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The Sphingidae are a family of
moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as suborder Heterocera, but the group is Paraphyly, paraphyletic with respect to butterflies (s ...
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 A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...
. It is best represented in the
tropics The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...
, but species are found in every region.Scoble, Malcolm J. (1995): ''The Lepidoptera: Form, Function and Diversity'' (2nd edition). Oxford University Press & Natural History Museum London. 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 Hummingbirds are birds native to the Americas and comprise the Family (biology), biological family Trochilidae. With approximately 366 species and 113 genus, genera, they occur from Alaska to Tierra del Fuego, but most species are found in Cen ...
s as to be reliably mistaken for them. Their narrow wings and streamlined abdomens are adaptations for rapid flight. The family was named by French zoologist
Pierre André Latreille Pierre André Latreille (; 29 November 1762 – 6 February 1833) was a French zoology, zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained hi ...
in 1802. Some hawk moths, such as the hummingbird hawk-moth or the
white-lined sphinx ''Hyles lineata'', also known as the white-lined sphinx, is a moth of the family Sphingidae. They are sometimes known as a "hummingbird moth" because of their bird-like size (2-3 inch wingspan) and flight patterns. As caterpillars, they have a ...
, hover in midair while they feed on
nectar Nectar is a viscous, sugar-rich liquid produced by Plant, plants in glands called nectaries, either within the flowers with which it attracts pollination, pollinating animals, or by extrafloral nectaries, which provide a nutrient source to an ...
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 Bats are flying mammals of the order Chiroptera (). With their forelimbs adapted as wings, they are the only mammals capable of true and sustained flight. Bats are more agile in flight than most birds, flying with their very long spread-out ...
s,
hoverflies Hoverflies, also called flower flies or syrphids, make up the insect family (biology), family Syrphidae. As their common name suggests, they are often seen Hover (behaviour), hovering or nectaring at flowers; the adults of many species feed main ...
, and these sphingids (an example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
). 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. Sphingids are some of the faster flying
insect Insects (from Latin ') are Hexapoda, hexapod invertebrates of the class (biology), class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (Insect morphology#Head, head, ...
s; some are capable of flying at over . They have wingspans from to over .


Description

Sphingid's antennae are generally not very feathery, even in males. They lack
tympanal organ A tympanal organ (or tympanic organ) is a hearing organ in insects, consisting of a tympanal membrane ( tympanum) stretched across a frame backed by an air sac and associated sensory neurons. Sounds vibrate the membrane, and the vibrations are s ...
s, but members of the group Choerocampini have hearing organs on their heads. They have a
frenulum A frenulum or frenum (: frenula or frena, from the Latin ''frēnulum'', "little bridle", the diminutive of ''frēnum'') is a small fold of tissue that secures the motion of a mobile organ in the body. In human anatomy Frenula on the human ...
and retinaculum to join hindwings and forewings. The thorax, abdomen, and wings are densely covered in scales. Some sphingids have a rudimentary
proboscis A proboscis () is an elongated appendage from the head of an animal, either a vertebrate or an invertebrate. In invertebrates, the term usually refers to tubular arthropod mouthparts, mouthparts used for feeding and sucking. In vertebrates, a pr ...
, but most have a very long one, which is used to feed on nectar from flowers. Most are
crepuscular In zoology, a crepuscular animal is one that is active primarily during the twilight period, being matutinal (active during dawn), vespertine (biology), vespertine/vespertinal (active during dusk), or both. This is distinguished from diurnalit ...
or
nocturnal Nocturnality is a ethology, behavior in some non-human animals characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnality, diurnal meaning the opposite. Nocturnal creatur ...
, but some species fly during the day. Both males and females are relatively long-lived (10 to 30 days). Prior to flight, most species shiver their flight muscles to warm them up, and, during flight, body temperatures may surpass . In some species, differences in form between the sexes is quite marked. For example, in the African species ''
Agrius convolvuli ''Agrius convolvuli'', the convolvulus hawk-moth, is a large hawk-moth. It is common throughout Europe, Asia, Africa, Australia and New Zealand, partly as a migrant. In New Zealand, it is also known as the kumara moth, and in the Māori language ...
'' (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. Pinhey, E. (1962): ''Hawk Moths of Central and Southern Africa''. Longmans Southern Africa, Cape Town. Females attract males with
pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...
s. The male may douse the female with a pheromone 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 diurnal. A number of species are known to be migratory, all in the
Sphingini Sphingini is a tribe of moths of the family Sphingidae. The tribe was described by Pierre André Latreille in 1802. Taxonomy *Genus '' Amphimoea'' *Genus '' Amphonyx'' *Genus ''Apocalypsis'' *Genus '' Ceratomia'' *Genus '' Cocytius'' *Genus '' ...
and Macroglossinae, and specially in the genera ''Agrius'', ''Cephonodes'', ''Macroglossum'', ''Hippotion'' and ''Theretra''.


Flight

In studies with ''
Manduca sexta ''Manduca sexta'' is a moth of the family Sphingidae present through much of the Americas. The species was first described by Carl Linnaeus in his 1763 ''Centuria Insectorum''. Commonly known as the Carolina sphinx moth and the tobacco hawk mo ...
'', 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 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 Johnston's organ is a collection of sensory cells found in the pedicel (the second segment) of the antennae in the class Insecta. Johnston's organ detects motion in the flagellum (third and typically final antennal segment). It consists of sc ...
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.


Life cycle

Most species are multivoltine, capable of producing several generations a year if weather conditions permit.Pittaway, A. R. (1993): ''The Hawkmoths of the Western Palaearctic''. Harley Books & Natural History Museum, London. Females lay translucent, greenish, flattened, smooth eggs, usually singly on the host plants. Egg development time varies highly, from three to 21 days. Sphingid
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...
s are medium to large in size, with stout bodies. They have five pairs of
proleg A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the Order (biology), order Lepidoptera, though they can also be found on larvae of insects such as symphyta, sawflies. In ...
s. 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 An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
. Many are cryptic greens and browns, and have
countershading Countershading, or Thayer's law, is a method of camouflage in which animal coloration, an animal's coloration is darker on the top or upper side and lighter on the underside of the body. This pattern is found in many species of mammals, reptile ...
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 The Great Sphinx of Giza is a limestone statue of a reclining sphinx, a mythical creature with the head of a human and the body of a lion. Facing east, it stands on the Giza Plateau on the west bank of the Nile in Giza, Egypt. The original sh ...
, 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 The foregut in humans is the anterior part of the alimentary canal, from the distal esophagus to the first half of the duodenum, at the entrance of the bile duct. Beyond the stomach, the foregut is attached to the abdominal walls by mesentery. ...
contents on attackers such as ants and
parasitoid In evolutionary ecology, a parasitoid is an organism that lives in close association with its host (biology), host at the host's expense, eventually resulting in the death of the host. Parasitoidism is one of six major evolutionarily stable str ...
s. Development rate depends on temperature, and to speed development, some northern and high-altitude species sunbathe. 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. 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 stage.


Food plants


Larvae

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 Nicotine is a natural product, naturally produced alkaloid in the nightshade family of plants (most predominantly in tobacco and ''Duboisia hopwoodii'') and is widely used recreational drug use, recreationally as a stimulant and anxiolytic. As ...
, as do several other related sphinx moths in the subfamilies
Sphinginae The Sphinginae are a subfamily of the hawkmoths (Sphingidae), moths of the order Lepidoptera. The subfamily was first described by Pierre André Latreille in 1802. Notable taxa include the pink-spotted hawkmoth (''Agrius cingulata''), being a ve ...
and Macroglossinae, but members of the
Smerinthinae The Smerinthinae are a subfamily of Sphingidae moths in the order Lepidoptera. Smerinthinae taxonomy *Tribe Ambulycini *Tribe Smerinthini *Tribe Sphingulini Sphingulini is a tribe of moths of the family Sphingidae The Sphingidae are a ...
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 stage.


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. Night-flying sphingids tend to prefer pale flowers with long corolla tubes and a sweet odor, a
pollination syndrome Pollination syndromes are suites of flower traits that have evolved in response to natural selection imposed by different Pollination, pollen vectors, which can be abiotic (wind and water) or biotic, such as birds, bees, flies, and so forth thro ...
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 ''Angraecum sesquipedale'' , also known as Darwin's orchid, Christmas orchid, Star of Bethlehem orchid, and king of the angraecums, is an epiphytic orchid in the genus ''Angraecum'' endemic to Madagascar. The orchid was first discovered by the Fre ...
''), a rare Malagasy flower with its nectar stored at the bottom of a tube, was described in 1822 by
Louis-Marie Aubert du Petit-Thouars Louis-Marie Aubert du Petit-Thouars (5 November 1758, Saint-Martin-de-la-Place, Bournois – 12 May 1831, Paris) was a French botanist known for his work collecting and describing orchids from the three islands of Madagascar, Mauritius and R� ...
, and later,
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
famously predicted there must be some specialized moth to feed from it:
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was an English naturalist, explorer, geographer, anthropologist, biologist and illustrator. He independently conceived the theory of evolution through natural selection; his 1858 pap ...
published a sort of "
wanted poster A wanted poster (or wanted sign) is a poster distributed to let the public know of a person whom authorities wish to apprehend. They generally include a picture of the person, either a photograph when one is available or of a facial composite ...
" (properly, a drawing in a book) of what this lepidopteran might look like, and, concurring with his colleague, added: The predicted sphingid was discovered 21 years later and described as a
subspecies In Taxonomy (biology), biological classification, subspecies (: subspecies) is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (Morphology (biology), morpholog ...
of the one African species studied by Wallace: ''
Xanthopan morganii praedicta ''Xanthopan'' is a monotypic genus of sphinx moth, with ''Xanthopan morganii'' (often misspelled as "''morgani''"), commonly called Morgan's sphinx moth, as its sole species. It is a very large Sphingidae, sphinx moth from Southern Africa (Zimbab ...
'', 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. 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

The Sphingidae is sometimes assigned its own exclusive superfamily, Sphingoidea, but is alternatively included with the more encompassing
Bombycoidea Bombycoidea is a superfamily of moths, including the silk moths, giant silk moths, sphinx moths, saturniids, and relatives. The superfamily Lasiocampoidea is a close relative and was historically sometimes merged in this group. After many ...
. Following Hodges (1971) two
subfamilies In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end botanical subfamily names with "-oideae", and zool ...
are accepted, namely the
Sphinginae The Sphinginae are a subfamily of the hawkmoths (Sphingidae), moths of the order Lepidoptera. The subfamily was first described by Pierre André Latreille in 1802. Notable taxa include the pink-spotted hawkmoth (''Agrius cingulata''), being a ve ...
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'', the privet hawk moth, is a moth found in most of the Palearctic realm. The species was Species description, first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae, 10th edition of ''Systema Naturae''. ...
(''Sphinx ligustri'') *
White-lined Sphinx ''Hyles lineata'', also known as the white-lined sphinx, is a moth of the family Sphingidae. They are sometimes known as a "hummingbird moth" because of their bird-like size (2-3 inch wingspan) and flight patterns. As caterpillars, they have a ...
(''Hyles lineata'') * 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'', the five-spotted hawkmoth, is a brown and gray hawk moth of the family Sphingidae. The caterpillar, often referred to as the tomato hornworm, can be a major pest in gardens; they get their name from a dark projection ...
(''Manduca quinquemaculata'') * Tobacco hornworm (''Manduca sexta'')


See also

* Sphingidae species list


References


External links


Sphingidae
Insecta.pro


Images of Sphingidae species in New Zealand

List of Sphingidae Types from Museum Witt München

Images from Flickr
{{Authority control Moth families Taxa named by Pierre André Latreille