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).
Nearctic, Palearctic, Oriental, Ethiopian, Neotropical, Australian
Larval head description:
Head mostly prognathous with 6 pairs of stemmata; AF2 absent; adfrontal sclerite elongate, extending to deeply incised epicranial notch.
Body setae on verrucae:
Body setae on chalazae:
Body setae on scoli:
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).
Pairs of thoracic legs:
Larval abdomen description:
Prolegs greatly reduced.
Pairs of abdominal legs:
Pairs of crochets per proleg:
multiserial, in transverse rows
Crochet arrangement description:
Anal comb on A10:
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:
Spines as modified cremaster:
Number of rows of tergal spines:
from 1-2 to scattered
Pupation within larval case.
Adult Abdomen Morphology
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 pregenital sexual scales:
Female accessory glands:
Female oviduct opening:
Female anterior apophyses originating:
originating from T8
Male pregenital sexual scales:
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 gland:
Adult abdomen description:
Abdomen with caudal rim of sternum 2a broadly U-shaped; caudal margin of female sternum 7 truncate.
phallotheca and aedeagus (phallus)
Adult Thorax Morphology
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.
Number of tibial spurs foreleg:
Number of tibial spurs midleg:
Number of tibial spurs hindleg:
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
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:
Forewing cell veins:
Forewing anal vein notation:
Forewing basal loop:
Number of Rs veins in forewing:
from 3 to 4
Number of M veins in forewing:
Forewing upper surface with microtrichia:
Hindwing anal vein notation:
Number of anal veins reaching margin:
Hindwing cell vein:
Number of M veins in hindwing:
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.
Forewings slender, L/W index 0.24-3.3; microtrichia usually present on all wing surfaces.
Hindwings nearly as broad as forewings; M with 3 branches; base of M usually preserved and forked; 1A + 2A with short basal fork.
Adult Head Morphology
Number of labial palp segments:
Labial palpus modification:
Labial palpi 3-segmented, relatively short, drooping, with erect bristles from apex of second segment.
Number of maxillary palp segments:
Number of chaetosomata:
Proboscis unscaled, short, ca. 1/2 the length of labial palpus.
Head vertex scaling:
Female scape description:
Scape smoothly scaled; pecten absent
Male scape description:
Scape smoothly scaled; pecten 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.
The depressed apex of the ovipositor and trunctate caudal margin of A7 (female), synapomorphies shared with its nearest relative. Crinopterygidae.
Presence of large, noncontiguous spines on the male valvae, in lieu of a pectinifer
Life History and Behavior
Life History: Immature Stages
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.
Evolution and Systematics
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.
Molecular Biology and Genetics
Statistics of barcoding coverage
Specimens with Sequences:631
Specimens with Barcodes:467
Species With Barcodes:75
Incurvariidae is a family of small primitive monotrysian moths in the order Lepidoptera. There are twelve genera recognised (Davis, 1999). Many species are leaf miners and much is known of their hostplants, excluding Paraclemensia acerifoliella (Fitch). The most familiar species in Europe are perhaps Incurvaria masculella and Phylloporia bistrigella. The narrow wings are held tightly along the body at rest and some species have very long antennae.
- "lepidoptera:Incurvariidae". www.leafmines.co.uk. Retrieved 2007-12-02.
- "HOSTS Search criteria: Lepidoptera Family: Incurvariidae". The Natural History Museum. Retrieved 2007-12-02.
- "Incurvaria masculella". UKmoths. Retrieved 2007-12-02.
- "Phylloporia bistrigella". UKmoths. Retrieved 2007-12-02.
- Davis, D.R. (1999). The Monotrysian Heteroneura. Ch. 6, pp. 65–90 in Kristensen, N.P. (Ed.). Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. Handbuch der Zoologie. Eine Naturgeschichte der Stämme des Tierreiches / Handbook of Zoology. A Natural History of the phyla of the Animal Kingdom. Band / Volume IV Arthropoda: Insecta Teilband / Part 35: 491 pp. Walter de Gruyter, Berlin, New York.
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