Tortricoidea is one of the most diverse superfamilies of the so-called microlepidoptera, second only to Gelechioidea in number of described species (Horak 1998Horak 1998:
Horak, M. 1998. The Tortricoidea, pp. 199-215. In : Kristensen, N. P. (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York.). It contains a single family, Tortricidae, which is composed of more than 11,300 described species assigned to approximately 1,150 genera (Gilligan et al. 2018Gilligan et al. 2018:
Gilligan, T. M., Baixeras, J., Brown, J. W. 2018. T@RTS: Online World Catalogue of the Tortricidae (Ver. 4.0). http://www.tortricid.net/catalogue.asp [accessed 8 October 2020].). The enigmatic Australian genus Heliocosma is sometimes assigned to the superfamily, but left unplaced to family. Tortricid adults are characterized by a combination of the following characters: vertex of head rough-scaled; scaling of lower fronsfrons:
the upper anterior portion of the head (in adults) 
short, appressed, and upwardly directed; proboscis well developed and unscaled; labial palpilabial palpi:
a pair of sensory appendages that project from the lower part of the head; usually covered in scales and three-segmented
three-segmented and typically directed horizontally or porrectporrect:
extending forward horizontally 
(but upcurved in some species), with the apical segment short and blunt; maxillary palpimaxillary palpi:
a pair of segmented appendages extending from the lower part of the head; usually smaller than the labial palpi and may be reduced or absent in some families
reduced; ocelliocelli:
adult head - a simple insect "eye" located dorsal to the compound eye; plural of ocellus 
usually present; chaetosematachaetosema:
a small sensory organ located above the compound eye in some families of Lepidoptera
always present; and ovipositor lobes in female genitalia flat (Horak 2006Horak 2006:
Horak, M. 2006. Olethreutine moths of Australia (Lepidoptera: Tortricidae). Monographs on Australian Lepidoptera, Vol. 10. 522 pp.). Although the structure of the female ovipositor is the only recognized adult apomorphy that unites the family, features of the larvae and pupae are diagnostic, or nearly so: D2 setae on a shared dorsaldorsal:
upper, to the top, on the back 
pinaculumpinaculum:
flattened sclerotized plates on a caterpillar that bear the setae
on A10; presence of a characteristic anal fork (although lost secondarily in many species with internal-feeding larvae); and pupae with two transverse rows of spines on the dorsumdorsum:
the lower (usually posterior) or inner margin 
of abdominal segments 3–6 (Powell and Brown 2012Powell and Brown 2012:
Powell, J. A., Brown, J. W. 2012. Tortricoidea, Tortricidae (part): Tortricinae (part): Sparganothini and Atteriini. In: Hodges, R. W. (ed.). The Moths of North America, fascicle 8.1. Wedge Entomological Research Foundation, Washington, D.C., 230 pp.). Tortricidae are divided into three subfamilies: Tortricinae, Olethreutinae, and Chlidanotinae (Horak 1998Horak 1998:
Horak, M. 1998. The Tortricoidea, pp. 199-215. In : Kristensen, N. P. (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York.). Whereas the monophyly of Tortricinae and Olethreutinae are well supported by morphological and molecular evidence, the monophyly of Chlidanotinae has been brought into question based on molecular data (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ).
Several of the taxa now recognized as tribes or subfamilies within Tortricidae have historically been considered distinct and separate families (e.g., Atteriidae, Sparganothidae, Ceracidae, Cochylidae, Eucosmidae). It wasn’t until the 1960s that consensus was reached that Tortricinae and Olethreutinae belong to a single family rather than representing different families. And as recently as 1983 (Powell 1983), Cochylina were still considered a family (Cochylidae or Phaloniidae) separate from Tortricidae. Tortricidae were catalogued by Brown (2005)Brown (2005):
Brown, J. W. 2005. Tortricidae (Lepidoptera) In : World Catalogue of Insects 5: 1-741. Apollo Books, Stenstrup, Denmark., and an updated online database of generic and species names along with their synonyms is available (Gilligan et al. 2018Gilligan et al. 2018:
Gilligan, T. M., Baixeras, J., Brown, J. W. 2018. T@RTS: Online World Catalogue of the Tortricidae (Ver. 4.0). http://www.tortricid.net/catalogue.asp [accessed 8 October 2020].).
Traditionally, Tortricoidea were considered sister to Cossoidea or the Cossoidea + Sesioidea lineage by various authors (Powell 1964bPowell 1964b:
Powell, J. A. 1964b. Biological and taxonomic studies on tortricine moths, with reference to the species in California. University of California Publications in Entomology. Vol. 32. 317 pp., Common 1990Common 1990:
Common, I. F. B. 1990. Moths of Australia. Melbourne University Publishing. 535 pp., Horak 1998Horak 1998:
Horak, M. 1998. The Tortricoidea, pp. 199-215. In : Kristensen, N. P. (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York.), based on the assumed shared ancestral internal-feeding habit of the larvae and similar morphological features of the pupae. However, it is now believed that ancestral tortricids were likely external leaf-rollers (Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ). An initial molecular study using five protein-coding nuclear genes to determine relationships within the Ditrysia failed to consistently place Tortricoidea as sister to any other superfamily, although it was consistently recovered as monophyletic (Regier et al. 2009Regier et al. 2009:
Regier, J. C., Zwick, A., Cummings, M. P., Kawahara, A. Y., Cho, S., Weller, S. J., Roe, A. D., Baixeras-Almela, J., Brown, J. W., Parr, C. S., Davis, D. R., Epstein, M. E., Hallwachs, W., Hausmann, A., Janzen, D. H., Kitching, I. J., Solis, M. A., Yen, S.-H., Bazinet, A., Mitter, C. 2009. Toward reconstructing the evolution of advanced moths and butterflies (Lepidoptera: Ditrysia): an initial molecular study. BMC Evolutionary Biology. 9(280): 1-21.). More recent studies found Tortricoidea to be sister to a clade comprised of Heliocosma + Galacticidae, with these two clades sister to a clade comprised of Zygaenoidea + Cossoidea + Sesiidae, all within the non-obtectomeran Apoditrysia (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574.). Tortricoidea appear to have diverged from their sister group approximately 120 mya and diversified approximately 97 mya along with the angiosperms in the Early-mid Cretaceous (Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ). Relationships within the superfamily remained mostly unresolved based on morphology, but recent molecular studies (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ) show well-supported, nested groups (tribes), many of which share various morphological and behavior traits. For example, within the subfamily Tortricinae, the tribes Archipini, Ceracini, Sparganothini, and Atteriini all share slender deciduous cornuticornutus:
spines used to anchor the male vesica in the female bursa during copulation
in the vesicavesica:
An eversible sheath in which the phallus (aedeagus) is extended through during mating. In many tortricids, cornuti are embedded in the vesica. 
; females lay large imbricate patches of eggs; and larvae are nearly all polyphagous (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Anzaldo et al. 2014Anzaldo et al. 2014:
Anzaldo, S.S., Dombroskie, J., Brown, J.W. 2014. Morphological variation, taxonomic distribution, and phylogenetic significance of cornuti in Tortricinae (Lepidoptera: Tortricidae)." Proceedings of the Entomological Society of Washington. 116 (1):1–31.). In the sister clade, comprised of Tortricini, Cnephasiini, and Cochylini, the cornuticornutus:
spines used to anchor the male vesica in the female bursa during copulation are non-deciduous; females lay small patches of eggs (2–6); and most species are relatively host specific. Nonetheless, the position of a few “orphan groups” (e.g., Mictopsichia, Arotrophora) remains unclear.
As presently defined, the subfamily Chlidanotinae consists of three tribes distributed mainly in the tropical regions of the planet. Although they share a number of characters with Tortricinae (see below), they are easily distinguished by a deep, dorso-longitudinal invagination of the valvaevalva:
an appendage flanking the intromittent organ that is used to clasp the female during copulation
holding distinctive hair-pencilshair-pencil:
specialized sex scales present in many male Lepidoptera that emit pheromones used in attracting a mate, courtship, and/or copulation. These can be present in many different locations in tortricids, including the abdomen, valvae, wings, or legs.
that arise from the eighth abdominal segment in the male genitalia (Tuck 1981Tuck 1981:
Tuck, K.R. 1981. A new genus of Chlidanotini (Lepidoptera: Tortricidae) from New Caledonia, with a key to the genera and checklist of species. Systematic Entomology. 6: 337-346., Brown 1990Brown 1990:
Brown, J. W. 1990. Description of a new genus in the Chlidanotini and review of phylogenetic relationships among chlidanotine tribes (Lepidoptera: Tortricidae: Chlidanotinae). Entomologica Scandinavica. 20: 439-448.). While morphology appears to support the monophyly of the three tribes assigned to Chlidanotinae, molecular data suggest that Polyorthini are actually the sister group to the remainder of Tortricidae, casting doubt on the monophyly of the subfamily.
Chlidanotinae and Tortricinae share the following characters that separate them from the Olethreutinae:
are usually articulated rather than fused in the male genitalia.
is usually connected with the female anterioranterior:
apophysesapophyses:
.
is almost always absent.Chlidanotinae can be separated from Tortricinae by a remarkable feature of the genitalia. In males of nearly all Chlidanotinae, the valvavalva:
an appendage flanking the intromittent organ that is used to clasp the female during copulation are large and membranous, deeply divided longitudinally at the costacosta:
the anterior margin of each wing 
into a pocket in which elongate hair-pencilshair-pencil:
specialized sex scales present in many male Lepidoptera that emit pheromones used in attracting a mate, courtship, and/or copulation. These can be present in many different locations in tortricids, including the abdomen, valvae, wings, or legs. from abdominal segment 8 reside (Tuck 1981Tuck 1981:
Tuck, K.R. 1981. A new genus of Chlidanotini (Lepidoptera: Tortricidae) from New Caledonia, with a key to the genera and checklist of species. Systematic Entomology. 6: 337-346., Horak 1984Horak 1984:
Horak, M. 1984. Assessment of taxonomically significant structures in Tortricinae (Lep., Tortricidae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft. 57: 3-64.).
Polyorthini contains approximately 150 species in 20 genera (Brown and Hoddle 2010Brown and Hoddle 2010:
Brown, J. W., Hoddle, M. S. 2010. A new species of Histura Razowski (Lepidoptera: Tortricidae: Polyorthini) from Guatemala attacking avocados ( Persea americana ) (Lauraceae). Proceedings of the Entomological Society of Washington. 102(1): 10-21.) with a pan-tropical distribution; greatest diversity is found in the Neotropical, Indomalayan, and Australian regions. The early stages are poorly known, but larvae appear to be leaf-rollers or stem borers, with a preference for Lauraceae (Brown et al. 2020bBrown et al. 2020b:
Brown, J. W., Gripenberg, S., Basset, Y., Calderoacute;n, O., Simon, I., Fernandez, C., Cedeno, M., Rivera, M. 2020b. Host Records for Tortricidae (Lepidoptera) Reared from Seeds and Fruits in Panama. Proceedings of the Entomological Society of Washington. 122(1): 12-24.). Synapomorphies for the tribe include the shape and structure of the valvaevalva:
an appendage flanking the intromittent organ that is used to clasp the female during copulation and anellusanellus:
a variably cup-shaped sclerotized sheet surrounding the base of the aedeagus
, the exceedingly long labial palpilabial palpi:
a pair of sensory appendages that project from the lower part of the head; usually covered in scales and three-segmented (with several exceptions), and the raised scale tufts of the forewingforewing:
the anterior wing
. Owing to the last character, combined with remarkable polymorphism in some species, many species of Polyortha (Polyorthini) and Acleris (Tortricini) were originally described in Peronea, now recognized as a synonym of Acleris.
Chlidanotini is represented by approximately 75 described species distributed throughout the Neotropics and Indomalayan/Australian regions, with diversity concentrated in the New World tropics (e.g., Razowski and Becker 2000Razowski and Becker 2000:
Razowski, J., Becker, V. O. 2000. A review of the New World Chlidanotini (Lepidoptera: Tortricidae). Revista Brasileira de Zoologia. 16: 1149-1182.). The early stages are poorly known, but reported larvae are fruit borers (e.g., Jayanthi and Verghese 2010Jayanthi and Verghese 2010:
Jayanthi, P. D. K., Verghese, A. 2010. Establishment of sapota seed borer, Trymalitis margarias Meyrick, an invasive species in India: exigencies involved in limiting the spread. Karnataka J. Agric. Sci. 23(1): 165., Bisane 2016Bisane 2016:
Bisane, K.D. 2016. Seasonal cyclicity and behavior of sapota seed borer, Trymalitis margarias Meyrick. Pest Management in Horticultural Ecosystems. 22(2): 129-133.) or gall inducers (e.g., Razowski and Giliomee 2014Razowski and Giliomee 2014:
Razowski, J., Giliomee, J. H. 2014. A new Afrotropical Chlidanotini species (Lepidoptera: Tortricidae) inhabiting galls of white milkwood, Sideroxylon inerme , in South Africa. African Entomology. 22: 796-800.). The tribe shares remarkably similar male and female genitalia and forewingforewing:
the anterior wing pattern elements with Hilarographini, suggesting that the two may be best considered a single tribe. Males in both tribes posses a pair of unique, sclerotizedsclerotized:
hardened; usually in reference to larval structures or adult genitalia 
, sociisocii:
a pair of lightly sclerotized setose lobes
-like structures in the male genitalia call hami; females have a characteristic “star-burst” signumsignum:
a sclerotized projection or patch on the interior of the corpus bursae
of numerous long spines from which a slender accessory bursa originates (Tuck 1981Tuck 1981:
Tuck, K.R. 1981. A new genus of Chlidanotini (Lepidoptera: Tortricidae) from New Caledonia, with a key to the genera and checklist of species. Systematic Entomology. 6: 337-346.); and females typically have two ancanthi in the frenulumfrenulum:
a single spine (males) or multiple spines (females) that project anteriorly from the base of the hindwing; used to couple the hindwings to the forewings in flight by means of a retinaculum on the forewing 
(Yang and Brown 2009Yang and Brown 2009:
Yang, A., Brown, J. W. 2009. Variation in the female frenulum in Tortricidae (Lepidoptera). Part 1. Chlidanotinae. Proceedings of the Entomological Society of Washington. 111: 743-751.). Molecular data strongly support the sister relationship of the two.
The pantropical tribe Hilarographini contains approximately 70 described species, most of which have brightly colored and distinctively patterned forewingsforewing:
the anterior wing. The early stages are poorly known, but the few discovered larvae are reported to be fruit or stem borers (e.g., Brown et al. 2014aBrown et al. 2014a:
Brown, J. W., Copeland, R. S., Aarvik, L., Miller, S. E., Rosati, M. E., Luke, Q. 2014a. Host Records for Fruit-Feeding Afrotropical Tortricidae (Lepidoptera). African Entomology. 22(2): 343-376., Brown et al. 2019bBrown et al. 2019b:
Brown, J.W., Basset, Y., Panmeng, M., Putnaul, S., Miller, S. E. 2019b. Host records for Tortricidae (Lepidoptera) reared from seeds and fruits in a Thailand rainforest. Proceedings of the Entomological Society of Washington. 121(4): 544-556.). Morphological synapomorphies include slender, smooth, upcurved palpi; large ocelliocelli:
adult head - a simple insect "eye" located dorsal to the compound eye; plural of ocellus ; and ciliate antennae in the male (Brown 1990Brown 1990:
Brown, J. W. 1990. Description of a new genus in the Chlidanotini and review of phylogenetic relationships among chlidanotine tribes (Lepidoptera: Tortricidae: Chlidanotinae). Entomologica Scandinavica. 20: 439-448.). As mentioned above, the male and female genitalia are extremely similar to those of Chlidanotini.
As presented defined, the subfamily Tortricinae comprises 10 tribes (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574.). Although long thought to be paraphyletic, recent molecular analyses provide strong support for the monophyly of Tortricinae. Brief summaries of the 10 tribes contained within the subfamily are given below (summarized from Horak 1984Horak 1984:
Horak, M. 1984. Assessment of taxonomically significant structures in Tortricinae (Lep., Tortricidae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft. 57: 3-64., Horak 1991Horak 1991:
Horak, M. 1991. Morphology, pp. 1-22. In : van der Geest, L. P. S., Evenhuis, H. H. (eds.), Tortricid Pests: Their Biology, Natural Enemies and Control. Elsevier, Amsterdam, The Netherlands., Horak 1998Horak 1998:
Horak, M. 1998. The Tortricoidea, pp. 199-215. In : Kristensen, N. P. (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York., Brown et al. 2011Brown et al. 2011:
Brown, J. W., Segura, R., Santiago-Jimenez, Q., Rota, J., Heard, T. A. 2011. Tortricid moths (Lepidoptera: Tortricidae) reared from the invasive weed Parkinsonia aculeata (Fabaceae), with comments on their host specificity, biology, and geographic distribution. Journal of Insect Science. 11(7): 1-17.).
The Phricanthini contains approximately 20 species in four genera that are distributed in Australia, Asia, and Madagascar. One species appears to have been introduced into the New World tropics and mainland Africa (Brown 2007Brown 2007:
Brown, J. W. 2007. Confirmation of the Old World species Phricanthes flexilineana (Walker, 1863) in the New World tropics (Lepidoptera: Tortricidae: Phricanthini). The Pan-Pacific Entomologist. 83(4): 352-357.). Members of this group feed exclusively on plants in the Dilleniaceae, an uncommon example of larval host specialization within the family. Synapomorphies for Phricanthini include the following: well-developed saccus in the male genitalia; erect, sclerotizedsclerotized:
hardened; usually in reference to larval structures or adult genitalia sociisocii:
a pair of lightly sclerotized setose lobes; inwardly projecting modified setae on the anterioranterior:
before, to the front, toward the head margin of the third and/or fourth terga of the male abdomen; larva with SD1, SD2, and spiracle on the same pinaculumpinaculum:
flattened sclerotized plates on a caterpillar that bear the setae on A1-8; and an upright egg.
The Schoenotenini consists of approximately 200 species that occur primarily in the Indomalayan and Australian regions. Larvae of the group have a variety of feeding habits. Adult synapomorphies include the unusual development of the M-vein within the forewingforewing:
the anterior wing cell and modified chaetosematachaetosema:
a small sensory organ located above the compound eye in some families of Lepidoptera.
The Ceracini is a small tribe of approximately 30 species in four genera. They are among the largest of all tortricids, with wingspans up to 60 mm. They are primarily Indomalayan in distribution, although some species occur in Japan and Korea. The larvae are leaf rollers. Synapomorphies for the tribe include a pupa without dorsaldorsal:
upper, to the top, on the back spines; the arrangement of the tarsal setae in the larvae; an ostiumostium:
see ostium bursae 
that is not connected with the anterioranterior:
before, to the front, toward the head apophysesapophyses:
apodemal rods used to extend the ovipositor; divided into the apophyses anteriores and apophyses posteriores in the female genitalia; adults that are large, colorful, and diurnal; and a female frenulumfrenulum:
a single spine (males) or multiple spines (females) that project anteriorly from the base of the hindwing; used to couple the hindwings to the forewings in flight by means of a retinaculum on the forewing with more than three acanthi (Monsalve et al. 2011Monsalve et al. 2011:
Monsalve, S., Dombroskie, J., Lam, W., Rota, J., Brown, J. W. 2011. Variation in the female frenulum in Tortricidae (Lepidoptera). Part 3. Tortricinae. Proceedings of the Entomological Society of Washington. 113: 335-370.).
Epitymbiini are found only in Australia where the approximately 100 members of the tribe feed in the leaf litter of Myrtaceae, primarily in eucalyptus forests. There is speculation that members of the Epitymbiini are simply archipines with specialized feeding habits, as no morphological characters have been identified to separate the two tribes.
The Archipini is the largest tribe within the Tortricinae, with more than 2,000 described species in approximately 230 genera (Brown 2005Brown 2005:
Brown, J. W. 2005. Tortricidae (Lepidoptera) In : World Catalogue of Insects 5: 1-741. Apollo Books, Stenstrup, Denmark.). Its members are present in all regions, but in contrast to many groups, they are considerably less diverse in the New World (Razowski 1997Razowski 1997:
Razowski, J. 1997. Generic composition of the New World Archipini (Lepidoptera, Tortricidae) with description of two new genera and two new species. Miscellanea Zoologicae. 20: 125-130.; Dombroskie and Sperling 2013Dombroskie and Sperling 2013:
Dombroskie, J. J., Sperling, F. A. H. 2013. Phylogeny of the tribe Archipini (Lepidoptera: Tortricidae: Tortricinae) and evolutionary correlates of novel secondary sexual structures. Zootaxa. 3729: 1-62.). Most are polyphagous leafrollers, and the tribe includes many pests, such as the light brown apple moth (Epiphyas postvittana), the oblique-banded leaf roller (Choristoneura rosaceana), the tea tortrix (Homona coffearia), and the notorious budworms (Choristoneura spp.) of North America. A proposed synapomorphy for the group is the dagger-shaped signumsignum:
a sclerotized projection or patch on the interior of the corpus bursae in the female corpus bursaecorpus bursae:
a dilated membranous sac at the anterior end of the bursa copulatrix
, but this structure has been independently lost in several genera. Another putative synapomorphy is the uncusuncus:
a sclerotized process which is fused to the posterodorsal margin of tergum IX
with apicoventral setae in the male, but this character is also present in Epitymbiini and Ceracini. As currently defined, the tribe may be paraphyletic (Horak 1984Horak 1984:
Horak, M. 1984. Assessment of taxonomically significant structures in Tortricinae (Lep., Tortricidae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft. 57: 3-64., Horak 1998Horak 1998:
Horak, M. 1998. The Tortricoidea, pp. 199-215. In : Kristensen, N. P. (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York.).
The Atteriini is a small tribe consisting of approximately 110 species in eight genera (Austin and Dombroskie 2020aAustin and Dombroskie 2020a:
Austin, K. A., Dombroskie, J. J. 2020a. New combinations in neotropical Archipini and Atteriini (Lepidoptera: Tortricidae: Tortricinae), with the description of a new genus. Proceedings of the Entomological Society of Washington. 122(1): 1-11.) restricted almost entirely to the neotropics. These moths are among the largest tortricids in the New World. Females of almost all members possess patches of conspicuous, modified scales on sternites 6–8, a possible synapomorphy (Brown et al. 2019aBrown et al. 2019a:
Brown, J. W., Dyer, L. A., Villamarin-Cortez, S., Salcido, D. 2019a. New larval host records for Tortricidae (Lepidoptera) from an Ecuadorian Andean cloud forest. Insecta Mundi. 1208.). Nearly 100 hosts in 31 plant families have been reported for four species of Anacrusis in Costacosta:
the anterior margin of each wing Rica (Brown et al. 2014bBrown et al. 2014b:
Brown, J. W., Janzen, D., Hallwachs, W. D., Zahiri, R., Hajibabaei, M., Hebert, P. N. D. 2014b. Cracking complex taxonomy of Costa Rican moths: Anacrusis Zeller (Lepidoptera: Tortricidae). Journal of the Lepidopterists#39; Society. 68: 248-263.); additional host plants for species in Ecuador have been reported (Brown et al. 2019aBrown et al. 2019a:
Brown, J. W., Dyer, L. A., Villamarin-Cortez, S., Salcido, D. 2019a. New larval host records for Tortricidae (Lepidoptera) from an Ecuadorian Andean cloud forest. Insecta Mundi. 1208.). Several species have been reared successfully on artificial diet in the lab (Brown et al. 2019aBrown et al. 2019a:
Brown, J. W., Dyer, L. A., Villamarin-Cortez, S., Salcido, D. 2019a. New larval host records for Tortricidae (Lepidoptera) from an Ecuadorian Andean cloud forest. Insecta Mundi. 1208.), and all species are likely polyphagous. Recent molecular studies support the sister relationship of Atteriini + Sparganothini (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ).
This tribe contains approximately 250 species, all but a few of which are restricted to the New World. With few exceptions, the larvae are polyphagous leafrollers. Sparganothini are characterized by male genitalia with large, densely scaled, lobed sociisocii:
a pair of lightly sclerotized setose lobes and a gnathosgnathos:
a narrow bandlike structure that joins the posterolateral edges of the tegumen and supports the anal tube
that is rudimentary or fused with the sociisocii:
a pair of lightly sclerotized setose lobes. The homology of the gnathosgnathos:
a narrow bandlike structure that joins the posterolateral edges of the tegumen and supports the anal tube in this tribe remains uncertain, and the structures in question are frequently referred to as secondary arms of the sociisocii:
a pair of lightly sclerotized setose lobes.
Cnephasiini contains approximately 250 species that are distributed throughout the Holarctic with a few species in south Asia, Australia, and New Zealand. The larvae of this group usually feed within webbed or spun leaves, but some tunnel in flowers (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574.). Synapomorphies for the group include the floricomous papillae analespapillae anales:
the female ovipositor lobes 
(i.e., with nail-head-like setae), a finely spined uncusuncus:
a sclerotized process which is fused to the posterodorsal margin of tergum IX, the absence of cornuticornutus:
spines used to anchor the male vesica in the female bursa during copulation, and pheromones based on 12-carbon chains (Roelofs and Brown 1982Roelofs and Brown 1982:
Roelofs, W. L., Brown, R. L. 1982. Pheromones and evolutionary relationships of Tortricidae. Annual Review of Ecology and Systematics. 13: 395-422., Safonkin 2007Safonkin 2007:
Safonkin, A. F. 2007. Pheromones and phylogenetic relations of leafrollers (Lepidoptera, Tortricidae). Zoologicheskii Zhurnal. 86(12): 1464-1467. [Entomological Review 87(9): 1238-1241]). The shape of the signumsignum:
a sclerotized projection or patch on the interior of the corpus bursae is extremely similar to that of Tortricini, and the floricomous ovipositor characteristic of Cnephasiini is present in a few putatively primitive Tortricini.
The 400 members of the Tortricini are primarily Holarctic and Indomalayan in distribution. The females oviposit eggs singly or in small clusters, and the larvae are leafrollers. Synapomorphies for the tribe include the complete loss of the uncusuncus:
a sclerotized process which is fused to the posterodorsal margin of tergum IX, the development of a subscaphium, the development of a brachiola, raised forewingforewing:
the anterior wing scales, and often extremely polymorphic forewingforewing:
the anterior wing patterns within the same species (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574.).
This is a large tribe with more than 1,670 described species. It includes the former tribe Euliini as subtribe Euliina, which remains paraphyletic (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. , Brown et al. 2020aBrown et al. 2020a:
Brown, J.W., Aarvik, L., Heikkilauml;, M., Brown, R., Mutanen, M. 2020a. A molecular phylogeny of Cochylina, with confirmation of its relationship to Euliina (Lepidoptera: Tortricidae). Systematic Entomology. 45: 160-174.). Cochylini occur in all faunal regions, but the greatest diversity is found in the Holarctic and Neotropical regions. Larvae are diverse, and include leaf rollers, internal feeders in roots, stalks, seeds, and a few species that appear to specialize on leaf litter (e.g., Anopina). A proposed synapomorphy for Euliina is the presence of a foreleg hair-pencilhair-pencil:
specialized sex scales present in many male Lepidoptera that emit pheromones used in attracting a mate, courtship, and/or copulation. These can be present in many different locations in tortricids, including the abdomen, valvae, wings, or legs. in the male (shared with Schoenotenini) (Brown 1990Brown 1990:
Brown, J. W. 1990. Description of a new genus in the Chlidanotini and review of phylogenetic relationships among chlidanotine tribes (Lepidoptera: Tortricidae: Chlidanotinae). Entomologica Scandinavica. 20: 439-448.), although it appears to have been secondarily lost in many genera.
The subtribe Cochylina is strongly supported as monophyletic by both molecular and morphological data. In the male genitalia the gnathosgnathos:
a narrow bandlike structure that joins the posterolateral edges of the tegumen and supports the anal tube is lost, the uncusuncus:
a sclerotized process which is fused to the posterodorsal margin of tergum IX is absent (except in the most primitive members), and the arms of the vinculumvinculum:
ventral section of the transverse ring in male genitalia 
are untied distally by membrane. The female genitalia usually have an accessory bursa, frequently lack differentiation between the corpus bursaecorpus bursae:
a dilated membranous sac at the anterior end of the bursa copulatrix and the ductus bursaeductus bursae:
a membranous tube connecting the ostium bursae to the corpus bursae
, and the corpus bursaecorpus bursae:
a dilated membranous sac at the anterior end of the bursa copulatrix often has complex sclerites, but never with a well-defined signumsignum:
a sclerotized projection or patch on the interior of the corpus bursae.
The subfamily Olethreutinae contains five tribes (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ), 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:
-aedeagus complex is usually fused in the male. is usually not connected with the female anterioranterior: apophysesapophyses:. is usually present in the hindwinghindwing:
.Morphological and molecular data strongly support the monophyly of Olethreutinae. Brief summaries of the five tribes included in the subfamily are given below (summarized from Horak 1991Horak 1991:
Horak, M. 1991. Morphology, pp. 1-22. In : van der Geest, L. P. S., Evenhuis, H. H. (eds.), Tortricid Pests: Their Biology, Natural Enemies and Control. Elsevier, Amsterdam, The Netherlands., Horak 1998Horak 1998:
Horak, M. 1998. The Tortricoidea, pp. 199-215. In : Kristensen, N. P. (ed.), Handbook of Zoology, Lepidoptera, Moths and Butterflies, Vol 1: Evolution, Systematics, and Biogeography. Walter de Gruyter, Berlin, New York., Horak 2006Horak 2006:
Horak, M. 2006. Olethreutine moths of Australia (Lepidoptera: Tortricidae). Monographs on Australian Lepidoptera, Vol. 10. 522 pp., Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574.).
Microcorsini is a small tribe consisting of approximately 45 described species in two genera (Cryptaspasma and Collogenes) with a pan-tropical distribution (Diakonoff 1959Diakonoff 1959:
Diakonoff, A. 1959. Revision of Cryptaspasma Walsingham, 1900 (Lepidoptera, Tortricidae). Zoologische Verhandelingen (Leiden). 43: 1-60.). The tribe is sister to the remaining Olethreutinae, and members retain some plesiomorphic characters of Tortricinae. The larvae are borers in hard nuts, seeds, and fruits (e.g., Gilligan et al. 2011bGilligan et al. 2011b:
Gilligan, T. M., Brown, J. W., Hoddle, M. S. 2011b. A new avocado pest in Central America (Lepidoptera: Tortricidae) with a key to Lepidoptera larvae threatening avocados in California. Zootaxa. 3137: 31-45.). Adults are remarkably uniform in appearance, with a somber brownish black forewingforewing:
the anterior wing with few distinct pattern elements. Synapomorphies for the group include anterioranterior:
before, to the front, toward the head apophysesapophyses:
apodemal rods used to extend the ovipositor; divided into the apophyses anteriores and apophyses posteriores connected to the sterigmasterigma:
the sclerotized region surrounding the female ostium bursae and a pair of large, hollow, cornucopia-shaped signasignum:
a sclerotized projection or patch on the interior of the corpus bursae in the female genitalia.
This is a large group, containing over 1,100 described species. Its members occur worldwide, although the greatest diversity is in southeast Asia and the Holarctic. The larvae are mainly leafrollers, and many genera exhibit conspicuous host fidelity to specific plant genera or families. There are no recognized synapomorphies that define the tribe, although hindwinghindwing:
the posterior wing veins M3 and CuA1 separate, and a sacculussacculus:
the ventral margin of the male valva 
with two clusters of spines serve to unite many genera. Diakonoff (1973)Diakonoff (1973):
Diakonoff, A. 1973. The Southeast Asia Olethreutini (Lepidoptera, Tortricidae). Zoologische Monographieen van het Rijksmuseum van Natuurlijke Historie, No. 1, Leiden. E. J. Brill. 700 pp. defined several subtribes in the group, which subsequently were considered distinct tribes by Nedoshivina (2016)Nedoshivina (2016):
Nedoshivina, S. V. 2016. Leafroller moths of the tribe Olethreutini (Lepidoptera, Tortricidae: Olethreutinae) of Russia. Korporaciya Technology Prodvizheniya Publishing, Ulyanovsk. 328 pp.. Horak (2006)Horak (2006):
Horak, M. 2006. Olethreutine moths of Australia (Lepidoptera: Tortricidae). Monographs on Australian Lepidoptera, Vol. 10. 522 pp. included the former tribe Gatesclarkeanini as the Gatesclarkeana group within the Olethreutini; a later molecular study synonymized Bactrini and Endothenini under Olethreutini as well (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574.).
This tribe consists of approximately 300 described species distributed worldwide, with greatest diversity in the Australasian and Indomalayan regions. The larvae are leaf-rollers or stem and root borers, often with restricted host preferences. There is no set of characters that defines the entire tribe, but potential synapomorphies include a forewingforewing:
the anterior wing with a falcatefalcate:
narrow, curved and pointed; sickle-shaped; shaped like a bird's beak 
apex, fused sociisocii:
a pair of lightly sclerotized setose lobes, deciduous cornuticornutus:
spines used to anchor the male vesica in the female bursa during copulation, and blade-like signasignum:
a sclerotized projection or patch on the interior of the corpus bursae in females (although the last two are most likely symplesiomorphies).
The Eucosmini is another large tribe, with approximately 1,650 described species. 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 synapomorphy for the tribe is the base of vein M2 bent towards the stalked base of veins M3 and CuA1 in the hindwinghindwing:
the posterior wing, but exceptions exist (Horak 2006Horak 2006:
Horak, M. 2006. Olethreutine moths of Australia (Lepidoptera: Tortricidae). Monographs on Australian Lepidoptera, Vol. 10. 522 pp.). Molecular data suggest Eucosmini is sister to Grapholitini (Regier et al. 2012Regier et al. 2012:
Regier J. C., Brown, J. W., Mitter, C., Baixeras, J., Cho, S., Cummings, P., Zwick, A. 2012. A molecular phylogeny for the leaf-roller moths (Lepidoptera: Tortricidae) and its implications for classification and life history evolution. PLoS ONE. 7(4): e35574., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ).
Grapholitini comprises nearly 1,000 species distributed worldwide. The larvae are nearly all borers in fruits, nuts, and seeds (e.g., Brown 2019Brown 2019:
Brown, J. W. 2019. Descriptions of four new species of fruit-feeding tortricid moths from Panama (Lepidoptera: Tortricidae). Tropical Lepidoptera Research. 29: 67-73., Brown et al. 2019aBrown et al. 2019a:
Brown, J. W., Dyer, L. A., Villamarin-Cortez, S., Salcido, D. 2019a. New larval host records for Tortricidae (Lepidoptera) from an Ecuadorian Andean cloud forest. Insecta Mundi. 1208., Brown et al. 2019bBrown et al. 2019b:
Brown, J.W., Basset, Y., Panmeng, M., Putnaul, S., Miller, S. E. 2019b. Host records for Tortricidae (Lepidoptera) reared from seeds and fruits in a Thailand rainforest. Proceedings of the Entomological Society of Washington. 121(4): 544-556.), and most are relatively host specific. The group includes many economically important fruit pests, such as the notorious and nearly cosmopolitan codling moth (Cydia pomonella), the Oriental fruit moth (Grapholita molesta), and the false codling moth (Thaumatotibia leucotreta). Morphological synapomorphies for the tribe include hindwinghindwing:
the posterior wing veins M2 and M3 widely separated at their base, and a reduced dorsaldorsal:
upper, to the top, on the back complex (i.e., uncusuncus:
a sclerotized process which is fused to the posterodorsal margin of tergum IX, sociisocii:
a pair of lightly sclerotized setose lobes, gnathosgnathos:
a narrow bandlike structure that joins the posterolateral edges of the tegumen and supports the anal tube) in the male genitalia. The larvae of several internal-feeding genera (e.g., Cydia, Cryptophlebia, Gymnandrosoma) lack an anal fork, the presence of which is characteristic of most tortricid caterpillars. Molecular data suggest that Grapholitini are sister to Eucosmini (Regier et al. 2013Regier et al. 2013:
Regier, J. C., Mitter, C., Zwick, A., Bazinet, A. L., Cummings, M. P., Kawahara, A. Y., Sohn, J. C., Zwickl, D. J., Cho, S., Davis, D. R., Baixeras, J., Brown, J., Parr, C., Weller, S., Lees, D. C., Mitter, K. T. 2013. A large-scale, higher-level, molecular phylogenetic study of the insect order lepidoptera (moths and butterflies). PLoS ONE. 8(3): e58568., Fagua et al. 2017Fagua et al. 2017:
Fagua G., Condamine, F. L., Horak, M., Zwick, A., Sperling, F. A. 2017. Diversification shifts in leafroller moths linked to continental colonization and the rise of angiosperms. Cladistics. 33: 449-466. ). A recent molecular phylogeny has been published for this tribe (Hu et al. 2023Hu et al. 2023:
Hu, G.-L., Brown, J.W., Heikkilä, M., Aarvik, L., Mutanen, M. 2023. Molecular phylogeny, divergence time, biogeography and trends in host plant usage in the agriculturally important tortricid tribe Grapholitini (Lepidoptera: Tortricidae: Olethreutinae). Cladistics. 39(5): 359-381.).
