MORPHOLOGY

Superfamily TORTRICOIDEA

The Superfamily TORTRICOIDEA currently includes only one family, the TORTRICIDAE. There are close to 9000 species worldwide, and over 1200 occur in the United States. The TORTRICOIDEA are most closely related to the COSSOIDEA (Common 1990), although to what extent is not certain.

Family TORTRICIDAE

The TORTRICIDAE includes three subfamilies, the Tortricinae, the Olethreutinae, and the Chlidanotinae. General characteristics of the adult are as follows:

• Adults have a 8-40 mm wingspan.
• The head is rough-scaled above.
• The antennae are less than 2/3rds the length of the forewing (or slightly more than half the length of the forewing).
• The antennae are usually filiform.
• Ocelli and chaetosemata are usually present.
• The proboscis is unscaled and well-developed.
• Labial palpi are rough-scaled and usually porrect, the third (or apical) segment is shorter than the second and usually horizontal.
• The base of the forewing in many males has a costal fold which contains an expandable hair pencil of scent scales.
• Many Olethreutinae and some Tortricinae have a pecten of hairs present near the base of Cu2 on the hindwing (see wing illustrations below).
• In the male, the frenulum is a single bristle which hooks beneath Sc on the forewing.
• In the female, the frenulum consists of three finer bristles which hook on a raised gropu of scales beneath Cu2 on the forewing.
• In males, a hair pencil is sometimes present on the hind tibia.

The following diagrams illustrate the venation of a "typical" moth from the family Tortricidae; both of these examples are from the subfamily Olethreutinae. (Illustrations adapted from Heinrich, 1923, following the nomenclature of Borror et al., 1989.)

Tortricid forewing (click for larger diagram)	Tortricid hindwing (click for larger diagram)

Subfamily Tortricinae

The subfamily Tortricinae contains 11 tribes (Scoble 1992), and its members are more widely distributed in the Southern Hemisphere. This is the Tortricid subfamily with the least specialized male genitalia, which is a characteristic of the group, as listed below. Two characteristics of the Tortricinae (from Scoble 1992) are:

• The antennae nearly always possess two rows of scales on each flagellar segment.
• The juxta and the aedeagus are usually articulated rather than fused in the male genitalia.

Some representatives of the subfamily are depicted below:

Epitymbia cosmota (Meyrick) 1886 Epitymbiini from Australia	Acleris subnivana (Walker) 1863 Tortricini from Connecticut

Please note that most authors agree that the Tortricinae is an unnatural group, so future publications may move some tribes out of the subfamily, however, currently the following tribes are included:

• Archipini
• Atteriini
• Ceracini
• Cnephasiini
• Cochylini
• Epitymbiini
• Euliini
• Phricanthini
• Schoenotenini
• Sparganothini
• Tortricini

Subfamily Olethreutinae

The subfamily Tortricinae contains 6 tribes (Scoble 1992), and its members are more widely distributed in the Northern Hemisphere. This is the largest of the three Tortricid subfamilies, containing at least 3/4 of all Tortricid species (Miller 1987). Two characteristics of the Olethreutinae (from Scoble 1992) are:

• In the male genitalia, the aedeagus is fused with the anellus and juxta.
• There is an "excavation" at the base of the valva in the male genitalia.

Some representatives of the subfamily are depicted below:

Olethreutes tiliana (Heinrich) 1923 Olethreutini from Ohio	Eucosma hyponomeutana (Walsingham) 1895 Eucosmini from Colorado

The following tribes are included in the Olethreutinae, which appears to be monophyletic:

• Bactrini
• Enarmoniini
• Eucosmini
• Gatesclarkeanini
• Grapholitini
• Microcorsini
• Olethreutini

Subfamily Chlidanotinae

The subfamily Chlidanotinae consists of three tribes which are found mainly in the Neotropics and Australia. Recently the Glyphipterigid-patterned Hilarographini have been placed in this subfamily, and it is worth noting that Hilarographa youngiella was described by Busck in 1922 from a specimen caught by Annette F. Braun in Washington. The main characteristic of the Chlidanotinae (from Scoble 1992) is:

• There is a deep, dorso-longitudinal invagination of each of the valvae in the male genitalia.

Some representatives of the subfamily are depicted below:

Trymalitis optima (Meyrick) 1911 Chlidanotini from Australia	Thaumatographa mesostigmatias Diakonoff 1977 Hilarographini from Tawian

The following tribes are included in the Chlidanotinae, which appears to be monophyletic:

• Chlidanotini
• Hilarographini
• Polyorthini