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Macroglossinae Smerinthinae Sphinginae

Diversity

About 200 genera, roughly 1,450 species

The Sphingidae
Sphingidae
are a family of moths (Lepidoptera), commonly known as hawk moths, sphinx moths, and hornworms; it includes about 1,450 species.[1] It is best represented in the tropics, but species are found in every region.[2] They are moderate to large in size and are distinguished among moths for their rapid, sustained flying ability.[2] Their narrow wings and streamlined abdomens are adaptations for rapid flight. Some hawk moths, such as the hummingbird hawk-moth or the 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 bats, hoverflies, and these sphingids[3] (an example of convergent evolution). Sphingids have been much 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.[3] Sphingids are some of the faster flying insects; some are capable of flying at over 5.3 m/s (12 miles per hour).[4] They have wingspans from 4 to over 10 cm [3.9 in].

Vine hawk moth larva (Hippotion celerio)

An example of the posterior "horn" seen on the tomato hornworm

Most species are multivoltine, capable of producing several generations a year if weather conditions permit.[5] Females lay translucent, greenish, flattened, smooth eggs, usually singly on the host plants. Egg development time varies highly, from three to 21 days.

A Hyles gallii
Hyles gallii
caterpillar seeking a place to pupate: the colour of the caterpillar darkens before pupation.

Sphingid caterpillars are medium to large in size, with stout bodies. They have five pairs of prolegs.[5] Usually, their bodies lack any hairs or tubercules, but most species have a "horn" at the posterior end,[2] which may be reduced to a button, or absent, in the final instar.[5] Many are cryptic greens and browns, and have countershading patterns to conceal them. Others are more conspicuously coloured, 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 larva usually holds its legs off the surface and tucks its head underneath (praying position), which, resembling the Egyptian Sphinx, gives rise to the name 'sphinx moth'.[5] Some tropical larvae are thought to mimic snakes.[2][6] Larvae are quick to regurgitate their sticky, often toxic, foregut contents on attackers such as ants and parasitoids.[5] Development rate depends on temperature, and to speed development, some northern and high-altitude species sunbathe.[5] Larvae burrow into soil to pupate, where they remain for 2–3 weeks before they emerge as adults. In some Sphingidae, the pupa has a free proboscis, rather than being fused to the pupal case as is most common in the Macrolepidoptera.[2] They have a cremaster at the tip of the abdomen.[5] Usually, they pupate off the host plant, in an underground chamber, among rocks, or in a loose cocoon.[5] In most species, the pupa is the overwintering stage.

A feeding hawk moth at Olympiaki Akti
Olympiaki Akti
(Greece)

Contents

1 Description

1.1 Behavior 1.2 Flight

2 Food plants

2.1 Larvae 2.2 Adults

3 Relationships and species 4 See also 5 References 6 External links

Description[edit]

Frenulum of a male comma nephele hawk moth as seen when the fore wing is pulled back. It pierces a retinaculum on vein Sc on the ventral side of the fore wing,[7] and couples the wings in flight for efficiency.[8] In males it consists of a single but compound bristle while in females it consists of several compound bristles.[9] A reduced frenulum is found in some Smerinthini.[8]

Antennae are generally not very feathery, even in the males.[2] They lack tympanal organs, but members of the group Choerocampini have hearing organs on their heads.[2] They have a frenulum and retinaculum to join hind wings and fore wings.[2] The thorax, abdomen, and wings are densely covered in scales. Some sphingids have a rudimental proboscis, but most have a very long one,[2] which is used to feed on nectar from flowers. Most are crepuscular or nocturnal, but some species fly during the day.[5] Both males and females are relatively long-lived (living 10 to 30 days).[5] Prior to flight, most species shiver their flight muscles to warm them up, and, during flight, body temperatures may surpass 40 °C (104 °F).[5] In some species, differences in form between the sexes is quite marked. For example, in the African species Agrius convolvuli
Agrius convolvuli
(the convolvulus or morning glory hawk moth), the antennae are thicker and wing markings more mottled in the male than in the female. Only males have both an undivided frenular hook and a retinaculum. Also, all male hawk moths have a partial comb of hairs along their antennae.[10] Females call males to them with pheromones. The male may douse the female with a pheromone[5] before mating. Behavior[edit] 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 Cephonodes hylas virescens (the Oriental bee hawk), Macroglossum hirundo, and Macroglossum trochilus, are diurnal.[10] Flight[edit] In studies with Manduca sexta, moths have dynamic flight sensory abilities 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 coriolis forces that are linearly proportional to the angular velocity of the body.[11] 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 Hz) and at twice the beat frequency. The relative magnitude of the two frequency responses enables the moth to distinguish rotation around the different principal axes, allowing for rapid course control during aerial maneuvers.[12] Food plants[edit] Larvae[edit]

Bee hawk moth (Cephonodes kingii), Crows Nest, New South Wales

Sphingid larvae tend to be specific feeders, rather than generalists.[5] Compared to similarly sized saturniids, sphingids eat soft young leaves of host plants with small toxic molecules, and chew and mash the food into very small bits.[13] Some species can tolerate quite high concentrations of specific toxins. Tobacco hornworms, Manduca sexta, detoxify and rapidly excrete nicotine, as do several other related sphinx moths in the subfamilies Sphinginae
Sphinginae
and Macroglossinae, but members of the Smerinthinae
Smerinthinae
that were tested are susceptible.[14] 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
Hyles euphorbiae
and Daphnis nerii, do sequester toxins from their hosts, but do not pass them on to the adult stage.[5] Adults[edit]

Hemaris thysbe
Hemaris thysbe
or hummingbird clearwing moth feeding on lantana flowers

Most adults feed on nectar, although a few tropical species feed on eye secretions, and the death's-head hawkmoth steals honey from bees.[5] Night-flying sphingids tend to prefer pale flowers with long corolla tubes and a sweet odour, a pollination syndrome known as 'sphingophily'.[3] 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.[3] Orchids frequently have such specific relations with hawk moths, and very long corolla tubes. The comet orchid, Angraecum sesquipedale, a rare Malagasy flower with its nectar stored at the bottom of a 30-cm-long tube, was described in 1822 by Louis-Marie Aubert du Petit-Thouars, and later, Charles Darwin famously predicted there must be some specialised animal to feed from it:

"[A. sesquipetale has] nectaries 11 and a half inches long [29 cm], with only the lower inch and a half [3.8 cm] 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
Madagascar
there must be moths with probosces capable of extension to a length of between 10 and 12 inches! [30 cm]"[15]

Alfred Russel Wallace
Alfred Russel Wallace
published a sort of "wanted poster" (properly, a drawing in a book)[16][unreliable source?] of what this butterfly might look like, and, concurring with his colleague, added:

"[The proboscis of a hawk moth] from tropical Africa ([Xanthopan] morganii) is seven inches and a half [19 cm]. A species having a proboscis two or three inches longer [7.6 cm] could reach the nectar in the largest flowers of Angræcum sesquipedale, whose nectaries vary in length from ten to fourteen inches [36 cm]. That such a moth exists in Madagascar
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."[17]

A possible species was discovered 21 years later, for which the hawk moth in question was found and described as a subspecies of the one mentioned by Wallace: Xanthopan morganii
Xanthopan morganii
praedicta,[18] for which, the subspecific name praedicta ("the predicted one") was given. The subspecies has, however, been subsequently declared as invalid due to its similarity to mainland species. Relationships and species[edit]

Oleander hawk moth (Daphnis nerii), in Mangaon, Maharashtra

A sphinx moth, subfamily Macroglossinae, in Cibodas Botanical Garden, Java

The Sphingidae
Sphingidae
is sometimes assigned its own exclusive superfamily, Sphingoidea, but is alternatively included with the more encompassing Bombycoidea. Following Hodges (1971) two subfamilies are accepted, namely the Sphinginae
Sphinginae
and Macroglossinae.[7] 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) Death's-head hawk moth (Acherontia atropos) Lime hawk moth (Mimas tiliae) Poplar hawk moth (Laothoe populi) Convolvulus
Convolvulus
hawk moth (Agrius convolvuli) Catalpa sphinx (Ceratomia catalpae) Hummingbird hawk-moth
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
Pandora sphinx moth
(Eumorpha pandorus) Tomato worm (Manduca quinquemaculata) Tobacco hornworm
Tobacco hornworm
(Manduca sexta)

See also[edit]

Sphingidae
Sphingidae
species list Twin-spotted Sphinx moth List of moths of India List of moths of Great Britain (Sphingidae)

References[edit]

^ van Nieukerken; et al. (2011). "Order Lepidoptera
Lepidoptera
Linnaeus, 1758. In: Zhang, Z.-Q. (Ed.) Animal
Animal
biodiversity: An outline of higher-level classification and survey of taxonomic richness" (PDF). Zootaxa. 3148: 212–221.  ^ a b c d e f g h i Scoble, Malcolm J. (1995): The Lepidoptera: Form, Function and Diversity (2nd edition). Oxford University Press & Natural History Museum London. ISBN 0-19-854952-0 ^ a b c d Kitching, Ian J (2002). "The phylogenetic relationships of Morgan's Sphinx, Xanthopan morganii
Xanthopan morganii
(Walker), the tribe Acherontiini, and allied long-tongued hawkmoths (Lepidoptera: Sphingidae, Sphinginae)". Zoological Journal of the Linnean Society. 135 (4): 471–527. doi:10.1046/j.1096-3642.2002.00021.x.  ^ Stevenson, R.; Corbo, K.; Baca, L.; Le, Q. (1995). "Cage size and flight speed of the tobacco hawkmoth Manduca sexta" (PDF). The Journal of Experimental Biology. 198 (Pt 8): 1665–1672. PMID 9319572. Retrieved 10 August 2012.  ^ a b c d e f g h i j k l m n o Pittaway, A. R. (1993): The hawkmoths of the western Palaearctic. Harley Books & Natural History Museum, London. ISBN 0-946589-21-6 ^ Hossie, Thomas; Sherratt, Thomas (August 2013). "Defensive posture and eyespots deter avian predators from attacking caterpillar models". Animal
Animal
Behaviour. 86 (2): 383–389. doi:10.1016/j.anbehav.2013.05.029. Retrieved 4 February 2014.  ^ a b Common, I.F.B. (1990). Moths of Australia. Leiden: Brill. p. 24. ISBN 9789004092273.  ^ a b Schreiber, Harald (1978). Dispersal Centres of Sphingidae (Lepidoptera) in the Neotropical Region. Dordrecht: Springer Netherlands. p. 18. ISBN 9789400999602.  ^ Messenger, Charlie (1997). "The Sphinx Moths (Lepidoptera: Sphingidae) of Nebraska". Transactions of the Nebraska Academy of Sciences (24): 91–93. Retrieved 18 April 2016.  ^ a b Pinhey, E. (1962): Hawk Moths of Central and Southern Africa. Longmans Southern Africa, Cape Town. ^ McNiell Alexander, R. (February 2007). "Antennae as Gyroscopes" (PDF). Science. 315: 771–772. doi:10.1126/science.1136840. PMID 17289963.  ^ Sane, S.; Dieudonné, A.; Willis, M.; Daniel, T. (February 2007). "Antennal mechanosensors mediate flight control in moths". Science. 315: 863–866. doi:10.1126/science.1133598. PMID 17290001.  ^ Bernays, E. A.; Janzen, D. H. (1988). "Saturniid and sphingid caterpillars: two ways to eat leaves". Ecology. 69 (4): 1153–1160. doi:10.2307/1941269.  ^ Wink, M.; Theile, Vera (2002). "Alkaloid tolerance in Manduca sexta and phylogenetically related sphingids (Lepidoptera: Sphingidae)". Chemoecology. 12: 29–46. doi:10.1007/s00049-002-8324-2.  ^ Darwin, Charles (1862): On the Various Contrivances by Which British and Foreign Orchids are Fertilised by Insects, and on the Good Effects of Intercrossing John Murray, London. HTML fulltext ^ "Image at perso.orange.fr". Retrieved 2011-10-18.  ^ Wallace, Alfred R. (1867). "Creation by law". Quarterly Journal of Science. 4: 470–488.  (p. 477) ^ Rothschild, Walter; Jordan, Karl (1903). "A revision of the lepidopterous family Sphingidae". Novitates Zoologicae. 9 (Supplement): 1–972. 

External links[edit]

Wikimedia Commons has media related to Sphingidae.

Wikispecies
Wikispecies
has information related to Sphingidae

European Butterflies and Moths Sphingidae
Sphingidae
at Lepidoptera.pro Images of Sphingidae
Sphingidae
of Costa Rica. Images of Sphingidae
Sphingidae
species in New Zealand List of Sphingidae
Sphingidae
Types from Museum Witt München. Flickr Images

v t e

Extant Lepidoptera
Lepidoptera
families

Kingdom: Animalia Phylum: Arthropoda Class: Insecta Subclass: Pterygota Infraclass: Neoptera Superorder: Endopterygota

Suborder Zeugloptera

Micropterigoidea

Micropterigidae
Micropterigidae
(mandibulate archaic moths)

Suborder Aglossata

Agathiphagoidea

Agathiphagidae (kauri moths)

Suborder Heterobathmiina

Heterobathmioidea

Heterobathmiidae

Suborder Glossata

Dacnonypha

Eriocranioidea

Eriocraniidae

Acanthoctesia

Acanthopteroctetoidea

Acanthopteroctetidae (archaic sun moths)

Lophocoronina

Lophocoronoidea

Lophocoronidae

Neopseustina

Neopseustoidea

Neopseustidae (archaic bell moths)

Exoporia

Hepialoidea

Anomosetidae Hepialidae
Hepialidae
(swift moths, ghost moths) Neotheoridae (Amazonian primitive ghost moths) Palaeosetidae (miniature ghost moths) Prototheoridae (African primitive ghost moths)

Mnesarchaeoidea

Mnesarchaeidae (New Zealand primitive moths)

H e t e r o n e u r a

M o n o t r y s i a

Incurvarioidea

Adelidae
Adelidae
(fairy longhorn moths) Cecidosidae Crinopterygidae Heliozelidae Incurvariidae Prodoxidae
Prodoxidae
(yucca moths)

Andesianoidea

Andesianidae (Andean endemic moths)

Nepticuloidea

Nepticulidae
Nepticulidae
(pigmy, or midget moths) Opostegidae
Opostegidae
(white eyecap moths)

Palaephatoidea

Palaephatidae (Gondwanaland moths)

Tischerioidea

Tischeriidae (trumpet leaf miner moths)

D i t r y s i a

Simaethistoidea

Simaethistidae

Tineoidea

Acrolophidae
Acrolophidae
(burrowing webworm moths) Arrhenophanidae Eriocottidae (Old World spiny-winged moths) Lypusidae Psychidae (bagworm moths) Tineidae
Tineidae
(fungus moths)

Gracillarioidea

Bucculatricidae
Bucculatricidae
(ribbed cocoon makers) Douglasiidae (Douglas moths) Gracillariidae Roeslerstammiidae

Yponomeutoidea

Acrolepiidae
Acrolepiidae
(false diamondback moths) Bedelliidae Glyphipterigidae
Glyphipterigidae
(sedge moths) Heliodinidae Lyonetiidae Plutellidae Yponomeutidae (ermine moths) Ypsolophidae

Gelechioidea

Autostichidae Batrachedridae Blastobasidae Coleophoridae
Coleophoridae
(case-bearers, case moths) Cosmopterigidae
Cosmopterigidae
(cosmet moths) Elachistidae
Elachistidae
(grass-miner moths) Gelechiidae
Gelechiidae
(twirler moths) Lecithoceridae
Lecithoceridae
(long-horned moths) Metachandidae Momphidae
Momphidae
(mompha moths) Oecophoridae
Oecophoridae
(concealer moths) Pterolonchidae Scythrididae
Scythrididae
(flower moths) Xyloryctidae
Xyloryctidae
(timber moths)

Galacticoidea

Galacticidae

Zygaenoidea

Heterogynidae Zygaenidae
Zygaenidae
(burnet, forester, or smoky moths) Himantopteridae Lacturidae Somabrachyidae Megalopygidae (flannel moths) Aididae Anomoeotidae Cyclotornidae Epipyropidae
Epipyropidae
(planthopper parasite moths) Dalceridae
Dalceridae
(slug caterpillars) Limacodidae
Limacodidae
(slug, or cup moths)

Cossoidea

Cossidae
Cossidae
(carpenter millers, or goat moths) Dudgeoneidae (dudgeon carpenter moths)

Sesioidea

Brachodidae (little bear moths) Castniidae
Castniidae
(castniid moths: giant butterfly-moths, sun moths) Sesiidae
Sesiidae
(clearwing moths)

Choreutoidea

Choreutidae
Choreutidae
(metalmark moths)

Tortricoidea

Tortricidae
Tortricidae
(tortrix moths)

Urodoidea

Urodidae
Urodidae
(false burnet moths)

Schreckensteinioidea

Schreckensteiniidae
Schreckensteiniidae
(bristle-legged moths)

Epermenioidea

Epermeniidae
Epermeniidae
(fringe-tufted moths)

Alucitoidea

Alucitidae (many-plumed moths) Tineodidae (false plume moths)

Pterophoroidea

Pterophoridae
Pterophoridae
(plume moths)

Whalleyanoidea

Whalleyanidae

Immoidea

Immidae

Copromorphoidea

Copromorphidae (tropical fruitworm moths) Carposinidae
Carposinidae
(fruitworm moths)

Hyblaeoidea

Hyblaeidae
Hyblaeidae
(teak moths)

Pyraloidea

Pyralidae
Pyralidae
(snout moths) Crambidae
Crambidae
(grass moth)

Thyridoidea

Thyrididae
Thyrididae
(picture-winged leaf moths)

Mimallonoidea

Mimallonidae (sack bearer moths)

Lasiocampoidea

Lasiocampidae
Lasiocampidae
(eggars, snout moths, or lappet moths)

Bombycoidea

Anthelidae
Anthelidae
(Australian lappet moth) Bombycidae
Bombycidae
(silk moths) Brahmaeidae
Brahmaeidae
(Brahmin moths) Carthaeidae (Dryandra moth) Endromidae
Endromidae
(Kentish glory and relatives) Eupterotidae Lemoniidae Saturniidae
Saturniidae
(saturniids) Sphingidae
Sphingidae
(hawk moths, sphinx moths and hornworms) Phiditiidae

Noctuoidea

Doidae Erebidae
Erebidae
(underwing, tiger, tussock, litter, snout, owlet moths) Euteliidae Noctuidae
Noctuidae
(daggers, sallows, owlet moths, quakers, cutworms, darts) Nolidae
Nolidae
(tuft moths) Notodontidae
Notodontidae
(prominents, kittens) Oenosandridae

Drepanoidea

Epicopeiidae
Epicopeiidae
(oriental swallowtail moths) Drepanidae
Drepanidae
(hook-tips)

Geometroidea

Sematuridae Uraniidae Geometridae (geometer moths)

Cimelioidea

Cimeliidae (gold moths)

Calliduloidea

Callidulidae
Callidulidae
(Old World butterfly-moths)

Superfamily unassigned

Millieriidae

Rhopalocera (butterflies)

Hedyloidea

Hedylidae
Hedylidae
(American moth-butterflies)

Hesperioidea

Hesperiidae (skippers)

Papilionoidea (true butterflies)

Lycaenidae
Lycaenidae
(gossamer-winged butterflies: blues, coppers and relatives) Nymphalidae
Nymphalidae
(brush-footed, or four-footed butterflies) Papilionidae (swallowtail butterflies) Pieridae
Pieridae
(whites, yellows, orangetips, sulphurs) Riodinidae
Riodinidae
(metalmarks)

Note: division Monotrysia
Monotrysia
is not a clade.

Taxonomy of the Lepidoptera Lists by region

Taxon identifiers

Wd: Q157683 BAMONA: Sphingidae BugGuide: 193 EoL: 905 EPPO: 1SPHIF Fauna Europaea: 6818 Fossilworks: 71815 GBIF: 8868 ITIS: 117293 NCBI: 7128 WoRMS: 989099

Authority control

LCCN: sh85126

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