Diversity description:

Approximately 1900 species grouped into 104 genera are known from all major regions of the world except Antarctica (DePrins, J and W. accessed 1 April 2009)

Geographic Range:

Nearctic, Palearctic, Oriental, Ethiopian, Neotropical, Australian, Oceanic Island

Texture:

smooth

Orientation:

flat

Egg mass pattern:

Eggs usually deposited singly on epidermis of host plant; rarely laid in a short transverse row of 4-8 contiguous eggs.

Description of egg morphology:

Usually oval to elliptical, partially flattened underneath and slightly convex above with nearly smooth to finely sculptured chorion, sometimes with upper and lower halves different in texture; less than 0.5 mm in length. Micropyle area finely reticulated, often reduced in size.

Secondary setae:

absent

Body setae on verrucae:

absent

Body setae on chalazae:

absent

Body setae on scoli:

absent

Larval body description:

Body usually not exceeding 10 mm in length, variable in color depending upon species and instar, from white to green, with some species becoming bright red prior to pupation, with or without darkly pigmented head and sclerotized plates.

Spinneret:

present

Thoracic glands:

absent

Thoracic legs:

present, absent

Larval Prothoracic L-group setae:

bisetose

Abdominal glands:

absent

Abdominal prolegs:

present, absent

Proleg configuration:

normal, odd

Proleg size:

short

Crochets:

uniserial, multiserial, arranged in circle

Anal comb on A10:

absent

Pupa type:

adecticous

Pupa description:

Head usually with a serrated or acute frontal process (cocoon cutter) from vertex, or smoothly rounded. Antennae extending to abdominal segment 6 (A6) to well beyond A10. Wings to A5-6. Tergal spines variably developed, usually small to minute in multiple, irregular rows or dense concentrations on A2 or 3-7 or 8; Phyllocnistis often with a few pairs of large tergal hooks on A2-7. A4-7 moveable in male, A4-6 in female (Mosher 1916). Cremaster highly variable, usually present as small paired spines, often with recurved apices. An accessory cremaster sometimes variably developed on sternum 7.

Pupal tergal spines:

present

Spines as modified cremaster:

present, absent

Cremaster:

present

Cocoon:

present

Reproductive system:

Ditrysian

Oviscapt (ovipositor):

non-piercing

Female genitalia description:

Oviscapt short, with usually 2 pairs of moderately short, rarely elongate apophyses; anterior apophyses sometimes reduced or absent. Ostium on S8 or at caudal margin; sterigma sometimes well sclerotized and variously modified. Ductus bursae usually elongate, slender, either membranous or partially sclerotized, expanding anteriorly into membranous corpus bursae; signa usually of 1-2 pairs, or sometimes absent.

Female corethrogyne:

absent

Female pregenital sexual scales:

absent

Female accessory glands:

one pair

Female oviduct opening:

below anus

Female bursa ostium opening:

on venter 8

Female anterior apophyses originating:

originating from T8

Male coremata:

present

Male pregenital sexual scales:

present, absent

Male genitalia description:

Uncus absent. Tegumen a relatively elongate, mostly membranous dorsal hood. Vinculum U- to Y - shaped, with often a short, rarely elongate saccus. Subscaphium frequently distinct, variably sclerotized. Gnathos absent. Transtilla present or absent. Valvae usually elongate and simple, rarely asymmetrical, sometimes lobed, spined, or with 1-2 pairs of pectinifers. Anellus usually membranous. Juxta absent. Aedoeagus usually elongate and slender; cornuti present or absent. Male genitalia with 4 pairs of muscles, M 1 absent (Kuznetsov and Stekol nikov, 1987).

Sternum 5:

without fenestra

Sternum 5 gland:

absent

Adult abdomen description:

Sternum 2 long and narrow, with long slender sternal apodemes continuing caudad as sternal rods ca. 0.8 the length of sternite. Segment 8 and sometimes 7 of male frequently with paired coremata; Sternum 8 of female often enlarged, particularly in Lithocolletinae; Tergum 8 sometimes with median sclerotization; corethrogyne absent.

Male has:

phallotheca and aedeagus (phallus)

Scale tufts:

absent

Epiphysis:

present

Adult thorax description:

Metafurca with anteromedial process elongate, relatively slender; furcal apophyses free, short to moderately long and downcurved, arising well caudad near secondary arms.

Forelegs:

normal

Leg description:

Legs with tibial spur pattern usually 0-2-4; epiphysis rarely absent.

Forewing length from base of forewing to the apex (mm):

from 2 to 10

Wing venation:

heteroneurous

Forewing anal vein notation:

A

Forewing basal loop:

absent

Forewing pterostigma:

absent

Forewing chorda:

present, absent

Forewing upper surface with microtrichia:

absent

Hindwing anal vein notation:

1A + 2A

Hindwing cell vein:

unforked

Hindwing pterostigma:

absent

Wing coupling:

with frenulum

Wing scales:

hollow

Forewing description:

Forewings slender; W/L ratio 0.9-0.25; venation variable with R/Rs 3- to 5-branched; Rs4 usually to costa, rarely to apex; Ml- to 3-branched; Cu 1 sometimes absent; discal cell elongate, extending 0.7-0.8 the length of forewing; accessory cell vestigial to absent; base of M vestigial to absent; intercalary cell absent; 1 A + 2 A usually present, sometimes vestigial, occasionally with minute basal fork; male retinaculum usually formed of curved scales from underneath slightly swollen base of Sc;

Hindwing description:

Hindwing lanceolate; index 0.1-0.2; female frenulum usually with 2 frenular setae, rarely 1; stout, composite pseudofrenular setae sometimes arising near apex of Sc in both sexes (Davis 1991); venation reduced, from 7 to 4 veins; discal cell often open.

Counter-tympanum:

absent

Abdomen tympanum:

absent

Thorax tympanum:

absent

Palp tympanum:

absent

Ocelli:

absent

Eyes:

smooth

Labial palpus:

porrect, upcurved

Labial palpus modification:

Labial palpi 3-segmented, usually upturned, to straight and drooping; segment 2 sometimes with ventral scale tuft.

Maxillary palpus:

present, minute

Proboscis:

present

Proboscis texture:

naked

Proboscis description:

Haustellum usually elongate, 1-2X length of labial palpi.

Mandibles:

absent

Head vertex scaling:

very rough, very smooth

Female antennae:

filiform

Male antennae:

filiform

Antennal sensillum:

Antennal sensillum present

Sensillum vesiculocladum:

absent

Asciod sensilla:

absent

General antennae description:

Antennae 0.8-1.75 the length of forewing; scape usually smooth, with or without pecten; flagellum filiform, with a single row of slender scales completely encircling each segment.

Adult head description:

Head vestiture variable, usually smooth (in Gracillariinae) with moderately broad scales from vertex directed forward and down over frons; similar in Lithocolletinae except with tufts of erect filiform and sometimes broader scales arising from occiput caudad to vertex. Eyes moderately large; interocular index 0.9-1.0.

Apomorphies:

Hypermetamorphic larval development (Kumata 1978)..  Early instar larvae specialized for sapfeeding in plant tissue; later instar(s) usually generalized tissue feeders, or sometimes nonfeeding silk spinning larvae in final instar..  Tissue feeding larval instars usually with prolegs absent on abdominal segment 6.

Gracilariidae (Gracilariid sp.) preys on:
Atriplex canescens
Atriplex polycarpa

Based on studies in:
USA: California, Southern California (Galls, Plant substrate)

This list may not be complete but is based on published studies.

Pupa life history description:

Pupation usually in a small, whitish silken cocoon either outside the mine of the host plant (most Gracillarinae) or within the mine (most Lithocolletinae and all Phyllocnistinae).

Larval food habits description:

Most species of Gracillariidae are leafminers, others mine the subepidermal layers of new growth stems and fruits or bore inside stems and galls (Davis 1991). Early sap-feeding instars with slashing mandibles typically feed in a horizontal plane, initiating slender, subepidermal serpentine mines while ingesting relatively little solid tissue. In some genera (e. g. Phyllonorycter) the sap-feeding instars instead initiate a blotch mine by devouring mostly spongy parenchymal cells (Watson 1956). Tissue-feeding instars possess typical chewing mandibles that enable these larvae to feed deeper into the host, usually ingesting the remainder of the spongy parenchyma and most of the palisade cells within the blotch. Later tissue-feeding instars of the most primitive genera (Gracillariinae) tend to exit the mine and feed externally, often in a rolled-over leaf.

Life history larvae:

Larval Gracillariidae undergo hypermetamorphic development (Kumata 1978; Davis 1987) with at least 2 distinct forms and habits: (1) an early, usually sap-feeding (Tragardh 1913) or flattened (Chambers 1877) form, often comprising the first 2-5 instars, with a depressed, apodal body with specialized, prognathous mouthparts and rudimentary spinneret, and (2) a later hypognathous, tissue-feeding or cylindrical form, possessing a more generalized, eruciform body with unspecialized, chewing mouthparts and legs; or a variously modified nonfeeding spinning form, with either a depressed or cylindrical body, mandibles and legs reduced or absent, but with a functional spinneret. An additional nonfeeding, transitional stage is also interposed between the final sap-feeding and single spinning instars of Chrysaster, Dendrorycter, and Marmara (De Gryse 1916; Kumata 1978). Moreover, in the latter two genera the quiescent instar exists in a pharate condition within the cuticle of the last sap-feeding instar. Later instar larvae of Gracillariidae are further characterized by 2 prespiracular (L) setae on T1 and crochets, if present, on A3-5 and 10 and almost always absent on A6 (present on A3-6 in Artifodina, Prophyllocnistis, and on A2-6 in Metriochroa psychotriella, Davis, 1994). Number of in stars varies from 4-11, depending upon genus and sometimes species (Fitzgerald and Simeone 1971).

Systematic and taxonomic history:

The Gracillariidae constitute the principal family of plant mining Lepidoptera. Fossil leafmines of Gracillariidae (Phyllocnistinae) are known from the early Cenomanian of the Dakota Formation in Kansas and Nebraska (ca. 97 m. y. a.; Davis 1994; Labandeira et al. 1994) near the beginning of the Angiosperm radiation. This currently represents the earliest fossil record of any ditrysian moth assignable to family. Three or four subfamilies are recognized:    Gracillariinae: Adults of most genera rest with anterior part of body raised at steep angle. Head usually smooth. Hindwings with base of Rs arched toward costa and approaching apex of Sc (Ely 1918; Kumata 1978). Abdomen of ♂ usually with S8 membranous; coremata usually present. Last instar larva with fully developed mandibles and legs, 5 coxal setae, and A9 with D group bisetose. Pupation usually outside of mine. Representative genera: Acrocercops, Aristaea, Caloptilia, Epicephala, Macrostola, Parectopa, Parornix, Philodoria, Stomphastis.    Lithocolletinae: Adults rest with body parallel to surface or with head end lowered. Head usually with occipital tufts of piliform scales. Hindwings with Rs nearly parallel to costal margin. Abdomen of ♂ with S8 sclerotized, elongate; coremata absent. Last instar larva usually with fully developed mandibles and legs (both reduced in Cameraria), 4 coxal setae, and A9 with D unisetose. Pupation normally within blotch mine. Representative genera: Cameraria, Chrysaster, Cremastobombycia, Leucanthiza, Phyllonorycter (= Lithocolletis).   Phyllocnistinae: Adults rest with body parallel to surface. Head smooth. Hindwings with Rs closely parallel to costal margin. Abdomen of ♂ with S8 membranous; coremata usually present. All feeding instars sap-feeding, creating elongate serpentine, subepidermal mines; last instar larva non-feeding with mandibles absent, legs reduced to unsegmented stubs, A9 with D bisetose. Pupation within slightly enlarged cavity at termination of mine. Representative genera: Cryphiomystis, Metriochroa, Phyllocnistis, Prophyllocnistis).   A fourth subfamily, OECOPHYLLYMBIINAE, has been redefined reently by Kumata (1998) to include all genera listed above within Phyllocnistinae except Phyllocnistis.

Fossil record:

Fossil records from J. and W. DePrins 2006:  Gracillariites lithuanicus Kozlov, 1987. Fossil in Eocene Amber (Lithuania).  Gracillariites mixtus Kozlov, 1987; Fossil in Eocene amber. (Baltic States).   Caloptilia sp. probably on a Quercus leaf (Miocene), U.S.A., Idaho, Latah Formation (Lewis 1969: 1210-1211, Opler 1973: 1322).  Cameraria sp. on Lithocarpus and Quercus simulata Knowlt. (middle to upper Miocene), U.S.A., Idaho, Thorn Creek (Opler 1973: 1321).  Phyllonorycter sp. on an unidentified fossil leaf (upper Eocene), Canada, British Columbia, White Lake Basin (Freeman 1965: 1069).  A leaf mine resembling Phyllocnistis has been reported on fossil Cedrela leaves from the early Eocene in the Sheridan Pass area southwest of Dubois, Wyoming, U.S.A. (Hickey and Hodges 1975: 718).  phyllocnistine leaf mines on a magnoliid dicot from the Dakota Formation of Kansas and Nebraska (U.S.A.) that date to the Early Cenomian (97 m.y.a.) (Labaneira et al, 1994, Davis 1994: 65).  Phyllonorycter sp. on Quercus hanibalii Dorf (late middle Miocene), U.S.A., Nevada, Cedar Mountains, Upper Goldyke (Opler 1973: 1321).  Phyllonorycter sp. on Populus trichocarpa var. ingrata (Jeps.) Parish (upper Miocene), U.S.A., Nevada, Stewart Valley (Opler 1973: 1321).

Collection Sites: world map showing specimen collection locations for Gracillariidae

Barcode of Life Data Systems (BOLD) Stats
                                                             

Specimen Records:	6,522
Specimens with Sequences:	4,456
Specimens with Barcodes:	3,947
Public Records:	915
Species:	750
Species With Barcodes:	556

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