Hedylidae

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Hedylidae
Macrosoma bahiata.jpg
Macrosoma bahiata
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Clade: Obtectomera
Superfamily: Papilionoidea
Family: Hedylidae
Guenée, 1857, nec. Bergh, 1895
Genus: Macrosoma
Hübner, 1818
Type species
Macrosoma tipulata
Hübner, 1818
Species

see List of species

Diversity
35 currently recognised species
Synonyms
  • EpirritaHübner, 1808 [unavailable name]
  • HedyleGuenée, 1857, type species Hedyle heliconiariaGuenée, 1857
  • PhellinodesGuenée, 1857, type species Phellinodes satellitiataGuenée, 1857
  • VenodesGuenée, 1857, type species Phellinodes satellitiataGuenée, 1857
  • MacrophilaWalker, 1862, type species Macrosoma tipulataHübner, 1818
  • HyphedyleWarren, 1894, type species Hedyle rubedinariaWalker, 1862
  • LasiopatesWarren, 1905, type species Lasiopates hyacinthinaWarren, 1905

Hedylidae, the "American moth-butterflies", is a family of insects in the order Lepidoptera, representing the superfamily Hedyloidea. They have traditionally been viewed as an extant sister group of the butterfly superfamily Papilionoidea, but a 2014 phylogenetic analysis has suggested Hedylidae is a subgroup of Papilionoidea, and not a sister group, and are more accurately referred to as butterflies rather than moths. [1] They are represented by a single Neotropical genus Macrosoma with 35 currently recognized species.

Contents

Taxonomy and systematics

Hedylidae were previously treated as a tribe of Geometridae: Oenochrominae, the "Hedylicae". [2] [3] Prout [4] considered they might even merit treatment as their own family. Scoble first considered them to be a hitherto unrecognised group of butterflies and also suggested Hedylidae might possibly constitute the sister group of the "true" butterflies (Papilionoidea), rather than of (Hesperioidea + Papilionoidea). Weintraub and Miller [5] argued against this placement (but see [6] ). In 1995, Weller and Pashley [7] found that molecular data did indeed place Hedylidae with the butterflies and a more comprehensive study in 2005 [8] based on 57 exemplar taxa, three genes and 99 morphological characters, recovered the genus Macrosoma as sister to the ("Papilionoidea" + Hesperioidea). However, the most recent phylogenetic analyses shows that skippers are true butterflies and therefore within the clade Papilionoidea, whereas the hedylids are a sister group that may be closely related to the obtectomeran moths. [9] This is contrary to some earlier studies that had shown both the skippers and hedylids as being nested within the Papilionoidea. [10] [11]

Since there are no obvious gaps between supposed species groups, according to basic morphological structure, Scoble (1986) synonymised the five pre-existing genera of Hedylidae (33 of which had been described in Phellinodes) into the single genus Macrosoma. [2] However, a phylogenetic analysis of all Macrosoma species is still needed.

Nomenclatural notes

In zoological nomenclature, numerous junior homonyms of Macrosoma (Hübner, 1818) exist, [12] (Macrosoma Leach 1819 (a reptile), Macrosoma de Haan 1826 (Odonata), Macrosoma Robineau-Desvoidy 1830 (Diptera: Muscidae), Macrosoma Brandt 1835 (Coelenterata), Macrosoma Hope 1837 (Coleoptera), Macrosoma Lioy 1864 or 1865 (Diptera: Sarcophagidae), Macrosoma Hammer 1979 [13] (Acarina: Oribatidae). To add to this potential confusion in lists of names, there exist two junior homonyms of Hedyle Guenée, 1857: Hedyle Bergh, 1895 (Opisthobranchia, "Acochlidioidea", Hedylopsidae; [14] currently placed in the genus Hedylopsis Thiele, 1931), [15] and Hedyle Malmgren 1865 (a polychaete worm). [16] The sea slug family name "Hedylidae Bergh, 1895" (type species Hedyle weberi Bergh, 1895) is thus also invalid.

Morphology and identification

The eggs of hedylid moths have an upright configuration and are variable in shape: in Macrosoma inermis they are particularly narrow and spindle-shaped, [17] resembling those of some Pieridae, and in the case of M. tipulata they are more barrel-shaped, [18] like certain Nymphalidae. The larvae resemble (probably convergently) those of Apaturinae. [17] Adult hedylids resemble geometer moths. They share many morphological and genetic characteristics with both the superfamilies Papilionoidea and the Hesperioidea. The abdomen is very long and slim, like many Neotropical butterflies of the subfamilies Ithomiinae and Heliconiinae, hence the name of one Macrosoma species "heliconiaria". Unlike other butterflies, however, the antennae are un-clubbed, but rather filiform or bipectinate. [19] Unlike the family Geometridae, in which they had been placed by the geometer expert L.B.Prout, hedylids lack tympanic organs at the base of the abdomen, but have them on the wings (see under Behaviour). Unlike other butterflies, however (except the unique case of the remarkable Australian skipper butterfly Euschemon rafflesia , whose males possess these structures), the single-spined frenulum and retinaculum are not lost or reduced in males, except in three Macrosoma species where there is no functional wing coupling system. The retinaculum is always lost in females, and the frenulum may be vestigial. [2] The family have been fully catalogued [17] and illustrated in an identification guide. [20]

Butterfly-like characteristics of Hedylidae

  1. "Mesoscutum" with "secondary line of weakness" near median "notal" wing process, [21] as in some representatives of Papilionidea and Hesperioidea (potentially unique butterfly character; [22]
  2. Mesothoracic aorta with horizontal chamber, as in other butterflies (not Papilionidae), but as also in Cossidae; [19] [22]
  3. Metathoracic "furca" resembling a blunt arrowhead; [2] this a variable but potentially unique character in butterflies; [22]
  4. Second median plate of forewing base lying partly under the base of vein "1A+2A", unlike the configuration in moths; [2]
  5. "Postspiracular bar" on first abdominal segment; [2]
  6. Female genitalic "anterior apophyses" reduced; [2]
  7. Male genitalia relatively "deep" dorso-ventrally; [2]
  8. Abdomen curved (especially in males), as in papilionoids; [2]
  9. Abdominal first tergal segment is strongly "pouched" (Scoble 1986; as also in Thyatirinae moths; [22]
  10. "Precoxal" sulcus joining "marginopleural" sulcus; [2]
  11. Male Foreleg pretarsus lost, thus fused into two elements [23] as in nymphalid butterflies, with the mid and hindlegs used for perching, but apparently redeveloped in hesperiids; [2]
  12. Egg upright, spindle-shaped and ribbed [24] as in some Pieridae (e.g. the orange tip butterfly), some other butterflies, and as in some moth groups also; [22]
  13. Larva with "anal comb", [25] as in some Hesperioidea (not however Megathyminae) and Pieridae, but not in other Papilionoidea except one species (and also independently in Tortricidae), that is used for propulsion of frass away from the caterpillar; [22]
  14. Caterpillar with horn-like processes and a "bifid" tail as in many Nymphalidae; [24] [26]
  15. Caterpillar with "secondary setae", as in Pieridae; [24]
  16. Ventral larval proleg "crochet" hooks not forming a complete circle, unlike configuration in hesperiids and papilionoids; [2]
  17. Pupa affixed to the substrate via a silken girdle around the 1st abdominal segment, [27] [28] like in Pieridae (as also in some Geometridae, especially the subfamily Sterrhinae (in which the girdle is around the abdomen), but lost in many Papilionoidea); [2]
  18. Pupal cocoon lost, as in papilionoids, and a few other groups of Lepidoptera; [2]
  19. "Temporal cleavage line" lost in the pupa (as in papilionoids). [2]

Distribution

Hedylidae range in North America south from central Mexico and in South America through the Amazon from southern Peru (where there are a full 26 species, [29] up to 12 at a single site: [30] to central Bolivia and southwestern Brazil [20] ). In the Caribbean, they also occur in Cuba, Jamaica, and Trinidad. [20] [30]

Behaviour

Hedylids are attracted to artificial lights, but occasionally some species can be found flying by day. [31] Thus, they may be involved in some mimicry complexes with Ithomiinae (e.g. the female only of Macrosoma lucivittata). [32] A few species are white [33] and resemble pierid butterflies (e.g. Macrosoma napiaria). Based on a study of Macrosoma heliconiaria, it has been found that hedylids have tympanic organs on their forewings for hearing [34] apparently homologous to the "Vogel's organ" in some Papilionoidea [35] that would help them evade bats at night. They have been shown to exhibit typical moth evasive behaviour towards bats such as erratic spiralling movements and dives. [36] The resting posture is often at a curious angle, [37] with the thorax tilted and the posterior edge of the hindwings nearly touching the substrate (Scoble, 1986). The larvae which lack the prominent horns in the first instar tend to rest on the midrib of the leaf and often skeletonise leaves or at either side produce an untidy patchwork of holes. [38] The elegant pupa is attached by a cremaster and silken girdle [39] and sometimes resembles a bird dropping. [40]

List of species

This list of species is largely based on phenetic characters. [17] [20]

Biology and host plants

The life history of Macrosoma heliconiaria was originally described from plants of Byttneria aculeata in Mexico. [31] This was a historical breakthrough into the biology of hedylids. In this study, Kendall commented notably "I thought the larvae might represent a satyr species, but when the first larva pupated I was sure it was a pierid. The first adult emerged as a complete surprise. The pupa...is secured by girdle and cremaster, not unlike a pierid". Macrosoma cascaria was later also reared on this plant in Panama. [24] More life histories are now known. From these data, known host plants span a wide range of (according to the APG II system) rosid dicotyledonous plants, including the rosid order Myrtales family Melastomataceae (genera Miconia , Conostegia , and Ossaea ), the eurosid I order Malpighiales, families Euphorbiaceae ( Croton ), and Malpighiaceae ( Byrsonima ), the eurosid II orders Sapindales, family Rutaceae ( Zanthoxylum ) and more commonly [41] Malvales, family Malvaceae, tribes: Bombacoideae ( Ochroma ), Malvoideae ( Hampea and also Hibiscus , [20] Byttnerioideae (Byttneria aculeata, Theobroma ) and Grewioideae ( Luehea ). The "green lizard caterpillar" Macrosoma tipulata [42] attacks an economically important local fruit tree "Cupuaçu" ( Theobroma grandiflorum ) in Brazil and can defoliate saplings; the biology of this species has been studied and illustrated in some detail. [18] The larva of this species lives about 15 days in 5 instars, the pupal stage lasts about 7 days and the adult lives about 10 days. M. tipulata and many other species can be found as adults through most of the year. [20]

DNA sequences

A few species have been sequenced for the mitochondrial genes "cytochrome oxidase I", and "ND1" and nuclear genes "Wingless" and "Ef-1?", [43] including Macrosoma semiermis. Some species are currently being barcoded. [44]

Cited literature

  1. Kawahara, Akito Y.; Breinholt, Jesse W. (2014-08-07). "Phylogenomics provides strong evidence for relationships of butterflies and moths". Proceedings of the Royal Society B: Biological Sciences. 281 (1788): 20140970. doi:10.1098/rspb.2014.0970. ISSN   0962-8452. PMC   4083801 . PMID   24966318.
  2. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 Scoble MJ (1986). "The structure and affinities of the Hedyloidea: a new concept of the butterflies". Bull. Br. Mus. (Nat. Hist.) Entomol. 53: 251–286.
  3. Prout LB (1910). "Lepidoptera Heterocera, Fam. Geometridae, Subfam. Oenochrominae". Genera Insectorum. 104: 1–119.
  4. Prout LB (1931). "The American Geometridae". The Macrolepidoptera of the World. 8: 1–144.
  5. Weintraub JD, Miller JS (1987). "The structure and affinities of the Hedyloidea: a new concept of butterflies". Cladistics. 3 (3): 299–304. doi:10.1111/j.1096-0031.1987.tb00512.x. S2CID   221574665.
  6. Scoble, Malcolm J. (1988). "Hedylidae: a response to Weintraub and Miller". Cladistics. 4 (1): 93–6. doi: 10.1111/j.1096-0031.1988.tb00470.x . PMID   34933497. S2CID   85579049.
  7. Weller SJ, Pashley DP (September 1995). "In search of butterfly origins". Molecular Phylogenetics and Evolution. 4 (3): 235–46. Bibcode:1995MolPE...4..235W. doi:10.1006/mpev.1995.1022. PMID   8845961.
  8. Wahlberg N, Braby MF, Brower AV, et al. (August 2005). "Synergistic effects of combining morphological and molecular data in resolving the phylogeny of butterflies and skippers". Proceedings of the Royal Society B. 272 (1572): 1577–86. doi:10.1098/rspb.2005.3124. PMC   1560179 . PMID   16048773.
  9. Heikkilä, Maria; Mutanen, Marko; Wahlberg, Niklas; Sihvonen, Pasi; Kaila, Lauri (2015). "Elusive ditrysian phylogeny: An account of combining systematized morphology with molecular data (Lepidoptera)". BMC Evolutionary Biology. 15 (1): 260. Bibcode:2015BMCEE..15..260H. doi: 10.1186/s12862-015-0520-0 . PMC   4654798 . PMID   26589618.
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  18. 1 2 Lourido, G.; Silva, N.M.; Motta, C.S. (2007). "Biological Parameters and Damage by Macrosoma tipulata Hübner (Lepidoptera: Hedylidae), in Cupuaçu tree [Theobroma grandiflorum (Wild ex Spreng Schum)] in Amazonas, Brazil". Neotropical Entomology. 36 (1): 102–106. doi: 10.1590/S1519-566X2007000100012 . PMID   17420867.
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  23. Ackery, P.R.; de Jong, R; Vane-Wright, R.I. (1999). "The Butterflies: Hedyloidea, Hesperioidea and Papilionoidae". In Kristensen, N.P. (ed.). Lepidoptera, Moths and Butterflies. Vol. 1: Evolution, Systematics, and Biogeography. Volume IV/Part 35: 491 pp. Berlin, New York: Walter de Gruyter. pp. 263–300.
  24. 1 2 3 4 Scoble, M.J.; Aiello, A. (1990). "Moth-like butterflies (Hedylidae: Lepidoptera): a summary, with comments on the egg". Journal of Natural History. 24 (1): 159-164. doi:10.1080/00222939000770101.
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  30. 1 2 Grados, J. (1998). Alonso, A.; F. Dallmeier (eds.). Biodiversity Assessment of the Lower Urubamba Region, Peru: Cashiriari-3 Well Site and the Camisea and Urubamba Rivers. SI/MAB Series #2. Washington, DC: Smithsonian Institution/MAB Biodiversity Program. pp. 119–120.
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  34. Organ of hearing
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Sources

Related Research Articles

<span class="mw-page-title-main">Lepidoptera</span> Order of insects including moths and butterflies

Lepidoptera or lepidopterans is an order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organisms, making it the second largest insect order with 126 families and 46 superfamilies, and one of the most widespread and widely recognizable insect orders in the world.

A common classification of the Lepidoptera involves their differentiation into butterflies and moths. Butterflies are a natural monophyletic group, often given the suborder Rhopalocera, which includes Papilionoidea, Hesperiidae (skippers), and Hedylidae. In this taxonomic scheme, moths belong to the suborder Heterocera. Other taxonomic schemes have been proposed, the most common putting the butterflies into the suborder Ditrysia and then the "superfamily" Papilionoidea and ignoring a classification for moths.

<span class="mw-page-title-main">Papilionoidea</span> Superfamily of butterflies

The superfamily Papilionoidea contains all the butterflies except for the moth-like Hedyloidea.

Butterfly evolution is the origin and diversification of butterflies through geologic time and over a large portion of the Earth's surface. The earliest known butterfly fossils are from the mid Eocene epoch, between 40-50 million years ago. Their development is closely linked to the evolution of flowering plants, since both adult butterflies and caterpillars feed on flowering plants. Of the 220,000 species of Lepidoptera, about 45,000 species are butterflies, which probably evolved from moths. Butterflies are found throughout the world, except in Antarctica, and are especially numerous in the tropics; they fall into eight different families.

<span class="mw-page-title-main">External morphology of Lepidoptera</span> External features of butterflies and moths

The external morphology of Lepidoptera is the physiological structure of the bodies of insects belonging to the order Lepidoptera, also known as butterflies and moths. Lepidoptera are distinguished from other orders by the presence of scales on the external parts of the body and appendages, especially the wings. Butterflies and moths vary in size from microlepidoptera only a few millimetres long, to a wingspan of many inches such as the Atlas moth. Comprising over 160,000 described species, the Lepidoptera possess variations of the basic body structure which has evolved to gain advantages in adaptation and distribution.

<i>Macrosoma albida</i> Species of butterfly

Macrosoma albida is moth-like butterfly described by William Schaus in 1901. It belongs to the family Hedylidae. Originally it belonged to the genus Hyphedyle.

Macrosoma albifascia is moth-like butterfly described by William Warren in 1904. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes. A subspecies with the name M. albifascia albifascia was defined by Warren. Another subspecies M. albifascia expedita was proposed by Louis Beethoven Prout in 1932.

Macrosoma albimacula is moth-like butterfly described by William Warren in 1900. It belongs to the family Hedylidae. Originally it belonged to the genus Hyphedyle.

Macrosoma albipannosa is moth-like butterfly described by Louis Beethoven Prout in 1916. It belongs to the family Hedylidae. Originally it belonged to the genus Hedyle. Malcolm J. Scoble combined it with Macrosoma in 1986.

Macrosoma albistria is moth-like butterfly described by Louis Beethoven Prout in 1916. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes. Malcolm J. Scoble combined it with Macrosoma in 1986.

Macrosoma amaculata is moth-like butterfly described by Malcolm J. Scoble in 1990. It belongs to the family Hedylidae.

<i>Macrosoma bahiata</i> Species of butterfly

Macrosoma bahiata is a moth-like butterfly species described by Cajetan von Felder and Alois Friedrich Rogenhofer in 1875. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes. Malcolm J. Scoble combined it with Macrosoma in 1986.

<i>Macrosoma cascaria</i> Species of butterfly

Macrosoma cascaria is moth-like butterfly described by William Schaus in 1901. It belongs to the family Hedylidae. Originally it belonged to the genus Hyphedyle. Malcolm J. Scoble combined it with Macrosoma in 1986.

<i>Macrosoma conifera</i> Species of butterfly

Macrosoma conifera is moth-like butterfly described by Warren in 1897. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes. Malcolm J. Scoble combined it with Macrosoma in 1986.

Macrosoma coscoja is moth-like butterfly described by Paul Dognin in 1900. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes. Malcolm J. Scoble combined it with Macrosoma in 1986.

Macrosoma costilunata is moth-like butterfly described by Louis Beethoven Prout in 1916. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes.

Macrosoma hedylaria is moth-like butterfly described by William Warren in 1894. It belongs to the family Hedylidae. Originally it belonged to the genus Phellinodes. Malcolm J. Scoble combined it with Macrosoma in 1986.

<i>Macrosoma heliconiaria</i> Species of butterfly

Macrosoma heliconiaria is moth-like butterfly described by Achille Guenée in 1857. It belongs to the family Hedylidae. Originally it belonged to the genus Hedyle. Malcolm J. Scoble combined it with Macrosoma in 1986. The species displays characteristics of both moths and butterflies and is believed to be the closest living ancestor to modern butterflies. Adaptions to avoid bat predation have given the species ultrasonic hearing and night vision through superposition optics.

<i>Macrosoma hyacinthina</i> Species of butterfly

Macrosoma hyacinthina is moth-like butterfly described by William Warren in 1905. It belongs to the family Hedylidae. Originally it belonged to the genus Lasiopates. Malcolm J. Scoble combined it with Macrosoma in 1986.

<i>Macrosoma intermedia</i> Species of butterfly

Macrosoma intermedia is moth-like butterfly described by Paul Dognin in 1911. It belongs to the Hedylidae family. Originally, it was in the genus Phellinodes.