The Heteroneura are 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. The frenulum, comprising one or more bristles arising from the humeral angle of the hind wing, is held in place by the retinaculum, a membranous hook or series of stiff, raised scales on the underside of the fore wing (Nielsen & Common 1991). Other synapomorphies of the Heteroneura include a reduction in branching of the Rs vein in the hind wing and loss of sclerotisation of the first abdominal sternite (Grimaldi & Engel 2005). Basal members of the clade possess plesiomorphic female genitalia with a single terminal opening used in both copulation and oviposition. Male genitalia often possess spine combs (pectinifers) on the valva in Nepticuloidea and Adeloidea but these are absent in Andesiana, Tischeriidae, Palaephatidae and Ditrysia (Davis & Gentili 2003).
Major subgroups of the Heteroneura 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. Larvae, where known, are miners in leaves or cambium and may construct mines up to one metre long. They lack legs and are long and slender, superficially resembling nematodes. Adults are most active in the late afternoon (Nielsen & Common 1991).
Andesiana is a genus of 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. Antennae are strongly dimorphic, being broadly bipectinate in males but filiform in females. Fore wing vestiture is largely grey with a metallic sheen and with a reticulate pattern of reddish-brown in the species A. lamellata and A. similis, and with a oblique, subapical white band in A. similis. The smallest species, A. brunnea (fore wing length 12 mm) is primarily dark reddish-brown. The genus Andesiana was originally described in the family Cossidae, which they resemble in their relatively large size (wing span up to 61 mm) and tendency of bodies and wings to become greasy in dried specimens, before examination of their genitalia excluded them from the ditrysian clade. Their larval biology is unkown but they are possibly stem borers like Cossidae (Davis & Gentili 2003).
The Tischeriidae are very small moths with a distinctive frontal tuft projecting over a triangular face. The front end of the body is held elevated above the substrate at rest with all legs held concealed under the body. Larvae are blotch leaf miners, usually on the upper side of leaves and most commonly on Fagaceae and Asteraceae.
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). Monophyly of this group may be questioned by the molecular phylogenetic analysis of Kawahara et al. (2019) which placed Metaphatus and Ptyssoptera closer to Tischeriidae but Palaephatus as sister to Ditrysia.
<==Heteroneura [Nepticulina] | i. s.: Irenicodes [Diplosariidae]P27 | `--I. eurychoraP27 | Coenoloba [Tineodinidae]P27 | `--C. obliteralisP27 |--NepticuloideaKP19 | |--NepticulidaeNB03 | `--OpostegidaeWRM02 | | i. s.: ParalopostegaNB03 | `--OposteginaeKP19 | |--Pseudopostega quadristrigellaKP19 | `--OpostegaWRM02 | |--O. gephyraeaNC91 | |--O. nonstrigellaP27 | |--O. quadristrigellaP27 | `--O. saliciellaWRM02 `--+--+--Andesiana Gentili 1989DG03 [Andesianidae, AndesianoideaKP19] | | |--*A. lamellata Gentili 1989DG03 | | |--A. brunnea Gentili 1989DG03 | | `--A. similis Gentili 1989DG03 | `--AdeloideaKP19 `--+--+--Tischeriidae [Tischerioidea]KP19 | | |--Coptotriche citrinipennellaKP19 | | `--TischeriaWRM02 | | |--T. badiiellaWRM02 | | |--T. citrinipennellaWRM02 | | |--T. quercitellaKP19 | | `--T. subcanalisG17 | `--+--Metaphatus ochraceusKP19 | `--PtyssopteraKP19 `--+--Palaephatidae [Palaephatoidea]WRM02 | |--Azaleodes microniphaNC91 | `--PalaephatusKP19 | |--P. dimorphusDG03 | |--P. falsusWRM02 | |--P. luteolusKP19 | `--P. nielseniKP19 `--DitrysiaWRM02
*Type species of generic name indicated
[DG03] Davis, D. R., & P. Gentili. 2003. Andesianidae, a new family of monotrysian moths (Lepidoptera: Andesianoidea) from South America. Invertebrate Systematics 17: 15–26.
[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).
[KP19] Kawahara, A. Y., D. Plotkin, M. Espeland, K. Meusemann, E. F. A. Toussaint, A. Donath, F. Gimnich, P. B. Frandsen, A. Zwick, M. dos Reis, J. R. Barber, R. S. Peters, S. Liu, X. Zhou, C. Mayer, L. Podsiadlowski, C. Storer, J. E. Yack, B. Misof & J. W. Breinholt. 2019. Phylogenomics reveals the evolutionary timing and pattern of butterflies and moths. Proceedings of the National Academy of Sciences of the USA 116 (45): 22657–22663.
[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.
[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.