The Sphingidae are a family of
moths
Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximately 160,000 species of moth, many of w ...
(
Lepidoptera
Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as “hornworms”; it includes about 1,450
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
. It is best represented in the
tropics
The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in
the Southern Hemisphere at S. The tropics are also referred to ...
, 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 hummingbirds 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 zoologist, specialising in arthropods. Having trained as a Roman Catholic priest before the French Revolution, Latreille was imprisoned, and only regained his freedom ...
in 1802.
Some hawk moths, such as the hummingbird hawk-moth
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 hummingbirds, as they feed on the nectar of tube-shaped flowers using their ...
or the white-lined sphinx, hover in midair while they feed on nectar
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
from flowers
A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechani ...
, so are sometimes mistaken for hummingbird
Hummingbirds are birds native to the Americas and comprise the biological family Trochilidae. With about 361 species and 113 genera, they occur from Alaska to Tierra del Fuego, but the vast majority of the species are found in the tropics aro ...
s. This hovering capability is only known to have evolved four times in nectar
Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
feeders: in hummingbirds, certain bats, hoverflies, 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 com ...
). 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 ]insect
Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three pairs ...
s; some are capable of flying at over .[
] They have wingspans from to over .
Description
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 membrane ( tympanum) stretched across a frame backed by an air sac and associated sensory neurons. Sounds vibrate the membrane, and the vibrations are sensed by ...
s, but members of the group Choerocampini have hearing organs on their heads.[ They have a ]frenulum
A frenulum (or frenum, plural: 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 h ...
and retinaculum
A retinaculum (plural ''retinacula'') is a band of thickened deep fascia around tendons that holds them in place. It is not part of any muscle. Its function is mostly to stabilize a tendon. The term retinaculum is New Latin, derived from the Latin ...
to join hindwings and forewings.[ The thorax, abdomen, and wings are densely covered in scales. Some sphingids have a rudimentary proboscis, 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, vespertine, or both. This is distinguished from diurnal and nocturnal behavior, where an animal is active during the hours of daylig ...
or nocturnal
Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite.
Nocturnal creatures generally have highly developed sens ...
, 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 or morning glory
Morning glory (also written as morning-glory) is the common name for over 1,000 species of flowering plants in the family Convolvulaceae, whose current taxonomy and systematics are in flux. Morning glory species belong to many genera, some of ...
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 hawkmoths 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 call males to them 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 bee hawk (''Cephonodes hylas
''Cephonodes hylas'', the coffee bee hawkmoth, pellucid hawk moth or coffee clearwing, is a moth of the family Sphingidae. The species was first described by Carl Linnaeus in 1771. A widely distributed moth, it is found in the Near East, Middle E ...
virescens''), '' Macroglossum hirundo'', and '' Macroglossum trochilus'', are diurnal.[
A number of species are known to be migratory, all in the Sphingini 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 Species description, first described by Carl Linnaeus in his 1763 ''Centuria Insectorum''.
Commonly known as the Carolina sphinx moth and ...
'', 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
In classical physics and special relativity, an inertial frame of reference (also called inertial reference frame, inertial frame, inertial space, or Galilean reference frame) is a frame of reference that is not undergoing any acceleration. ...
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 scol ...
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 Sym ...
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 Lepidoptera, though they can also be found on other larval insects such as sawflies and a few other types of 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, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...
.[ Many are cryptic greens and browns, and have ]countershading
Countershading, or Thayer's law, is a method of camouflage in which 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, reptiles, birds, fish, and ...
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 larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
The ...
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 directly from west to east, it stands on the Giza Plateau on the west bank of the Nile in Giza, E ...
, 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 is the anterior part of the alimentary canal, from the mouth to the duodenum at the entrance of the bile duct. Beyond the stomach, the foregut is attached to the abdominal walls by mesentery. The foregut arises from the endoderm, deve ...
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 the soil to pupate, where they remain for two to three 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.][ 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
Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subst ...
molecule
A molecule is a group of two or more atoms held together by attractive forces known as chemical bonds; depending on context, the term may or may not include ions which satisfy this criterion. In quantum physics, organic chemistry, and bioch ...
s, and chew and mash the food into very small bits. Some species can tolerate quite high concentrations of specific toxins. Tobacco hornworm
''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 m ...
s (''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 tribe was described b ...
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
''Hyles euphorbiae'', the spurge hawk-moth, is a European moth of the family Sphingidae. This hawk moth is used as an agent of biological pest control against the noxious weed leafy spurge (''Euphorbia virgata''), but usually only in conjunction ...
'' and ''Daphnis nerii
''Daphnis nerii'', the oleander hawk-moth or army green moth, is a moth of the family Sphingidae. It was described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''.
Distribution
''Daphnis nerii'' is a large hawk-moth found in wi ...
'', 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
The name death's-head hawkmoth refers to any of three moth species of the genus ''Acherontia'' (''Acherontia atropos'', ''Acherontia styx'' and ''Acherontia lachesis''). The former species is found in Europe and throughout Africa, the latter t ...
s steal 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 pollen vectors, which can be abiotic (wind and water) or biotic, such as birds, bees, flies, and so forth through a process c ...
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, Bournois – 12 May 1831, Paris) was an eminent French botanist known for his work collecting and describing orchids from the three islands of Madagascar, Mauritius and Réunion.
Introd ...
, and later, Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
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 a British naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution through natural se ...
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 biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics (morphology), but that can successfully interbreed. Not all species ...
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 sphinx moth from Southern Africa (Zimbabwe, Zambia, ...
'', 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. morgani praedicta'' and the African subspecies ''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
SUPERFAMILY is a database and search platform of structural and functional annotation for all proteins and genomes. It classifies amino acid sequences into known structural domains, especially into SCOP superfamilies. Domains are functional, str ...
, Sphingoidea, but is alternatively included with the more encompassing Bombycoidea
Bombycoidea is a superfamily of moths. It contains the silk moths, emperor moths, sphinx moths
The Sphingidae are a family of moths ( Lepidoptera) called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars ...
. 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 subfamily botanical names with "-oideae", and zoologi ...
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 first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''.
Description
It has a wingspan (generally deflexed ...
(''Sphinx ligustri'')
* White-lined Sphinx (''Hyles lineata'')
* Death's-head hawk moth (''Acherontia atropos'')
* Lime hawk moth (''Mimas tiliae'')
* Poplar hawk moth
''Laothoe populi'', the poplar hawk-moth, is a moth of the family Sphingidae. The species was first described by Carl Linnaeus in his 1758 10th edition of ''Systema Naturae''. It is found throughout the Palearctic region and the Near East and ...
(''Laothoe populi'')
* Convolvulus hawk moth (''Agrius convolvuli'')
* Catalpa sphinx (''Ceratomia catalpae'')
* Hummingbird hawk-moth
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 hummingbirds, as they feed on the nectar of tube-shaped flowers using their ...
(''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'' 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 m ...
(''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