
File:Macroglossum stellatarum (Hummingbird Hawk) larva UK.jpg|Larva of same, Hampshire, UK, while on prepupational walkabout
</gallery>

===Pupae===
The [[pupa]]e are pale brownish with a prominent, keeled [[proboscis]], and two sharp spines at the end of the [[Chrysalis|cremaster]]. They are enclosed in loose silken cocoons among the host plant debris or on the ground among [[leaf litter]].<ref name="pittaway"/>
<gallery mode="packed">
File:Hummingbird Hawk pupa 1.jpg|Pupa (2 weeks old) of hummingbird hawk
</gallery>

===Adults===
The forewings are brown, with black wavy lines across them, while the hindwings are orange with a black edge. The abdomen is quite broad, with a fan-tail of [[seta]]e at the end. The wingspan is {{convert|40|-|45|mm}}.

In the southern parts of its range, the hummingbird hawk-moth is highly active even when temperatures are high, and thoracic temperatures above {{convert|45|C|F}} have been measured.<ref name="herrera"/> This is among the highest recorded for hawk-moths, and near the limit for insect muscle activity.

<gallery heights="150px" mode="packed">
File:Macroglossum stellatarum - Schönbrunn.jpg|Closeup of its compound eye
File:Macrogl Stellat.jpg|The wing action is frozen in this photo by using [[electronic flash]]. This picture was shot in [[Hanko, Finland]], [[60th parallel north|latitude 60°N]].
File:Hummingbird hawk-moth straw streched.JPG|With [[proboscis]] extended, drinking nectar from a flower
File:Macroglossum stellatarum01(js).jpg|Nectaring on ''Verbena''
File:Taubenschwaenzchen Zeitlupe - Hummingbird Hawk-Moth Slow Motion.ogv|Slow motion video
</gallery>

==Behaviour==
Its long [[proboscis]] ({{cvt|25–28|mm|1}})<ref name=":0">{{Cite journal |last=Faucheux |first=Michel J. |date=2013-03-01 |title=Sensillum types on the proboscis of the Lepidoptera: a review |journal=Annales de la Société Entomologique de France |series=Nouvelle Série |volume=49 |issue=1 |pages=73–90 |doi=10.1080/00379271.2013.767514 |s2cid=84400239 |issn=0037-9271}}</ref> and its hovering behavior, accompanied by an audible humming noise, make it look remarkably like a [[hummingbird]] while feeding on flowers. Like hummingbirds, it feeds on flowers which have tube-shaped [[Corolla (flower)|corollae]].<ref name=":0" /> It should not be confused with the moths called hummingbird moths in North America, genus ''[[Hemaris]]'', members of the same family and with similar appearance and behavior.
The resemblance to hummingbirds is an example of [[convergent evolution]]. It flies during the day, especially in bright sunshine, but also at dusk,<ref name="herrera">
{{cite journal |last1=Herrera |title=Activity pattern and thermal biology of a day-flying hawkmoth (''Macroglossum stellatarum'') under Mediterranean summer conditions |journal=Ecological Entomology |volume=17 |pages=52–56 |year=1992 |doi=10.1111/j.1365-2311.1992.tb01038.x |first1=Carlos M. |bibcode=1992EcoEn..17...52H |hdl=10261/44693 |s2cid=85320151 |hdl-access=free}}</ref> dawn, and even in the rain, which is unusual for even diurnal hawkmoths.<ref name="pittaway">
{{cite book |last=Pittaway |first=A. R. |title=The Hawkmoths of the Western Palaearctic |publisher=Harley Books |location=London |year=1993 |isbn=978-0-946589-21-0}}</ref> ''M. stellatarum'' engages in free hovering flight, which allows more maneuverability and control than fixed-wing flight, despite high energetic cost.<ref name=":2">{{Cite journal |last1=Yao |first1=J. |last2=Yeo |first2=K. S. |date=2019-05-30 |title=Free hovering of hummingbird hawkmoth and effects of wing mass and wing elevation |journal=Computers & Fluids |volume=186 |pages=99–127 |doi=10.1016/j.compfluid.2019.04.007 |s2cid=145969157 |issn=0045-7930}}</ref><ref name=":1" /> Like many large insects, it relies upon [[Johnston's organ]]s for body positioning information.<ref>{{Cite journal |last1=Kelber |first1=Almut |last2=Sane |first2=Sanjay P. |last3=Foster |first3=James J. |last4=Stöckl |first4=Anna |last5=Dahake |first5=Ajinkya |date=2018-02-04 |title=Independent control of slow and fast flight manoeuvres in insects: the roles of vision and antennal mechanoreception |url=https://www.biorxiv.org/content/10.1101/222448v1 |journal=bioRxiv |volume=7 |pages=222448 |doi=10.1101/222448 |pmid=30526849 |doi-access=free |pmc=6303104 |access-date=2019-05-30 |archive-date=2020-10-30 |archive-url=https://web.archive.org/web/20201030135931/https://www.biorxiv.org/content/10.1101/222448v1 |url-status=live }}</ref>

== Vision ==
[[File:Macroglossum stellatarum on flowers - nectaring process.png|thumb|400px|Nectaring process on ''Buddleja'' (5 frames/second)]]
The hummingbird hawkmoth's visual abilities have been studied extensively, and they have demonstrated a relatively good ability to learn colours.<ref name="kelber">{{cite journal |last=Kelber |first=Almut |title=Colour learning in the hawkmoth ''Macroglossum stellatarum'' |journal=[[Journal of Experimental Biology]] |volume=199 |issue=5 |pages=1127–1131 |year=1996 |doi=10.1242/jeb.199.5.1127 |url=http://jeb.biologists.org/cgi/reprint/199/5/1127.pdf |pmid=9318956 |bibcode=1996JExpB.199.1127K |access-date=October 21, 2011 |archive-date=September 17, 2023 |archive-url=https://web.archive.org/web/20230917235817/https://journals.biologists.com/jeb/article/199/5/1127/7284/Colour-Learning-in-the-Hawkmoth-Macroglossum |url-status=live }}</ref> They have a [[trichromatic]] visual system, and are most sensitive to wavelength in the range of 349–521&nbsp;nm. They have been shown to discriminate a wavelength difference as small as 1–2&nbsp;nm between sources. This discrimination is even more precise than ''[[Western honey bee|Apis mellifera]]'', or the western honey bee. Among other flower visitors, their visual system is similar to ''[[Papilio xuthus]]'', or the Asian swallowtail butterfly, and ''[[Deilephila elpenor]]'', the nocturnal elephant hawkmoth.<ref>{{Cite journal |last1=Rodríguez-Gironés |first1=Miguel A. |last2=Kelber |first2=Almut |last3=Telles |first3=Francismeire J. |date=2016-02-15 |title=Wavelength discrimination in the hummingbird hawkmoth ''Macroglossum stellatarum'' |journal=Journal of Experimental Biology |volume=219 |issue=4 |pages=553–560 |doi=10.1242/jeb.130484 |issn=0022-0949 |pmid=26747900 |doi-access=free}}</ref><ref name=":1" /> Their food preference is based mainly on visual identification, while ''D.&nbsp;elpenor'' preference relies upon olfactory identification.<ref name=":1" /> Compared to ''D.&nbsp;elpenor'', ''M.&nbsp;stellatarum'' have a much smaller number of [[Ommatidium|ommatidia]], but a larger optic lobe volume to provide more visual processing tissue.<ref name=":1" />

==Habitat and host plants==
[[File:Sphingidae - Macroglossum stellatarum - Habitat.JPG|thumb|right|One of the preferred habitats of ''M.&nbsp;stellatarum'' ([[woodland edge]] with [[red valerian]])]]

Hummingbird hawk-moths can be easily seen in gardens, parks, meadows, bushes, and [[woodland edge]], where the preferred food plants grow ([[honeysuckle]], [[red valerian]] and many others).<ref>{{Cite web |url=https://www.rspb.org.uk/birds-and-wildlife/wildlife-guides/birdwatching/how-to-identify-birds/hummingbirds-and-hawkmoths/ |title=Hummingbird Hawk Moth &#124; Moth Migration |access-date=2022-04-25 |archive-date=2022-05-20 |archive-url=https://web.archive.org/web/20220520133139/https://www.rspb.org.uk/birds-and-wildlife/wildlife-guides/birdwatching/how-to-identify-birds/hummingbirds-and-hawkmoths |url-status=live }}</ref>

Their larvae usually feed on [[Galium|bedstraws]] or [[Rubia|madder]]s (''Rubia'') but have been recorded on other [[Rubiaceae]] and ''[[Centranthus]]'', ''[[Stellaria]]'', and ''[[Epilobium]]''.<ref name="pittaway"/>

Adults are particularly fond of nectar-rich flowers with a long and narrow [[Calyx (botany)|calyx]], since they can then take advantage of their long [[proboscis]] and avoid competition from other insects.<ref name="pittaway" /> Flowers with longer tubes typically present the feeding animal a higher nectar reward.<ref name=":3">{{Cite journal |last1=Johnson |first1=Steven D. |last2=Moré |first2=Marcela |last3=Amorim |first3=Felipe W. |last4=Haber |first4=William A. |last5=Frankie |first5=Gordon W. |last6=Stanley |first6=Dara A. |last7=Cocucci |first7=Andrea A. |last8=Raguso |first8=Robert A. |date=January 2017 |editor-last=Nicolson |editor-first=Sue |title=The long and the short of it: a global analysis of hawkmoth pollination niches and interaction networks |journal=Functional Ecology |volume=31 |issue=1 |pages=101–115 |doi=10.1111/1365-2435.12753 |pmc=5363726 |pmid=28344378|bibcode=2017FuEco..31..101J }}</ref> Proboscis length is thought to have been evolutionarily impacted by the length of flower feeding tubes.<ref name=":3" /> Examples of such plants include ''[[Centranthus]]'', ''[[Jasminum]]'', ''[[Buddleia]]'', ''[[Nicotiana]]'', ''[[Primula]]'', ''[[Viola (plant)|Viola]]'', ''[[Syringa]]'', ''[[Verbena]]'', ''[[Echium]]'', ''[[Phlox]]'', and ''[[Stachys]]''.<ref name="pittaway"/> They are reported to trap-line, that is, to return to the same flower beds at about the same time each day.

==In culture==
Hummingbird hawk-moths have been seen as a lucky [[omen]]. In particular, a swarm of the moths was seen flying across the [[English Channel]] on [[D-Day]], the day of the [[Normandy landings]] in the [[Second World War]].<ref>{{cite web |title=Hummingbird moths colonise UK |url=https://www.bbc.co.uk/news/science-environment-36474577 |publisher=BBC |access-date=10 June 2016 |date=10 June 2016 |archive-date=10 June 2016 |archive-url=https://web.archive.org/web/20160610092905/http://www.bbc.co.uk/news/science-environment-36474577 |url-status=live }}</ref> These moths, along with other moths, are in the family Sphingidae because their larvae were thought to resemble the Egyptian Sphinx.<ref>{{Cite web |url=http://www.pbs.org/wnet/nature/blog/featured-creature-hummingbird-hawk-moth/ |title=Featured Creature: Hummingbird Hawk-Moth |website=pbs.org |date=11 August 2015 |access-date=2019-05-30 |archive-date=2019-06-12 |archive-url=https://web.archive.org/web/20190612101734/http://www.pbs.org/wnet/nature/blog/featured-creature-hummingbird-hawk-moth/ |url-status=live }}</ref> The [[They Might Be Giants]] song [[The Else|"The Bee of the Bird of the Moth"]] is about the hummingbird moth.

==References==
{{Reflist}}

==Further reading==
*{{cite journal |last1=Telles |first1=Francismeire Jane |last2=Lind |first2=Olle |last3=Henze |first3=Miriam Judith |title=Out of the blue: the spectral sensitivity of hummingbird hawkmoths |journal=Journal of Comparative Physiology A |date=20 February 2014 |volume=200 |issue=6 |pages=537–546 |doi=10.1007/s00359-014-0888-0 |pmid=24553915 |s2cid=10608475}}
*{{cite journal |last1=Farina |first1=WM |last2=Varjú |first2=D |last3=Zhou |first3=Y |year=1994 |title=The regulation of distance to dummy flowers during hovering flight in the hawk moth ''Macroglossum stellatarum'' |journal=Journal of Comparative Physiology A |volume=174 |issue=2 |pages=239–274 |doi=10.1007/bf00193790 |s2cid=39694791}}

==External links==
{{Commons category|Macroglossum stellatarum}}
* [https://www.ukmoths.org.uk/species/macroglossum-stellatarum/ "69.010 BF1984 Hummingbird Hawk-moth ''Macroglossum stellatarum'' (Linnaeus, 1758)"]. ''UKMoths''. Retrieved December 12, 2018.
* ''[https://www.leps.it/indexjs.htm?SpeciesPages/MacroStellat.htm Moths and Butterflies of Europe and North Africa]''
* {{cite web |last=Pittaway |first=A. R. |date=2018 |url=http://tpittaway.tripod.com/sphinx/m_ste.htm |title=''Macroglossum'' Scopoli, 1777 |website=Sphingidae of the Western Palaearctic |access-date=December 12, 2018}}
* ''[https://lepiforum.org/wiki/page/Macroglossum_Stellatarum Lepiforum e. V.]''
* [https://www.hummingbirdsplus.org/hummingbird-moth-facts/ ''Hummingbird Moth - Species info & Facts'']

{{taxonbar |from=Q369034}}

[[Category:Macroglossum]]
[[Category:Moths described in 1758]]
[[Category:Moths of Africa]]
[[Category:Moths of Asia]]
[[Category:Moths of Europe]]
[[Category:Moths of Iceland]]
[[Category:Moths of Japan]]
[[Category:Taxa named by Carl Linnaeus]]
[[Category:Articles containing video clips]]
[[Category:Animal migration]]