Sphingidae

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Hawk moths
Temporal range: Miocene–Recent
Convolvulus hawk-moth (Agrius convolvuli) 2.jpg
Convolvulus hawk-moth, Agrius convolvuli
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Superfamily: Bombycoidea
Family: Sphingidae
Latreille, 1802
Type species
Sphinx ligustri
Subfamilies
Diversity
About 200 genera, roughly 1,450 species

The Sphingidae are a family of moths commonly called sphinx moths, also colloquially known as hawk moths, with many of their caterpillars known as 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 agile and sustained flying ability, similar enough to that of hummingbirds as to be reliably mistaken for them. [2] Their narrow wings and streamlined abdomens are adaptations for rapid flight. The family was named by French zoologist Pierre André Latreille in 1802.

Contents

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 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 (19 km/h). [4] They have wingspans from 4 cm (1+1⁄2 in) to over 10 cm (4 in).

Description

Sphingid's antennae are generally not very feathery, even in 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 hindwings and forewings. [2] The thorax, abdomen, and wings are densely covered in scales. Some sphingids have a rudimentary 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 (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 (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. [6] Females attract males with pheromones. The male may douse the female with a pheromone [5] 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. [6]

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. [7]

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 Coriolis forces that are linearly proportional to the angular velocity of the body. [8] 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. [9]

Vine hawk-moth larva (Hippotion celerio) Hawk moth caterpillar.jpg
Vine hawk-moth larva (Hippotion celerio)
An example of the posterior "horn" seen on the tomato hornworm Tomato hornworm.jpg
An example of the posterior "horn" seen on the tomato hornworm

Life cycle

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 caterpillar seeking a place to pupate: the color of the caterpillar darkens before pupation. Hyles-gallii-caterpillar.jpg
A Hyles gallii caterpillar seeking a place to pupate: the color of the caterpillar darkens before pupation.
Hyles euphorbiae pupa Sphingidaepupa.JPG
Hyles euphorbiae pupa

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 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". [5] Some tropical larvae are thought to mimic snakes. [2] [10] 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 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. [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.

Food plants

Larvae

C. kingii, Crows Nest Bee hawk moth newspaper.jpg
C. kingii , Crows Nest

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. [11] 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 and Macroglossinae, but members of the Smerinthinae that were tested are susceptible. [12] 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. [5]

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. [5] Night-flying sphingids tend to prefer pale flowers with long corolla tubes and a sweet odor, 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-centimetre-long (12 in) 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:

[A. sesquipetale has] nectaries 11 and a half inches long [29 cm], with only the lower inch and a half [4 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! [25 and 30 cm] [13]

Alfred Russel Wallace published a sort of "wanted poster" (properly, a drawing in a book) [14] of what this lepidopteran 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 [8 cm] could reach the nectar in the largest flowers of Angraecum 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. [15]

The predicted sphingid was discovered 21 years later and described as a subspecies of the one African species studied by Wallace: Xanthopan morganii praedicta , [16] 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. [17] 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

A sphinx moth, subfamily Macroglossinae, in Cibodas Botanical Garden, Java Hawk moth.jpg
A sphinx moth, subfamily Macroglossinae, in Cibodas Botanical Garden, Java

The 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 and Macroglossinae. [18] Around 1,450 species of hawk moths are classified into around 200 genera. Some of the best-known hawk moth species are:

See also

Related Research Articles

<i>Nicotiana</i> Genus of flowering plants in the nightshade family Solanaceae

Nicotiana is a genus of herbaceous plants and shrubs in the family Solanaceae, that is indigenous to the Americas, Australia, Southwestern Africa and the South Pacific. Various Nicotiana species, commonly referred to as tobacco plants, are cultivated as ornamental garden plants. N. tabacum is grown worldwide for the cultivation of tobacco leaves used for manufacturing and producing tobacco products, including cigars, cigarillos, cigarettes, chewing tobacco, dipping tobacco, snuff, and snus.

<i>Angraecum</i> Genus of orchids

Angraecum, also known as comet orchid, is a genus of the family Orchidaceae native to tropical and South Africa, as well as Sri Lanka. It contains 223 species.

<span class="mw-page-title-main">Hummingbird hawk-moth</span> Species of moth

The hummingbird hawk-moth 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 long proboscis while hovering in the air; this resemblance is an example of convergent evolution.

<span class="mw-page-title-main">Death's-head hawkmoth</span> Species of moth

The name death's-head hawkmoth refers to any of three moth species of the genus Acherontia. The former species is found throughout Africa and in Europe, the latter two are Asian; most uses of the common name refer to the African species. These moths are easily distinguishable by the vaguely human skull-shaped pattern of markings on the thorax. They are large nocturnal moths with brown and yellow or orange coloring, and all three species are fairly similar in size, coloration and life cycle.

<i>Manduca sexta</i> Species of moth

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.

<span class="mw-page-title-main">Predictive power</span> Ability of a scientific theory to generate testable predictions

The concept of predictive power, the power of a scientific theory to generate testable predictions, differs from explanatory power and descriptive power in that it allows a prospective test of theoretical understanding.

<i>Manduca quinquemaculata</i> Species of moth

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 on their posterior end and their use of tomatoes as host plants. Tomato hornworms are closely related to the tobacco hornworm Manduca sexta. This confusion arises because caterpillars of both species have similar morphologies and feed on the foliage of various plants from the family Solanaceae, so either species can be found on tobacco or tomato leaves. Because of this, the plant on which the caterpillar is found does not indicate its species.

<i>Manduca</i> Genus of moths

Manduca is a genus of moths in the family Sphingidae, the hawkmoths. The genus is used as a model in the biological sciences. The tobacco hornworm and the tomato hornworm in particular have been well studied. The genus was erected by Jacob Hübner in 1807.

<i>Agrius convolvuli</i> Species of moth

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 as hīhue.

<i>Hemaris</i> Genus of moths

Hemaris is a genus of sphinx moths in the subfamily Macroglossinae, which is native to the Holarctic. Their main host plants are herbs and shrubs of the teasel and honeysuckle families. Moths in genus Hemaris are known collectively as clearwing moths or hummingbird moths in the US and Canada and bee hawk-moths in Britain. The related Old World hummingbird hawk-moths, genus Macroglossum, are similar in appearance and habits. Both genera have tails that are provided with an expansile truncated tuft of hairs, but only Hemaris has the disc of the wings transparent, as these scales are dropped soon after eclosion.

<i>Hemaris thysbe</i> Species of moth

Hemaris thysbe, the hummingbird clearwing, is a moth of the family Sphingidae (hawkmoths). Coloration varies between individuals, but typically the moth is olive green and burgundy on its back, and white or yellow and burgundy on the underside. Its wings are transparent with a reddish-brown border. It has light-colored legs, which combined with the lack of striping on the underside is diagnostic. Beating its wings rapidly, H. thysbe hovers to collect nectar from a variety of flowers. The combination of its appearance and its behavior commonly leads to it being confused with a hummingbird or bumblebee.

<i>Hyles lineata</i> Species of moth

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 and flight patterns.

<span class="mw-page-title-main">Sphinginae</span> Subfamily of moths

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, being a very common and recognizable species, the death's-head hawkmoths of Silence of the Lambs fame, and Xanthopan morganii with its enormous proboscis.

<i>Angraecum sesquipedale</i> Species of orchid

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 French botanist Louis-Marie Aubert du Petit-Thouars in 1798, but was not described until 1822. It is noteworthy for its long spur and its association with the naturalist Charles Darwin, who surmised that the flower was pollinated by a then undiscovered moth with a proboscis whose length was unprecedented at the time. Darwin's prediction went unverified for 21 years after his death, until just such a moth was discovered and his conjecture vindicated. The story of its postulated pollinator has come to be seen as one of the celebrated predictions of the theory of evolution.

<i>Xanthopan</i> Genus of moths

Xanthopan is a monotypic genus of sphinx moth, with Xanthopan morganii, commonly called Morgan's sphinx moth, as its sole species. It is a very large sphinx moth from Southern Africa and Madagascar. Little is known about its biology, though the adults have been found to visit orchids and are one of the main pollinators of several of the Madagascar endemic baobab (Adansonia) species, Adansonia perrieri or Perrier's baobab.

<i>Macroglossum glaucoptera</i> Species of moth

Macroglossum glaucoptera, the dark hummingbird hawkmoth, is a moth of the family Sphingidae. It was described by Arthur Gardiner Butler in 1875. It is known from Sri Lanka, Thailand, southern China, Vietnam, Malaysia (Peninsular), Indonesia and the Philippines (Mindanao). Single specimen recorded from Papua New Guinea.

<i>Coelonia solani</i> Species of moth

Coelonia solani is a moth of the family Sphingidae. It is known from Mauritius, Réunion, Madagascar and the Comoro Islands. It is a pollinator of some species of baobab in Madagascar, including Adansonia za.

<span class="mw-page-title-main">Nectar spur</span> Nectar spur, secrets

A nectar spur is a hollow extension of a part of a flower. The spur may arise from various parts of the flower: the sepals, petals, or hypanthium, and often contain tissues that secrete nectar (nectaries). Nectar spurs are present in many clades across the angiosperms, and are often cited as an example of convergent evolution.

<i>Angraecum longicalcar</i> Species of plant

Angraecum longicalcar, also known as the long spurred Angraecum, is a large critically endangered orchid endemic to the Central Highlands of Madagascar. This lithophytic species is noteworthy for its exceptionally long 40 cm nectar spur, rivalling that of the famous Darwins' orchid , and as such may have the longest spur of any orchid species.

References

  1. ↑ 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: 212–221. doi:10.11646/zootaxa.3148.1.41.
  2. 1 2 3 4 5 6 7 8 9 Scoble, Malcolm J. (1995): The Lepidoptera: Form, Function and Diversity (2nd edition). Oxford University Press & Natural History Museum London. ISBN   0-19-854952-0
  3. 1 2 3 4 Kitching, Ian J (2002). "The phylogenetic relationships of Morgan's Sphinx, 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 .
  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. doi:10.1242/jeb.198.8.1665. PMID   9319572 . Retrieved 10 August 2012.
  5. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Pittaway, A. R. (1993): The Hawkmoths of the Western Palaearctic. Harley Books & Natural History Museum, London. ISBN   0-946589-21-6
  6. 1 2 Pinhey, E. (1962): Hawk Moths of Central and Southern Africa. Longmans Southern Africa, Cape Town.
  7. ↑ Holloway, J.D. (1987). "Family Sphingidae". The Moths of Borneo. Southene Sdn Bhd.
  8. ↑ McNiell Alexander, R. (February 2007). "Antennae as Gyroscopes" (PDF). Science. 315 (5813): 771–772. doi:10.1126/science.1136840. PMID   17289963. S2CID   118833201. Archived from the original (PDF) on 2016-03-03. Retrieved 2014-12-06.
  9. ↑ Sane, S.; Dieudonné, A.; Willis, M.; Daniel, T. (February 2007). "Antennal mechanosensors mediate flight control in moths". Science. 315 (5813): 863–866. Bibcode:2007Sci...315..863S. CiteSeerX   10.1.1.205.7318 . doi:10.1126/science.1133598. PMID   17290001. S2CID   2429129.
  10. ↑ 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. S2CID   53263767.
  11. ↑ Bernays, E. A.; Janzen, D. H. (1988). "Saturniid and sphingid caterpillars: two ways to eat leaves". Ecology . 69 (4): 1153–1160. Bibcode:1988Ecol...69.1153B. doi:10.2307/1941269. JSTOR   1941269. S2CID   56310810.
  12. ↑ Wink, M.; Theile, Vera (2002). "Alkaloid tolerance in Manduca sexta and phylogenetically related sphingids (Lepidoptera: Sphingidae)". Chemoecology. 12 (1): 29–46. Bibcode:2002Checo..12...29W. doi:10.1007/s00049-002-8324-2. S2CID   41120103.
  13. ↑ 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
  14. ↑ Beccaloni, George (April 2010). "Darwin and Wallace's Predictions Come True | The Alfred Russel Wallace Website". wallacefund.info. Archived from the original on 2018-10-10. Retrieved 2020-08-05.
  15. ↑ Wallace, Alfred R. (1867). "Creation by law". Quarterly Journal of Science. 4: 470–488. (p. 477)
  16. ↑ Rothschild, Walter; Jordan, Karl (1903). "A revision of the lepidopterous family Sphingidae". Novitates Zoologicae. 9 (Supplement): 1–972.
  17. ↑ Netz, Christoph; Renner, Susanne S. (2017). "Long-spurred Angraecum orchids and long-tongued sphingid moths on Madagascar: A time-frame for Darwin's predicted Xanthopan/Angraecum coevolution". Biological Journal of the Linnean Society. 122 (Supplement): 469–478. doi:10.1093/biolinnean/blx086.
  18. ↑ Common, I.F.B. (1990). Moths of Australia . Leiden: Brill. p.  24. ISBN   9789004092273.
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