Opostega salaciella, from here.

Belongs within: Glossata.
Contains: Nepticulidae, Incurvarioidea, Tineoidea, Yponomeutoidea, Gelechioidea, Gracillariidae, Phalaena, Tortricidae, Sesioidea, Cossoidea, Zygaenoidea, Alucitoidea, Pterophoridae, Copromorphoidea, Thyrididae, Pyraloidea, Bombycoidea, Noctuoidea, Rhopalocera, Drepanidae, Geometroidea.

The Heteroneura is a major clade of Lepidoptera, basally characterised by the coupling of the fore and hind wings (though this has been lost in some derived subgroups). Major subgroups include the leaf-mining Nepticuloidea, some species of which are among the smallest lepidopterans (Grimaldi & Engel 2005). Nepticuloids include the Opostegidae, which are usually white with a dark pattern and have their wings coupled by long, curved pseudofrenular bristles curving around a ventrally expanded M vein in the fore wing (Nielsen & Common 199). Andesiana is a genus of relatively large, heavy-bodied moths found in southern South America, characterised by an elongate second labial palpal segment and a pheromone-dispersing pouch on the male tibia associated with a femoral hair pencil (Davis & Gentili 2003). Tischeria includes very small, brown or blackish-grey moths with a distinctive frontal tuft projecting over a triangular face. The Palaephatidae are small to very small moths found in South America and Australia whose larvae start life mining in leaves of Proteaceae and later live in shelters formed by joining adjacent leaves (Nielsen & Common 1991).

The clade Ditrysia includes the majority of the Heteroneura and indeed some 98% of the species of Lepidoptera, united by the presence of an internal passage between the copulatory tract and the oviduct. Members include the Simaethistiidae, a small, poorly known group of basal ditrysians found in eastern Asia and Australia. Major clades within the Ditrysia include the Apoditrysia, characterised by the presence of two specialised apodemes with short, large bases on the abdominal sternum. Within the Apoditrysia, members of the Obtectomera have pupae with abdominal segments 1–4 immobile and without a dorsal row of spines on the tergites. The Epermeniidae are a group of very small moths with narrow, almost parallel-sided forewings bearing raised scales along the dorsum. The Choreutidae are a group of very small to small moths, often with metallic patterning, that walk in a characteristic jerky manner and often curl their wings at rest (Nielsen & Common 1991). The Urodidae, false burnet moths, are dull-coloured moths found in Europe and the Americas whose pupa is contained within an open-mesh cocoon that may hang suspended on a long thread beneath a leaf. The Galacticidae are medium-sized moths found in the Old World, characterised by relatively short antennae (much shorter than the fore wing) with the flagellum scaled on the dorsal surface only. Members of the superfamilies Alucitoidea and Pterophoroidea, plume moths, often have the fore and hind wings deeply cleft into multiple plumes. Pterophoroidea usually possess vein CuP in the fore wing whereas this vein is absent in Alucitoidea (Nielsen & Common 1991).

The Macrolepidoptera is a clade containing, as its name suggests, many (but not all) of the larger species of Lepidoptera. It is characterised by the complete loss of vein CuP and the larval crochets being in a crescentic series on the mesal side of the proleg. A number of macrolepidopterans also possess sound-receptive tympanal organs, thought differences in location and structure imply non-homology: Drepanidae and Geometroidea have tympanal organs on the abdomen, Noctuoidea have them on the metathorax, usually with counter-tympanal cavities on the base of the abdomen that may have a prominent hood, and Bombycoidea lack tympanal organs (Grimaldi & Engel 2005). Tympanal organs are also absent in the Calliduloidea, a small group of diurnal moths with simple antennae and wings held back to back at rest, resembling small butterflies (Common 1970).

Synapomorphies (from Grimaldi & Engel 2005): Female genitalia with gonopore separate from ovipore, no connection between the two tracts; reduction in branching of Rs veins in hind wing; wings coupled with frenulum and retinaculum; loss of abdominal sternum 1.

<==Heteroneura [Nepticulina]
    |  i. s.: Irenicodes [Diplosariidae] P27
    |           `--I. eurychora P27
    |         Coenoloba [Tineodinidae] P27
    |           `--C. obliteralis P27
    |--Andesiana Gentili 1989 [Andesianidae, Andesianoidea] DG03
    |    |--*A. lamellata Gentili 1989 DG03
    |    |--A. brunnea Gentili 1989 DG03
    |    `--A. similis Gentili 1989 DG03
    |--Nepticuloidea WRM02
    |    |--Nepticulidae NB03
    |    `--Opostegidae NB03
    |         |--Paralopostega NB03
    |         `--Opostega WRM02
    |              |--O. gephyraea NC91
    |              |--O. nonstrigella P27
    |              |--O. quadristrigella P27
    |              `--O. saliciella WRM02
    `--+--Incurvarioidea WRM02
       `--+--+--Tischeria [Tischeriidae, Tischerioidea] WRM02
          |  |    |--T. badiiella WRM02
          |  |    |--T. citrinipennella WRM02
          |  |    `--T. subcanalis G17
          |  `--Palaephatidae [Palaephatoidea] WRM02
          |       |--Ptyssoptera AY04
          |       |--Azaleodes micronipha NC91
          |       `--Palaephatus WRM02
          |            |--P. dimorphus DG03
          |            `--P. falsus WRM02
          `--Ditrysia [Papilionina, Tineina, Tineites, Tortrices] WRM02
               |  i. s.: Azinis hilarella WM66
               |         Apoprogonidae NC91
               |         Epicopeia DS73 [Epicopeiidae NC91]
               |           |--E. caroli DS73
               |           `--E. polydora DS73
               |         Cerostoma Latreille 1802 L02
               |           |--*C. dorsatum [=Ypsolophus dorsatus] L02
               |           `--C. persicella F92
               |--Simaethistiidae GE05
               |--Tineoidea GE05
               `--+--Yponomeutoidea GE05
                  |--Gelechioidea GE05
                  |--Gracillariidae GE05
                  `--Apoditrysia [Phalaenites, Pyralites] GE05
                       |  i. s.: Oligostigma [Hypocampidae] WM66
                       |           `--O. orbitalis WM66
                       |         Spilomelidae WM66
                       |           |--Lepyrodes geometralis WM66
                       |           `--Zebronia WM66
                       |                |--Z. abdicalis WM66
                       |                `--Z. perspicualis WM66
                       |         Phalaena L02
                       |--Urodidae [Urodoidea] GE05
                       |--Schrecksteiniidae [Schreckensteinoidea] GE05
                       |--Tortricidae GE05
                       |--Homadaula NC91 [Galacticidae, Galacticoidea GE05]
                       |    `--H. myriospila NC91
                       |--+--Sesioidea GE05
                       |  |--Cossoidea GE05
                       |  `--Zygaenoidea GE05
                       |--+--Alucitoidea GE05
                       |  `--Pterophoroidea [Pterophorii] GE05
                       |       |--Pterophoridae GE05
                       |       `--Orneodes Latreille 1802 L02 [Orneodidae P27]
                       |            |--*O. hexadactylus [=Pterophorus hexadactylus] L02
                       |            |--O. objurgatella B88
                       |            `--O. phricodes P27
                       |--Epermeniidae [Epermenioidea] GE05
                       |    |--Gnathifera eurybias NC91
                       |    |--Paraepermenia santaliella NC91
                       |    |--Epermenia [Epermeniinae] NC91
                       |    |    |--E. eurybias C70
                       |    |    `--E. exilis NC91
                       |    `--Ochromolopis [Ochromolopinae] NC91
                       |         |--O. cornutifera NC91
                       |         `--O. eurybias C74
                       |--Choreutidae GE05
                       |    |--Asterivora lampadias NC91
                       |    |--Saptha exanthista NC91
                       |    |--Tebenna NC91
                       |    |    |--T. bradleyi M83
                       |    |    `--T. micalis NC91
                       |    `--Choreutis NC91
                       |         |--C. basalis NC91
                       |         |--C. bjerkandrella P27
                       |         `--C. lampadias C70
                       `--Obtectomera [Pyralidina] GE05
                            |  i. s.: Hyblaea IR05 [Hyblaeidae, Hyblaeoidea NC91]
                            |           |--H. ibidias NC91
                            |           `--H. pleura IR05
                            |--Copromorphoidea GE05
                            |--Whalleyaniidae GE05
                            |--Thyrididae GE05
                            |--Pyraloidea GE05
                            |--Immiidae [Immoidea] GE05
                            |    |--Birthana cleis NC91
                            |    `--Imma NC91
                            |         |--I. acosma NC91
                            |         |--I. lyrifera NC91
                            |         |--I. stilbiota NC91
                            |         `--I. vaticina NC91
                            `--Macrolepidoptera [Bombycina, Bombycites] GE05
                                 |  i. s.: Cypra delicatula [incl. Cozistra membranacea] M86
                                 |         Artaxa M86
                                 |           |--A. lutea M86
                                 |           `--A. paraneura Meyrick 1886 M86
                                 |--Bombycoidea GE05
                                 |--Noctuoidea GE05
                                 `--+--Rhopalocera GE05
                                    |--+--Drepanidae GE05
                                    |  `--Geometroidea GE05
                                    `--Calliduloidea DS73
                                         |--Pterothysanidae DS73
                                         `--Cleis C70 [Calliduliidae GE05]
                                              `--C. scotti C70

*Type species of generic name indicated


[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales & G. S. Taylor. 2004. Insects ‘Down Under’—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216–234.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[C70] Common, I. F. B. 1970. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 765–866. Melbourne University Press.

[C74] Common, I. F. B. 1974. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 98–107. Melbourne University Press.

[DG03] Davis, D. R., & P. Gentili. 2003. Andesianidae, a new family of monotrysian moths (Lepidoptera: Andesianoidea) from South America. Invertebrate Systematics 17: 15–26.

[DS73] Dickens, M., & E. Storey. 1973. The World of Moths. Macmillan Publishing Co., Inc.: New York.

[F92] Fan Z. 1992. Key to the Common Flies of China 2nd ed. Science Press: Beijing.

[G17] Girault, A. A. 1917. Descriptiones stellarum novarum. Privately published (reprinted Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 80–101).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[IR05] Idris, A. B., & A. Rizki. 2005. Notes on the tribe Ephialtini (Hymenoptera: Ichneumonidae: Pimplinae) of Malaysia. Serangga 10 (1–2): 111–126.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[M86] Meyrick, E. 1886. On some Lepidoptera from the Fly River. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 241–258.

[NC91] Nielsen, E. S., & I. F. B. Common. 1991. Lepidoptera (moths and butterflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 817–915. Melbourne University Press: Carlton (Victoria).

[NB03] Nieukerken, E. J. van, & C. van der Berg. 2003. A new Stigmella feeding on Urticaceae from Guam: first records of Nepticulidae (Lepidoptera) from Micronesia and Polynesia. Invertebrate Systematics 17: 27–37.

[P27] Philpott, A. 1927. The maxillae in the Lepidoptera. Transactions and Proceedings of the New Zealand Institute 57: 721–746.

[WM66] Wallace, A. R., & F. Moore. 1866. List of lepidopterous insects collected at Takow, Formosa, by Mr. R. Swinhoe. Proceedings of the Zoological Society of London 1866: 355–365.

[WRM02] Wiegmann, B. M., J. C. Regier & C. Mitter. 2002. Combined molecular and morphological evidence on the phylogeny of the earliest lepidopteran lineages. Zoologica Scripta 31 (1): 67–81.

Last updated: 22 March 2020.

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