View source for Sphingidae

{{Short description|Family of insects}}
{{Distinguish|Sphindidae}}
{{More citations needed|date=September 2025}}
{{For-multi|a complete species list of this family|Sphingidae species list|the ''Miraculous: Tales of Ladybug & Cat Noir'' villain known as Hawk Moth|List of Miraculous: Tales of Ladybug & Cat Noir characters#Gabriel Agreste / Hawk Moth / Shadow Moth / Monarch|the 1920s British four-seat cabin monoplane|de Havilland Hawk Moth}}
{{Automatic taxobox
| name = Hawk moths
| fossil_range = {{fossil range|Miocene|Recent}}
| image = Convolvulus hawk-moth (Agrius convolvuli) 2.jpg
| image_caption = [[Convolvulus hawk-moth]], ''Agrius convolvuli''
| taxon = Sphingidae
| authority = [[Pierre André Latreille|Latreille]], 1802
| type_species = ''[[Sphinx ligustri]]''
| type_species_authority = [[Carl Linnaeus|Linnaeus]], [[Systema Naturae|1758]]
| diversity = About 200 genera, [[List of Sphingidae species|roughly 1,450 species]]
| subdivision_ranks = Subfamilies
| subdivision = *[[Macroglossinae (moth)|Macroglossinae]]
*[[Smerinthinae]]
*[[Sphinginae]]
}}

The '''Sphingidae''' are a family of [[moth]]s commonly called '''sphinx moths''', also colloquially known as '''hawk moths''', with many of their caterpillars known as '''hornworms'''. It includes about 1,450 [[species]].<ref>{{cite journal| author=van Nieukerken| title=Order Lepidoptera Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal biodiversity: An outline of higher-level classification and survey of taxonomic richness | journal=Zootaxa| volume=3148| year=2011| pages=212–221| url=http://mapress.com/zootaxa/2011/f/zt03148p221.pdf|display-authors=etal| doi=10.11646/zootaxa.3148.1.41 }}</ref> It is best represented in the [[tropics]], but species are found in every region.<ref name=scoble>Scoble, Malcolm J. (1995): ''The Lepidoptera: Form, Function and Diversity'' (2nd edition). Oxford University Press & Natural History Museum London. {{ISBN|0-19-854952-0}}</ref> They are moderate to large in size and are distinguished among moths for their agile and sustained flying ability, similar enough to that of [[hummingbird]]s as to be reliably mistaken for them.<ref name=scoble/> Their narrow wings and streamlined abdomens are adaptations for rapid flight. The family was named by French zoologist [[Pierre André Latreille]] in 1802.

Some hawk moths, such as the [[hummingbird hawk-moth]] or the [[Hyles lineata|white-lined sphinx]], hover in midair while they feed on [[nectar]] from flowers, so are sometimes mistaken for hummingbirds. This hovering capability is only known to have evolved four times in nectar feeders: in hummingbirds, certain [[bat]]s, [[hoverfly|hoverflies]], and these sphingids<ref name= kitching>{{cite journal | last1 = Kitching | first1 = Ian J | year = 2002 | title = The phylogenetic relationships of Morgan's Sphinx, ''Xanthopan morganii'' (Walker), the tribe Acherontiini, and allied long-tongued hawkmoths (Lepidoptera: Sphingidae, Sphinginae) | journal = [[Zoological Journal of the Linnean Society]] | volume = 135 | issue = 4| pages = 471–527 | doi = 10.1046/j.1096-3642.2002.00021.x | doi-access = free }}</ref> (an example of [[convergent evolution]]). Sphingids have been studied for their flying ability, especially their ability to move rapidly from side to side while hovering, called "swing-hovering" or "side-slipping". This is thought to have evolved to deal with ambush predators that lie in wait in flowers.<ref name=kitching/>

Sphingids are some of the faster flying [[insect]]s; some are capable of flying at over {{cvt|5.3|m/s|km/h}}.<ref name= "Stevenson1995">{{Cite journal |last1= Stevenson |first1= R. |last2=Corbo |first2=K. |last3=Baca |first3= L. |last4= Le |first4= Q. |title= Cage size and flight speed of the tobacco hawkmoth ''Manduca sexta'' |journal= The Journal of Experimental Biology |volume= 198 |issue= Pt 8 |pages= 1665–1672 |year=1995 |doi= 10.1242/jeb.198.8.1665 |pmid= 9319572|bibcode= 1995JExpB.198.1665S |url=http://jeb.biologists.org/content/198/8/1665.full.pdf |access-date= 10 August 2012}}</ref> They have wingspans from {{cvt|4|cm|in|frac=2}} to over {{cvt|10|cm|in|frac=2}}.

==Description==

Sphingid's antennae are not very feathery, even in males.<ref name=scoble/> They lack [[tympanal organ]]s, but members of the group Choerocampini have hearing organs on their heads.<ref name=scoble/> They have a [[Wing coupling (Lepidoptera anatomy)#frenulum|frenulum]] and [[Retinaculum (moth)|retinaculum]] to join hindwings and forewings.<ref name=scoble/> The thorax, abdomen, and wings are densely covered in scales. Some sphingids have a rudimentary [[proboscis]], but most have a very long one,<ref name=scoble/> which is used to feed on nectar from flowers. Most are [[crepuscular]] or [[nocturnal]], but some species fly during the day.<ref name=pittaway/> Both males and females are relatively long-lived (10 to 30 days).<ref name=pittaway/> Prior to flight, most species shiver their flight muscles to warm them up, and, during flight, body temperatures may surpass {{cvt|40|°C}}.<ref name=pittaway/>

In some species, [[sexual dimorphism|differences in form between the sexes]] is quite marked. For example, in the African species ''[[Agrius convolvuli]]'' (the convolvulus or morning glory hawk-moth), males have thicker antennae and more mottled wing markings than females. Only males have both an undivided frenular hook and a retinaculum. Only males have a partial comb of hairs along with their antennae.<ref name=pinhey>[[Elliot Pinhey|Pinhey, E.]] (1962): ''Hawk Moths of Central and Southern Africa''. Longmans Southern Africa, Cape Town.</ref> Females attract males with [[pheromone]]s. The male may douse the female with a pheromone<ref name= pittaway/> before mating.

===Behavior===
Some species fly only for short periods either around dusk or dawn, while other species only appear later in the evening and others around midnight, but such species may occasionally be seen feeding on flowers during the day. A few common species in Africa, such as the Oriental beehawk (''[[Cephonodes hylas]] virescens''), ''[[Macroglossum hirundo]]'', and ''[[Macroglossum trochilus]]'', are [[Diurnality|diurnal]].<ref name=pinhey/>

A number of species are known to be migratory, all in the [[Sphingini]] and [[Macroglossinae (moth)|Macroglossinae]], and specially in the genera ''Agrius'', ''Cephonodes'', ''Macroglossum'', ''Hippotion'' and ''Theretra''.<ref>{{cite web|url=http://www.mothsofborneo.com/part-3/sphingidae/sphingidae.php|website=The Moths of Borneo|title=Family Sphingidae|last=Holloway|first=J.D.|date=1987|publisher=Southene Sdn Bhd}}</ref>

===Flight===
In studies with ''[[Manduca sexta]]'', moths have dynamic flight sensory capabilities due to their antennae. The antennae are vibrated in a plane so that when the body of the moth rotates during controlled aerial maneuvers, the antennae are subject to the [[inertial force|inertial]] [[Coriolis effect|Coriolis forces]] that are linearly proportional to the angular velocity of the body.<ref>{{Cite journal |url=http://www.case.edu/artsci/biol/willis/Publications/Alexander_2007.pdf |title=Antennae as Gyroscopes |last=McNiell Alexander |first=R. |date=February 2007 |journal=Science |doi=10.1126/science.1136840 |pmid=17289963 |volume=315 |issue=5813 |pages=771–772 |s2cid=118833201 |access-date=2014-12-06 |archive-date=2016-03-03 |archive-url=https://web.archive.org/web/20160303222315/http://www.case.edu/artsci/biol/willis/Publications/Alexander_2007.pdf |url-status=dead }}</ref> The Coriolis forces cause deflections of the antennae, which are detected by the [[Johnston's organ]] at the base of each antenna, with strong frequency responses at the beat frequency of the antennae (around 25&nbsp;Hz) and at twice the beat frequency. The relative magnitude of the two frequency responses enables the moth to distinguish rotation around the different [[Aircraft principal axes#Principal axes|principal axes]], allowing for rapid course control during aerial maneuvers.<ref>{{Cite journal|title=Antennal mechanosensors mediate flight control in moths |author=Sane, S. |author2=Dieudonné, A.|author3=Willis, M. |author4=Daniel, T. |date=February 2007 |journal=Science |doi=10.1126/science.1133598 |pmid=17290001 |volume=315 |issue=5813 |pages=863–866|bibcode=2007Sci...315..863S |citeseerx=10.1.1.205.7318 |s2cid=2429129}}</ref>

[[Image:Hawk moth caterpillar.jpg|thumb|[[Vine hawk-moth]] larva (''Hippotion celerio'')]]
[[File:Tomato hornworm.jpg|thumb|An example of the posterior "horn" seen on the [[Manduca quinquemaculata|tomato hornworm]]]]

==Life cycle==
Most species are multivoltine, capable of producing several generations a year if weather conditions permit.<ref name=pittaway>Pittaway, A. R. (1993): ''The Hawkmoths of the Western Palaearctic''. Harley Books & Natural History Museum, London. {{ISBN|0-946589-21-6}}</ref> Females lay translucent, greenish, flattened, smooth eggs, usually singly on the host plants. Egg development time varies highly, from three to 21 days.

[[Image:Hyles-gallii-caterpillar.jpg|thumb|A ''[[Hyles gallii]]'' caterpillar seeking a place to pupate: the color of the caterpillar darkens before pupation.]]
[[File:Sphingidaepupa.JPG|thumb|''[[Hyles euphorbiae]]'' pupa]]
Sphingid [[caterpillar]]s are medium to large in size, with stout bodies. They have five pairs of [[proleg]]s.<ref name=pittaway/> Usually, their bodies lack any hairs or tubercules, but most species have a "horn" at the posterior end,<ref name=scoble/> which may be reduced to a button, or absent, in the final [[instar]].<ref name=pittaway/> Many are [[crypsis|cryptic]] greens and browns, and have [[countershading]] patterns to conceal them. Others are more conspicuously colored, typically with white spots on a black or yellow background along the length of the body. A pattern of diagonal slashes along the side is a common feature. When resting, the caterpillar usually holds its legs off the surface and tucks its head underneath (praying position), which, resembling the [[Great Sphinx of Giza]], gives rise to the name "sphinx moth".<ref name=pittaway/> Some tropical larvae are thought to mimic snakes.<ref name=scoble/><ref>{{cite journal|last=Hossie|first=Thomas|author2=Sherratt, Thomas |title=Defensive posture and eyespots deter avian predators from attacking caterpillar models|journal=Animal Behaviour|date=August 2013|volume=86|issue=2|pages=383–389|doi=10.1016/j.anbehav.2013.05.029|s2cid=53263767}}</ref> Larvae are quick to regurgitate their sticky, often toxic, [[foregut]] contents on attackers such as ants and [[parasitoid]]s.<ref name=pittaway/> Development rate depends on temperature, and to speed development, some northern and high-altitude species sunbathe.<ref name=pittaway/> Larvae burrow into the soil to pupate, where they remain for two to three weeks before they emerge as adults.

In some sphingids, the pupa has a free proboscis, rather than being fused to the pupal case as is most common in the macrolepidoptera.<ref name=scoble/> They have a [[Chrysalis|cremaster]] at the tip of the abdomen.<ref name=pittaway/> Usually, they pupate off the host plant, in an underground chamber, among rocks, or in a loose cocoon.<ref name=pittaway/> In most species, the pupa is the overwintering stage.

==Food plants==

===Larvae===
[[Image:Bee hawk moth newspaper.jpg|thumb|''[[Cephonodes kingii|C. kingii]]'', [[Crows Nest, New South Wales|Crows Nest]]]]

Sphingid larvae tend to be specific feeders, rather than generalists.<ref name=pittaway/> Compared to similarly sized [[Saturniidae|saturniids]], sphingids eat soft young leaves of host plants with small [[toxin|toxic molecule]]s, and chew and mash the food into very small bits.<ref>{{cite journal|last1=Bernays |first1=E. A. |last2=Janzen |first2=D. H. |year=1988 |title=Saturniid and sphingid caterpillars: two ways to eat leaves |journal=[[Ecology (journal)|Ecology]] |volume=69 |issue=4 |pages=1153–1160 |doi=10.2307/1941269 |jstor=1941269 |bibcode=1988Ecol...69.1153B |s2cid=56310810}}</ref> Some species can tolerate quite high concentrations of specific toxins. [[Tobacco hornworm]]s (''Manduca sexta'') detoxify and rapidly excrete [[nicotine]], as do several other related sphinx moths in the subfamilies [[Sphinginae]] and [[Macroglossinae (moth)|Macroglossinae]], but members of the [[Smerinthinae]] that were tested are susceptible.<ref>{{cite journal |last1=Wink |first1=M. |last2=Theile |first2=Vera |year=2002 |title=Alkaloid tolerance in ''Manduca sexta'' and phylogenetically related sphingids (Lepidoptera: Sphingidae) |journal=Chemoecology |volume=12 |issue=1 |pages=29–46 |doi=10.1007/s00049-002-8324-2 |bibcode=2002Checo..12...29W |s2cid=41120103}}</ref> The species that are able to tolerate the toxin do not sequester it in their tissues; 98% was excreted. However, other species, such as ''[[Hyles euphorbiae]]'' and ''[[Daphnis nerii]]'', do sequester toxins from their hosts, but do not pass them on to the adult stage.<ref name=pittaway/>

===Adults===

Most adults feed on nectar, although a few tropical species feed on eye secretions, and the [[death's-head hawkmoth]] steals honey from bees.<ref name=pittaway/> Night-flying sphingids tend to prefer pale flowers with long corolla tubes and a sweet odor, a [[pollination syndrome]] known as "sphingophily".<ref name=kitching/> Some species are quite general in visitations, while others are very specific, with the plant only being successfully pollinated by a particular species of moth.<ref name=kitching/> Orchids frequently have such specific relations with hawk moths and very long corolla tubes. The comet orchid (''[[Angraecum sesquipedale]]''), a rare [[Madagascar|Malagasy]] flower with its nectar stored at the bottom of a {{convert|30|cm|in|adj=mid|-long}} tube, was described in 1822 by [[Louis-Marie Aubert du Petit-Thouars]], and later, [[Charles Darwin]] famously predicted there must be some specialized moth to feed from it:

{{blockquote|[''A. sesquipetale'' has] nectaries 11 and a half inches long [{{cvt|11.5|in|cm|0|disp=out}}], with only the lower inch and a half [{{cvt|1.5|in|cm|0|disp=out}}] filled with very sweet nectar [...] it is, however, surprising, that any insect should be able to reach the nectar: our English sphinxes have probosces as long as their bodies, but in Madagascar, there must be moths with probosces capable of extension to a length of between 10 and 12 inches! [{{cvt|10|and|12|in|cm|0|disp=out}}]<ref>Darwin, Charles (1862): ''[[Fertilisation of Orchids|On the Various Contrivances by Which British and Foreign Orchids are Fertilised by Insects, and on the Good Effects of Intercrossing]]'' John Murray, London. [http://darwin-online.org.uk/content/frameset?itemID=F800&viewtype=text&pageseq=1 HTML fulltext]
</ref>}}

[[Alfred Russel Wallace]] published a sort of "[[wanted poster]]" (properly, a drawing in a book)<ref>{{Cite web|last=Beccaloni|first=George|date=April 2010|title=Darwin and Wallace's Predictions Come True {{!}} The Alfred Russel Wallace Website|url=http://wallacefund.info/darwin-and-wallaces-predictions-come-true-0|access-date=2020-08-05|website=wallacefund.info|archive-date=2018-10-10|archive-url=https://web.archive.org/web/20181010084150/http://wallacefund.info/darwin-and-wallaces-predictions-come-true-0|url-status=dead}}</ref> of what this lepidopteran might look like, and, concurring with his colleague, added:

{{blockquote|[The proboscis of a hawk moth] from tropical Africa (''[Xanthopan] morganii'') is seven inches and a half [{{cvt|7.5|in|cm|0|disp=out}}]. A species having a proboscis two or three inches longer [{{cvt|3|in|cm|0|disp=out}}] could reach the nectar in the largest flowers of ''Angraecum sesquipedale'', whose nectaries vary in length from ten to fourteen inches [{{cvt|14|in|cm|0|disp=out}}]. That such a moth exists in Madagascar may be safely predicted, and naturalists who visit that island should search for it with as much confidence as astronomers searched for the planet [[Neptune]], – and they will be equally successful.<ref>{{Cite journal |last=Wallace |first=Alfred R. |date=1867 |title=Creation by law |url=http://www.wku.edu/~smithch/wallace/S140.htm |journal= Quarterly Journal of Science |volume=4 |pages=470–488 }} (p. 477)</ref>}}

The predicted sphingid was discovered 21 years later and described as a [[subspecies]] of the one African species studied by Wallace: ''[[Xanthopan morganii praedicta]]'',<ref>{{cite journal |last1=Rothschild |first1=Walter |last2=Jordan |first2=Karl |year=1903 |title=A revision of the lepidopterous family Sphingidae |journal=Novitates Zoologicae |volume=9 |issue=Supplement |pages=[https://archive.org/details/revisionoflepido01roth/page/n142 1]–972 |url=https://archive.org/details/revisionoflepido01roth }}</ref> for which, the subspecific name ''praedicta'' ("the predicted one") was given. The Madagascan individuals had a pink, rather than white, breast and abdomen and a black apical line on the forewing, broader than in mainland specimens. Molecular clock models using either rate- or fossil-based calibrations imply that the Madagascan subspecies ''X. m. praedicta'' and the African subspecies ''X. m. morgani'' diverged 7.4 ± 2.8 Mya (million years ago), which overlaps the divergence of ''A. sesquipedale'' from its sister, ''A. sororium'', namely 7.5 ± 5.2 Mya.<ref>{{cite journal |last1=Netz |first1=Christoph |last2=Renner |first2=Susanne S.|year=2017 |title=Long-spurred ''Angraecum'' orchids and long-tongued sphingid moths on Madagascar: A time-frame for Darwin's predicted ''Xanthopan''/''Angraecum'' coevolution |journal=Biological Journal of the Linnean Society |volume=122 |issue=Supplement |pages=469–478 |doi=10.1093/biolinnean/blx086 }}</ref> Since both these orchids have extremely long spurs, longspurs likely existed before that and were exploited by long-tongued moths similar to ''Xanthopan morganii praedicta''. The long geological separation of subspecies ''morgani'' and ''praedicta'' matches their morphological differences in the color of the breast and abdomen.

==Relationships and species==
[[File:Hawk moth.jpg|thumb|A sphinx moth, subfamily [[Macroglossinae (moth)|Macroglossinae]], in [[Cibodas Botanical Garden]], [[Java]]]]
The Sphingidae is sometimes assigned its own exclusive [[Taxonomic rank|superfamily]], Sphingoidea, but is alternatively included with the more encompassing [[Bombycoidea]]. Following Hodges (1971) two [[Subfamily|subfamilies]] are accepted, namely the [[Sphinginae]] and [[Macroglossinae (moth)|Macroglossinae]].<ref name="comm">{{cite book|last1=Common |first1=I.F.B. |title=Moths of Australia |url=https://archive.org/details/mothsaustralia00comm |url-access=limited |date=1990 |publisher=Brill |location=Leiden |isbn=9789004092273 |page=[https://archive.org/details/mothsaustralia00comm/page/n25 24]}}</ref> Around 1,450 species of hawk moths are classified into around 200 genera. Some of the best-known hawk moth species are:
* [[Privet hawk moth]] (''Sphinx ligustri'')
* [[White-lined Sphinx]] (''Hyles lineata'')
* [[Acherontia atropos|Death's-head hawk moth]] (''Acherontia atropos'')
* [[Lime hawk moth]] (''Mimas tiliae'')
* [[Poplar hawk moth]] (''Laothoe populi'')
* [[Convolvulus hawk moth]] (''Agrius convolvuli'')
* [[Catalpa sphinx]] (''Ceratomia catalpae'')
* [[Hummingbird hawk-moth]] (''Macroglossum stellatarum'')
* [[Elephant hawk moth]] (''Deilephila elpenor'')
* [[Vine hawk moth]] (''Hippotion celerio'')
* [[Spurge hawk moth]] (''Hyles euphorbiae'')
* [[Oleander hawk moth]] (''Daphnis nerii'')
* [[Pandora sphinx moth]] (''Eumorpha pandorus'')
* [[Tomato worm]] (''Manduca quinquemaculata'')
* [[Tobacco hornworm]] (''Manduca sexta'')

==See also==
* [[Sphingidae species list]]

==References==
{{Reflist|colwidth=33em}}

==External links==
{{Commons category}}
{{Wikispecies}}
* [http://insecta.pro/family/sphingidae/ Sphingidae]. Insecta.pro
* [http://janzen.sas.upenn.edu/caterpillars/chckradufam/sphiadufam/sph1.html Images of Sphingidae of Costa Rica] {{Webarchive|url=https://web.archive.org/web/20190720122244/http://janzen.sas.upenn.edu/caterpillars/chckradufam/sphiadufam/sph1.html |date=2019-07-20 }}
* [http://www.landcareresearch.co.nz/resources/identification/animals/large-moths/image-gallery/sphingidae Images of Sphingidae species in New Zealand]; {{Webarchive|url=https://web.archive.org/web/20200809224452/https://www.landcareresearch.co.nz/resources/identification/animals/large-moths/image-gallery/sphingidae |date=2020-08-09 }}
* [http://www.insecta-web.org/cgi-bin/MWM/db_typen/display.pl?tname=family&fam=Sphingidae List of Sphingidae Types from Museum Witt München]
* [https://www.flickr.com/search/?q=sphingidae Images from Flickr]

{{Lepidoptera}}
{{Taxonbar |from=Q157683}}
{{Authority control}}

[[Category:Sphingidae| ]]
[[Category:Moth families]]
[[Category:Taxa named by Pierre André Latreille]]