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Sphingidae

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.

Hawk moths
Temporal range: Miocene–Recent
Convolvulus hawk-moth, Agrius convolvuli
Scientific classification
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

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 (19 km/h).[4] They have wingspans from 4 cm (1+12 in) to over 10 cm (4 in).

Description edit

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), 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.[6] 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 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 edit

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

Life cycle edit

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 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 edit

Larvae edit

 
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 edit

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 edit

 
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 edit

References edit

  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. ^ 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
  3. ^ a b c d 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. ^ 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
  6. ^ a b 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). (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). . 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.

External links edit

  •   Media related to Sphingidae at Wikimedia Commons
  •   Data related to Sphingidae at Wikispecies
  • Sphingidae. Insecta.pro
  • Images of Sphingidae of Costa Rica 2019-07-20 at the Wayback Machine
  • Images of Sphingidae species in New Zealand 2020-08-09 at the Wayback Machine
  • List of Sphingidae Types from Museum Witt München
  • Images from Flickr

sphingidae, confused, with, sphindidae, complete, species, list, this, family, species, list, miraculous, tales, ladybug, noir, villain, known, hawk, moth, list, miraculous, tales, ladybug, noir, characters, hawk, moth, family, moths, commonly, called, sphinx,. Not to be confused with Sphindidae For a complete species list of this family see Sphingidae species list For the Miraculous Tales of Ladybug amp Cat Noir villain known as Hawk Moth see List of Miraculous Tales of Ladybug amp Cat Noir characters Hawk Moth 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 Andre Latreille in 1802 Hawk mothsTemporal range Miocene Recent PreꞒ Ꞓ O S D C P T J K Pg N Convolvulus hawk moth Agrius convolvuli Scientific classification Domain Eukaryota Kingdom Animalia Phylum Arthropoda Class Insecta Order Lepidoptera Superfamily Bombycoidea Family SphingidaeLatreille 1802 Type species Sphinx ligustriLinnaeus 1758 Subfamilies Macroglossinae Smerinthinae Sphinginae Diversity About 200 genera roughly 1 450 species 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 19 km h 4 They have wingspans from 4 cm 1 1 2 in to over 10 cm 4 in Contents 1 Description 1 1 Behavior 1 2 Flight 2 Life cycle 3 Food plants 3 1 Larvae 3 2 Adults 4 Relationships and species 5 See also 6 References 7 External linksDescription editSphingid 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 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 6 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 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 edit 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 nbsp Vine hawk moth larva Hippotion celerio nbsp An example of the posterior horn seen on the tomato hornwormLife cycle editMost 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 nbsp A Hyles gallii caterpillar seeking a place to pupate the color of the caterpillar darkens before pupation nbsp 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 editLarvae edit nbsp 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 edit 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 edit nbsp 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 Privet hawk moth Sphinx ligustri White lined Sphinx 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 Tobacco hornworm Manduca sexta See also editSphingidae species listReferences edit 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 a b c d e f g h i Scoble Malcolm J 1995 The Lepidoptera Form Function and Diversity 2nd edition Oxford University Press amp 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 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 doi 10 1242 jeb 198 8 1665 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 amp Natural History Museum London ISBN 0 946589 21 6 a b Pinhey E 1962 Hawk Moths of Central and Southern Africa Longmans Southern Africa Cape Town Holloway J D 1987 Family Sphingidae The Moths of Borneo Southene Sdn Bhd 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 Sane S Dieudonne 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 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 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 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 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 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 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 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 Common I F B 1990 Moths of Australia Leiden Brill p 24 ISBN 9789004092273 External links edit nbsp Media related to Sphingidae at Wikimedia Commons nbsp Data related to Sphingidae at Wikispecies Sphingidae Insecta pro Images of Sphingidae of Costa Rica Archived 2019 07 20 at the Wayback Machine Images of Sphingidae species in New Zealand Archived 2020 08 09 at the Wayback Machine List of Sphingidae Types from Museum Witt Munchen Images from Flickr Retrieved from https en wikipedia org w index php title Sphingidae amp oldid 1192912148, wikipedia, wiki, book, books, library,

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