We present an updated list of the members of the subtribe Cochylina (Tortricidae) in North America north of Mexico. We summarize the proposed changes in the classification since the end of 1978. We propose revised status for two genera, Rolandylis Gibeaux, 1985 and Thyraylia Walsingham, 1897. We propose eleven revised combinations: Saphenista parvimaculana (Walsingham, 1879), unplaced Cochylina omphacitis (Meyrick, 1912) [Cochylis], unplaced Cochylina voxcana (Kearfott, 1907) [Phalonia], Thyraylia bana (Kearfott, 1907), Thyraylia rhodites (Meyrick, 1912), Thyraylia bunteana (Robinson, 1869), Thyraylia discana (Kearfott, 1907), Thyraylia cricota (Meyrick, 1912), Thyraylia gunniana (Busck, 1907), Thyraylia hollandana (Kearfott, 1907), and Thyraylia nana (Haworth, ). We propose four revised combinations: Rolandylis fusca Pogue, 2001, Rolandylis maiana (Kearfott, 1907), Rolandylis catalonica Gibeaux, 1985, Rolandylis virilia Pogue, 2001; and three new synonymies: Aethes ziscana Kearfott with A. bomonana (Kearfott), Henricus edwarsiana (Walsingham) with H. contrastana (Kearfott), and Phtheochroa pecosana (Kearfott, 1907) with Phtheochroa cartwrightana (Kearfott, 1907). The described fauna includes 20 genera and 137 species, yet it is likely that this region of North America includes two to three times that many species; at least six new genera are defined, but not yet formally described.
The most comprehensive, contemporary list of the North American members of the family “Cochylidae” was provided by Powell (1983) in his contribution to the Check list of the Lepidoptera of America North of Mexico (Hodges et al. 1983). In 1986, a PhD thesis completed by Michael Pogue at the University of Minnesota provided a thorough review of the classification of the North American fauna at the generic level, including manuscript descriptions of several new genera and many new combinations. The phylogenetic analysis portion of the thesis was published by Pogue and Mickevich (1990), but the descriptive and taxonomic portions were not. As a consequence, the nomenclatural changes and descriptions were not “formally” proposed, and the new names and actions remained unavailable. Brown (2005) compiled the first list of Cochylini worldwide, and his treatment of the North American fauna basically followed that developed by Pogue (1986). The new genera recognized by Pogue (1986) were identified as “Cochylini new genus 1,” “Cochylini new genus 2,” and so forth in order to provide a more meaningful taxonomic framework for the fauna, i.e., this action seemed a better alternative to leaving the associated species “unplaced.”
Taxonomic changes proposed by Razowski (2009, 2011), Metzler & Albu (2013), and others over the last decade resulted in the generic reassignments of many species. Many of the changes proposed by Pogue (1986) were formalized through various publications (e.g., Pogue 2001, Brown 2005), and new species were added to the faunal list through introduction (e.g., Agapeta zoegana) (Powell et al. 2000) or description of the native fauna (e.g., Metzler & Forbes 2012, Brown 2013, Metzler & Albu 2013). In a phylogenetic analysis of Tortricidae based entirely on molecular data, Regier et al. (2012) revealed that Cochylini are a monophyletic lineage embedded within the larger Euliini and should be treated as the subtribe Cochylina until further evidence contradicts this proposed change in classification. Because so much information has accumulated since Powell (1983), we deem it advisable to recap the current list of species from North America and their generic assignments.
Materials and Methods
Consistent with the Moths of North America (MONA) project, we treat the fauna of Canada, the United States, and Greenland. This geographical region is ca. 99% congruent with the Nearctic ecozone. Notable differences are the southern tip of peninsular Florida, which is Neotropical, and most of central Mexico south to approximately 18° north latitude, which is considered Nearctic.
We examined the original descriptions and genital preparations of all species of Cochylina that might be found within the study region. Whenever possible, we examined genitalia of types with the proviso that not all types are extant. If types are not available, we examined the genitalia of specimens overwhelmingly considered to be conspecific. In some cases, a judgment of generic assignment seemed to be a matter of convenience rather than based on scientific facts, thus a few species are considered unplaced pending more evidence. We know from experience that many species cannot be placed to genus without examination of the genitalia, thus our conservative approach is justified. While we recognize it is inconvenient to have species names without generic assignments, it would introduce imprecision to pretend we have information not available. Combinations without merit will not help settle the confusion in Cochylina classification.
We examined the literature for species descriptions and revised generic assignments published subsequent to Brown (2005). We followed the most recently published combinations, although as more species are examined and described, the combinations are subject to change.
Most of the new genera recognized (but not formally described) by Pogue (1986) and subsequently used by Brown (2005) are retained for stability in the classification. We emphasize that the manuscript names in Pogue (1986) are not published, we do not publish them here, and they are not available. We use original orthography in the spelling of species names.
For ease of use, and because the phylogeny of Cochylina is uncertain, the genera and synonyms are listed in alphabetical order, and the species and synonyms are listed in alphabetical order within each genus. We created an index to genus and species names. Detailed literature references to the original description, type locality, and disposition of type specimen(s) can be found in Brown (2005) and Gilligan et al. (2012). Not all misspellings from the literature are included here. Brown (2005) should be consulted for variations of species names found in much of the literature. We did not attempt to find all spellings previously used. Footnotes provide details for the placement of certain taxa subsequent to Pogue (1986) and indicate areas of ambiguity or disagreement that require further investigation.
List of genera and associated species from North America north of Mexico and Greenland
Cochylina Guenée, 1845 (sensu Regier et al. 2012)
Aethes Billberg, 1920
Argyridia Stephens, 1852
Chlidonia Hübner, 
Chrosis Guenée, 1845
Cirriaethes Razowski, 1962, subgenus
Coecaethes Obraztsov, 1943, subgenus
Dapsilia Hübner, 
LoxoperaWalsingham, 1900 (emendation of Lozopera)
Lozopera Stephens, 1829
Phalonia Hübner, 
Phelonia; Stephens, 1834 (misspelling of Phalonia)
angulata (Robinson, 1869)
angustana (Clemens, 1860)
augustana; (Powell, 1983) (misspelling of angustana)
dorsimaculana (Robinson,1869)(unnecessary replacement name for angustana)
argentilimitana (Robinson, 1869)
labeculana (Robinson, 1869)1
atomosana (Busck, 1907)
baloghi Sabourin & Metzler, 2002
biscana (Kearfott, 1907)
bomonana (Kearfott, 1907)
cyamitis (Meyrick, 1912) (unnecessary replacement name for bomonana)
cyanitis Razowski, 2000 (misspelling of cyamitis)fabicola (Meyrick, 1912) (unnecessary replacement name for ziscana)
ziscana (Kearfott, 1907), new synonym2
deutschiana (Zetterstedt, 1839)
chalcana (Packard, 1866)
fuscostriana Razowski, 1997 (misspelling of fuscostrigana)
fuscostrigana (Clemens, 1864)
lutulentana (Herrich-Schäffer, 1856)
murciana Caradja, 1916 fernaldana (Walsingham, 1879)
floccosana (Walker, 1863)
confusana (Robinson, 1869)
flaccosana; (Powell, 1983) (misspelling of floccosana)
heleniana Razowski, 1997
intactana (Walsingham, 1879)
interruptofasciata (Robinson, 1869)
louisiana (Busck, 1907)
matheri Sabourin & Miller, 2002
matthewcruzi Sabourin & Vargo, 2002
monera Razowski, 1886 (nominal species is extralimital)
ssp. septentrionalis Razowski,1997 (described from Canada)
mymara Razowski, 1997
obliquana (Kearfott, 1907)
patricia Metzler, 2000
promptana (Robinson, 1869)
rana (Busck, 1907)
razowskii Sabourin & Miller, 2002
rutilana (Hübner, ) (nominal species is extra-limital)
ssp. canadiana Razowski, 1997 (described from Canada)
interruptana (Klemensiewicz, 1907)
purpurella (Coquebert de Montbret, 1801)
roridana (Mann, 1867)
ssp. tatricana (Adamszewski, 1936) (extra-limital)
seriatana (Zeller, 1875)
sexdentata Sabourin & Miller, 2002
smeathmanniana (Fabricius, 1781)
sonorae (Walsingham, 1884)
spartinana (Barnes & McDunnough, 1916)
terriae Sabourin & Miller, 2002
vachelliana (Kearfott, 1907)
westratei Sabourin & Miller, 2002
Agapeta Hübner, 1822
Aapeta Anonymous, 1990 (misspelling of Agapeta)
Agapete Hübner 1825 (misspelling of Agapeta)
Euxanthis Hübner, 
Apapeta Razowski, 1977
Xanthosetia Stephens, 1829
Xanthosetia Stephens, 1829 (unnecessary redescription)
Cochylidia Obraztsov, 1956
Cochylis Treitschke, 1829
Acornutia Obraztsov, 1944
Brevicornutia Razowski, 1960
ChochylisDuponchel, 1836 (misspelling of Cochylis)
Cochylichroa Obraztsov & Swatschek, 1958
Conchlis Razowski, 2011 (misspelling of Conchylis)
Conchyli Sodoffsky, 1837 (misspelling of Cochylis)
Longicornutia Razowski, 1960
Neocochylis Razowski, 1960 (described as a subgenus of Cochylis) (extra-limital)
Paracochylis Razowski, 1960 (described as a subgenus of Cochylis) (extra-limital)
Pontoturania Obratzsov, 1943
arthuri Dang, 1984
aurorana (Kearfott, 1907)3
avita Razowski, 1997
bucera Razowski, 1997
carmelana (Kearfott, 1907)
caulocatax Razowski, 1984
dormitoria Razowski, 1997
dubitana (Hübner, 1796)
formonana (Kearfott, 1907)4
hoffmanana (Kearfott, 1907)3
baryzela (Meyrick, 1912) (unnecessary replacement name for toxcana)
hofmanana Razowski, 1997 (misspelling of hoffmanana)
magnaedoeagana Gibeaux, 19855
marloffiana (Busck, 1907)
nonlavana (Kearfott, 1907)
telifera (Meyrick, 1912) (unnecessary replacement name for zoxcana)
toxcana (Kearfott, 1907)
zoxcana (Kearfott, 1907)
hospes (Walsingham, 1884)3
parallelana (Walsingham, 1879)
ringsi Metzler, 20007
temerana (Busck, 1907)3
transversana (Walsingham, 1879)8
viscana (Kearfott, 1907)3
yinyangana Metzler, 2012
Eugnosta Hübner, 9
Argyrolepia Stephens, 1829
Carolella Busck, 193910
Pharmacis Hübner, 1823
Safra Walker, 1863
argyroplaca (Meyrick, 1931)
beevorana (Comstock, 1940)
bimaculana (Robinson, 1869)11
brownana Metzler & Forbes, 2012
busckana (Comstock, 1939)
deceptana (Busck, 1907)
erigeronana (Riley, 1881)
mexicana (Busck, 1907)
sartana (Hübner, 1823)11
willettana (Comstock, 1939)
Eupinivora Brown, 2013
ponderosae Brown, 2013
Gynnidomorpha Turner, 1916
Pierca Razowski, 1977 (misspelling of Piercea)
Piercea Filipjev, 1940
romonana (Kearfott, 1908)12
Henricus Busck, 1943
Heinrichia Busck, 1939
Irazona Razowski, 196413
Lorita Busck, 1939
Phalonidia Le Marchand, 1933
Brevisociaria Obraztsov, 1943
Platphalonidia Razowski, 1985
basiochreana (Kearfott, 1907)4
elderana (Kearfott, 1907)4
felix (Walsingham, 1895)15
latipunctana (Walsingham, 1879)16
lepidana (Clemens, 1860)16
memoranda Razowski, 1997
ontariana Razowski, 1997
Phtheochroa Stephens, 1829
Arce Joannis, 1919
Durrantia Razowski, 1960 (described as a subgenus of Hysterosia) (extra-limital)
Hysterosia Stephens, 1852
Idiographis Lederer, 1859
Parahysterosia Razowski, 1960 (described as a subgenus of Hysterosia) (extra-limital)
Phteochroa Caradja, 1926 (misspelling of Phtheochroa)
Ptheochroa Razowski, 1964 (misspelling of Phtheochroa)
Propira Durrant, 1914
Trachysmia Guenée, 1845
aegrana (Walsingham, 1879)
aureoalbida (Walsingham, 1895)
baracana (Busck, 1907)
birdana (Busck, 1907)
canariana (Barnes & Busck, 1920)
cartwrightana (Kearfott, 1907)
pecosana Kearfott, 1907, new synonym18
fulviplicana (Walsingham, 1879)
huachucana (Kearfott, 1907)
modestana (Busck, 1907)
perspicuana (Barnes & Busck, 1920)
riscana (Kearfott, 1907)
terminana (Busck, 1907)
villana (Busck, 1907)
vitellinana (Zeller, 1875)
vulneratana (Zetterstedt, 1839)
waracana (Kearfott, 1907)
Platphalonia Razowski, 2011
albertae (Razowski, 1997)19
dangi (Razowski, 1997)19
magdalenae Metzler & Albu, 2013
Rolandylis Gibeaux, 1985, revised combination20
“ Clothoa” (unavailable name)21
fusca Pogue, 2001, revised combination22
maiana (Kearfott, 1907), revised combination22
catalonica Gibeaux, 1985, revised combination5, 22
virilia Pogue, 2001, revised combination22
Rudenia Razowski, 1985
Saphenista Walsingham, 1914
Spinipogon Razowski, 1967
resthavenensisMetzler & Sabourin, 2002
thes Razowski & Becker, 1983
Thyraylia Walsingham, 1897 revised status24
Thyralia Walsingham 1914 (misspelling of Thyraylia)
bana (Kearfott, 1907), new combination
bunteana (Robinson, 1869), new combination
discana (Kearfott, 1907), new combination
gunniana (Busck, 1907), new combination hollandana (Kearfott, 1907), new combination nana (Haworth, ) new combination29
albidana (Walker, 1866)6
altocorsicana (Petry, 1904)
carneana (Guenée, 1845)
cruentana (Guenée, 1845)
ochreoalbana (Walker, 1863)
pallidana (Herrich-Schäffer, 1847) (uninomial)
pallidana (Herrich-Schäffer, 1851)
pumillana (Herrich-Schäffer, 1847) (uninomial)
winniana (Kearfott, 1905)
baboquivariana Kearfott, 1907 [Tortrix]
dilutana Walsingham, 1879 [Cochylis]
foxcana Kearfott, 1907 [Phalonia]
fulvotinctana Walsingham, 1884 [Conchylis]
glaucofuscana Zeller, 1875 [Conchylis]
imitabilis Razowski, 1997 [Platphalonidia]26
punctadiscana Kearfott, 1908 [Phalonia]27
voxcana Kearfott, 1907 [Phalonia]28
Alphabetical list of species, their synonyms, and their associated genera from North America north of Mexico.
Aapeta (misspelling of Agapeta)
Acornutia (synonym of Cochylis)
acropeda, Cochylina new genus 6 (“Poterioparvus”)
Agapete (misspelling of Agapeta)
Arce (synonym of Phtheochroa)
Argyridia (synonym of Aethes)
Argyrolepia (synonym of Eugnosta)
“Atroposia” (Cochylina new genus 1) (unavailable name)
baboquivariana, Cochylina unplaced
Brevicornutia (synonym of Cochylis)
Brevisociaria (synonym of Phalonidia)
“Cagiva” (Cochylina new genus 2) (unavailable name)
Carolella (synonym of Eugnosta)
cephalanthana, Cochylina new genus 2 (“Cagiva”)
Chlidonia (synonym of Aethes)
Chochylis (mispelling of Cochylis)
Chrosis (synonym of Aethes)
Cirriaethes (synonym of Aethes)
“ Clothoa” (synonym of Rolandylis)
Cochylichroa (synonym of Cochylis)
Coecaethes (synonym of Aethes)
Conchlis (misspelling of Cochylis)
Conchyli (misspelling of Cochylis)
Conchylis (misspelling of Cochylis)
“Cybilla” (Cochylina new genus 3) (unavailable name)
Dapsilia (synonym of Aethes)
dilutana, Cochylina unplaced
Durrantia (synonym of Phtheochroa)
Euxanthis (synonym of Agapeta)
foxcana, Cochylina unplaced
fulvotinctana, Cochylina unplaced
glaucofuscana, Cochylina unplaced
grandis, Cochylina new genus 4 (“Honca”)
Heinrichia (synonym of Henricus)
“Honca” (Cochylina new genus 4) (unavailable name)
hubbardana, Cochylina new genus 3 (“Cybilla”)
Hysterosia (synonym of Phtheochroa)
Idiographis (synonym of Phtheochroa)
imitabilis, Cochylina unplaced
Irazona (synonym of Henricus)
liquida, Cochylina unplaced
Longicornutia (synonym of Cochylis)
Loxopera (synonym of Aethes)
Lozopera (synonym of Aethes)
Neocochylis (synonym of Cochylis)
“Nycthia” (Cochylina new genus 5)
oenotherana, Cochylina new genus 1 (“Atroposia”)
omphacitis, Cochylina unplaced
Paracochylis (synonym of Cochylis)
Parahysterosia (synonym of Phtheochroa)
Phalonia (synonym of Aethes)
Pharmacis (synonym of Eugnosta)
Phelonia (synonym of Aethes)
Phteochroa (misspelling of Phtheochroa)
Pierca (synonym of Gynnidomorpha)
Piercea (synonym of Gynnidomorpha)
pimana, Cochylina new genus 5 (“Nycthia”)
Platphalonidia (synonym of Phalonidia)
Pontoturamia (synonym of Cochylis)
“ Poterioparvus” (Cochylina new genus 6) (unavailable name)
Propira (synonym of Phtheochroa)
Ptheochroa (misspelling of Phtheochroa)
punctadiscana, Cochylina unplaced
Safra (synonym of Eugnosvta)
Thyralia (misspelling of Thyraylia)
Trachybyris (synonym of Eugnosta)
Trachybyrsis (synonym of Eugnosta)
Trachysmia (synonym of Phtheochroa)
voxcana, Cochylina unplaced
wiscana, Cochylina new genus 6 (“Poterioparvus”)
Xanthosetia (synonym of Agapeta)
yuccatana, Cochylina new genus 5 (“Nycthia”)
The taxonomy of the subtribe Cochylina is poorly known. Although the number of described species worldwide increases every year, mostly as a result of the efforts of Józef Razowski, Polish Academy of Sciences, Krakow and his co-authors, the number of described species from North America has not increased dramatically since Powell (1983).
Michael G. Pogue (pers. comm.) repeated anecdotal information provided by J. F. G. Clarke that approximately 2/3 of the species from North America are undescribed, suggesting that the North American fauna may include more than 300 species. Identifying specimens of Cochylina is problematic because of the number of undescribed species, lack of diagnostic illustrations, and a list of current names.
The frustration of not knowing the identity of an individual specimen of a “Cochylina” in the sea of named and unnamed species is an unresolved issue. Jason Dombroskie (pers. comm.), who is a consultant for Bug Guide (2013), finds that users are not satisfied if they do not receive a specific identification based on photographs of specimens that are neither spread nor dissected. Many species of Cochylina cannot be placed to genus without examination of the genitalia (Brown 2006). Identification is no less complicated because many species of Cochylina, even within a single genus, e.g., Aethes Billberg, 1820, often have forewing patterns that are incongruous. Many species' descriptions do not include illustrations of adults.
Photographs on Moth Photographers Group (2013) are useful for guidance on the habitus of many species. Genitalic illustrations of the types for many species are lacking. Information about descriptions, with references to literature, subsequent to Brown (2005), along with illustrations of type specimens is available at Gilligan et al. (2013). We caution that without examination aided by microscopy, many species simply cannot be identified.
We thank Jerry Powell for offering insights into preparation of the Cochylidae section of the Hodges (1983) Check List. We especially appreciate the tolerance of our spouses and colleagues during our emotional outbreaks while attempting to identify the bewildering number of undescribed and cryptic species in this group of moths. We thank the following for useful comments on the manuscript; Michael Sabourin, Joaquin Baixeras, and one anonymous reviewer.
 1 Aethes labeculana was listed as a separate species by Pogue (1986); it was synonymized with A. argentilimitana by Sabourin et al. (2002).
 2 The synonymy of Aethes ziscana with A. bomonana is based on examination of photographs of the types and male genitalia of both species.
 3 These species were placed in “Recavicula” by Pogue (1986).
 4 These species were listed as “incertae sedis” by Pogue (1986).
 5 This name is supposedly based on a mislabeled specimen (Razowski, 2002).
 6 Cochylis albidana was listed as a synonym of Cochylis dubitana by Pogue (1986).
 7 The date was incorrectly listed as 1999 by Brown (2005).
 8 Cochylis transversana was placed in Saphenista by Pogue (1986).
 9 Trachybyrsis Meyrick, 1927 was listed as a separate genus by Brown (2005) and Aarvik (2010). It was listed as a synonym of Eugnosta by Razowski (2011). Because species of Trachybyrsis occur in Africa rather than in our faunal region, we make no further comments.
 10 Carolella was synonymized with Eugnosta by Razowski (2009).
 11 This species was placed in Carolella by Pogue (1986) and Brown (2005). The combination listed here was created by Razowski (2009).
 12 Gynnidomorpha romonana was listed in Saphenista by Pogue (1986).
 13 Irazona was treated as a separate genus by Pogue (1986) and synonymized with Henricus by Razowski (1997).
 14 Henricus edwardsiana is synonymized with H. contrastana based on examination of the types of both species.
 15 Phalonidia felix was placed in Saphenista by Pogue (1986), it was placed in Platphalonidia by Razowski (1985) and Brown (2005), and it was placed in Phalonidia by Razowski (2011).
 16 These species were listed in Saphenista by Pogue (1986).
 17 The relationship between lepidana, plummeriana, schwarziana, straminoides, and zaracana was variously presented (Pogue 1986, Razowski 1997, Brown 2005). We examined photographs of the types of lepidana, plummeriana, schwarziana, straminoides, and zaracana. We propose that straminoides is not conspecific with the others.
 18 Pogue (1986) listed Phtheochroa pecosana as a synonym of P. cartwrightana. Razowski (1997) did not include pecosana in the synonymy of cartwightana. Brown (2005) listed cartwrightana and pecosana as separate species. We validate the synonymy proposed by Pogue (1986).
 19 These species were listed in Platphalonidia by Brown (2005). When Razowski (2011) transferred P. felix, the type species of Platphalonidia, to Phalonidia the action resulted in the synonymy of Platphalonidia with Phalonidia. However, all other species of Plaphalonidia were left without a generic assignment. Metzler & Albu (2013) transferred some of the unplaced species to Platphalonia. More analysis is needed.
 20 Rolandylis was synonymized with Cochylis by Razowski (2009), and restated (2011). We consider Rolandylis to be a separate genus.
 21 These genera were proposed by Pogue (1986) in his unpublished Ph.D. thesis and used without descriptions or designation of type species in a phylogenetic analysis by Pogue & Mickevitch (1990). We retain the names here for the sake of continuity and clarity, but we do not publish the names here.
 22 These species were listed in Rolandylis by Brown (2005).
 23 Saphenista parvimaculana was listed in Platphalonidia by Brown (2005). When Razowski (2011) transferred P. felix, the type species of Platphalonidia, to Phalonidia the action resulted in the synonymy of Platphalonidia with Phalonidia. However, all other species of Plaphalonidia were left without a generic assignment. We follow Pogue (1986), and we place P. parvimaculana in Saphenista. More analysis is needed.
 24 Thyraylia was listed as a synonym of Cochylis by Razowski (1997, 2009, 2011). However, Pogue (1986) and Brown (2005) both listed Thyraylia as a separate genus. We follow the latter concept.
 25 Clepsis listerana, listed as “incertae sedis” in Pogue (1986), is in the tribe Archipini (Tortricidae).
 26 Platphalonidia imitabilis was listed in Platphalonidia by Brown (2005). When Razowski (2011) transferred felix, the type species of Platphalonidia, to Phalonidia the action resulted in the synonymy of Platphalonidia with Phalonidia. However, all other species of Plaphalonidia were left withut a generic assignment. We propose that more study is needed before P. imitabilis can be placed in a genus.
 27 Phalonia punctadiscana was listed in “Recavicula” (unavailable name) in Pogue (1986). It was listed in “Cochylini Unplaced Species” by Brown (2005).
 28 Phalonia voxcana was listed as a synonym of Thyraylia hollandana by Pogue (1986), placed in Cochylis in Razowski (1997) and Brown (2005). The genitalia illustrated by Razowski (1997) are congeneric with Thyraylia. Our examination of the wings of the types of hollandana and voxcana indicate they are not conspecific.