Gracillariidae is an important

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...

in the order

Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...

and the principal family of

leaf miner A leaf miner is any one of numerous species of insects in which the larval stage lives in, and eats, the leaf tissue of plants. The vast majority of leaf-mining insects are moths ( Lepidoptera), sawflies ( Symphyta, the mother clade of wasp ...

s that includes several economic, horticultural or recently invasive pest

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

such as the

horse-chestnut leaf miner The horse-chestnut leaf miner (''Cameraria ohridella'') 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.

, ''Cameraria ohridella''.

Taxonomy and systematics

There are 98 described

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

of Gracillariidae (see below). A complete checklist is available of all currently recognised species. There are many undescribed species in the tropics but there is also an online catalogue of Afrotropical described speci

the South African fauna is quite well known. Although Japanese and Russian authors have recognised additional subfamilies, there are three currently recognised subfamilies, Phyllocnistinae of which is likely to be basal. In this subfamily, the primitive genus ''Prophyllocnistis'' from

Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...

feeds on the plant genus '' Drimys'' (Winteraceae), and has leaf mines structurally similar in structure to fossils (see "Fossils"). While there have been some recent

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...

-based studies of

Palaearctic The Palearctic or Palaearctic is the largest of the eight biogeographic realms of the Earth. It stretches across all of Eurasia north of the foothills of the Himalayas, and North Africa. The realm consists of several bioregions: the Euro-Sibe ...

species there is need for a satisfactory modern global

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

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 mine A leaf miner is any one of numerous species of insects in which the larval stage lives in, and eats, the leaf tissue of plants. The vast majority of leaf-mining insects are moths ( Lepidoptera), sawflies ( Symphyta, the mother clade of wasp ...

rs as

caterpillars Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Symp ...

, 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.

Life history

The first to fifth-

instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow or ass ...

larvae are flattened and possess specialised mouthparts

adapted In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the po ...

for feeding on

sap Sap is a fluid transported in xylem cells (vessel elements or tracheids) or phloem sieve tube elements of a plant. These cells transport water and nutrients throughout the plant. Sap is distinct from latex, resin, or cell sap; it is a separa ...

. Older-instar

larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...

e are cylindrical and have normal chewing mouthparts for feeding on plant tissue within the

s, and have a fully functional

silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...

-producing organ, the "spinneret". Some genera have an intermediate stage in this remarkable hyper

metamorphosis Metamorphosis is a biological process by which an animal physically develops including birth or hatching, involving a conspicuous and relatively abrupt change in the animal's body structure through cell growth and differentiation. Some inse ...

(Davis and Robinson, 1999).

Larval hostplants

Many host plants are known, generally dicotyledonous trees or shrubs. Patterns of hostplant shifting have been inferred for many

United Kingdom The United Kingdom of Great Britain and Northern Ireland, commonly known as the United Kingdom (UK) or Britain, is a country in Europe, off the north-western coast of the continental mainland. It comprises England, Scotland, Wales and North ...

species in the genus ''Phyllonorycter'' and its sister genus ''Cameraria''. A recent

DNA sequencing DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...

study mainly of

species has shown that the burst of

evolutionary Evolution is change in the heredity, heritable Phenotypic trait, characteristics of biological populations over successive generations. These characteristics are the Gene expression, expressions of genes, which are passed on from parent to ...

adaptive radiation In evolutionary biology, adaptive radiation is a process in which organisms diversify rapidly from an ancestral species into a multitude of new forms, particularly when a change in the environment makes new resources available, alters biotic int ...

occurred long after that of the larval hostplants, rather than demonstrating a tight

coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well ...

ary process.

Fossils

The family is an old one, with fossil Phyllocnistinae mines known from 97-million-year-old rocks in

Kansas Kansas () is a state in the Midwestern United States. Its capital is Topeka, and its largest city is Wichita. Kansas is a landlocked state bordered by Nebraska to the north; Missouri to the east; Oklahoma to the south; and Colorado to the ...

and

Nebraska Nebraska () is a state in the Midwestern region of the United States. It is bordered by South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Kansas to the south; Colorado to the southwe ...

. There are other fossil mines known from rocks of

Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...

and

Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...

age. There are also two adult moths known from

Lithuania Lithuania (; lt, Lietuva ), officially the Republic of Lithuania ( lt, Lietuvos Respublika, links=no ), is a country in the Baltic region of Europe. It is one of three Baltic states and lies on the eastern shore of the Baltic Sea. Lithuania ...

n or

Baltic amber The Baltic region is home to the largest known deposit of amber, called Baltic amber or succinite. It was produced sometime during the Eocene epoch, but exactly when is controversial. It has been estimated that these forests created more than 1 ...

age: '' Gracillariites lithuanicus'' Kozlov, 1987 and '' G. mixtus'' Kozlov, 1987.

Subfamilies and genera

Gracillariidae phylogeny has been revised in 2017 and is now containing eight subfamilies: * Acrocercopinae Kawahara & Ohshima, 2016 *

Gracillariinae Gracillariinae are a subfamily of moths which was described by Henry Tibbats Stainton in 1854. The subfamilies of Gracillariidae differ by the adult moth resting posture.(Davis and Robinson, 1999) Most Gracillariinae rest with the front of the ...

Stainton, 1854 *

Lithocolletinae Lithocolletinae is a subfamily of insects in the moth family Gracillariidae. It is distributed worldwide, with most species in temperate regions.Davis, D. R. and J. De Prins. (2011)Systematics and biology of the new genus ''Macrosaccus'' with d ...

Stainton, 1854 *

Marmarinae Marmarinae is a subfamily of moth Moths are a paraphyletic group of insects that includes all members of the order Lepidoptera that are not butterflies, with moths making up the vast majority of the order. There are thought to be approximatel ...

Kawahara & Ohshima, 2016 * Oecophyllembiinae Réal & Balachowsky, 1966 * Ornixolinae Kuznetzov & Baryshnikova, 2001 * Parornichinae Kawahara & Ohshima, 2016 *

Phyllocnistinae Phyllocnistinae is a subfamily of insects in the moth family Gracillariidae.Müller-Rutz, 1920, this species was described from Switzerland. Larvae were recorded feeding on ''

Salix Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist so ...

''. 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 ''

'' 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 ''Phyllonorycter'' is a genus of moths in the family Gracillariidae. Diversity The genus comprises about 400 species, with a worldwide distribution. The vast majority of species are found in the temperate regions, with about 257 species describe ...

'' *''"Lithocolletis" italica'' Herrich-Schäffer, 1855, this species was described from Italy. The present taxonomic status is unknown. *''"Ornix" jyngipennella'' Heydenreich, 1851, ''

nomen nudum In taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published with an adequate descr ...

''. *''"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 ''Pistacia terebinthus'' also called the terebinth and the turpentine tree, is a deciduous tree species of the genus ''Pistacia'', native to the Mediterranean region from the western regions of Morocco and Portugal to Greece and western and s ...

''. *''"Ornix" quercella'' Müller-Rutz, 1934, this species was described from Switzerland. Larvae were probably bred from a mine on a ''

Quercus An oak is a tree or shrub in the genus ''Quercus'' (; Latin "oak tree") of the beech family, Fagaceae. There are approximately 500 extant species of oaks. The common name "oak" also appears in the names of species in related genera, notably '' ...

'' 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 ''

'' species. ''

References

*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

Tree of LifeNatural History Museum hosts databaseNatural History Museum Lepidoptera genus databaseAfrotropical ChecklistGlobal Taxonomic Database of Gracillariidae (Lepidoptera)Fauna Europaea GracillariidaeFauna Europaea : Experts

on the UF / IFAS Featured Creatures Web site
''Marmara gulosa'', citrus peelminer
on the UF / IFAS Featured Creatures Web site {{Authority control Moth families Leaf miners Taxa named by Henry Tibbats Stainton