Gracillariidae

Gracillariidae is an important family of insects in the order Lepidoptera and the principal family of leaf miners that includes several economic, horticultural or recently invasive pest species such as the horse-chestnut leaf miner, Cameraria ohridella.

## Contents

* 1 Taxonomy and systematics
* 2 Distribution
* 3 Identification
* 4 Life history
* 5 Larval hostplants
* 6 Fossils
* 7 Subfamilies and genera
* 8 References
* 9 External links

## Taxonomy and systematics

There are 98 described genera of Gracillariidae (see below). A complete checklist is available of all currently recognised species.[1] There are many undescribed species in the tropics but there is also an online catalogue of Afrotropical described species;[2] the South African fauna is quite well known. Although Japanese and Russian authors have recognised additional subfamilies,[1] there are three currently recognised subfamilies, Phyllocnistinae of which is likely to be basal. In this subfamily, the primitive genus Prophyllocnistis from Chile feeds on the plant genus Drimys (Winteraceae), and has leaf mines structurally similar in structure to fossils (see "Fossils").[3] While there have been some recent DNA sequence-based studies of Palaearctic species[4][5] there is need for a satisfactory modern global phylogenetic framework for the subfamilies of Gracillariidae. Some genera are very large, e.g. Acrocercops, Caloptilia, Cameraria, Epicephala and Phyllonorycter.

## Distribution

Gracillariidae occur in all terrestrial regions of the world except Antarctica.

## Identification

These generally small (wingspan 5–20 mm) moths are leaf miners as caterpillars,[6] which can provide a useful means of identification, especially if the hostplant is known. The subfamilies differ by the adult moth resting posture (Davis and Robinson, 1999). Most Gracillariinae rest with the front of the body steeply raised; Lithocolletinae and Phyllocnistinae rest with the body parallel to the surface; in Lithocolletinae often with the head lowered.

Some additional features can be seen under close examination. The antennal flagellomeres of adults have two rows of scales: a basal row with large scales covering an apical row with smaller scales. The male genitalia has only four pair of muscles and lacks a gnathos. The female ovipositor is short and laterally flattened.[7]

## Life history

Most larval Gracillariidae undergo hypermetamorphosis, meaning there is a major change within the larval stage. Early instars have a flattened head and body, modified mandibles, and lack functional spinnerets or legs; these feed on sap.[7][8] Later instars are cylindrical with round heads, and have chewing mouthparts, legs and functional spinnerets; these feed on plant tissue.[7][8] Some species additionally have a spinning instar that does not feed, and others have a non-feeding instar before this spinning instar.[8]

The Brazilian Spinivalva gaucha is an exception to this trend, lacking a sap-feeding stage.[9]

Gracillariidae have various lifestyles. Some larvae mine leaves while others instead roll leaves, and there are also larvae which mine other plant parts or form galls.[8]

## Larval hostplants

Many host plants are known, generally dicotyledonous trees or shrubs.[10] Patterns of hostplant shifting have been inferred for many United Kingdom species in the genus Phyllonorycter and its sister genus Cameraria.[5] A recent DNA sequencing study mainly of Palaearctic species has shown that the burst of evolutionary adaptive radiation occurred long after that of the larval hostplants, rather than demonstrating a tight coevolutionary process.[4]

## Fossils

The family is an old one, with fossil Phyllocnistinae mines known from 97-million-year-old rocks in Kansas and Nebraska.[11] There are other fossil mines known from rocks of Eocene and Miocene age.[1] There are also two adult moths known from Lithuanian or Baltic amber of Eocene age: Gracillariites lithuanicus Kozlov, 1987 and G. mixtus Kozlov, 1987.[1]

## Subfamilies and genera

Phylogeny of Gracillariidae

| Gracillariidae  |  |  |  |  |  |  |  |  |  |  |

Lithocolletinae

|

Phyllocnistinae

|  |  |

Marmarinae

|

|  |  |

Oecophyllembiinae

|

Acrocercopinae

|  |  |

Gracillariinae

|

|  |  |

Parornichinae

|

Callicercopinae

Ornixolinae

from Li et al. 2022
Horse-chestnut leaf miner (Cameraria ohridella) tree damage in Parma, Italy

Gracillariidae phylogeny has been revised in 2017 and is now containing eight subfamilies:[12]

* Acrocercopinae Kawahara & Ohshima, 2016
* Callicercopinae Li, Ohshima & Kawahara, 2022
* Gracillariinae Stainton, 1854
* Lithocolletinae Stainton, 1854
* Marmarinae Kawahara & Ohshima, 2016
* Oecophyllembiinae Réal & Balachowsky, 1966
* Ornixolinae Kuznetzov & Baryshnikova, 2001
* Parornichinae Kawahara & Ohshima, 2016
* Phyllocnistinae Herrich-Schäffer, 1857

Unplaced species

* "Ornix" blandella Müller-Rutz, 1920, this species was described from Switzerland. Larvae were recorded feeding on Salix. The present taxonomic status is unknown.
* "Gracilaria" confectella Walker, 1864
* "Gracilaria" delicatulella Walker, 1864
* "Phyllonorycter" fennicella Hering, 1924, this species was described from Finland. The larval hostplant is probably a Salix species. The present taxonomic status is unknown, but is probably a junior subjective synonym of Lithocolletis viminetorum or Lithocolletis salictella.
* "Lithocolletis" graeseriella Sorhagen, 1900, see Phyllonorycter
* "Lithocolletis" italica Herrich-Schäffer, 1855, this species was described from Italy. The present taxonomic status is unknown.
* "Ornix" jyngipennella Heydenreich, 1851, nomen nudum.
* "Lithocolletis" lativitella Sorhagen, 1900, this species was described from Germany. Larvae were recorded feeding on Sorbus aria and Pyrus scandinavica. The present taxonomic status is unknown. It might be a synonym of Tinea lantanella Schrank, 1802.
* "Lithocolletis" norvegicella Strand, 1919, this species was described from Norway. The present taxonomic status is unknown.
* "Gracillaria" pistaciella Rondani, 1876, this species was described from Italy. Larvae were recorded feeding on Pistacia terebinthus.
* "Ornix" quercella Müller-Rutz, 1934, this species was described from Switzerland. Larvae were probably bred from a mine on a Quercus species. The present taxonomic status is unknown.
* "Phyllonorycter" sessilifoliella Hering, 1957, this species was recorded from southern France, where it was said to have been reared on a Quercus species. nomen nudum

## References

1. ↑ 1.0 1.1 1.2 1.3 Prins, Willy De; Prins, Jurate De (1998-01-01) (in English). World Catalogue of Insects. 6. ISBN 978-87-88757-64-4. https://www.wikidata.org/wiki/Q98651418. Retrieved 2021-12-16.
2. ↑ [1]
3. ↑ , Wikidata Q110143832
4. ↑ 4.0 4.1 , Wikidata Q42036997
5. ↑ 5.0 5.1 , Wikidata Q57381916
6. ↑ "Gracillariidae". http://www.leafmines.co.uk/html/gracillariidae.htm.
7. ↑ 7.0 7.1 7.2 Kawahara, Akito Y.; Plotkin, David; Ohshima, Issei; Lopez-Vaamonde, Carlos; Houlihan, Peter R.; Breinholt, Jesse W.; Kawakita, Atsushi; Xiao, Lei et al. (2017). "A molecular phylogeny and revised higher-level classification for the leaf-mining moth family G racillariidae and its implications for larval host-use evolution" (in en). Systematic Entomology 42 (1): 60–81. doi:10.1111/syen.12210. ISSN 0307-6970. Bibcode: 2017SysEn..42...60K. https://resjournals.onlinelibrary.wiley.com/doi/10.1111/syen.12210.
8. ↑ 8.0 8.1 8.2 8.3 Li, Xuankun; St Laurent, Ryan; Earl, Chandra; Doorenweerd, Camiel; van Nieukerken, Erik J.; Davis, Donald R.; Johns, Chris A.; Kawakita, Atsushi et al. (2022). "Phylogeny of gracillariid leaf-mining moths: evolution of larval behaviour inferred from phylogenomic and Sanger data" (in en). Cladistics 38 (3): 277–300. doi:10.1111/cla.12490. ISSN 0748-3007. PMID 34710244. https://onlinelibrary.wiley.com/doi/10.1111/cla.12490.
9. ↑ Brito, Rosangela; Gonçalves, Gislene; Vargas, Hector; Moreira, Gilson (2013-04-17). "A new Brazilian Passiflora leafminer: Spinivalva gaucha, gen. n., sp. n. (Lepidoptera, Gracillariidae, Gracillariinae), the first gracillariid without a sap-feeding instar" (in en). ZooKeys (291): 1–26. doi:10.3897/zookeys.291.4910. ISSN 1313-2970. PMID 23794860. PMC 3677287. Bibcode: 2013ZooK..291....1B. https://zookeys.pensoft.net/article/3920/.
10. ↑ "HOSTS - the Hostplants and Caterpillars Database at the Natural History Museum". http://internt.nhm.ac.uk/jdsml/research-curation/projects/hostplants/list.dsml?searchPageURL=browse.dsml&Family=Gracillariidae&Genus=&Species=&Country=&sort=Family.
11. ↑ , Wikidata Q24564423
12. ↑ Kawahara, A. Y.; Plotkin, D.; Ohshima, I.; Lopez-Vaamonde, C.; Houlihan, P. R.; Breinholt, J. W.; Davis, D. R.; Kumata, T. et al. (2017). "A molecular phylogeny and revised higher-level classification for the leaf-mining moth family Gracillariidae and its implications for larval host-use evolution". Systematic Entomology 42 (1): 60–81. doi:10.1111/syen.12210. Bibcode: 2017SysEn..42...60K.

* Davis, D.R, and Robinson, G.S. (1999). The Tineoidea and Gracillarioidea. In: Kristensen, N.P. (ed.), Lepidoptera, Moths and Butterflies, 1: Evolution, Systematics, and Biogeography. Handbuch der Zoologie 4 (35): 91-117. Walter de Gruyter. Berlin, New York.

## External links

| Wikimedia Commons has media related to Gracillariidae.

|

| Wikispecies has information related to Gracillariidae

|

* Tree of Life
* Natural History Museum hosts database
* Natural History Museum Lepidoptera genus database
* Afrotropical Checklist
* Global Taxonomic Database of Gracillariidae (Lepidoptera)
* Fauna Europaea Gracillariidae
* Fauna Europaea : Experts
* Caloptilia azaleella, azalea leafminer on the UF / IFAS Featured Creatures Web site
* Marmara gulosa, citrus peelminer on the UF / IFAS Featured Creatures Web site

* v
* t
* e

Extant Lepidoptera families

* Kingdom: Animalia
* Phylum: Arthropoda
* Class: Insecta
* Subclass: Pterygota
* Infraclass: Neoptera
* Superorder: Endopterygota

Suborder Zeugloptera

| Micropterigoidea|

Micropterigidae (mandibulate archaic moths)

|

Suborder Aglossata

| Agathiphagoidea|

Agathiphagidae (kauri moths)

|

Suborder Heterobathmiina

| Heterobathmioidea|

Heterobathmiidae

|

Suborder Glossata

| Dacnonypha|

| Eriocranioidea|

* Eriocraniidae

|

Acanthoctesia|

| Acanthopteroctetoidea|

* Acanthopteroctetidae (archaic sun moths)

|

Lophocoronina|

| Lophocoronoidea|

Lophocoronidae

|

Neopseustina|

| Neopseustoidea|

* Aenigmatineidae
* Neopseustidae (archaic bell moths)

|

Exoporia|

| Hepialoidea|

* Anomosetidae
* Hepialidae (swift moths, ghost moths)
* Neotheoridae (Amazonian primitive ghost moths)
* Palaeosetidae (miniature ghost moths)
* Prototheoridae (African primitive ghost moths)

|

Mnesarchaeoidea|

* Mnesarchaeidae (New Zealand primitive moths)

| M

| Adeloidea|

Adelidae (fairy longhorn moths)
Cecidosidae
Heliozelidae
Incurvariidae
Prodoxidae (yucca moths)

|

Andesianoidea|

* Andesianidae (Andean endemic moths)

Nepticuloidea|

Nepticulidae (pigmy, or midget moths)
Opostegidae (white eyecap moths)

Palaephatoidea|

Palaephatidae (Gondwanaland moths)

Tischerioidea|

Tischeriidae (trumpet leaf miner moths)

| Simaethistoidea|

Simaethistidae

|

Tineoidea|

Acrolophidae (burrowing webworm moths)
Arrhenophanidae
Eriocottidae (Old World spiny-winged moths)
Lypusidae
Psychidae (bagworm moths)
Tineidae (fungus moths)

Gracillarioidea|

Bucculatricidae (ribbed cocoon makers)
Douglasiidae (Douglas moths)
Gracillariidae
Roeslerstammiidae

Yponomeutoidea|

Acrolepiidae (false diamondback moths)
Bedelliidae
Glyphipterigidae (sedge moths)
Heliodinidae
Lyonetiidae
Plutellidae
Yponomeutidae (ermine moths)
Ypsolophidae

Gelechioidea|

Autostichidae
Batrachedridae
Blastobasidae
Coleophoridae (case-bearers, case moths)
Cosmopterigidae (cosmet moths)
Elachistidae (grass-miner moths)
Gelechiidae (twirler moths)
Lecithoceridae (long-horned moths)
Metachandidae
Momphidae (mompha moths)
Oecophoridae (concealer moths)
Pterolonchidae
Scythrididae (flower moths)
Xyloryctidae (timber moths)

Galacticoidea|

Galacticidae

Zygaenoidea|

Heterogynidae
Zygaenidae (burnet, forester, or smoky moths)
Himantopteridae
Lacturidae
Somabrachyidae
Megalopygidae (flannel moths)
Aididae
Anomoeotidae
Cyclotornidae
Epipyropidae (planthopper parasite moths)
Dalceridae (slug caterpillars)
Limacodidae (slug, or cup moths)

Cossoidea|

Cossidae (carpenter millers, or goat moths)
Dudgeoneidae (dudgeon carpenter moths)

Sesioidea|

Brachodidae (little bear moths)
giant butterfly-moths, sun moths)
Sesiidae (clearwing moths)

Choreutoidea|

Choreutidae (metalmark moths)

Tortricoidea|

Tortricidae (tortrix moths)

Urodoidea|

Urodidae (false burnet moths)

Schreckensteinioidea|

Schreckensteiniidae (bristle-legged moths)

Epermenioidea|

Epermeniidae (fringe-tufted moths)

Alucitoidea|

Alucitidae (many-plumed moths)
Tineodidae (false plume moths)

Pterophoroidea|

Pterophoridae (plume moths)

Whalleyanoidea|

Whalleyanidae

Immoidea|

Immidae

Copromorphoidea|

Copromorphidae (tropical fruitworm moths)
Carposinidae (fruitworm moths)

Thyridoidea|

Thyrididae (picture-winged leaf moths)

Calliduloidea|

Callidulidae (Old World butterfly-moths)

Papilionoidea
(butterflies)|

Papilionidae (swallowtail butterflies)
Hedylidae (American moth-butterflies)
Hesperiidae (skippers)
Pieridae (whites, yellows, orangetips, sulphurs)
Riodinidae (metalmarks)
blues, coppers and relatives)
Nymphalidae (brush-footed, or four-footed butterflies)

Hyblaeoidea|

Hyblaeidae (teak moths)

Pyraloidea|

Pyralidae (snout moths)
Crambidae (grass moth)

Mimallonoidea|

Mimallonidae (sack bearer moths)

Lasiocampoidea|

Lasiocampidae (eggars, snout moths, or lappet moths)

Bombycoidea|

Anthelidae (Australian lappet moth)
Apatelodidae (American silkworm moths)
Bombycidae (silk moths)
Brahmaeidae (Brahmin moths)
Carthaeidae (Dryandra moth)
Endromidae (Kentish glory and relatives)
Eupterotidae
Phiditiidae
Saturniidae (saturniids)
Sphingidae (hawk moths, sphinx moths and hornworms)

Noctuoidea|

Erebidae (underwing, tiger, tussock, litter, snout, owlet moths)
Euteliidae
Noctuidae (daggers, sallows, owlet moths, quakers, cutworms, darts)
Nolidae (tuft moths)
Notodontidae (prominents, kittens)
Oenosandridae

Drepanoidea|

Drepanidae (hook-tips)
Cimeliidae (gold moths)
Doidae

Geometroidea|

Sematuridae
Pseudobistonidae
Epicopeiidae (oriental swallowtail moths)
Uraniidae
Geometridae (geometer moths)

Superfamily unassigned|

Millieriidae

Note: division Monotrysia is not a clade.

* Taxonomy of the Lepidoptera
* Lists by region

Wikidata ☰ Q721100 entry

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