The Tortricoidea is one of the most diverse superfamilies in the microlepidoptera,
second only to the Gelechioidea in number of described species (Horak 1998). It
contains a single family, Tortricidae, which is composed of more than 9,800 species in
approximately 1,050 genera.
Tortricid adults are characterized
by a combination of the following characters: head rough-scaled above; scaling of
lower frons short, appressed and upwardly directed; proboscis well developed and
unscaled; labial palpi three-segmented and generally held horizontally or porrect,
with apical segment short and blunt; maxillary palpi reduced; ocelli and chaetosema
present; ovipositor lobes flat (Horak 2006). The structure of the female ovipositor
is the only single apomorphy that unites the entire family.
Tortricidae is divided into three
subfamilies: Tortricinae, Olethreutinae, and Chlidanotinae (Horak 1998). The family
has been recently catalogued (Brown 2005), and an updated online database of
generic and species names along with their synonyms is available
(T@RTS; Baixeras et al. 2008).
Phylogenetic relationships surrounding the Tortricidae are only recently investigated.
Tortricids are included in the non-obtectomeran Apoditrysia, and have been hypothesized
to be related to the Cossoidea or the Cossoidea + Sesioidea by various authors (Common 1990, Horak 1998).
A recent molecular study using five protein-coding nuclear
genes to determine relationships within the Ditrysia failed to consistently place the
Tortricoidea within the Lepidoptera, although the superfamily was consistently recovered as
monophyletic (Regier et al. 2009).
The subfamily Tortricinae contains 11 tribes (Horak 1998). Both the Tortricinae and
Chlidanotinae share a combination of the following characters that separate them from
- 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
- The sterigma is usually connected with the female anterior apophyses.
- A cubital pecten is rarely present.
- Most pheromones consist of a 14-carbon chain.
Although they share many of the same characters, the Chlidanotinae can be separated from the
Tortricinae by the structure of the male and female genitalia.
Epitymbia cosmota (Meyrick)
Epitymbiini from Australia
Acleris subnivana (Walker)
Tortricini from Connecticut
Most authors agree that the Tortricinae is paraphyletic, and the status of this subfamily may
be better understood with future molecular study. A brief summary of the 11 tribes contained within
the subfamily is given below (summarized from Horak 1991 & 1998, Brown et al. 2011).
The Phricanthini contains approximately 20 species that are distributed in Australia, Asia, and Madagascar.
Members of this group feed exclusively on plants in the Dilleniaceae, an uncommon example of larval host
specialization within the family. Synapomorphies within the group include the following: sclerotized
socii; inwardly projecting modified setae on male abdomen; larva with SD1, SD2, and spiracle on
the same pinaculum on A1-8.
The Archipini is the largest tribe within the Tortricinae, with more than 1,600 described species in 150 genera.
Its members are present in all regions but are uncommon in the neotropics. Most are polyphagous leafrollers,
and this tribe contains many pests, such as the light brown apple moth (Epiphyas postvittana Walker).
A synapomorphy for the group is the dagger-shaped signum in the female corpus bursae.
The Epitymbiini is found only in Australia where its approximately 100 members feed on eucalyptus
leaf litter (Myrtaceae). There is speculation that members of the Epitymbiini are simply archipines
with specialized feeding habits, as no morphological characters separate the two tribes.
The Schoenotenini consists of approximately 200 species that occur primarily in the Oriental and
Australian regions. Larvae of the group have a variety of feeding habits. Adult synapomorphies include
the development of the M-vein within the forewing cell and modified chaetosemata.
The Atteriini is a small tribe consisting of roughly 50 species. Most are Neotropical, and little
is known regarding their larval habits. Synapomorphies for the group include elaborate scaling on the
Most of the 670 members of this tribe occur in the Neotropics. Like the Atteriini, the larval
stages of most species are poorly known. A possible synapomorphy for the tribe is the presence of a hair
pencil on the base of the male femur.
This tribe contains approximately 240 members that occur mostly in the New World. The larvae are
polyphagous leafrollers. Male Sparganothini are characterized by large, scaled, lobed socii and
a gnathos that is rudimentary or fused with the socii.
This is a large tribe with more than 1,000 described species. The Cochylini occur in all faunal regions, but are
primarily Holarctic and Neotropical. The larvae are internal feeders in roots, stalks, and seeds.
Synapomorphies for the group include wing pattern and venation and the structure of the male genitalia.
The Cnephasiini contains approximately 250 species that are distributed throughout the Holarctic. The larvae of this
group feed within webbed or spun leaves. Synapomorphies for the group include a spined uncus, floricomous ovipositor,
band-shaped signum, and a 12-carbon pheromone (usually restricted to olethreutines).
The Ceracini is a small tribe of approximately 30 species. They are primarily Oriental, occuring in
Japan and Malaysia. The larvae are leaf rollers. Synapomorphies for the tribe include a pupa without
dorsal spines, the arrangement of the tarsal setae in the larvae, and an ostium that is not connected
with the anterior apophyses in the female genitalia.
The 400 members of the Tortricini are primarily Holarctic and Oriental. The females oviposit eggs singly, and
the larvae are leafrollers. Synapomorphies for the group include a modified juxta-anal tube complex.
The subfamily Olethreutinae contains 6 tribes (Horak 2006), and its members are more
widely distributed in the Northern Hemisphere. This is the second largest of the three
Tortricid subfamilies. Several characteristics of the Olethreutinae are:
- The antennae nearly always possess one row of scales on each flagellar segment.
- The juxta-caulis-aedeagus complex is usually fused in the male.
- The sterigma is usually not connected with the female anterior apophyses.
- A cubital pecten is usually present.
- Most pheromones consist of a 12-carbon chain.
Olethreutes tiliana (Heinrich)
Olethreutini from Ohio
Eucosma hyponomeutana (Walsingham)
Eucosmini from Colorado
Most authors agree that the Olethreutinae is monophyletic. A brief summary of the 6 tribes contained within
the subfamily is given below (summarized from Horak 1991, 1998, & 2006).
The Microcorsini is a small tribe consisting of approximately 45 species in two genera (Cryptaspasma and Collogenes).
This is the most basal group within the Olethreutinae. The larvae are borers in hard nuts. Synapomorphies for the group
includes anterior apophyses connected to the sterigma and horn-shaped signa.
This tribe contains over 150 species that are distributed worldwide. The larvae are stem borers in Juncaceae, Cyperaceae,
and Gramineae. Synapomorphies for the group include a modified sacculus and anteriorly invaginated sterigma. Dang (1990) included the
former tribe Endotheniini in the Bactrini.
This is a large group, containing over 1,100 described species. Its members occur worldwide, although the greatest diversity is in SE Asia
and the Holarctic. The larvae are mainly leafrollers. There are no synapomophies that define the entire tribe, although a hindwing with
veins M3 and CuA1 separate and a sacculus with two clusters of spines unite some species. Horak (2006) included the former tribe
Gatesclarkeanini as the Gatesclarkeana group within the Olethreutini.
The Eucosmini is another large tribe, with approximately 1,650 described species. Their greatest diversity is in the Holarctic, although
members of the tribe occur worldwide. The larvae are leaf-rollers, leaf-webbers, or stem and root borers. One potential synapomophy for
the tribe is the base of vein M2 bent towards the stalked base of veins M3 and CuA1 in the hindwing.
This tribe consists of approximately 300 species. The Enarmoniini is distributed worldwide, although their greatest diversity is
in the Australian and Oriental regions. The larvae are leaf-rollers or stem and root borers. There is no set of characters that
defines the entire tribe; potential synapomorphies include forewing shape, fused socii, and deciduous cornuti.
The Grapholitini contains approximately 900 species with a worldwide distribution. The larvae are borers in fruits, roots, and shoots, and
this group contains many pest species. Synapomorphies for the tribe include a hindwing with veins M2 and M3 widely separated at the base
and a reduced dorsal complex in the male genitalia.
The subfamily Chlidanotinae consists of three tribes found mainly in the
Neotropics and Australia. They share a combination of characters with the Tortricinae,
although structures of the genitalia, such as a deep dorso-longitudinal invagination
of the valvae in the male genitalia, can be used to separate the Chlidanotinae.
Trymalitis optima (Meyrick)
Chlidanotini from Australia
Thaumatographa mesostigmatias Diakonoff
Hilarographini from Tawian
The following tribes are included in the Chlidanotinae, which appears to be
The Polyorthini contains approximately 140 species that are distributed primarily throughout the Neotropics and
Oriental/Australian regions. The larvae are leaf-rollers or stem borers. Synapomorphies for the tribe include
the shape and structure of the valvae and anellus.
Around 75 species are contained in this tribe, which is distributed throughout the Neotropics and
Oriental/Australian regions. The larvae are internal feeders. The group is defined by modified, distally
compressed antennae as well as wing venation.
The pantropical Hilarographini contains approximately 70 species. The known larvae are stem borers. Synapomorphies
include slender, smooth, upcurved palpi, large ocelli, and ciliate male antennae.
Baixeras, J., J. W. Brown and T. M. Gilligan. 2008. T@RTS: Online World Catalogue of the Tortricidae (Version 1.2.1). http://www.tortricidae.com/catalogue.asp.
Brown, J. W. 2005. Tortricidae (Lepidoptera) In World Catalogue of Insects 5:1-741. Apollo Books, Stenstrup, Denmark.
Brown, J. W., J. Baixeras and C. Mitter. 2011. Taxon: Tortricidae. Leptree.net. http://www.leptree.net/lep_taxon_page/Tortricidae/view.
Common, I. F. B. 1990. Moths of Australia. Melbourne University Publishing. 535 pp.
Horak, M. 1991. Morphology, pp. 1-22. In L. P. S. van der Geest, H. H. Evenhuis (eds.), Tortricid Pests: Their Biology, Natural Enemies and Control. Elsevier, Amsterdam, The Netherlands.
Horak, M. 1998. The Tortricoidea, pp. 199-215. In N. P. Kristensen (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York.
Horak, M. 2006. Olethreutine moths of Australia (Lepidoptera: Tortricidae). Monographs on Australian Lepidoptera, Vol. 10. 522 pp.
Regier, J. C., A. Zwick, M. P. Cummings, A. Y. Kawahara, S. Cho, S. J. Weller, A. D. Roe, J. Baixeras-Almela, J. W. Brown, C. S. Parr, D. R. Davis, M. E. Epstein, W. Hallwachs, A. Hausmann, D. H. Janzen, I. J. Kitching, M. A. Solis, S.-H. Yen, A. Bazinet and C. Mitter. 2009. Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary Biology 9: 280.