About 200 genera,
roughly 1,450 species
Sphingidae are a family of moths (Lepidoptera), commonly known as
hawk moths, sphinx moths, and hornworms; it includes about 1,450
species. It is best represented in the tropics, but species are
found in every region. They are moderate to large in size and are
distinguished among moths for their rapid, sustained flying
ability. 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 (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.
Sphingids are some of the faster flying insects; some are capable of
flying at over 5.3 m/s (12 miles per hour). 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. Females lay
translucent, greenish, flattened, smooth eggs, usually singly on the
host plants. Egg development time varies highly, from three to 21
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. Usually, their bodies lack any
hairs or tubercules, but most species have a "horn" at the posterior
end, which may be reduced to a button, or absent, in the final
instar. 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'. Some tropical larvae are thought to mimic snakes.
Larvae are quick to regurgitate their sticky, often toxic, foregut
contents on attackers such as ants and parasitoids. Development
rate depends on temperature, and to speed development, some northern
and high-altitude species sunbathe. Larvae burrow into soil to
pupate, where they remain for 2–3 weeks before they emerge as
In some Sphingidae, the pupa has a free proboscis, rather than being
fused to the pupal case as is most common in the Macrolepidoptera.
They have a cremaster at the tip of the abdomen. Usually, they
pupate off the host plant, in an underground chamber, among rocks, or
in a loose cocoon. In most species, the pupa is the overwintering
A feeding hawk moth at
Olympiaki Akti (Greece)
2 Food plants
3 Relationships and species
4 See also
6 External links
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, and couples the wings in flight for
efficiency. In males it consists of a single but compound bristle
while in females it consists of several compound bristles. A
reduced frenulum is found in some Smerinthini.
Antennae are generally not very feathery, even in the males. They
lack tympanal organs, but members of the group Choerocampini have
hearing organs on their heads. They have a frenulum and retinaculum
to join hind wings and fore wings. The thorax, abdomen, and wings
are densely covered in scales. Some sphingids have a rudimental
proboscis, but most have a very long one, which is used to feed on
nectar from flowers. Most are crepuscular or nocturnal, but some
species fly during the day. Both males and females are relatively
long-lived (living 10 to 30 days). Prior to flight, most species
shiver their flight muscles to warm them up, and, during flight, body
temperatures may surpass 40 °C (104 °F).
In some species, differences in form between the sexes is quite
marked. For example, in the African species
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.
Females call males to them with pheromones. The male may douse the
female with a pheromone before mating.
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.
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. 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.
Bee hawk moth (Cephonodes kingii), Crows Nest, New South Wales
Sphingid larvae tend to be specific feeders, rather than
generalists. 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. 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
Macroglossinae, but members of the
Smerinthinae that were tested are
susceptible. 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
Hemaris thysbe or hummingbird clearwing moth feeding on lantana
Most adults feed on nectar, although a few tropical species feed on
eye secretions, and the death's-head hawkmoth steals honey from
bees. Night-flying sphingids tend to prefer pale flowers with long
corolla tubes and a sweet odour, a pollination syndrome known as
'sphingophily'. 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. 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
"[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 there must be
moths with probosces capable of extension to a length of between 10
and 12 inches! [30 cm]"
Alfred Russel Wallace
Alfred Russel Wallace published a sort of "wanted poster" (properly, a
drawing in a book)[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 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."
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 praedicta, 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
Oleander hawk moth (Daphnis nerii), in Mangaon, Maharashtra
A sphinx moth, subfamily Macroglossinae, in Cibodas Botanical Garden,
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,
Sphinginae and Macroglossinae. 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 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
Pandora sphinx moth (Eumorpha pandorus)
Tomato worm (Manduca quinquemaculata)
Tobacco hornworm (Manduca sexta)
Sphingidae species list
Twin-spotted Sphinx moth
List of moths of India
List of moths of Great Britain (Sphingidae)
^ van Nieukerken; et al. (2011). "Order
Lepidoptera Linnaeus, 1758.
In: Zhang, Z.-Q. (Ed.)
Animal biodiversity: An outline of higher-level
classification and survey of taxonomic richness" (PDF). Zootaxa. 3148:
^ 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
Xanthopan morganii (Walker), the tribe Acherontiini,
and allied long-tongued hawkmoths (Lepidoptera: Sphingidae,
Sphinginae)". Zoological Journal of the Linnean Society. 135 (4):
^ 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 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.
^ 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.
^ 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
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European Butterflies and Moths
Sphingidae at Lepidoptera.pro
Sphingidae of Costa Rica.
Sphingidae species in New Zealand
Sphingidae Types from Museum Witt München.
Micropterigidae (mandibulate archaic moths)
Agathiphagidae (kauri moths)
Acanthopteroctetidae (archaic sun moths)
Neopseustidae (archaic bell moths)
Hepialidae (swift moths, ghost moths)
Neotheoridae (Amazonian primitive ghost moths)
Palaeosetidae (miniature ghost moths)
Prototheoridae (African primitive ghost moths)
Mnesarchaeidae (New Zealand primitive moths)
Adelidae (fairy longhorn moths)
Prodoxidae (yucca moths)
Andesianidae (Andean endemic moths)
Nepticulidae (pigmy, or midget moths)
Opostegidae (white eyecap moths)
Palaephatidae (Gondwanaland moths)
Tischeriidae (trumpet leaf miner moths)
Acrolophidae (burrowing webworm moths)
Eriocottidae (Old World spiny-winged moths)
Psychidae (bagworm moths)
Tineidae (fungus moths)
Bucculatricidae (ribbed cocoon makers)
Douglasiidae (Douglas moths)
Acrolepiidae (false diamondback moths)
Glyphipterigidae (sedge moths)
Yponomeutidae (ermine moths)
Coleophoridae (case-bearers, case moths)
Cosmopterigidae (cosmet moths)
Elachistidae (grass-miner moths)
Gelechiidae (twirler moths)
Lecithoceridae (long-horned moths)
Momphidae (mompha moths)
Oecophoridae (concealer moths)
Scythrididae (flower moths)
Xyloryctidae (timber moths)
Zygaenidae (burnet, forester, or smoky moths)
Megalopygidae (flannel moths)
Epipyropidae (planthopper parasite moths)
Dalceridae (slug caterpillars)
Limacodidae (slug, or cup moths)
Cossidae (carpenter millers, or goat moths)
Dudgeoneidae (dudgeon carpenter moths)
Brachodidae (little bear moths)
Castniidae (castniid moths: giant butterfly-moths, sun moths)
Sesiidae (clearwing moths)
Choreutidae (metalmark moths)
Tortricidae (tortrix moths)
Urodidae (false burnet moths)
Schreckensteiniidae (bristle-legged moths)
Epermeniidae (fringe-tufted moths)
Alucitidae (many-plumed moths)
Tineodidae (false plume moths)
Pterophoridae (plume moths)
Copromorphidae (tropical fruitworm moths)
Carposinidae (fruitworm moths)
Hyblaeidae (teak moths)
Pyralidae (snout moths)
Crambidae (grass moth)
Thyrididae (picture-winged leaf moths)
Mimallonidae (sack bearer moths)
Lasiocampidae (eggars, snout moths, or lappet moths)
Anthelidae (Australian lappet moth)
Bombycidae (silk moths)
Brahmaeidae (Brahmin moths)
Carthaeidae (Dryandra moth)
Endromidae (Kentish glory and relatives)
Sphingidae (hawk moths, sphinx moths and hornworms)
Erebidae (underwing, tiger, tussock, litter, snout, owlet moths)
Noctuidae (daggers, sallows, owlet moths, quakers, cutworms, darts)
Nolidae (tuft moths)
Notodontidae (prominents, kittens)
Epicopeiidae (oriental swallowtail moths)
Geometridae (geometer moths)
Cimeliidae (gold moths)
Callidulidae (Old World butterfly-moths)
Hedylidae (American moth-butterflies)
Lycaenidae (gossamer-winged butterflies: blues, coppers and relatives)
Nymphalidae (brush-footed, or four-footed butterflies)
Papilionidae (swallowtail butterflies)
Pieridae (whites, yellows, orangetips, sulphurs)
Monotrysia is not a clade.
Taxonomy of the Lepidoptera
Lists by region
Fauna Europaea: 6818