Gracillariidae | |
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Privet leaf miner, Gracillaria syringella (Fabricius, 1794) | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Lepidoptera |
Superfamily: | Gracillarioidea |
Family: | Gracillariidae Stainton, 1854 |
Subfamilies | |
Genera - see "Subfamilies and genera" | |
Diversity [1] | |
About 101 genera and 1,866 species |
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.
There are 98 described genera of Gracillariidae (see below). A complete checklist is available of all currently recognised species. [2] There are many undescribed species in the tropics but there is also an online catalogue of Afrotropical described species [3] ; the South African fauna is quite well known. Although Japanese and Russian authors have recognised additional subfamilies, [2] 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"). [4] While there have been some recent DNA sequence-based studies of Palaearctic species [5] [6] 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.
Gracillariidae occur in all terrestrial regions of the world except Antarctica.
These generally small (wingspan 5–20 mm) moths are leaf miners as caterpillars, [7] 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. [8]
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. [8] [9] Later instars are cylindrical with round heads, and have chewing mouthparts, legs and functional spinnerets; these feed on plant tissue. [8] [9] Some species additionally have a spinning instar that does not feed, and others have a non-feeding instar before this spinning instar. [9]
The Brazilian Spinivalva gaucha is an exception to this trend, lacking a sap-feeding stage. [10]
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. [9]
Many host plants are known, generally dicotyledonous trees or shrubs. [11] Patterns of hostplant shifting have been inferred for many United Kingdom species in the genus Phyllonorycter and its sister genus Cameraria. [6] 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. [5]
The family is an old one, with fossil Phyllocnistinae mines known from 97-million-year-old rocks in Kansas and Nebraska. [12] There are other fossil mines known from rocks of Eocene and Miocene age. [2] There are also two adult moths known from Lithuanian or Baltic amber of Eocene age: Gracillariites lithuanicus Kozlov, 1987 and G. mixtus Kozlov, 1987. [2]
Phylogeny of Gracillariidae |
from Li et al. 2022 |
Gracillariidae phylogeny has been revised in 2017 and is now containing eight subfamilies: [13]
Unplaced species
Hypermetamorphosis, or heteromorphosis, is a term used mainly in entomology; it refers to a class of variants of holometabolism, that is to say, complete insect metamorphosis. Hypermetamorphosis is exceptional in that some instars, usually larval instars, are functionally and visibly distinct from the rest. The differences between such instars usually reflect transient stages in the life cycle; for instance, one instar might be mobile while it searches for its food supply, while the following instar immediately sheds its locomotory organs and settles down to feed until it is fully grown and ready to change into the reproductive stage, which in turn, does not have the same nutritional requirements as the larvae.
The horse-chestnut leaf miner is a leaf-mining moth of the family Gracillariidae. The horse-chestnut leaf miner was first observed in North Macedonia in 1984, and was described as a new species in 1986. Its larvae are leaf miners on the common horse-chestnut. The horse-chestnut leafminer was first collected and inadvertently pressed in herbarium sheets by the botanist Theodor von Heldreich in central Greece in 1879.
Philodoria molokaiensis is a moth of the family Gracillariidae. It was first described by Otto Herman Swezey in 1928. It is endemic to the Hawaiian island of Molokai.
Dextellia is a genus of moths in the family Gracillariidae.
Phyllonorycter loxozona is a moth of the family Gracillariidae. It is known from South Africa and Uganda. The record for Kenya is a misidentification of Cameraria torridella.
Borboryctis euryae is a moth of the family Gracillariidae. It is known from Japan.
Neolithocolletis pentadesma is a moth of the family Gracillariidae. It is known from Indonesia (Java), Malaysia, the Philippines (Luzon) and the Seychelles.
Caloptilia cecidophora is a moth of the family Gracillariidae. It is known from Japan and Taiwan.
Phyllonorycter lucidicostella, lesser maple leaf blotch miner, is a moth of the family Gracillariidae. It is known from Ontario and Québec in Canada and Alabama, Connecticut, Illinois, Kentucky, Pennsylvania, Maine, Michigan, New York, Vermont and North Carolina in the United States.
Cameraria aesculisella is a moth of the family Gracillariidae. It is known from the United States.
Cameraria macrocarpae is a moth of the family Gracillariidae. It is known only from Manitoba, Canada.
Porphyrosela hardenbergiella is a moth of the family Gracillariidae. First identified in a suburban garden in Epsom, Auckland, New Zealand in 1955 feeding on Hardenbergia, the species was likely introduced from Australia, although no members of the species have been found there.
Lithocolletinae is a subfamily of insects in the moth family Gracillariidae. It is distributed worldwide, with most species in temperate regions.
Gracillariinae are a subfamily of moths which was described by Henry Tibbats Stainton in 1854.
Phyllocnistinae is a subfamily of insects in the moth family Gracillariidae.
Acrocercopinae is a subfamily of moths described by Akito Yuji Kawahara and Issei Ohshima in 2016.
Marmarinae is a subfamily of moths described by Akito Yuji Kawahara and Issei Ohshima in 2016.
Parornichinae is a subfamily of moths described by Vladimir Ivanovitsch Kuznetzov and Svetlana Vladimirovna Baryshnikova in 2001.
Oecophyllembiinae is a subfamily of moths described by Pierre Réal and Alfred Serge Balachowsky in 1966.
Callicercopinae are a subfamily of moths which was described by Li, Ohshima & Kawahara in 2022.