Diversity description:

The Incurvariidae is a small, cosmopolitan family of approximately 100 described species and 11 genera. It is best represented in the Australian (endemic genus Perthida and several other undescribed genera and species, Nielsen and Common 1991) and Palearctic (Alloclemensia. Excurvaria, Incurvaria, Paraclemensia, Procacitas, and Subclemensia, Kozlov 1987, Kuprijanov 1992, 1993, Nielsen 1981) regions. It is poorly represented in Africa (Protaephagus capensis Scoble 1980), South America (Basileura, Simacauda Nielsen and Davis 1981, Parra and Ibarra 1994a), and North America (endemic Vespina quercivora*, Davis 1972) and one species each of Alloclemensia and Paraclemensia (Davis 1983, Nielsen 1981, 1982).

Geographic Range:

Nearctic, Palearctic, Oriental, Ethiopian, Neotropical, Australian

Orientation:

flat

Number of stemmata:

from 6

Larval head description:

Head mostly prognathous with 6 pairs of stemmata; AF2 absent; adfrontal  sclerite elongate, extending to deeply incised epicranial notch.

Secondary setae:

absent

Body setae on verrucae:

absent

Body setae on chalazae:

absent

Body setae on scoli:

absent

Spinneret:

present

Larval thorax description:

Thorax with prespiracular sclerite bearing spiracle usually separate from pronotum. Sterna1 plates reduced, coxae separate. Legs well developed; tarsi sometimes with an enlarged, squamiform seta adjacent to claw (Davis 1987).

Thoracic glands:

absent

Thoracic legs:

present

Pairs of thoracic legs:

from 3

Larval abdomen description:

Prolegs greatly reduced.

Abdominal glands:

absent

Abdominal prolegs:

present

Pairs of abdominal legs:

from 4

Proleg size:

vestigial

Pairs of crochets per proleg:

from 1

Crochets:

multiserial, in transverse rows

Crochet arrangement description:

Crochets present on A3-6 and arranged in a uniserial, transverse band; A10 usually without crochets but present in bi- to triserial rows in Excurvaria, Paraclemensia, and Vespina.

Anal comb on A10:

absent

Pupa type:

adecticous, exarate

Pupa description:

Length of pupa 4-10 mm. Vertex smooth. Haustellum short, usually extending to end of labial palpi. Wings extending to A5-7 movable in male, A2-6 in female(Common 1990). A2-8 with scattered tergal spines (Jensen 1932), reduced to 1-2 rows in some genera (Ross 1958). Pupation occurs within the larval case, either attached to the host or on the ground in leaf litter. All species are believed univoltine.

Pupal tergal spines:

present

Spines as modified cremaster:

absent

Cremaster:

present

Number of rows of tergal spines:

from 1-2 to scattered

Cocoon:

present

Cocoon description:

Pupation within larval case.

Reproductive system:

Monotrysian

Oviscapt (ovipositor):

piercing

Female genitalia description:

Female genitalia: Ovipositor with a depressed, acute apex usually bearing minute, lateral serrations. Spermatheca without lateral lagena. Ductus bursae often with internal sclerotizations; signa absent.

Female corethrogyne:

absent

Female pregenital sexual scales:

absent

Female accessory glands:

one pair

Female oviduct opening:

in cloaca

Female anterior apophyses originating:

originating from T8

Male coremata:

absent

Male pregenital sexual scales:

absent

Male genitalia description:

Uncus usually indistinct and shallowly bilobed, consisting of an elongate rodlike, caudal appendage in Alloclemensia. Vinculum well developed, 1-2x the length of valvae, U-to V-shaped. Valvae without a distinct pectinifer but with flattened spines often arranged in 1-3 groups. Juxta with 1-2 pairs of usually elongate caudal arms partially enclosing aedoeagus. Aedoeagus tubular, usually with 1 or more large cornuti.

Sternum 5:

without fenestra

Sternum 5 gland:

absent

Adult abdomen description:

Abdomen with caudal rim of sternum 2a broadly U-shaped; caudal margin of female sternum 7 truncate.

Male has:

phallotheca and aedeagus (phallus)

Scale tufts:

absent

Epiphysis:

present, absent

Adult thorax description:

Ventral half of lateral cervical sclerites broad, with lateral angles only slightly produced. Metafurca similar to Adelidae, with well developed dorsal apophyses arising perpendicular from anteromedial process, free from secondary arms.

Forelegs:

normal

Number of tibial spurs foreleg:

from 0

Number of tibial spurs midleg:

from 2

Number of tibial spurs hindleg:

from 4

Leg description:

Legs with tibial spur pattern of 0-2-4; epiphyses either present or absent.

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

from 3.5 to 9.0

Wing venation??description:

Radius with 4-5 branches, Rs4 to costa above apex; accessory cell present; M with 2-3 branches; base of M faint or absent within cell; 1A + 2A with short basal fork:

Wing venation:

heteroneurous

Forewing cell veins:

unforked

Forewing anal vein notation:

1A+2A

Forewing basal loop:

present

Forewing pterostigma:

absent

Forewing chorda:

present

Number of Rs veins in forewing:

from 3 to 4

Number of M veins in forewing:

from 3

Forewing upper surface with microtrichia:

present

Hindwing anal vein notation:

1A+2A

Number of anal veins reaching margin:

from 1

Hindwing cell vein:

forked

Number of M veins in hindwing:

from 3

Hindwing pterostigma:

absent

Wing coupling:

present, with frenulum

Wing coupling description:

Male frenulum a single stout bristle in male accompanied by several smaller, costal setae; female with usually an undifferentiated, basal series of much smaller costal setae. Male retinaculum either a long costal fold or a short flap extending under base of Sc.

Wing scales:

hollow

Forewing description:

Forewings slender, L/W index 0.24-3.3; microtrichia usually present on all wing surfaces.

Hindwing description:

Hindwings nearly as broad as forewings; M with 3 branches; base of M usually preserved and forked; 1A + 2A with short basal fork.

Counter-tympanum:

absent

Abdomen tympanum:

absent

Thorax tympanum:

absent

Palp tympanum:

absent

Ocelli:

absent

Eyes:

smooth

Number of labial palp segments:

from 3

Labial palpus modification:

Labial palpi 3-segmented, relatively short, drooping, with erect  bristles from apex of second segment.

Maxillary palpus:

present, folded

Number of maxillary palp segments:

from 5

Number of chaetosomata:

from 0

Proboscis:

reduced

Proboscis texture:

naked

Proboscis description:

Proboscis unscaled, short, ca. 1/2 the length of labial palpus.

Mandibles:

reduced

Head vertex scaling:

very rough

Female antennae:

bipectinate, filiform

Female scape description:

Scape smoothly scaled; pecten absent

Male scape description:

Scape smoothly scaled; pecten absent

Sensillum vesiculocladum:

absent

Asciod sensilla:

absent

General antennae description:

Antennae 0.4-0.8 the length of forewing; scape smoothly scaled; pecten present; flagellum usually filiform, bipectinnate in male Incurvaria; annuli usually indistinct with scales scattered over flagellomeres.

Adult head description:

Head usually entirely rough with piliform scales; frons smooth in Paraclemensia, with broader, laminate scales. Eyes moderately reduced; interocular index 0.6-1.0; cornea without interfacetal microsetae.

Other diagnostic characters:

The depressed apex of the ovipositor and trunctate caudal margin of A7 (female), synapomorphies shared with its nearest relative. Crinopterygidae.

Apomorphies:

Presence of large, noncontiguous spines on the male valvae, in lieu of a pectinifer

Adult behavior:

diurnal

Larval food habits description:

Hosts include members of the Aceraceae, Betulaceae, Caprifoliaceae, Cornaceae, Ericaceae, Fagaceae, Myrtaceae, Proteaceae, and Rosaceae (Common 1990. Heath and Pelham-Clinton 1976, Scoble 1980).

Life history larvae:

Eggs are frequently inserted into the underside of the host leaf where the first instar develops an irregular blotch mine (Dziurzynski 1958). On completion of the mining stage, most larvae cut through the upper and lower epidermal layers of the mine and construct an oval,  lenticular case of the two sections by sewing the edges together. At this stage, leaf damage becomes most obvious as nearly circular holes begin to appear on the host, sometimes accompanied by circular patches of skeletonized leaf (Jensen 1932). Dragging the case, the larva continues to feed externally on its host (Alloclemensia, Basileura, Paraclemensia, Vespina; Davis 1972, Nielsen 1981,  Parra & Ibarra 1994a, Ross 1958) or descends often to the ground to feed on dead or partially withered leaves (Incurvaria). As the larva develops, it enlarges the case by cutting oval sections from leaves and adding these to the top and bottom of the smaller, older case. A  few genera (e. g.. Perthida, Protaephagus) continue to mine throughout their larval development and do not abandon the mine to construct a case until ready to pupate (Common 1969, Scoble 1980).

Description of egg life history:

Eggs are frequently inserted into the underside of the host leaf where the first instar develops an irregular blotch mine.

Systematic and taxonomic history:

Based in part on the shared synapomorphies of the depressed apex of the oviscapt and truncate caudal margin of the abdominal segment 7 in the female, the sister group of the Incurvariidae appears to be the monotypic family Crinopterigidae.

Barcode of Life Data Systems (BOLD) Stats
                                        

Specimen Records:	161	Public Records:	7
Specimens with Sequences:	101	Public Species:	3
Specimens with Barcodes:	29	Public BINs:	2
Species:	65
Species With Barcodes:	6

Access Published & Released Data: Download FASTA File

Collection Sites: world map showing specimen collection locations for Incurvariidae